Reproducing Five Motor Behaviors in a Salamander Robot With Virtual Muscles and a Distributed CPG Controller Regulated by Drive Signals and Proprioceptive Feedback

Diverse locomotor behaviors emerge from the interactions between the spinal central pattern generator (CPG), descending brain signals and sensory feedback. Salamander motor behaviors include swimming, struggling, forward underwater stepping, and forward and backward terrestrial stepping. Electromyographic and kinematic recordings of the trunk show that each of these five behaviors is characterized by specific patterns of muscle activation and body curvature. Electrophysiological recordings in isolated spinal cords show even more diverse patterns of activity. Using numerical modeling and robotics, we explored the mechanisms through which descending brain signals and proprioceptive feedback could take advantage of the flexibility of the spinal CPG to generate different motor patterns. Adapting a previous CPG model based on abstract oscillators, we propose a model that reproduces the features of spinal cord recordings: the diversity of motor patterns, the correlation between phase lags and cycle frequencies, and the spontaneous switches between slow and fast rhythms. The five salamander behaviors were reproduced by connecting the CPG model to a mechanical simulation of the salamander with virtual muscles and local proprioceptive feedback. The main results were validated on a robot. A distributed controller was used to obtain the fast control loops necessary for implementing the virtual muscles. The distributed control is demonstrated in an experiment where the robot splits into multiple functional parts. The five salamander behaviors were emulated by regulating the CPG with two descending drives. Reproducing the kinematics of backward stepping and struggling however required stronger muscle contractions. The passive oscillations observed in the salamander's tail during forward underwater stepping could be reproduced using a third descending drive of zero to the tail oscillators. This reduced the drag on the body in our hydrodynamic simulation. We explored the effect of local proprioceptive feedback during swimming and forward terrestrial stepping. We found that feedback could replace or reduce the need for different drives in both cases. It also reduced the variability of intersegmental phase lags toward values appropriate for locomotion. Our work suggests that different motor behaviors do not require different CPG circuits: a single circuit can produce various behaviors when modulated by descending drive and sensory feedback

Axial dynamics during locomotion in vertebrates: lesson from the salamander

Much of what we know about the flexibility of the locomotor networks in vertebrates is derived from studies examining the adaptation of limb movements during stepping in various conditions. However, the body movements play important roles during locomotion: they produce the thrust during undulatory locomotion and they help to increase the stride length during legged locomotion. In this chapter, we review our current knowledge about the flexibility in the neuronal circuits controlling the body musculature during locomotion. We focus especially on salamander because, as an amphibian, this animal is able to display a rich repertoire of aquatic and terrestrial locomotor modes

Decoding the mechanisms of gait generation in salamanders by combining neurobiology, modeling and robotics

Vertebrate animals exhibit impressive locomotor skills. These locomotor skills are due to the complex interactions between the environment, the musculo-skeletal system and the central nervous system, in particular the spinal locomotor circuits. We are interested in decoding these interactions in the salamander, a key animal from an evolutionary point of view. It exhibits both swimming and stepping gaits and is faced with the problem of producing efficient propulsive forces using the same musculo-skeletal system in two environments with significant physical differences in density, viscosity and gravitational load. Yet its nervous system remains comparatively simple. Our approach is based on a combination of neurophysiological experiments, numerical modeling at different levels of abstraction, and robotic validation using an amphibious salamander-like robot. This article reviews the current state of our knowledge on salamander locomotion control, and presents how our approach has allowed us to obtain a first conceptual model of the salamander spinal locomotor networks. The model suggests that the salamander locomotor circuit can be seen as a lamprey-like circuit controlling axial movements of the trunk and tail, extended by specialized oscillatory centers controlling limb movements. The interplay between the two types of circuits determines the mode of locomotion under the influence of sensory feedback and descending drive, with stepping gaits at low drive, and swimming at high driv

Where are we in understanding salamander locomotion: biological and robotic perspectives on kinematics

Salamanders have captured the interest of biologists and roboticists for decades because of their ability to locomote in different environments and their resemblance to early representatives of tetrapods. In this article, we review biological and robotic studies on the kinematics (i.e., angular profiles of joints) of salamander locomotion aiming at three main goals: (i) to give a clear view of the kinematics, currently available, for each body part of the salamander while moving in different environments (i.e., terrestrial stepping, aquatic stepping, and swimming), (ii) to examine what is the status of our current knowledge and what remains unclear, and (iii) to discuss how much robotics and modeling have already contributed and will potentially contribute in the future to such studies

A simulation model of the locomotion controllers for the nematode Caenorhabditis elegans

This paper presents a simple yet biologicallygrounded model of the C. elegans neural circuit for forward locomotive control. The model considers a limited subset of the C. elegans nervous system, within a minimal two-dimensional environment. Despite its reductionist approach, this model is sufficiently rich to generate patterns of undulations that are reminiscent of the biological worm’s behaviour and qualitatively similar to patterns which have been shown to generate locomotion in a model of a richer physical environment. Interestingly, and contrary to conventional wisdom about neural circuits for motor control, our results are consistent with the conjecture that the worm may be relying on feedback from the shape of its body to generate undulations that propel it forward or backward

Design of artificial neural oscillatory circuits for the control of lamprey- and salamander-like locomotion using evolutionary algorithms

This dissertation investigates the evolutionary design of oscillatory artificial neural networks for the control of animal-like locomotion. It is inspired by the neural organ¬ isation of locomotor circuitries in vertebrates, and explores in particular the control of undulatory swimming and walking. The difficulty with designing such controllers is to find mechanisms which can transform commands concerning the direction and the speed of motion into the multiple rhythmic signals sent to the multiple actuators typically involved in animal-like locomotion. In vertebrates, such control mechanisms are provided by central pattern generators which are neural circuits capable of pro¬ ducing the patterns of oscillations necessary for locomotion without oscillatory input from higher control centres or from sensory feedback. This thesis explores the space of possible neural configurations for the control of undulatory locomotion, and addresses the problem of how biologically plausible neural controllers can be automatically generated.Evolutionary algorithms are used to design connectionist models of central pattern generators for the motion of simulated lampreys and salamanders. This work is inspired by Ekeberg's neuronal and mechanical simulation of the lamprey [Ekeberg 93]. The first part of the thesis consists of developing alternative neural controllers for a similar mechanical simulation. Using a genetic algorithm and an incremental approach, a variety of controllers other than the biological configuration are successfully developed which can control swimming with at least the same efficiency. The same method is then used to generate synaptic weights for a controller which has the observed biological connectivity in order to illustrate how the genetic algorithm could be used for developing neurobiological models. Biologically plausible controllers are evolved which better fit physiological observations than Ekeberg's hand-crafted model. Finally, in collaboration with Jerome Kodjabachian, swimming controllers are designed using a developmental encoding scheme, in which developmental programs are evolved which determine how neurons divide and get connected to each other on a two-dimensional substrate.The second part of this dissertation examines the control of salamander-like swimming and trotting. Salamanders swim like lampreys but, on the ground, they switch to a trotting gait in which the trunk performs a standing wave with the nodes at the girdles. Little is known about the locomotion circuitry of the salamander, but neurobiologists have hypothesised that it is based on a lamprey-like organisation. A mechanical sim¬ ulation of a salamander-like animat is developed, and neural controllers capable of exhibiting the two types of gaits are evolved. The controllers are made of two neural oscillators projecting to the limb motoneurons and to lamprey-like trunk circuitry. By modulating the tonic input applied to the networks, the type of gait, the speed and the direction of motion can be varied.By developing neural controllers for lamprey- and salamander-like locomotion, this thesis provides insights into the biological control of undulatory swimming and walking, and shows how evolutionary algorithms can be used for developing neurobiological models and for generating neural controllers for locomotion. Such a method could potentially be used for designing controllers for swimming or walking robots, for instance

トカゲ様歩行に内在する全身自由度の統御原理に関する研究

要約のみTohoku University石黒章夫課

From lamprey to salamander: an exploratory modeling study on the architecture of the spinal locomotor networks in the salamander

The evolutionary transition from water to land required new locomotor modes and corresponding adjustments of the spinal "central pattern generators" for locomotion. Salamanders resemble the first terrestrial tetrapods and represent a key animal for the study of these changes. Based on recent physiological data from salamanders, and previous work on the swimming, limbless lamprey, we present a model of the basic oscillatory network in the salamander spinal cord, the spinal segment. Model neurons are of the Hodgkin-Huxley type. Spinal hemisegments contain sparsely connected excitatory and inhibitory neuron populations, and are coupled to a contralateral hemisegment. The model yields a large range of experimental findings, especially the NMDA-induced oscillations observed in isolated axial hemisegments and segments of the salamander Pleurodeles waltlii. The model reproduces most of the effects of the blockade of AMPA synapses, glycinergic synapses, calcium-activated potassium current, persistent sodium current, and -current. Driving segments with a population of brainstem neurons yields fast oscillations in the in vivo swimming frequency range. A minimal modification to the conductances involved in burst-termination yields the slower stepping frequency range. Slow oscillators can impose their frequency on fast oscillators, as is likely the case during gait transitions from swimming to stepping. Our study shows that a lamprey-like network can potentially serve as a building block of axial and limb oscillators for swimming and stepping in salamanders