GEOTROPISM AND MUSCLE TENSION IN HELIX

1. The snail Helix aspersa Müller, is negatively geotropic during the daytime, but positive or indifferent at night. 2. The precision of geotropic orientation is a function of the gravity component acting on the body. 3. The rate of geotropic locomotion is also determined by the gravity component (sine of the angle of inclination). 4. The rate of upward movement is increased 1.51 times at 45° inclination by loading the snail with one-half its weight. No such increase is seen in loaded snails creeping on a horizontal surface. 5. Moderate centrifugation results in orientation and locomotion towards the center of rotation. 6. A response analogous to the homostrophic reflex occurs when a backward pull to right or to left is exerted on the shell. Bilaterally equal tension applied to the shell causes locomotion along a path parallel and opposite to the direction of the pull. 7. All the observations go to show that the stimulus for geotropic orientation and locomotion is tension of the body muscles produced by the downward pull of gravity, and that the stimulus is received by the proprioreceptors of these muscles. Otolith apparatus and analogous organs, when present, may assist in the response, but they do not seem to be requisite in all cases. Since the precision of orientation and the rate of locomotion are functions of the gravity component acting on the body, the muscle tension theory of the geotropic reactions accords fully with Loeb''s tropism doctrine for animals.     

THE REACTION OF NEREIS VIRENS TO UNILATERAL TENSION OF ITS MUSCULATURE

1. The anterior segments of Nereis are oriented reflexly by passive unilateral tension of the posterior musculature. 2. The afferent impulses of the homostrophic reflex rise from any part of the worm and are conducted forward by way of the ventral nerve cord. 3. The efferent impulses flow out from the brain and anterior two or three ventral ganglia. 4. The homostrophic reflex may be partially or wholly masked by stereotropism.     

MUSCLE TENSION AND REFLEXES IN THE EARTHWORM

1. By the use of preparations of earthworm in which the cutaneous receptors have been anesthetized with a solution of M/8 MgCl₂, it is shown that peristalsis can be initiated by tension alone. 2. The receptors of the tension reflex are the intermyal sensory cells of the ventral region of the body wall. 3. It is concluded that Straub obtained the tension reflex because his preparations contained the intermyal receptors; Budington was unable to observe the tension reflex in any preparation from which the intermyal receptors had been removed. 4. Intermyal receptors are the receptors of the following reaction: Passive unilateral tension of the posterior part of an earthworm induces active homolateral tension of the musculature of the anterior segments, and results in the course of progress being brought into line with the enforced orientation of the tail. This reaction is termed the homostrophic reflex. 5. The receptors for the reaction are distributed throughout the entire length of the worm, the effectors are limited to the anterior 15 to 20 segments. The impulse is conducted by the ventral nerve cord. 6. The interaction of the homostrophic reflex and tropisms is considered.     

Biologically inspired control of physically simulated bipeds

Summary The basic rhythmicity underlying animal locomotion is created by dedicated neural structures called central pattern generators (CPGs). We describe the implementation of such structures in simulation and their successful use for the control of bipedal walking. Artificial evolution (in the form of genetic algorithms) is used as the optimisation procedure. Two CPG types are illustrated, the more advanced of which being based on recent theoretical findings on the nature of neural architectures required to drive animal locomotion. It is shown that CPGs in conjunction with simple reflex responses as well as an appropriate mechanical implementation of the biped are capable of producing stable walking patterns on planar surfaces. This finding corroborates circumstantial experimental evidence that limited bipedal locomotion is possible without the employment of higher level control centres.     

Elegant‐crested Tinamous Eudromia elegans do not synchronize head and leg movements during head‐bobbing

Head‐bobbing is the fore–aft movement of the head relative to the body during terrestrial locomotion in birds. It is considered to be a behaviour that helps to stabilize images on the retina during locomotion, yet some studies have suggested biomechanical links between the movements of the head and legs. This study analysed terrestrial locomotion and head‐bobbing in the Elegant‐crested Tinamou Eudromia elegans at a range of speeds by synchronously recording high‐speed video and ground reaction forces in a laboratory setting. The results indicate that the timing of head and leg movements are dissociated from one another. Nonetheless, head and neck movements do affect stance duration, ground reaction forces and body pitch and, as a result, the movement of the centre of mass in head‐bobbing birds. This study does not support the hypothesis that head‐bobbing is itself constrained by terrestrial locomotion. Instead, it suggests that visual cues are the primary trigger for head‐bobbing in birds, and locomotion is, in turn, constrained by a need for image stabilization and depth perception.     

An Approach for Adaptive Limbless Locomotion Using a CPG-Based Reflex Mechanism

Animals＇ free movement in natural environments has attracted many researchers to explore control methods for bio-inspired robots. This paper presents a novel reflex mechanism based on a Central Pattern Generator （CPG） for adaptive locomotion of limbless robots. First, inspired by the concept of reflex arc, the reflex mechanism is designed on a connectionist CPG model. Since the CPG model inspired by the spinal cord of lampreys is developed at the neuron level, it provides a possible natural solution for sensory reflex integration. Therefore, sensory neurons that bridge the external stimuli and the CPG model, together with the concept of reflex arc, are utilized for designing the sensory reflex mechanism. Then, a border reflex and a body reflex are further developed and applied on the ends and the middle part of a limbless robot, respectively. Finally, a ball hitting scenario and a corridor passing scenario are designed to verify the proposed method. Results of simulations and on-site experiments show the feasibility and effectiveness of the reflex mechanism in realizing fast response and adaptive limbless locomotion.     

ON STEREOTROPISM IN TENEBRIO LARV?

Larvæ of Tenebrio while creeping show homostrophic responses, and stereotropic orientation to lateral contacts. Homostrophic orientation is inhibited by stereotropism. Both depend upon the anterior portion of the central nervous system. Stereotropic orientation due to unilateral contact, particularly at the anterior end, persists briefly after the cessation of the contact. Equal posterior bilateral contact of the body obliterates stereotropic bending. Unequal posterior bilateral contacts lead to orientation through an angle roughly proportional to the differences in contact areas. Functional symmetry in such responses is not disturbed by asymmetrical distribution of the body "hairs." The stereotropic orientation undergoes reversal of direction, central in origin, when the stimulation is sufficiently intense. Stereotropic response, leading to maintained lateral contact with a surface or to bending when the end of such a surface has been passed, is inhibited by a definite intensity of light. These findings (1) round out the demonstration that stereotropism is truly of a tropistic character, and (2) make possible the understanding of conduct in a case involving the participation of contact stimulation, phototropism, temperature, and homostrophy.     

The mucus producing glands and the distribution of the cilia of the pulmonate slug Limax pseudoflavus

The gross anatomy and histochemistry of the mucus-producing glands of Limax pseudoflavus Evans were investigated. The body mucus can be divided into three areas. The dorsal body surface is covered with a sulphated acid mucopolysaccharide/protein mixture secreted largely by five cell types. The pedal mucus is a mixture of neutral mucopolysaccharide from the suprapedal gland. The dorsal and pedal mucus sheets are separated by the peripodal groove whose cells secrete a weakly acid mucus. The duct of the suprapedal gland, the epidermis around the pneumostome, the ventral surface of the peripodal groove and the centre of the underside of the foot are ciliated. The dorsal and pedal mucus remain stationary relative to the body and the substrate respectively and the only rejection currents seen in the mucus are around the pneumostome.
It is suggested that the pedal mucus is formed by the mixture of the products of the suprapedal gland and the mucoprotein secreting gland in the leading edge of the foot, thus producing a mucus suitable for locomotion. Many areas of the animal (e.g. the head, pneumostome, sole and the leading edge of the foot) are capable of producing both a fluid (neutral or weakly acid) and a viscous (acid) mucus. It is postulated that such an arrangement allows for both adhesion and lubrication at different times.     

Convergent evolution and locomotion through complex terrain by insects, vertebrates and robots

Arthropods are the most successful members of the animal kingdom largely because of their ability to move efficiently through a range of environments. Their agility has not been lost on engineers seeking to design agile legged robots. However, one cannot simply copy mechanical and neural control systems from insects into robotic designs. Rather one has to select the properties that are critical for specific behaviors that the engineer wants to capture in a particular robot. Convergent evolution provides an important clue to the properties of legged locomotion that are critical for success. Arthropods and vertebrates evolved legged locomotion independently. Nevertheless, many neural control properties and mechanical schemes are remarkably similar. Here we describe three aspects of legged locomotion that are found in both insects and vertebrates and that provide enhancements to legged robots. They are leg specialization, body flexion and the development of a complex head structure. Although these properties are commonly seen in legged animals, most robotic vehicles have similar legs throughout, rigid bodies and rudimentary sensors on what would be considered the head region. We describe these convergent properties in the context of robots that we developed to capture the agility of insects in moving through complex terrain.     

Functional morphology of the muscles in Philodina sp. (Rotifera: Bdelloidea)

Whole-mounts of Philodina sp., a bdelloid rotifer, were stained with fluorescent-labeled phalloidin to visualize the musculature. Several different muscle types were identified including incomplete circular bands, coronal retractors and foot retractors. Based on the position of the larger muscle bands in the body wall, their function during creeping locomotion and tun formation was inferred. Bdelloid creeping begins with the contraction of incomplete circular muscle bands against the hydrostatic pseudocoel, resulting in an anterior elongation of the body. One or more sets of ventral longitudinal muscles then contract bringing the rostrum into contact with the substrate, where it presumably attaches via adhesive glands. Different sets of ventral longitudinal muscles, foot and trunk retractors, function to pull the body forward. These same longitudinal muscle sets are also used in `tun' formation, in which the head and foot are withdrawn into the body. Three sets of longitudinal muscles supply the head region (anterior head segments) and function in withdrawal of the corona and rostrum. Two additional pairs of longitudinal muscles function to retract the anterior trunk segments immediately behind the head, and approximately five sets of longitudinal retractors are involved in the withdrawal of the foot and posterior toes. To achieve a greater understanding of rotifer behavior, it is important to elucidate the structural complexity of body wall muscles in rotifers. The utility of fluorescently-labeled phalloidin for the visualization of these muscles is discussed and placed in the context of rotifer functional morphology.     

Mantisbot is a robotic model of visually guided motion in the praying mantis

Insects use highly distributed nervous systems to process exteroception from head sensors, compare that information with state-based goals, and direct posture or locomotion toward those goals. To study how descending commands from brain centers produce coordinated, goal-directed motion in distributed nervous systems, we have constructed a conductance-based neural system for our robot MantisBot, a 29 degree-of-freedom, 13.3:1 scale praying mantis robot. Using the literature on mantis prey tracking and insect locomotion, we designed a hierarchical, distributed neural controller that establishes the goal, coordinates different joints, and executes prey-tracking motion. In our controller, brain networks perceive the location of prey and predict its future location, store this location in memory, and formulate descending commands for ballistic saccades like those seen in the animal. The descending commands are simple, indicating only 1) whether the robot should walk or stand still, and 2) the intended direction of motion. Each joint's controller uses the descending commands differently to alter sensory-motor interactions, changing the sensory pathways that coordinate the joints' central pattern generators into one cohesive motion. Experiments with one leg of MantisBot show that visual input produces simple descending commands that alter walking kinematics, change the walking direction in a predictable manner, enact reflex reversals when necessary, and can control both static posture and locomotion with the same network.     

From Spontaneous Motor Activity to Coordinated Behaviour: A Developmental Model

In mammals, the developmental path that links the primary behaviours observed during foetal stages to the full fledged behaviours observed in adults is still beyond our understanding. Often theories of motor control try to deal with the process of incremental learning in an abstract and modular way without establishing any correspondence with the mammalian developmental stages. In this paper, we propose a computational model that links three distinct behaviours which appear at three different stages of development. In order of appearance, these behaviours are: spontaneous motor activity (SMA), reflexes, and coordinated behaviours, such as locomotion. The goal of our model is to address in silico four hypotheses that are currently hard to verify in vivo: First, the hypothesis that spinal reflex circuits can be self-organized from the sensor and motor activity induced by SMA. Second, the hypothesis that supraspinal systems can modulate reflex circuits to achieve coordinated behaviour. Third, the hypothesis that, since SMA is observed in an organism throughout its entire lifetime, it provides a mechanism suitable to maintain the reflex circuits aligned with the musculoskeletal system, and thus adapt to changes in body morphology. And fourth, the hypothesis that by changing the modulation of the reflex circuits over time, one can switch between different coordinated behaviours. Our model is tested in a simulated musculoskeletal leg actuated by six muscles arranged in a number of different ways. Hopping is used as a case study of coordinated behaviour. Our results show that reflex circuits can be self-organized from SMA, and that, once these circuits are in place, they can be modulated to achieve coordinated behaviour. In addition, our results show that our model can naturally adapt to different morphological changes and perform behavioural transitions.     

Aquatic and terrestrial locomotory energetics in a toad and a turtle: a search for generalisations among ectotherms

Murray short-necked turtles were trained to walk on a motorised treadmill and to swim in a recirculating flume. Through filmed records, the frequency of limb movement and the time that thrust was directed against the substrate were measured. The animals wore masks when walking and accessed air when swimming from a ventilated capsule placed on top of the water surface. Measurement of the exhalant O(2) and CO(2) levels from these devices enabled the measurement of metabolic rates. Equivalent data were obtained from swimming and hopping cane toads, although repeatable measures of limb frequency and contact times were not obtained due to the intermittent form of locomotion in this species. Comparing the cost of transport, the energy required to transport a mass of animal over a unit distance, with other animals showed that toads do not have a cheap form of terrestrial locomotion, but turtles do; turtles use half the cost predicted from their body mass. This economy of locomotion is consistent with what is known about turtle muscle, the mechanics of their gait, and the extremely long contact time for a limb with the substrate. Swimming in toads is energetically expensive, whereas turtles, on the basis of mass, use about the same energy to transport a unit mass as an equivalent-size fish. The data were compared with the predictions of the Kram-Taylor hypothesis for locomotory scaling, and walking turtles were found to provide a numerical fit. The data show that both terrestrial and aquatic locomotory energetics in toads are generally higher than predictions on the basis of mass, whereas in turtles they are lower.     

Predictability of visual perturbation during locomotion: implications for corrective efference copy signaling

In guiding adaptive behavior, efference copy signals or corollary discharge are traditionally considered to serve as predictors of self-generated sensory inputs and by interfering with their central processing are able to counter unwanted consequences of an animal??s own actions. Here, in a speculative reflection on this issue, we consider a different functional role for such intrinsic predictive signaling, namely in stabilizing gaze during locomotion where resultant changes in head orientation in space require online compensatory eye movements in order to prevent retinal image slip. The direct activation of extraocular motoneurons by locomotor-related efference copies offers a prospective substrate for assisting self-motion derived sensory feedback, rather than being subtracted from the sensory signal to eliminate unwanted reafferent information. However, implementing such a feed-forward mechanism would be critically dependent on an appropriate phase coupling between rhythmic propulsive movement and resultant head/visual image displacement. We used video analyzes of actual locomotor behavior and basic theoretical modeling to evaluate head motion during stable locomotion in animals as diverse as Xenopus laevis tadpoles, teleost fish and horses in order to assess the potential suitability of spinal efference copies to the stabilization of gaze during locomotion. In all three species, and therefore regardless of aquatic or terrestrial environment, the head displacements that accompanied locomotor action displayed a strong correlative spatio-temporal relationship in correspondence with a potential predictive value for compensatory eye adjustments. Although spinal central pattern generator-derived efference copies offer appropriately timed commands for extraocular motor control during self-generated motion, it is likely that precise image stabilization requires the additional contributions of sensory feedback signals. Nonetheless, the predictability of the visual consequences of stereotyped locomotion renders intrinsic efference copy signaling an appealing mechanism for offsetting these disturbances, thus questioning the exclusive role traditionally ascribed to sensory-motor transformations in stabilizing gaze during vertebrate locomotion.     

Burrowing,Locomotion and Other Movements of the Echiuran Ochetostoma caudex

Burrowing, iocomotory and other movements of the echiuran Ochetostoma caudex have been examined and discussed. A continuous body cavity enables the worm to undergo peristaltic waves to pump water through the burrow without causing locomotion. The animal is capable of both forward and backward locomotion in its burrow. During forward locomotion, retrograde peristaltic waves are utilized which advance the animal in a step-wise fashion. Pressure changes within the coelom during burrowing, locomotion and during irrigation movements have been measured with the use of electronic recording techniques and the results interpreted in relation to direct visual observation. The structural and functional specializations for burrowing are discussed and compared with the activities of Priapulus caudatus, Sipunculus nudus and Bonellia viridis.     

Chinchilla (Chinchilla lanigera) behavioral responses to a visual signal preceding handling

Zoos use ambassador animals in educational programs featuring close contact with humans. Chinchillas (Chinchilla lanigera) at the Saint Louis Zoo are retrieved for programs by a keeper wearing brown handling gloves, but green cleaning gloves are worn during normal husbandry when physical contact with the animal is only incidental. The chinchillas’ primary keeper anecdotally reported more reactivity and movement from chinchillas when approached with handling gloves. Animals’ behavioral reactions to the presence of humans often include locomotion and vigilance, but these responses may be attenuated by predictability. To investigate these behaviors, handling trials involving brief contact attempts with both cleaning and handling gloves were filmed. Results indicated that chinchillas responded to disturbances by moving, jumping, and adopting more alert body postures. Surprisingly, movement was recorded in longer durations when the keeper attempted to touch the animals with cleaning gloves. This higher arousal may indicate that the animal was not expecting to be handled, yet an attempt to do so was being made. This reaction provides evidence that potentially aversive events should be reliably and consistently signaled.     

The burrowing of Priapulus caudatus

An account is given of the way in which Priapulus caudatus burrows in the muddy sea-bed in which it lives. Three phases are distinguishable in the muscular activity which is responsible for locomotion. During the first phase the animal is able to feed and defaecate, during the second the proboscis becomes invaginated, and during the third the animal moves forward. The power for locomotion is provided by contraction of the longitudinal and circular muscles of the body wall, not, as has been suggested previously, by the retractor muscles of the praesoma. Invagination of the proboscis is apparently stimulated by the arrival of a wave of contraction in the body wall musculature, propagated from the trunk.
In general the animal burrows in a way common for soft-bodied animals; the anterior and posterior extremities acting in turn as "terminal" and "penetration" anchors in the substratum. The muscular activities of the larva are limited by the presence of a lorica which encases the trunk, and the animal's powers of movement at this stage are very restricted.     

Coupling sensory inputs to the appropriate motor responses: new aspects of cerebellar function

It is known that proprioceptive signals modify the spatial organization of the postural reflexes, thus leading to body stability. The neurophysiological basis of this phenomenon are at present unknown. The present report documents that, in decerebrate cat, body-to-head rotation in the horizontal plane modified the preferred response direction to labyrinthine stimulation of the forelimb extensor triceps brachii. Such direction resulted always perpendicular to the longitudinal body axis of the animal, whatever its relative position with respect to the head could be. The rotation of the preferred response direction of the triceps was greatly reduced by functional inactivation of the ipsilateral cerebellar vermis. On the other hand, following body-to-head displacement, the preferred response directions of the corresponding P-cells tended, on the average, to rotate in the same direction and by the same angle as the body. We propose that the neck input finely tunes parallel vestibular channels, endowed with different spatial and temporal properties, impinging upon P-cells, thus modifying their responses to animal tilt and, as a consequence, the spatial properties of VS reflexes. It is possible that, by a similar mechanism, the cerebellum may contribute to the changes in reference frame occurring in sensorimotor transformations of reflex and voluntary nature.     

An allometric model of home range formation explains the structuring of animal communities exploiting heterogeneous resources

Understanding and predicting the composition and spatial structure of communities is a central challenge in ecology. An important structural property of animal communities is the distribution of individual home ranges. Home range formation is controlled by resource heterogeneity, the physiology and behaviour of individual animals, and their intra‐ and interspecific interactions. However, a quantitative mechanistic understanding of how home range formation influences community composition is still lacking. To explore the link between home range formation and community composition in heterogeneous landscapes we combine allometric relationships for physiological properties with an algorithm that selects optimal home ranges given locomotion costs, resource depletion and competition in a spatially‐explicit individual‐based modelling framework. From a spatial distribution of resources and an input distribution of animal body mass, our model predicts the size and location of individual home ranges as well as the individual size distribution (ISD) in an animal community. For a broad range of body mass input distributions, including empirical body mass distributions of North American and Australian mammals, our model predictions agree with independent data on the body mass scaling of home range size and individual abundance in terrestrial mammals. Model predictions are also robust against variation in habitat productivity and landscape heterogeneity. The combination of allometric relationships for locomotion costs and resource needs with resource competition in an optimal foraging framework enables us to scale from individual properties to the structure of animal communities in heterogeneous landscapes. The proposed spatially‐explicit modelling concept not only allows for detailed investigation of landscape effects on animal communities, but also provides novel insights into the mechanisms by which resource competition in space shapes animal communities.