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Two Australian frogs were introduced to New Zealand over 100 years ago. Since their introduction they have become widespread and well established with Litoria ewingii being more prevalent in alpine and cooler areas of New Zealand, while Litoria raniformis is found in more temperate coastal areas. Very little physiological data exists for these frogs and aside from their distribution not much is known about them in New Zealand. Thus the effects of thermal acclimation and temperature change on respiration and locomotion were examined in these two exotic frogs. The more terrestrial and alpine dwelling L. ewingii was found to be able to thermally acclimate its respiration performance, where respiration was highest when acclimation temperature matched test temperature. It was also able to thermally acclimate its locomotory performance, jumping significantly further at lower temperatures, however, acclimation to high temperatures impacted its jump performance at cold temperatures. The frog L. raniformis was found to only be able to thermally acclimate its respiration and locomotion to high temperatures, as performance at low temperatures was often poor. The data shows that L. ewingii is a cold temperate frog rather than a warm habitat frog, while L. raniformis is an animal adapted to warm temperatures. From this we can begin to see the effect that temperature has on the physiology of these two exotic frogs and the major role that temperature may be playing in driving the differences seen in the distribution of these two species in New Zealand.

Fishes are the most speciose group of living vertebrates, making up more than half of extant vertebrate diversity. They have evolved a wide array of swimming modes and body forms, including the batoid elasmobranchs, the dorsoventrally flattened skates and rays, which swim via oscillations or undulations of a broad pectoral fin disc. In this work I offer insights into locomotion by an undulatory batoid, freshwater stingray Potamotrygon orbignyi (Castelnau, 1855), combining studies of live animals, physical models, and preserved specimens. In Chapter 1, I quantify the three-dimensional kinematics of the P. orbignyi pectoral fin during undulatory locomotion, analyzing high-speed video to reconstruct three-dimensional pectoral fin motions. A relatively small portion (~25%) of the pectoral fin undulates with significant amplitude during swimming. To swim faster, stingrays increase the frequency, not the amplitude of propulsive motions, similar to the majority of studied fish species. Intermittently during swimming, a sharp, concave-down lateral curvature occurred at the fin margin; as the fin was cupped against the pressure of fluid flow this curvature is likely to be actively controlled. Chapter 2 employs a simple physical model of an undulating fin to examine the ground effects that stingrays may experience when swimming near a substrate. Previous research considering static air- and hydrofoils indicated that near-substrate locomotion offers a benefit to propulsion. Depending on small variations in swimming kinematics, undulating fins can swim faster near a solid boundary, but can also experience significant increases (~25%) in cost-of-transport. In Chapter 3, I determine how pectoral and pelvic fin locomotion are combined in P. orbignyi during augmented punting, a hybrid of pectoral and pelvic fin locomotion sometimes employed as stingrays move across a substrate. The timing of pectoral and pelvic fin motions is linked, indicating coordination of thrust production. Chapter 4 discusses pectoral fin structure and morphological variations within the fin, correlating morphology with the swimming kinematics observed in Chapter 1. Passive and active mechanisms may stiffen the anterior fin to create the stable leading edge seen during swimming; stingrays have converged on several structural features (fin ray segmentation and branching) shared by actinopterygian fishes.

Biological research over the past several decades has elucidated some of the mechanisms behind highly mobile, efficient, and robust locomotion in insects such as the cockroach. Roboticists have used this information to create biologically-inspired machines capable of running, jumping, and climbing robustly over a variety of terrains. To date, little work has been done to develop an at-scale insect-inspired robot capable of similar feats, due to limitations in fabrication, actuation, and electronics integration at small scales. This thesis addresses these challenges, focusing on the mechanical design and fabrication of a sub-2g walking robot, the Harvard Ambulatory MicroRobot (HAMR). The development of HAMR includes modeling and parameter selection for a two degree of freedom leg powertrain that enables locomotion. In addition, a design inspired by pop-up books that enables fast and repeatable assembly of the miniature walking robot is presented. Finally, a method to drive HAMR resulting in speeds up to 37cm/s is presented, along with simple control schemes. / Engineering and Applied Sciences

published_or_final_version / Chemistry / Doctoral / Doctor of Philosophy

No description available.

The stopping of an earlier intended action is best explained in a race between a go process and a stop process (Logan & Cowan, 1984). The finish line, to which each process races, has been likened to a point of no return, specifically one that marks the onset of a final ballistic (unstoppable) process. Of note is the typical relation of reduced go probabilities and faster go latencies at shorter signal onset asynchronies (SOAs). (The SOA is the time interval between presentation of the go signal and presentation of the stop signal.) We report, in some cases, sub-maximal surface electromyograms (EMGs) at onset when trying to stop a maximal speeded action. These data indicate reduced synaptic drive to reach the motor pools as a result of earlier stopping effects and, as such, hold important implications for a theory of control. First, we interpret these data to suggest that the point of no return is phantom. Sub-maximal EMGs indicate a point in the control stream beyond which some EMG will be later observed but, importantly, they fail to mark the onset of a final ballistic process if, once breached, the same process remains subject to further effects of stopping. The alternative interpretation, however, that of a final ballistic process that receives sub-maximal input which results in sub-maximal output (i.e., EMG onset) cannot be ruled out from these data. We used the Hoffmann (H) reflex to probe further the mechanism of control for stopping a voluntary action. The H-reflex, an involuntary reflex that is taken as an index of spinal control, is relevant to the control of stopping because it is typically facilitated a short time before EMG onset. In other words, it provides a window of control within which a final ballistic process would otherwise be expected to locate. Thus, we interpret the effects of stopping on the H-reflex before EMG onset as strong evidence against a final ballistic process. Second, while the race model can explain the relation between the go probabilities, the go latencies and the SOAs, it fails to explain the sub-maximal EMG onsets that describe that same action in some cases. We submit a mechanism of excitatory-inhibitory interaction at all times up to the motor pool to explain both sets of empirical data. The viability of this theory is demonstrated using computer analyses.

Recently it has been suggested that the use of neuromechanical simulations could be used to further our understanding of the neural control mechanisms involved in the control of animal locomotion. The models used to carry out these neuromechanical simulations typically consist of a representation of the neural control systems involved in walking and a representation of the mechanical locomotor apparatus. These separate models are then integrated to produce motion of the locomotor apparatus based on signals that are generated by the neural control models. Typically in past neuromechanical simulations of human walking the parameters of the neural control model have been specifically chosen to produce a walking pattern that resembles the normal human walking pattern as closely as possible. Relatively few of these studies have systematically tested the effect of manipulating the control parameters on the walking pattern that is produced by the locomotor apparatus. The goal of this thesis was to develop models of the locomotor control system and the human locomotor apparatus and systematically manipulate several parameters of the neural control system and determine what effects these parameters would have on the walking pattern of the mechanical model. Specifically neural control models were created of the Central Pattern Generator (CPG), feedback mechanisms from muscle spindles and contact sensors that detect when the foot was contact with the ground. Two models of the human locomotor apparatus were used to evaluate the outputs of the neural control systems; the first was a rod pendulum, which represented a swinging lower-limb, while the second was a 5-segment biped model, which included contact dynamics with the ground and a support system model to maintain balance. The first study of this thesis tested the ability of a CPG model to control the frequency and amplitude of the pendulum model of the lower-limb, with a strictly feedforward control mechanism. It was found that the frequency of the pendulum’s motion was directly linked (or entrained) to the frequency of the CPG’s output. It was also found that the amplitude of the pendulum’s motion was affected by the frequency of the CPG’s output, with the greatest amplitude of motion occurring when the frequency of the CPG matched the pendulum’s natural frequency. The effects of altering several other parameters of the pendulum model, such as the initial angle, the magnitude of the applied viscous damping or the moment arms of the muscles, were also analyzed. The second study again used the pendulum model, and added feedback to the neural control model, via output from simulated muscle spindles. The output from these spindle models was used to trigger a simulated stretch reflex. It was found that the addition of feedback led to sensory entrainment of the CPG output to the natural frequency of the pendulum. The effects of altering the muscle spindle’s sensitivity to length and velocity changes were also examined. The ability of this type of feedback system to respond to mechanical perturbations was also analyzed. The third and fourth studies used a biped model of the musculoskeletal system to assess the effects of altering the parameters of the neural control systems that were developed in the first two studies. In the third study, the neural control system consisted only of feedforward control from the CPG model. It was found that the walking speed of the biped model could be controlled by altering the frequency of the CPG’s output. It was also observed that variability of the walking pattern was decreased when there was a moderate level of inhibition between the CPGs of the left and right hip joints. The final study added feedback from muscle receptors and from contact sensors with the ground. It was found that the most important source of feedback was from the contact sensors to the extensor centres of the CPG. This feedback increased the level of extensor activity and produced significantly faster walking speeds when compared to other types of feedback. This thesis was successful in testing the effects of several control parameters of the neural control system on the movement of mechanical systems. Particularly important findings included the importance of connectivity between the CPGs of the left and right hip joints and positive feedback regarding the loading of the limb for establishing an appropriate forward walking speed. It is hoped that the models developed in this thesis can form the basis of future neuromechanical models and that the simulations carried out in this thesis help provide a better understanding of the interactions between neural and mechanical systems during the control of locomotion.

How behaviors are generated by neural circuits is one of the central questions in neurobiology. Under standard culture conditions, Caenorhabditis elegans travel by propagating sinusoidal waves, moving primarily forward, punctuated by brief runs of backing. How these behaviors are generated and altered is not well understood. Using a combination of behavioral analyses and neuronal imaging, I reveal that an activity imbalance between cholinergic A- and B-motoneurons is the key determinant of directional locomotion. Furthermore, heterotypic gap junctions that couple command interneurons and motoneurons of the backward motor circuit, mediated by innexins UNC-7 in AVA and UNC-9 in A-motoneurons, respectively, establish the B>A activity pattern required for forward movement. Loss of this coupling results in both the hyperactivation of AVA backward interneurons revealing the unregulated, endogenous activity of A-motoneurons. With equal A-motoneuron activity levels as B-motoneurons, innexin mutant animals exhibit irregular body bending (kinking) instead of executing forward motion, as well as increased backing. Through a genetic screen, I identified two stomatin-like proteins as regulators of innexin UNC-9 activity that affect C. elegans’ directional movement. The loss of function of stomatin-like unc-1 leads to the same kinker phenotype as unc-7 or unc-9 mutants. Like UNC-9, UNC-1 functions primarily in the A-motoneurons to allow forward motion, suggesting that UNC-1 is required for effective UNC-7-UNC-9 coupling between AVA and A-motoneurons. Dominant mutations in UNC-1, and another stomatin-like protein STO-6, exhibit genetic interactions with these innexin mutants. These mutations partially restore the forward movement of unc-7 mutants, in an UNC-9-dependent manner, indicating that they regulate UNC-9 channel activity in motoneurons to re-establish the B>A-motoneuron activity pattern in the absence of heterotypic gap junctions between interneurons and motoneurons. These studies describe a role of gap junctions as regulators of circuit dynamics by establishing an imbalanced motoneuron activity pattern that favors forward motion, which can be modulated by upper layer inputs. This study also identifies stomatin-like regulators of innexin hemichannel and gap junction function. Future work will focus on understanding mechanisms through which these stomatins regulate the activity of specific innexin channels in C. elegans motoneurons, as well as their contribution to the dynamic output of the C. elegans motor circuit.

No description available.