Roll Perturbations Research Articles

The role of mice tails in response to external and self-generated balance perturbations on the roll plane.

Chordate tails exhibit considerable morphological and functional diversity, with variations in length, diameter and texture adapted to various ecological roles. While some animals, including humans, have lost or reduced their tails, many vertebrates retain and use their tails for activities such as balancing, climbing, and escaping predators. This study investigates how laboratory mice (Mus musculus) use their tails to maintain balance when dealing with external and self-generated perturbations. Mice crossed platforms of different widths, while responding to roll-plane tilts. Our findings show that mice swing their tails to counteract external roll perturbations, generating angular momentum to stabilize themselves. Mice were also found to use active (dynamic stabilizer) and passive (counterweight) tail movement strategies when locomoting on narrow platforms. The results suggest that the tail is a core component of mouse locomotion, especially in challenging balancing conditions.

Linear and non-linear stability analysis of double-diffusive Hadley-Prats flow through horizontal porous layer with non-uniform thermal and solutal gradients

A scientific model is devised to understand the onset of double-diffusion Hadley-Prats flow through an infinite parallel permeable channel with non-uniform thermal, solutal gradients, and concentration based internal heat generation under the influence of convection conditions is considered. Darcy’s model is implemented in the context of a porous medium, which is characterized as isotropic and homogeneous. A linear and nonlinear stability analysis is conducted with the help of energy functional method and longitudinal roll perturbations are examined. The emerged dimensionless eigenvalue problem in the linear and nonlinear stability analysis is solved numerically by utilizing fourth order Runge-Kutta method (RK-4) method and shooting scheme. The critical values of wave and vertical thermal Rayleigh numbers are examined. A comprehensive analysis of solutions concerning the onset of convection mechanism is conducted. The produced results are discussed for the various values of emerged physical parameters. It is evident that, the presence of non-uniform thermal and solutal gradients and the internal heat generation mechanism causes the significant variations in the critical values of vertical thermal Rayleigh number in the channel regime.

A Novel 3D Path Following Scheme for a Class of Underactuated Underwater Vehicles with Unconventional Actuation

A 3D path following algorithm involving a novel motion control methodology is investigated for use on a new class of autonomous underwater vehicle (AUV). Unlike standard path following AUV control algorithms for underactuated vehicles, where roll perturbations are avoided, this unconventional algorithm utilizes active roll control to achieve a desired heading. The desired heading is calculated using traditional look-ahead techniques, and is achieved by performing a roll rotation to align the body-fixed yaw plane with the desired heading, and then a yaw rotation is performed to achieve that desired heading. It is demonstrated that this new path following algorithm achieves good tracking results, but experiences complications for increased forward velocities. It is also demonstrated that the complications at higher velocities can be overcome with adaptive adjustment of the look-ahead algorithm parameters.

An Output Feedback Approach for Regulation of 5-DOF Offshore Cranes With Ship Yaw and Roll Perturbations

In practice, offshore cranes are effective transportation tools used on ships. Different from land-fixed cranes, offshore cranes work in the <italic xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink">noninertial</i> frame, which are usually affected by different disturbances. Therefore, the control problem of offshore cranes is much more <italic xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink">difficult</i> . Till now, only few control methods have been proposed for offshore cranes, which are usually designed for <italic xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink">planar</i> offshore cranes, while the more practical three-dimensional (3-D) movements are <italic xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink">ignored</i> . Considering these facts, in this article, we focus on the control problem of a 5 degrees of freedom (DOF) offshore crane in 3-D space with persistent ship yaw and roll perturbations and propose an effective <italic xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink">output feedback</i> control method. Specifically, we first present an elaborate coordinate transformation method to deal with ship perturbations. Then an energy-like function is constructed based on the transformed model, and then by designing some auxiliary signals, an output feedback method is designed with rigorous mathematical analysis to prove the closed-loop asymptotic stability results. As far as we know, the proposed method is the <italic xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink">first</i> control method designed for 5-DOF offshore cranes <italic xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink">without</i> any linearization, which <italic xmlns:mml="http://www.w3.org/1998/Math/MathML" xmlns:xlink="http://www.w3.org/1999/xlink">only</i> uses output signals. Finally, experimental results are included to verify the performance of the proposed method.

Modulation of Flight Muscle Recruitment and Wing Rotation Enables Hummingbirds to Mitigate Aerial Roll Perturbations.

Both biological and artificial fliers must contend with aerial perturbations that are ubiquitous in the outdoor environment. Flapping fliers are generally least stable but also most maneuverable around the roll axis, yet our knowledge of roll control in biological fliers remains limited. Hummingbirds are suitable models for linking aerodynamic perturbations to flight control strategies, as these small, powerful fliers are capable of remaining airborne even in adverse wind conditions. We challenged hummingbirds to fly within a steady, longitudinally (streamwise) oriented vortex that imposed a continuous roll perturbation, measured wing kinematics and neuromotor activation of the flight muscles with synchronized high-speed video and electromyography and used computational fluid dynamics (CFD) to estimate the aerodynamic forces generated by observed wing motions. Hummingbirds responded to the perturbation with bilateral differences in activation of the main flight muscles while maintaining symmetry in most major aspects of wing motion, including stroke amplitude, stroke plane angle, and flapping frequency. Hummingbirds did display consistent bilateral differences in subtler wing kinematic traits, including wing rotation and elevation. CFD modeling revealed that asymmetric wing rotation was critical for attenuating the effects of the perturbation. The birds also augmented flight stabilization by adjusting body and tail posture to expose greater surface area to upwash than to the undesirable downwash. Our results provide insight into the remarkable capacity of hummingbirds to maintain flight control, as well as bio-inspiration for simple yet effective control strategies that could allow robotic fliers to contend with unfamiliar and challenging real-world aerial conditions.

Identification of In-Flight Wingtip Folding Effects on the Roll Characteristics of a Flexible Aircraft

Wingtip folding is a means by which an aircraft’s wingspan can be extended, allowing designers to take advantage of the associated reduction in induced drag. This type of device can provide other benefits if used in flight, such as flight control and load alleviation. In this paper, the authors present a method to develop reduced order flight dynamic models for in-flight wingtip folding, which are suitable for implementation in real-time pilot-in-the-loop simulations. Aspects such as the impact of wingtip size and folding angle on aircraft roll dynamics are investigated along with failure scenarios using a time domain aeroservoelastic framework and an established system identification method. The process discussed in this paper helps remove the need for direct connection of complex physics based models to engineering flight simulators and the need for tedious programming of large look-up-tables in simulators. Instead, it has been shown that a generic polynomial model for roll aeroderivatives can be used in small roll perturbation conditions to simulate the roll characteristics of an aerodynamic derivative based large transport aircraft equipped with varying fold hinge lines and tip deflections. Moreover, the effects of wing flexibility are also considered.

Modulation of Flight Muscle Recruitment and Wing Rotation Enables Hummingbirds to Mitigate Aerial Roll Perturbations

Both biological and artificial fliers must contend with aerial perturbations that are ubiquitous in the outdoor environment. Flapping fliers are generally least stable, but also the most maneuverable in roll, yet roll control in biological fliers remains less well understood. Hummingbirds are suitable models for linking aerodynamic perturbations to flight control strategies, as these small, powerful fliers are capable of remaining airborne even in adverse airflows. We challenged hummingbirds to fly within a longitudinally oriented vortex that imposed a continuous roll perturbation, measured wing kinematics and neuromotor activation of the major flight muscles with synchronized high-speed video and electromyography and used computational fluid dynamics (CFD) to estimate the aerodynamic forces generated by wing motions. Hummingbirds responded to the perturbation using bilaterally different activation of the main flight muscles and maintained symmetry in most major aspects of wing motion including stroke amplitude, stroke plane angle, and flapping frequency. However, hummingbirds also displayed consistent bilateral differences in subtle wing kinematics traits, including wing rotation and elevation. CFD modeling implicate asymmetric responses in wing rotation as important for generating the necessary stabilizing torques, suggesting that intrinsic wing muscles play a critical role in aerodynamic control. The birds also augment flight stabilization by adjusting body and tail posture to expose greater surface area to upwash than to the undesirable downwash. Our results provide insight into the remarkable capacity of hummingbirds to maintain flight control and bio-inspiration for simple yet effective control strategies for robotic fliers to contend with unfamiliar and challenging real-word aerial conditions.

Controlling roll perturbations in fruit flies.

Owing to aerodynamic instabilities, stable flapping flight requires ever-present fast corrective actions. Here, we investigate how flies control perturbations along their body roll angle, which is unstable and their most sensitive degree of freedom. We glue a magnet to each fly and apply a short magnetic pulse that rolls it in mid-air. Fast video shows flies correct perturbations up to 100° within 30 ± 7 ms by applying a stroke-amplitude asymmetry that is well described by a linear proportional-integral controller. For more aggressive perturbations, we show evidence for nonlinear and hierarchical control mechanisms. Flies respond to roll perturbations within 5 ms, making this correction reflex one of the fastest in the animal kingdom.

Streamwise vortices destabilize swimming bluegill sunfish (Lepomis macrochirus).

In their natural environment, fish must swim stably through unsteady flows and vortices, including vertical vortices, typically shed by posts in a flow, horizontal cross-flow vortices, often produced by a step or a waterfall in a stream, and streamwise vortices, where the axis of rotation is aligned with the direction of the flow. Streamwise vortices are commonly shed by bluff bodies in streams and by ships' propellers and axial turbines, but we know little about their effects on fish. Here, we describe how bluegill sunfish use more energy and are destabilized more often in flow with strong streamwise vorticity. The vortices were created inside a sealed flow tank by an array of four turbines with similar diameter to the experimental fish. We measured oxygen consumption for seven sunfish swimming at 1.5 body lengths (BL)s(-1) with the turbines rotating at 2 Hz and with the turbines off (control). Simultaneously, we filmed the fish ventrally and recorded the fraction of time spent maneuvering side-to-side and accelerating forward. Separately, we also recorded lateral and ventral video for a combination of swimming speeds (0.5, 1.5 and 2.5 BLs(-1)) and turbine speeds (0, 1, 2 and 3 Hz), immediately after turning the turbines on and 10 min later to test for accommodation. Bluegill sunfish are negatively affected by streamwise vorticity. Spills (loss of heading), maneuvers and accelerations were more frequent when the turbines were on than in the control treatment. These unsteady behaviors, particularly acceleration, correlated with an increase in oxygen consumption in the vortex flow. Bluegill sunfish are generally fast to recover from roll perturbations and do so by moving their pectoral fins. The frequency of spills decreased after the turbines had run for 10 min, but was still markedly higher than in the control, showing that fish partially adapt to streamwise vorticity, but not completely. Coping with streamwise vorticity may be an important energetic cost for stream fishes or migratory fishes.

Influence of Turbulence on MAV Roll Perturbations

There are significant challenges in the control of fixed-wing Micro Air Vehicles (MAVs) in high turbulence environments. Birds can sustain stable flight in such environments by obtaining flow information through mechanoreceptors embedded in their wings. Inspired by nature's flyers, an investigation into replicating the function of mechanoreceptors with commercially available pressure sensors is presented. Implementation requires an in-depth understanding of the level of correlation that exists between pressure variations over the wing and the roll perturbation of the MAV. This paper investigates the variation in correlation and coherence along a representative wing-chord and wing-span of a MAV. Highest correlation and coherence is found to exist in the vicinity of the leading edge, with significant perturbations which are evident up to ∼35Hz for a 0.49m wingspan MAV.

Feed-forward and visual feed-back control of head roll orientation in wasps (Polistes humilis, Vespidae, Hymenoptera)

Flying insects keep their visual system horizontally aligned, suggesting that gaze stabilization is a crucial first step in flight control. Unlike flies, hymenopteran insects such as bees and wasps do not have halteres that provide fast, feed-forward angular rate information to stabilize head orientation in the presence of body rotations. We tested whether hymenopteran insects use inertial (mechanosensory) information to control head orientation from other sources, such as the wings, by applying periodic roll perturbations to male Polistes humilis wasps flying in tether under different visual conditions indoors and in natural outdoor conditions. We oscillated the thorax of the insects with frequency-modulated sinusoids (chirps) with frequencies increasing from 0.2 to 2 Hz at a maximal amplitude of 50 deg peak-to-peak and maximal angular velocity of ±245 deg s(-1). We found that head roll stabilization is best outdoors, but completely absent in uniform visual conditions and in darkness. Step responses confirm that compensatory head roll movements are purely visually driven. Modelling step responses indicates that head roll stabilization is achieved by merging information on head angular velocity, presumably provided by motion-sensitive neurons and information on head orientation, presumably provided by light level integration across the compound eyes and/or ocelli (dorsal light response). Body roll in free flight reaches amplitudes of ±40 deg and angular velocities greater than 1000 deg s(-1), while head orientation remains horizontal for most of the time to within ±10 deg. In free flight, we did not find a delay between spontaneous body roll and compensatory head movements, and suggest that this is evidence for the contribution of a feed-forward control to head stabilization.

154 SENSORY BIAS PREDICTS POSTURAL STABILITY, ANXIETY, AND COGNITIVE PERFORMANCE IN HEALTHY ADULTS WALKING IN NOVEL DISCORDANT CONDITIONS

We designed a gait training study that presented combinations of visual flow and support surface manipulations to investigate the response of healthy adults to novel discordant sensorimotor conditions. We aimed to determine whether a relationship existed between subjects visual dependence and their scores on a collective measure of anxiety, cognition, and postural stability in a new discordant environment presented at the conclusion of training (Transfer Test). A treadmill was mounted to a motion base platform positioned 2 m behind a large visual screen. Training consisted of three walking sessions, each within a week of the previous visit, that presented four 5-minute exposures to various combinations of support surface and visual scene manipulations, all lateral sinusoids. The conditions were scene translation only, support surface translation only, simultaneous scene and support surface translations in-phase, and simultaneous scene and support surface translations 180 out-of-phase. During the Transfer Test, the trained participants received a 2-minute novel exposure. A visual sinusoidal roll perturbation, with twice the original flow rate, was superimposed on a sinusoidal support surface roll perturbation that was 90 out of phase with the scene. A high correlation existed between normalized torso translation, measured in the scene-only condition at the first visit, and a combined measure of normalized heart rate, stride frequency, and reaction time at the transfer test. Results suggest that visually dependent participants experience decreased postural stability, increased anxiety, and increased reaction times compared to their less visually dependent counterparts when negotiating novel discordant conditions.

216 GAIT TRAINING IMPROVES PERFORMANCE IN HEALTHY ADULTS EXPOSED TO NOVEL DISCORDANT CONDITIONS

After they return to Earth, astronauts experience sensorimotor disturbances that disrupt their ability to walk. We have previously shown that training with a variety of sensorimotor adaptive challenges enhances the capability of adapting to novel sensorimotor conditions. We are currently developing a sensorimotor adaptability (SA) training program designed to facilitate recovery of function after gravitational transitions. The purpose of this study was to determine whether trained subjects could transfer learned skills from one discordant visuo-proprioceptive environment to another. During three sessions, subjects walked at 2.5 km/h on a treadmill mounted on a motion base platform. Ten subjects trained with a combination of lateral treadmill translation and superimposed sinusoidal lateral optic flow that was presented on a large screen positioned in front of them. Ten controls completed the same training schedule while viewing only the forward optic flow with no visual or physical oscillation. Twenty minutes after the final training session, all subjects completed a 2-minute trial with a novel combination of visual and treadmill roll perturbations not previously experienced during the training (Transfer Test). Compared to the untrained group, participants who received SA training showed faster reaction times and, based on a composite score derived from stride frequency, heart rate, and reaction time, an overall enhanced performance. Our results showed that an SA training program can improve overall walking performance when subjects are exposed to novel incongruent sensory environments. This training has application for both enhancing adaptive responses in astronauts and reducing fall and injury risk in the elderly.

Control of roll and pitch motion during multi-directional balance perturbations

Does the central nervous system (CNS) independently control roll and pitch movements of the human body during balance corrections? To help provide an answer to this question, we perturbed the balance of 16 young healthy subjects using multi-directional rotations of the support surface. All rotations had pitch and roll components, for which either the roll (DR) or the pitch (DP) component were delayed by 150 ms or not at all (ND). The outcome measures were the biomechanical responses of the body and surface EMG activity of several muscles. Across all perturbation directions, DR caused equally delayed shifts (150 ms) in peak lateral centre of mass (COM) velocity. Across directions, DP did not cause equally delayed shifts in anterior-posterior COM velocity. After 300 ms however, the vector direction of COM velocity was similar to the ND directions. Trunk, arm and knee joint rotations followed this roll compared to pitch pattern, but were different from ND rotation synergies after 300 ms, suggesting an intersegmental compensation for the delay effects. Balance correcting responses of muscles demonstrated both roll and pitch directed components regardless of axial alignment. We categorised muscles into three groups: pitch oriented, roll oriented and mixed based on their responses to DR and DP. Lower leg muscles were pitch oriented, trunk muscles were roll oriented, and knee and arm muscles were mixed. The results of this study suggest that roll, but not pitch components, of balance correcting movement strategies and muscle synergies are separately programmed by the CNS. Reliance on differentially activated arm and knee muscles to correct roll perturbations reveals a dependence of the pitch response on that of roll, possibly due to biomechanical constraints, and accounts for the failure of DP to be transmitted equally in time across all limbs segments. Thus it appears the CNS preferentially programs the roll response of the body and then adjusts the pitch response accordingly.

Spatio-Temporal Separation of Roll and Pitch Balance-Correcting Commands in Humans

This study was designed to provide evidence for the hypothesis that human balance corrections in response to pitch perturbations are controlled by muscle action mainly about the ankle and knee joints, whereas balance corrections for roll perturbations are controlled predominantly by motion about the hip and lumbro-sacral joints. A dual-axis rotating support surface delivered unexpected random perturbations to the stance of 19 healthy young adults through eight different directions in the pitch and the roll planes and three delays between pitch and roll directions. Roll delays with respect to pitch were no delay, a short 50-ms delay of roll with respect to pitch movements, (chosen to correspond to the onset time of leg muscle stretch reflexes), and a long 150-ms delay between roll and pitch movements (chosen to shift the time when trunk roll velocity peaks to the time when trunk peak pitch velocity normally occurs). Delays of stimulus roll with respect to pitch resulted in delayed roll responses of the legs, trunk, arms, and head consistent with stimulus delay without any changes in roll velocity amplitude. Delayed roll perturbations induced only small changes in the pitch motion of the legs and trunk; however, major changes were seen in the time when roll motion of the trunk was arrested. Amplitudes and directional sensitivity of short-latency (SL) stretch reflexes in ankle muscles were not altered with increasing roll delay. Small changes to balance correcting responses in ankle muscles were observed. SL stretch reflexes in hip and trunk muscles were delayed, and balance-correcting responses in trunk muscles became split into two distinct responses with delayed roll. The first of these responses was small and had a directional responsiveness aligned more along the pitch plane. The main, larger, response occurred with an onset and time-to-peak consistent with the delay in trunk roll displacement and its directional responsiveness was roll oriented. The sum of the amplitudes of these two types of balance-correcting responses remained constant with roll delay. These results support the hypothesis that corrections of the body's pitch and roll motion are programmed separately by neural command signals and provide insights into possible triggering mechanisms. The evidence that lower leg muscle balance-correcting activity is hardly changed by delayed trunk roll also indicates that lower leg muscle activity is not predominant in correcting roll motion of the body. Lower leg and trunk muscle activity appears to have a dual action in balance corrections. In trunk muscles the main action is to correct for roll perturbations and the lesser action may be an anticipatory stabilizing reaction for pitch perturbations. Likewise, the small changes in lower leg muscle activity may result from a generalized stabilizing reaction to roll perturbations, but the main action is to correct for pitch perturbations.

Transition in Homogeneously Heated Inclined Plane Parallel Shear Flows

The transition of internally heated inclined plane parallel shear flows is examined numerically for the case of finite values of the Prandtl number Pr. We show that as the strength of the homogeneously distributed heat source is increased the basic flow loses stability to two-dimensional perturbations of the transverse roll type in a Hopf bifurcation for the vertical orientation of the fluid layer, whereas perturbations of the longitudinal roll type are most dangerous for a wide range of the value of the angle of inclination. In the case of the horizontal inclination transverse roll and longitudinal roll perturbations share the responsibility for the prime instability. Following the linear stability analysis for the general inclination of the fluid layer our attention is focused on a numerical study of the finite amplitude secondary travelling-wave solutions (TW) that develop from the perturbations of the transverse roll type for the vertical inclination of the fluid layer. The stability of the secondary TW against three-dimensional perturbations is also examined and our study shows that for Pr=0.71 the secondary instability sets in as a quasi-periodic mode, while for Pr=7 it is phase-locked to the secondary TW. The present study complements and extends the recent study by Nagata and Generalis (2002) in the case of vertical inclination for Pr=0.

Vestibular influences on human postural control in combinations of pitch and roll planes reveal differences in spatiotemporal processing

The present study examined the influence of bilateral peripheral vestibular loss (BVL) in humans on postural responses to multidirectional surface rotations in the pitch and roll planes. Specifically, we examined the effects of vestibular loss on the directional sensitivity, timing, and amplitude of early stretch, balance correcting, and stabilizing reactions in postural leg and trunk muscles as well as changes in ankle torque and trunk angular velocity following multidirectional rotational perturbations of the support surface. Fourteen normal healthy adults and five BVL patients stood on a dual axis rotating platform which rotated 7.5 degrees at 50 degrees/s through eight different directions of pitch and roll combinations separated by 45 degrees. Directions were randomized within a series of 44 perturbation trials which were presented first with eyes open, followed by a second series of trials with eyes closed. Vestibular loss did not influence the range of activation or direction of maximum sensitivity for balance correcting responses (120-220 ms). Response onsets at approximately 120 ms were normal in tibialis anterior (TA), soleus (SOL), paraspinals (PARAS), or quadriceps muscles. Only SOL muscle activity demonstrated a 38- to 45-ms delay for combinations of forward (toe-down) and roll perturbations in BVL patients. The amplitude of balance correcting responses in leg muscles between 120 and 220 ms was, with one exception, severely reduced in BVL patients for eyes open and eyes closed conditions. SOL responses were decreased bilaterally for toe-up and toe-down perturbations, but more significantly reduced in the downhill (load-bearing) leg for combined roll and pitch perturbations. TA was significantly reduced bilaterally for toe-up perturbations, and in the downhill leg for backward roll perturbations. Forward perturbations, however, elicited significantly larger TA activity in BVL between 120 and 220 ms compared to normals, which would act to further destabilize the body. As a result of these changes in response amplitudes, BVL patients had reduced balance correcting ankle torque between 160 and 260 ms and increased torque between 280 and 380 ms compared to normals. There were no differences in the orientation of the resultant ankle torque vectors between BVL and normals, both of which were oriented primarily along the pitch plane. For combinations of backward (toe-up) and roll perturbations BVL patients had larger balance correcting and stabilizing reactions (between 350 and 700 ms) in PARAS than normals and these corresponded to excessive trunk pitch and roll velocities. During roll perturbations, trunk velocities in BVL subjects after 200 ms were directed along directions different from those of normals. Furthermore, roll instabilities appeared later than those of pitch particularly for backward roll perturbations. The results of the study show that combinations of roll and pitch surface rotations yield important spatiotemporal information, especially with respect to trunk response strategies changed by BVL which are not revealed by pitch plane perturbations alone. Our results indicate that vestibular influences are earlier for the pitch plane and are directed to leg muscles, whereas roll control is later and focused on trunk muscles.

Open inflation from quantum cosmology with a strong non-minimal coupling

We propose a mechanism of quantum creation of the open Universe in the observable range of values of Ω. This mechanism is based on the no-boundary quantum state with the Hawking–Turok instanton applied to the model with a strong non-minimal coupling of the inflaton field. We develop the slow roll perturbation expansion for the instanton solution and obtain a non-trivial contribution to the classical instanton action. The interplay of this classical contribution with the loop effects due to quantum effective action generates the probability distribution peak with necessary parameters of the inflation stage without invoking any anthropic considerations. In contrast with a similar mechanism for closed models, existing only for the tunneling quantum state of the Universe, the observationally justified open inflation originates from the no-boundary cosmological wavefunction.

Directional sensitivity of stretch reflexes and balance corrections for normal subjects in the roll and pitch planes.

A large body of evidence has been collected which describes the response parameters associated with automatic balance corrections in man to perturbations in the pitch plane. However, perturbations to human stance can be expected from multiple directions. The purpose of the present study was to describe the directional sensitivities of muscle responses re-establishing disturbed stance equilibrium in normal subjects. The contributions of stretch reflex and automatic balance-correcting responses to balance control, and concomitant biomechanical reactions, were examined for combinations of pitch and roll perturbations of the support surface. More specifically, muscle responses, initial head accelerations and trunk velocities were analyzed with the intention of identifying possible origins of directionally specific triggering signals and to examine how sensory information is used to modulate triggered balance corrections with respect to direction. Fourteen healthy adults were required to stand on a dual-axis rotating platform capable of delivering rotational perturbations with constant amplitude (7.5 degrees ) and velocity (50 degrees /s) through multiple directions in the pitch and roll planes. Each subject was randomly presented with 44 support surface rotations through 16 different directions separated by 22.5 degrees first under eyes-open, and then, for a second identical set of rotations, under eyes-closed conditions. Bilateral muscle activities from tibialis anterior, soleus, lateral quadriceps and paraspinals were recorded, averaged across direction, and areas calculated over intervals with significant bursts of activity. Trunk angular velocity and ankle torque data were averaged over intervals corresponding to significant biomechanical events. Stretch reflex (intervals of 40-100, 80-120 ms) and automatic balance-correcting responses (120-220, 240-340 ms) in the same muscle were sensitive to distinctly different directions. The directions of the maximum amplitude of balance-correcting activity in leg muscles were oriented along the pitch plane, approximately 180 degrees from the maximum amplitude of stretch responses. Ankle torques for almost all perturbation directions were also aligned along the pitch plane. Stretch reflexes in paraspinal muscles were tuned along the 45 degrees plane but at 90 degrees to automatic balance corrections and 180 degrees to unloading responses in the same muscle. Stretch reflex onsets in paraspinal muscles were observed at 60 ms, as early as those of soleus muscles. In contrast, unloading reflexes in released paraspinal muscles were observed at 40 ms for perturbations which caused roll of the trunk towards the recorded muscle. Onsets of trunk roll velocities were earlier and more rapid than those observed for pitch velocities. Trunk pitch occurred for pure roll directions but not vice versa. When considered together, early stretch and unloading of paraspinals, and concomitant roll and pitch velocities of the trunk requiring a roll-and-pitch-based hip torque strategy, bring into question previous hypotheses of an ankle-based trigger signal or ankle-based movement strategies for postural balance reactions. These findings are compatible with the hypothesis that stretch-, force- and joint-related proprioceptive receptors at the level of the trunk provide a directionally sensitive triggering mechanism underlying a, minimally, two-stage (pitch-based leg and pitch-and-roll-based trunk) balance-correcting strategy. Accelerometer recordings from the head identified large vertical linear accelerations only for pitch movements and angular roll accelerations only during roll perturbations with latencies as early as 15 ms. Thus, it appears that balance corrections in leg and trunk muscles may receive strong, receptor-dependent (otolith or vertical canal) and directionally sensitive amplitude-modulating input from vestibulospinal signals.

Resistance reflex that maintains upright head posture in the flesh fly neobellieria bullata (Sarcophagidae)

In flesh flies Neobellieria bullata, we investigated a resistance reflex that maintains upright head posture around the roll axis relative to the thorax. The gain of the reflex depends upon the fly's behavioral state: moving flies immediately correct 90 % of the amplitude of experimentally imposed roll perturbations, returning the head almost to the fully upright position; motionless flies allow perturbations to persist for minutes before correcting only 70 % of perturbation amplitude. To investigate the role of various neural pathways, we examined the control of head posture after sectioning relevant propriosensory or motor nerves. Excision of the prosternal chordotonal organ causes no decrements in the control of head posture. Unilateral deafferentation of a cervical propriosensory organ, the prosternal organ, induces roll towards the cut side. Unilateral section of the frontal nerve, a mixed motor nerve that supplies the neck depressors and levators, leads to unilateral deficits in correcting perturbations towards the contralateral side. After bilateral propriosensory or frontal motor nerve section, approximately 40 % of perturbation amplitude is still corrected. To determine the contributions of the passive elastic properties of the neck skeleto-muscular system, flies were tested under reversible nitrogen anesthesia. They immediately corrected 40 % of perturbation amplitude. Taken together, the results demonstrate that passive elasticity plus active prosternal nerve afference to contralateral depressors innervated by the frontal nerve in combination constitute a sufficient and necessary reflex loop to control head roll posture.