The halteres of the blowfly Calliphora

The movement of the halteres during fixed flight was video recorded under stroboscopic illumination phase coupled to the wing beat. The halteres swing in a rounded triangular manner through an angle of almost 80° in vertical planes tilted backwards from the transverse plane by ca. 30° (Figs. 1, 2).The physics of the halteres are described in terms of a general formula for the force acting onto the endknob of the moving haltere during rotations and linear accelerations of the fly (Eq. 1). On the basis of the experimentally determined kinematics of the haltere, the primary forces and the forces dependent on angular velocity and on angular acceleration are calculated (Figs. 3, 4).Three distinct types of angular velocity dependent (Coriolis) forces are generated by rotations about 3 orthogonal axes. Thus, in principle one haltere could detect all rotations in space (Fig. 6).The angular acceleration dependent forces have the same direction and frequency as the Coriolis forces, but they are shifted in phase by 90°. Thus, they could be evaluated in parallel and independently from the Coriolis forces. They are, however, much smaller than the Coriolis forces for oscillation frequencies of the fly up to 20 Hz (Fig. 5). From these considerations it is concluded that Coriolis forces play the major role in detecting body rotations.     

Haltere-mediated equilibrium reflexes of the fruit fly, Drosophila melanogaster.

Flies display a sophisticated suite of aerial behaviours that require rapid sensory-motor processing. Like all insects, flight control in flies is mediated in part by motion-sensitive visual interneurons that project to steering motor circuitry within the thorax. Flies, however, possess a unique flight control equilibrium sense that is encoded by mechanoreceptors at the base of the halteres, small dumb-bell-shaped organs derived through evolutionary transformation of the hind wings. To study the input of the haltere system onto the flight control system, I constructed a mechanically oscillating flight arena consisting of a cylindrical array of light-emitting diodes that generated the moving image of a 30 degrees vertical stripe. The arena provided closed-loop visual feedback to elicit fixation behaviour, an orientation response in which flies maintain the position of the stripe in the front portion of their visual field by actively adjusting their wing kinematics. While flies orientate towards the stripe, the entire arena was swung back and forth while an optoelectronic device recorded the compensatory changes in wing stroke amplitude and frequency. In order to reduce the background changes in stroke kinematics resulting from the animal's closed-loop visual fixation behaviour, the responses to eight identical mechanical rotations were averaged in each trial. The results indicate that flies possess a robust equilibrium reflex in which angular rotations of the body elicit compensatory changes in both the amplitude and stroke frequency of the wings. The results of uni- and bilateral ablation experiments demonstrate that the halteres are required for these stability reflexes. The results also confirm that halteres encode angular velocity of the body by detecting the Coriolis forces that result from the linear motion of the haltere within the rotating frame of reference of the fly's thorax. By rotating the flight arena at different orientations, it was possible to construct a complete directional tuning map of the haltere-mediated reflexes. The directional tuning of the reflex is quite linear such that the kinematic responses vary as simple trigonometric functions of stimulus orientation. The reflexes function primarily to stabilize pitch and yaw within the horizontal plane.     

Within-wingbeat damping: dynamics of continuous free-flight yaw turns in Manduca sexta

Free-flight body dynamics and wing kinematics were collected from recordings of continuous, low-speed, multi-wingbeat yaw turns in hawkmoths (Manduca sexta) using stereo videography. These data were used to examine the effects of rotational damping arising from interactions between the body rotation and flapping motion (flapping counter-torque, FCT) on continuous turning. The moths were found to accelerate during downstroke, then decelerate during upstroke by an amount consistent with FCT damping. Wing kinematics related to turning were then analysed in a simulation of hawkmoth flight; results were consistent with the observed acceleration–deceleration pattern. However, an alternative wing kinematic which produced more continuous and less damped accelerations was found in the simulation. These findings demonstrate that (i) FCT damping is detectable in the dynamics of continuously turning animals and (ii) FCT-reducing kinematics do exist but were not employed by turning moths, possibly because within-wingbeat damping simplifies control of turning by allowing control systems to target angular velocity rather than acceleration.     

Haltere activity in a flightless hippoboscid fly, Crataerina pallida

The halteres of the subapterous fly parasite of swifts, Crataerina pallida, retain a full complement of sensilla. They beat during and for many minutes after wing extension, leg movements and other forms of activity. They can also be caused to beat by a variety of visual and mechanical stimuli, including sound pulses at up to 2 kHz, for several minutes in the absence of other movements. Fed flies show markedly reduced locomotory responsiveness compared to unfed flies, but the halteres of both groups appear to be equally responsive. Haltere extirpation or inactivation does not appear to reduce ocomotory responsiveness. The possibility that haltere activity depresses responsiveness is discussed.     

Measurement of six degrees of freedom head kinematics in impact conditions employing six accelerometers and three angular rate sensors (6aω configuration)

The ability to measure six degrees of freedom (6 DOF) head kinematics in motor vehicle crash conditions is important for assessing head-neck loads as well as brain injuries. A method for obtaining accurate 6 DOF head kinematics in short duration impact conditions is proposed and validated in this study. The proposed methodology utilizes six accelerometers and three angular rate sensors (6aω configuration) such that an algebraic equation is used to determine angular acceleration with respect to the body-fixed coordinate system, and angular velocity is measured directly rather than numerically integrating the angular acceleration. Head impact tests to validate the method were conducted using the internal nine accelerometer head of the Hybrid III dummy and the proposed 6aω scheme in both low (2.3?m/s) and high (4.0?m/s) speed impact conditions. The 6aω method was compared with a nine accelerometer array sensor package (NAP) as well as a configuration of three accelerometers and three angular rate sensors (3aω), both of which have been commonly used to measure 6 DOF kinematics of the head for assessment of brain and neck injuries. The ability of each of the three methods (6aω, 3aω, and NAP) to accurately measure 6 DOF head kinematics was quantified by calculating the normalized root mean squared deviation (NRMSD), which provides an average percent error over time. Results from the head impact tests indicate that the proposed 6aω scheme is capable of producing angular accelerations and linear accelerations transformed to a remote location that are comparable to that determined from the NAP scheme in both low and high speed impact conditions. The 3aω scheme was found to be unable to provide accurate angular accelerations or linear accelerations transformed to a remote location in the high speed head impact condition due to the required numerical differentiation. Both the 6aω and 3aω schemes were capable of measuring accurate angular displacement while the NAP instrumentation was unable to produce accurate angular displacement due to double numerical integration. The proposed 6aω scheme appears to be capable of measuring accurate 6 DOF kinematics of the head in any severity of impact conditions.     

Head kinematics in mini-sled tests of foam padding: relevance of linear responses from free motion headform (FMH) testing to head angular responses

The revised Federal Motor Vehicle Safety Standard (FMVSS) No. 201 specifies that the safety performance of vehicle upper interiors is determined from the resultant linear acceleration response of a free motion headform (FMH) impacting the interior at 6.7 m/s. This study addresses whether linear output data from the FMH test can be used to select an upper interior padding that decreases the likelihood of rotationally induced brain injuries. Using an experimental setup consisting of a Hybrid III head-neck structure mounted on a mini-sled platform, sagittal plane linear and angular head accelerations were measured in frontal head impacts into foam samples of various stiffness and density with a constant thickness (51 mm) at low (approximately 5.0 m/s), intermediate (approximately 7.0 m/s), and high (approximately 9.6 m/s) impact speeds. Provided that the foam samples did not bottom out, recorded peak values of angular acceleration and change in angular velocity increased approximately linearly with increasing peak resultant linear acceleration and value of the Head Injury Criterion (HIC36). The results indicate that the padding that produces the lowest possible peak angular acceleration and peak change in angular velocity without causing high peak forces is the one that produces the lowest possible HIC36 without bottoming out in the FMH test.     

Statocysts in crabs: short-term control of locomotion and long-term monitoring of hydrostatic pressure

Crabs show well-coordinated locomotion. They have proprioceptors similar to those of lobsters, but they differ in terms of their balancing systems and their condensed nervous system, which allows rapid interganglionic conduction. Typically they exhibit dynamically stable locomotion with a highly developed semicircular canal system that codes angular acceleration in each of three orthogonal planes (horizontal and vertical at 45 degrees and 135 degrees to the pitching plane). Left and right interneurons each code one direction of angular acceleration, carrying information between the brain and the thoracic ganglia. Cell A codes head-up vertical plane angular accelerations. Cell B codes rotations in the horizontal plane. Interneurons C and D code headdown vertical plane information, carrying it ipsilaterally and contralaterally respectively. These interneurons have a central role in locomotion. They are activated and have their responsiveness to angular acceleration enhanced before and during locomotion. Such simple activation pathways point to how an angular-acceleration-controlled robot (CRABOT) could be constructed. Hydrostatic pressure information carried by the thread hairs, which also sense angular acceleration, is filtered out from direct pathways onto the interneurons, but spectral analysis shows that it still has an influence via central pathways. Long-term recordings from equilibrium interneurons in free-walking crabs taken from the wild into constant conditions show tidally changing frequencies     

Haltere and visual inputs sum linearly to predict wing (but not gaze) motor output in tethered flying Drosophila

In the true flies (Diptera), the hind wings have evolved into specialized mechanosensory organs known as halteres, which are sensitive to gyroscopic and other inertial forces. Together with the fly''s visual system, the halteres direct head and wing movements through a suite of equilibrium reflexes that are crucial to the fly''s ability to maintain stable flight. As in other animals (including humans), this presents challenges to the nervous system as equilibrium reflexes driven by the inertial sensory system must be integrated with those driven by the visual system in order to control an overlapping pool of motor outputs shared between the two of them. Here, we introduce an experimental paradigm for reproducibly altering haltere stroke kinematics and use it to quantify multisensory integration of wing and gaze equilibrium reflexes. We show that multisensory wing-steering responses reflect a linear superposition of haltere-driven and visually driven responses, but that multisensory gaze responses are not well predicted by this framework. These models, based on populations, extend also to the responses of individual flies.     

Multi-day Longitudinal Assessment of Physical Activity and Sleep Behavior Among Healthy Young and Older Adults Using Wearable Sensors

ObjectivesThe number of elderly people is growing rapidly and aging is found to affect activities of daily living. Older adults are found to perform less physical activity when compared to younger ones. In the perspective of movement behavior, it is not well understood how are elderly different from younger ones. It is not known whether they produce only low frequency movement accelerations or the overall number of movements produced are reduced in elderly. It is also not known how elderly and younger ones perform movement transitions throughout the duration of a day and during night-time sleep.Material and methodsIn this study, 10 healthy young and 10 healthy old participants wore inertial measurement unit at their lower back for 3-days. The 24 hours of day were divided into four 6 hour time zones and transitions made by young and elderly were investigated. All participants performed their regular daily activities unhindered and longitudinal multi-day signals for acceleration and angular velocity were analyzed. Time-frequency analysis was performed using wavelet transform and frequency content of each movement performed was computed.ResultsWe found that both young and older adults performed significantly more low amplitude movements than medium and high amplitude movements. Healthy young adults produced significantly more movements at 1.1 Hz than older adults. Healthy young adults were also found to have produced significantly smaller number of transitions in the mid-phases of sleep. They were also found to produce significantly larger accelerations during night-time sleep transitions than their older counterparts.ConclusionThe advantages of collecting longitudinal data about human movement and sleep transition data can lead us to important clinical diagnosis. The information from longitudinal assessment can help develop lifestyle interventions for disease prevention, monitoring of chronic diseases to prevent or slow disease progression among elderly people.     

Cardiac rhythms in chick embryos during hatching

Avian embryos develop within a hard eggshell which permits the measurement of heart rate while maintaining an adequate gas exchange through the chorioallantoic membrane. Heart rate has been determined from cardiogenic signals detected either noninvasively, semi-invasively or invasively with various transducers. Firstly, we reviewed these previously-developed methods and experimental results on heart rate fluctuations in prenatal embryos. Secondly, we presented new findings on the development of heart rate fluctuations during the last stages of incubation, with emphasis on the perinatal period, which remained to be studied. Three patterns of acceleration of the instantaneous heart rate were unique to the external pipping period: irregular intermittent large accelerations, short-term repeated large accelerations and relatively long-lasting cyclic small accelerations. Besides these acceleration patterns, respiratory arrhythmia, which comprimised oscillating patterns with a period of 1-1.5 s, appeared during the external pipping period. Furthermore, additional oscillating patterns with a period of 10-15 min were found in some externally pipped embryos.     

A general Newtonian simulation of an n-segment open chain model

This paper presents a set of general Newtonian equations which govern the simulation of movement of a body represented by n open chain links. The input for the simulation consisted of the joint moment of force histories, lengths, masses and moments of inertia, the initial absolute angular displacements and velocities and, for the fixed or constrained axis of the nth segment, the acceleration history. Angular accelerations were then determined by solving n linear equations simultaneously, and angular velocities and displacements determined by integrating forwards. The final output was in the form of a graphical display of the linked figure. Applications of the simulation were demonstrated using three-segment representations of movements of the upper and lower extremities and a five-segment representation of a jump. Good agreement was achieved between the displayed angular displacements for the original and simulated movements. The potential for varying the input data has been examined and the implications of anticipating the effects of changed torques, inertial characteristics including attached prosthetic or sports implements and/or the initial conditions for a movement are discussed.     

Negative regulation of dorsoventral signaling by the homeotic gene Ultrabithorax during haltere development in Drosophila.

Growth and patterning during Drosophila wing development are mediated by signaling from its dorsoventral (D/V) organizer. In the metathorax, wing development is essentially suppressed by the homeotic selector gene Ultrabithorax (Ubx) to mediate development of a pair of tiny balancing organs, the halteres. Here we show that expression of Ubx in the haltere D/V boundary down-regulates its D/V organizer signaling compared to that of the wing D/V boundary. Somatic loss of Ubx from the haltere D/V boundary thus results in the formation of a wing-type D/V organizer in the haltere field. Long-distance signaling from this organizer was analyzed by assaying the ability of a Ubx(-) clone induced in the haltere D/V boundary to effect homeotic transformation of capitellum cells away from the boundary. The clonally restored wing D/V organizer in mosaic halteres not only enhanced the homeotic transformation of Ubx(-) cells in the capitellum but also caused homeotic transformation of even Ubx(+) cells in a genetic background known to induce excessive cell proliferation in the imaginal discs. In addition to demonstrating a non-cell-autonomous role for Ubx during haltere development, these results reveal distinct spatial roles of Ubx during maintenance of cell fate and patterning in the halteres.     

Coordination of wing motion and walking suggests common control of zigzag motor program in a male silkworm moth

The male silkworm moth, Bombyx mori, exhibits a zigzagging pattern as it walks upwind to pheromones. This species usually does not fly, but obvious wing-beating accompanies the pheromone-mediated walking. Males supported by a `sled', after having their legs removed, also moved upwind in a pheromone plume along zigzagging tracks, indicating that wing-generated thrust and torque result in locomotory paths similar to those observed from walking moths. Using a high-speed video system we investigated the correlation between the wing movements and zigzag walking. The wing ipsilateral to the direction of the turn showed a greater degree of retraction with respect to the contralateral wing. The timing of the wing retraction pattern was synchronized with changes of direction in the walking track. Coordination of wing movements and walking pattern was not dependent on visual feedback or sensory feedback generated from neck movements associated with turning. The results presented here, taken together with our previous studies of descending interneurons suggest that the coordination of wing movements with the walking pattern may result from the activity of a set of identified interneurons descending from the brain to the thoracic ganglia and/or may be coordinated by coupling of oscillating circuits for walking and wing beating. Accepted: 15 May 1997     

Structure and Function of the Elasmobranch Auditory System

Behavioral evidence indicates that sharks detect underwatersound at frequencies up to 1000 Hz, and that certain low frequencysignals attract sharks from large distances. It appears thatthe adequate stimulus for "sound detecting" systems of the sharkis panicle motion, as opposed to fluctuations in sound pressure.The elasmobranch ear consists of the three semi-circular canalsfor detecting angular accelerations, and otolith organs fordetecting linear motion and accelerations due to gravity. Twoof these organs, the sacculus and macula neglecta, have beenshown to be responsive to vibratory motion, with the maculaneglecta having best sensitivity to vertical movements. A directvibrational pathway exists to the macula neglecta from the parietalfossa of the dorsal chondrocranium. It is not clear at present,however, whether it is the inner ear or the lateral line systemwhich is responsible for hearing. Both detection systems aretheoretically capable of providing information to the brainabout sound source location using non-parallel arrays of directionallysensitive hair-cell receptors. Recent theories of underwatersound localization by fishes and sharks suggest that the abilityto detect a vertical displacement component of an acoustic signal(e. g., via the macula neglecta) is necessary for instantaneouslocation decisions. It is not known, however, whether the sharkslocalize by processing information about various aspects ofthe sound field simultaneously (in parallel), or whether thesound field is sampled successively at different points in space.Clearly, more experimental work on the physiology of elasmobranchacoustic behavior is called for.     

Tripping Elicits Earlier and Larger Deviations in Linear Head Acceleration Compared to Slipping

Slipping and tripping contribute to a large number of falls and fall-related injuries. While the vestibular system is known to contribute to balance and fall prevention, it is unclear whether it contributes to detecting slip or trip onset. Therefore, the purpose of this study was to investigate the effects of slipping and tripping on head acceleration during walking. This information would help determine whether individuals with vestibular dysfunction are likely to be at a greater risk of falls due to slipping or tripping, and would inform the potential development of assistive devices providing augmented sensory feedback for vestibular dysfunction. Twelve young men were exposed to an unexpected slip or trip. Head acceleration was measured and transformed to an approximate location of the vestibular system. Peak linear acceleration in anterior, posterior, rightward, leftward, superior, and inferior directions were compared between slipping, tripping, and walking. Compared to walking, peak accelerations were up to 4.68 m/s² higher after slipping, and up to 10.64 m/s² higher after tripping. Head acceleration first deviated from walking 100-150ms after slip onset and 0-50ms after trip onset. The temporal characteristics of head acceleration support a possible contribution of the vestibular system to detecting trip onset, but not slip onset. Head acceleration after slipping and tripping also appeared to be sufficiently large to contribute to the balance recovery response.     

Effects of luminance, size, and angular velocity on the recognition of nonlocomotive prey models by the praying mantis

Adult females of the mantis Tenodera angustipennis were presented with the "nonlocomotive" prey model, a static rectangle with two lines oscillating regularly at its sides, generated on a computer display. The models were varied in rectangle luminance (black, gray, and light gray), rectangle height (0.72, 3.6, and 18 mm), rectangle width (0.72, 3.6, and 18 mm), and angular velocity of oscillating lines (65°, 260°, and 1040°/s) to examine their effects on prey recognition. Before striking the model, the mantis sometimes showed peering movements that involved swaying its body from side to side. The black model of medium size (both height and width) elicited higher rates of fixation, peering, and strike responses than the large, small, or gray model. The model of medium angular velocity elicited a higher strike rate than that of large or small angular velocity, but angular velocity had little effect on fixation and peering. We conclude that mantises respond to a rectangle in deciding whether to fixate, and to both rectangle and lines in deciding whether to strike after fixation. Received: September 2, 1999 / Accepted: March 21, 2000     

Kinematic diversity suggests expanded roles for fly halteres

The halteres of flies are mechanosensory organs that provide information about body rotations during flight. We measured haltere movements in a range of fly taxa during free walking and tethered flight. We find a diversity of wing–haltere phase relationships in flight, with higher variability in more ancient families and less in more derived families. Diverse haltere movements were observed during free walking and were correlated with phylogeny. We predicted that haltere removal might decrease behavioural performance in those flies that move them during walking and provide evidence that this is the case. Our comparative approach reveals previously unknown diversity in haltere movements and opens the possibility of multiple functional roles for halteres in different fly behaviours.