Influence of gravitoinertial force level on the subjective vertical during recumbent yaw axis body tilt.

We tilted recumbent subjects at various angles about their yaw (foot to head) axis and had them indicate the direction of their subjective vertical and apparent head midline about the same axis. One set of tests was conducted during parabolic flight maneuvers where the background gravitoinertial acceleration varied from 0 to 1.8g. The blindfolded subjects (n = 6) were tested supine and at tilts of 60 degrees and 30 degrees left and right about their horizontal long body axis. They used a gravity neutral joystick to indicate their subjective vertical or their head midline continuously from the high force through the 0g portions of parabolas. In 0g, all subjects felt supine and oriented the joystick perpendicular to their body when indicating the subjective vertical. This points to strong influences of the symmetric somatic touch and pressure cues from the apparatus on orientation when the otolith organs are unloaded. In contrast to the settings in 0g, settings of the subjective vertical in 1g and 1.8g varied as a function of body orientation. However, the settings did not differ between 1g and 1.8g test conditions. Subjective vertical judgments were also made by subjects (n = 11) in the Brandeis slow rotation room, with the room stationary and rotating at a speed that produced a 2g resultant of gravitational and centrifugal acceleration. There were no differences between settings of the subjective vertical made in 1g and 2g. The similarity of 1g and hyper-g settings during recumbent yaw tilts, both in parabolic flight and in the rotating room, contrasts with the previously observed, strong influence which force levels above 1g have on settings of the subjective vertical during tilt of the body in pitch or roll. The findings for all three axes are consistent with a recently developed model of static spatial orientation.

Mechanisms of human static spatial orientation.

We have developed a tri-axial model of spatial orientation applicable to static 1g and non-1g environments. The model attempts to capture the mechanics of otolith organ transduction of static linear forces and the perceptual computations performed on these sensor signals to yield subjective orientation of the vertical direction relative to the head. Our model differs from other treatments that involve computing the gravitoinertial force (GIF) vector in three independent dimensions. The perceptual component of our model embodies the idea that the central nervous system processes utricular and saccular stimuli as if they were produced by a GIF vector equal to 1g, even when it differs in magnitude, because in the course of evolution living creatures have always experienced gravity as a constant. We determine just two independent angles of head orientation relative to the vertical that are GIF dependent, the third angle being derived from the first two and being GIF independent. Somatosensory stimulation is used to resolve our vestibular model's ambiguity of the up-down directions. Our otolith mechanical model takes into account recently established non-linear behavior of the force-displacement relationship of the otoconia, and possible otoconial deflections that are not co-linear with the direction of the input force (cross-talk). The free parameters of our model relate entirely to the mechanical otolith model. They were determined by fitting the integrated mechanical/perceptual model to subjective indications of the vertical obtained during pitch and roll body tilts in 1g and 2g force backgrounds and during recumbent yaw tilts in 1g. The complete data set was fit with very little residual error. A novel prediction of the model is that background force magnitude either lower or higher than 1g will not affect subjective vertical judgments during recumbent yaw tilt. These predictions have been confirmed in recent parabolic flight experiments.

Waveform dynamics of spermatozeugmata during the transfer from paternal to maternal individuals of Membranipora membranacea.

Analysis of standard (60 frames/s) and high-speed (200 frames/s) video records revealed that unencapsulated sperm aggregates (spermatozeugmata) of the gymnolaemate bryozoan Membranipora membranacea spontaneously generate at least three types of waveforms: small amplitude, large amplitude, and reverse. All three waveforms significantly differed from one another in amplitude. Additionally, small- and large-amplitude waveforms propagated from the base to the tip of axonemes, whereas the reverse waveform propagated from the tip to the base of axonemes. Small-amplitude waveforms, which were generated most frequently by spermatozeugmata in the paternal perivisceral coelom and in the water column after spawning, produced almost no curvature of the axoneme. Large-amplitude waveforms were produced by spermatozeugmata in the water column and within lophophores. Reverse waveforms were produced while spermatozeugmata moved tail-end forward through the paternal tentacles during spawning and after spermatozeugmata had contacted the intertentacular organ (ITO), a tubular structure that spermatozeugmata pass through to enter the maternal coelom and that eggs pass through to enter the seawater. The production of reverse waveforms by spermatozeugmata after reaching the ITO may be evidence for a behavioral response of bryozoan sperm to conspecific maternal individuals.

Analysis of human postural responses to recoverable falls.

We studied the kinematics and kinetics of human postural responses to "recoverable falls." To induce brief falling we used a Hold and Release (H&R) paradigm. Standing subjects actively resisted a force applied to their sternum. When this force was quickly released they were suddenly off balance. For a brief period, approximately 125 ms, until restoring forces were generated to shift the center of foot pressure in front of the center of mass, the body was in a forward fall acted on by gravity and ground support forces. We were able to describe the whole-body postural behavior following release using a multilink inverted pendulum model in a regime of "small oscillations." A three-segment model incorporating upper body, upper leg, and lower leg, with active stiffness and damping at the joints was fully adequate to fit the kinematic data from all conditions. The significance of our findings is that in situations involving recoverable falls or loss of balance the earliest responses are likely dependent on actively-tuned, reflexive mechanisms yielding stiffness and damping modulation of the joints. We demonstrate that haptic cues from index fingertip contact with a stationary surface lead to a significantly smaller angular displacement of the torso and a more rapid recovery of balance. Our H&R paradigm and associated model provide a quantifiable approach to studying recovery from potential falling in normal and clinical subjects.

Haptic stabilization of posture: changes in arm proprioception and cutaneous feedback for different arm orientations.

Postural sway during quiet stance is attenuated by actively maintained contact of the index finger with a stationary surface, even if the level of applied force (<1 N) cannot provide mechanical stabilization. In this situation, changes in force level at the fingertip lead changes in center of foot pressure by approximately 250 ms. These and related findings indicate that stimulation of the fingertip combined with proprioceptive information about the hand and arm can serve as an active sensor of body position relative to the point of contact. A geometric analysis of the relationship between hand and torso displacement during body sway led to the prediction that arm and hand proprioceptive and finger somatosensory information about body sway would be maximized with finger contact in the plane of body sway. Therefore, the most postural stabilization should be possible with such contact. To test this analysis, subjects touched a laterally versus anteriorly placed surface while in each of two stances: the heel-to-toe tandem Romberg stance that reduces medial-lateral stability and the heel-to-heel, toes-outward, knees-bent, "duck stance" that reduces fore-aft stability. Postural sway was always least with finger contact in the unstable plane: for the tandem stance, lateral fingertip contact was significantly more effective than frontal contact, and, for the duck stance, frontal contact was more effective than lateral fingertip contact. Force changes at the fingertip led changes in center of pressure of the feet by approximately 250 ms for both fingertip contact locations for both test stances. These results support the geometric analysis, which showed that 1) arm joint angles change by the largest amount when fingertip contact is maintained in the plane of greatest sway, and 2) the somatosensory cues at the fingertip provide both direction and amplitude information about sway when the finger is contacting a surface in the unstable plane.