Properties of the internal representation of gravity inferred from spatial-direction and body-tilt estimates.
One of the key questions in spatial perception is whether the brain has a common representation of gravity that is generally accessible for various perceptual orientation tasks. To evaluate this idea, we compared the ability of six tilted subjects to indicate earth-centric directions in the dark with a visual and an oculomotor paradigm and to estimate their body tilt relative to gravity. Subjective earth-horizontal and -vertical data were collected, either by adjusting a visual line or by making saccades, at 37 roll-tilt angles across the entire range. These spatial perception responses and the associated body-tilt estimates were subjected to a principal-component analysis to describe their tilt dependence. This analysis allowed us to separate systematic and random errors in performance, to disentangle the effects of task (horizontal vs. vertical) and paradigm (visual vs. oculomotor) in the space-perception data, and to compare the veridicality of space perception and the sense of self-tilt. In all spatial-orientation tests, whether involving space-perception or body-tilt judgments, subjects made considerable systematic errors which mostly betrayed tilt underestimation [Aubert effect (A effect)] and peaked near 130 degrees tilt. However, the A effect was much smaller in body-tilt estimates than in spatial pointing, implying that the underlying signal processing must have been different. Pointing results obtained with the visual and the oculomotor paradigm were not identical either, but these differences, which were task-related (horizontal vs. vertical), were subtle in comparison. The tilt-dependent pattern of random errors (noisy scatter) was almost identical in visual and oculomotor pointing results, showing a steep monotonic increase with tilt angle, but was again clearly different in the body-tilt estimates. These findings are discussed in the context of a conceptual model in an attempt to explain how the different patterns of systematic and random errors in external-space and self-tilt perception may come about. The scheme proposes that basically similar computational mechanisms, working with different settings, may be responsible. (+info)
Postural control in the lamprey: A study with a neuro-mechanical model.
The swimming lamprey normally maintains the dorsal-side-up orientation due to activity of the postural control system driven by vestibular organs. Commands for postural corrections are transmitted from the brain stem to the spinal cord mainly by the reticulospinal (RS) pathways. As shown in previous studies, RS neurons are activated by contralateral roll tilt, they exhibit a strong dynamic response, but much weaker static response. Here we test a hypothesis that decoding of these commands in the spinal cord is based on the subtraction of signals in the left and right RS pathways. In this study, we used a neuro-mechanical model. An intact lamprey was mounted on a platform that restrained its postural activity but allowed lateral locomotor undulations to occur. The activity in the left and right RS pathways was recorded by implanted electrodes. These natural biological signals were then used to control an electrical motor rotating the animal around its longitudinal axis toward the stronger signal. It was found that this "hybrid" system automatically stabilized a normal orientation of the lamprey in the gravitational field. The system compensated for large postural disturbances (lateral tilt up to +/-180 degrees ) due to wide angular zones of the gravitational sensitivity of RS neurons. In the nonswimming lamprey, activity of RS neurons and their vestibular responses were considerably reduced, and the system was not able to stabilize the normal orientation. However, the balance could be restored by imposing small oscillations on the lamprey, which elicited additional activation of the vestibular organs. This finding indicates that head oscillations caused by locomotor movements may contribute to postural stabilization. In addition to postural stabilization, the neuro-mechanical model reproduced a number of postural effects characteristic of the lamprey: 1) unilateral eye illumination elicited a lateral tilt ("dorsal light response") due to a shift of the equilibrium point in the vestibular-driven postural network; 2) removal of one labyrinth resulted in a loss of postural control due to an induced left-right asymmetry in the vestibulo-reticulospinal reflexes, which 3) could be compensated for by asymmetrical visual input. The main conclusion of the present study is that natural supraspinal commands for postural corrections in the roll plane can be effectively decoded on the basis of subtraction of the effects of signals delivered by the left and right RS pathways. Possible mechanisms for this transformation are discussed. (+info)
Context-specific adaptation of the vertical vestibuloocular reflex with regard to gravity.
We determined whether head position with regard to gravity is an important context for angular vestibuloocular reflex (aVOR) gain adaptation. Vertical aVOR gains were adapted with monkeys upright or on side by rotating the animals about an interaural axis in phase or out of phase with the visual surround for 4 h. When aVOR gains were adapted with monkeys upright, gain changes were symmetrical when tested in either on-side position (23 +/- 7%; mean +/- SD). After on-side adaptation, however, gain changes were always larger when animals were tested in the same on-side position in which they were adapted. Gain changes were 43 +/- 16% with ipsilateral side down and 9 +/- 8% with contralateral side down. The context-specific effects of head position on vertical aVOR gain were the same whether the gain was increased or decreased. The data indicate that vertical aVOR gain changes are stored in the context of the head orientation in which changes were induced. This association could be an important context for expressing the adapted state of the aVOR gain during vertical head movement. (+info)
Stereopsis outweighs gravity in the control of the eyes.
The eyes are controlled by multiple brain circuits, some phylogenetically old and some new, whose aims may conflict. Old otolith reflexes counterroll the eyes when the head tilts relative to gravity. Newer vergence mechanisms coordinate the eyes to aid stereoptic vision. We show that counterroll hinders stereopsis, weakly when you look into the distance but strongly when you look near. The resolution of this conflict is that counterroll virtually vanishes when monkeys look close, i.e., stereopsis overrides gravity-driven reflexes but only on near gaze. This balance between gyroscopic and stereoptic mechanisms explains many other puzzling features of primate gaze control, such as the weakness of our otolith-ocular reflexes even during far viewing and the strange geometry of the primate counterpitch reflex, which rolls the eyes clockwise when monkeys look leftward while their heads are tipped nose up, but rolls them counterclockwise when the monkeys look rightward, and reverses this pattern when the head is tipped nose down. (+info)
Changes in root cap pH are required for the gravity response of the Arabidopsis root.
Although the columella cells of the root cap have been identified as the site of gravity perception, the cellular events that mediate gravity signaling remain poorly understood. To determine if cytoplasmic and/or wall pH mediates the initial stages of root gravitropism, we combined a novel cell wall pH sensor (a cellulose binding domain peptide-Oregon green conjugate) and a cytoplasmic pH sensor (plants expressing pH-sensitive green fluorescent protein) to monitor pH dynamics throughout the graviresponding Arabidopsis root. The root cap apoplast acidified from pH 5.5 to 4.5 within 2 min of gravistimulation. Concomitantly, cytoplasmic pH increased in columella cells from 7.2 to 7.6 but was unchanged elsewhere in the root. These changes in cap pH preceded detectable tropic growth or growth-related pH changes in the elongation zone cell wall by 10 min. Altering the gravity-related columella cytoplasmic pH shift with caged protons delayed the gravitropic response. Together, these results suggest that alterations in root cap pH likely are involved in the initial events that mediate root gravity perception or signal transduction. (+info)
Gravitoinertial force magnitude and direction influence head-centric auditory localization.
We measured the influence of gravitoinertial force (GIF) magnitude and direction on head-centric auditory localization to determine whether a true audiogravic illusion exists. In experiment 1, supine subjects adjusted computer-generated dichotic stimuli until they heard a fused sound straight ahead in the midsagittal plane of the head under a variety of GIF conditions generated in a slow-rotation room. The dichotic stimuli were constructed by convolving broadband noise with head-related transfer function pairs that model the acoustic filtering at the listener's ears. These stimuli give rise to the perception of externally localized sounds. When the GIF was increased from 1 to 2 g and rotated 60 degrees rightward relative to the head and body, subjects on average set an acoustic stimulus 7.3 degrees right of their head's median plane to hear it as straight ahead. When the GIF was doubled and rotated 60 degrees leftward, subjects set the sound 6.8 degrees leftward of baseline values to hear it as centered. In experiment 2, increasing the GIF in the median plane of the supine body to 2 g did not influence auditory localization. In experiment 3, tilts up to 75 degrees of the supine body relative to the normal 1 g GIF led to small shifts, 1--2 degrees, of auditory setting toward the up ear to maintain a head-centered sound localization. These results show that head-centric auditory localization is affected by azimuthal rotation and increase in magnitude of the GIF and demonstrate that an audiogravic illusion exists. Sound localization is shifted in the direction opposite GIF rotation by an amount related to the magnitude of the GIF and its angular deviation relative to the median plane. (+info)
Effect of dark pretreatment on the kinetics of response of barley pulvini to gravistimulation and hormones.
Starch in pulvinus amyloplasts of barley (Hordeum vulgare cv Larker) disappears when 45-day-old, light-grown plants are given 5 days of continuous darkness. The effect of this loss on the pulvinus graviresponse was evaluated by following changes in the kinetics of response during the 5-day dark period. Over 5 days of dark pretreatment, the lag to initial graviresponse and the subsequent half-time to maximum steady state bending rate increased significantly while the maximum bending rate did not change. The change in response to applied indoleacetic acid (100 micromolar) plus gibberellic acid (10 micromolar) without gravistimulation, under identical dark pretreatments, was used as a model system for the response component of gravitropism. Dark pretreatment did not change the lag to initial response following hormone application to vertical pulvini, but both the maximum bending rate and the half-time to the maximum rate were significantly reduced. Also, after dark pretreatment, significant bending responses following hormone application were observed in vertical segments with or without added sucrose, while gravistimulation produced a response only if segments were given sucrose. These results indicate that starch-filled amyloplasts are required for the graviresponse of barley pulvini and suggest that they function in the stimulus perception and signal transduction components of gravitropism. (+info)