Minimizing the Negative Flavor Attributes and Evaluating Consumer Acceptance of Chocolate Fortified with Peanut Skin Extracts.

In recent years, there has been increased interest in antioxidant-rich products by consumers wanting to enhance the health benefits of their diet. Chocolate has been identified as a natural source of antioxidant compounds, which resulted in the development of polyphenol-enriched chocolate products that are now available commercially. This study investigated the use of phenolic compounds extracted from peanut skins as a novel antioxidant source for the enrichment of milk chocolate. The extracts were encapsulated with maltodextrin to lessen their bitterness. Antioxidant potential of the encapsulated peanut skin extracts was evaluated by the 2,2-diphenyl-1-picrylhydrazl radical quenching assay. Encapsulated peanut skins were found to have a corrected Trolox equivalency of 31.1 µmol/g of chocolate up to 0.8% (w/w). To produce a product with an antioxidant content similar to that of dark chocolate yet which maintained the milder flavor of milk chocolate, the best estimate threshold of encapsulated peanut skin extract in chocolate was 0.9 % (w/w) based on the standard method (American Society of Testing Materials; ASTM E-679). Consumer liking of milk chocolate enhanced by adding subthreshold (0.8 % (w/w)) inclusion levels of encapsulated peanut skin extract was found to be at parity with milk chocolate as a control.

Static and dynamic properties of vergence-induced reduction of ocular counter-roll in near vision.

We have examined the characteristics of vergence-induced reduction of ocular counter-roll in near vision. Monkeys were trained to make convergent and divergent refixations with the head and body either upright or in various roll orientations. During near viewing requiring 17 degrees horizontal vergence, we found that static binocular torsion was suppressed by about 68% (averaged over both eyes, two monkeys and both near target locations). This result is in accordance with a previous study in which binocular torsion was quantified based on the displacement planes of eye positions in far and near viewing. Latency and duration of the change in torsional eye position depended (for each eye differently) on body roll and the depth plane of fixation. For instance, during convergent refixations in left-ear-down orientations, the latencies of the left eye were smaller and the durations were longer than those of the right eye. However, both eyes reached their final positions required to fixate the second visual target at roughly the same time. The different dynamics of the two eyes is explained by the fact that each eye rotated temporally when the eyes converged, a pattern named binocular extension of Listing's law. Coming from or aiming at a common torsional value (normal ocular counter-roll) in convergent or divergent refixations, the required torsion differs in the two eyes. The brain compensates for these differences by adjusting the dynamics of each eye's movement.

Direction specific error patterns during continuous tracking of the subjective visual vertical.

The aim of this study was to characterize the error pattern of continuously tracking the perceived earth-vertical during roll rotations from upright to right or left ear-down and from right or left ear-down to upright. We compared the tracking responses of two paradigms, which either continuously activated the otoliths organs alone (constant velocity tilt) or both the otolith organs and the semicircular canals (constant acceleration tilt). The tracking responses of the subjective visual vertical showed characteristic differences depending on starting position and tilt direction relative to gravity. The error patterns in the constant-velocity and constant-acceleration tilt paradigm were reversed. Estimations during tracking, when otolith information was continuously changing, were more precise compared to estimations following fast tilts to fixed roll tilt positions. We conclude that the central processing underlying these perceptual tracking responses requires, besides the otolith input, information from the vertical semicircular canals.

Common reference system for estimation of the postural and subjective visual vertical.

When tilted subjects are asked to set a luminous line to the perceived earth-vertical in a dark surrounding, they systematically underestimate the true direction of earth-vertical at large tilt angles, a phenomenon first described by Aubert (A-phenomenon). At small tilt angles, subjects usually overestimate the direction of earth-vertical. Overestimation has been first reported by Müller, who termed the notion of E-phenomenon. Since these first reports, this rather remarkable error behavior has been studied extensively. The prevailing notion in most earlier studies was that the erroneous estimation of verticality results from otolith signals, which are thought to represent the major input for spatial orientation, and their interaction with somatosensory signals. To bring the subjects into tilted positions, most investigators used slow tilt velocities or waited for some time to prevent interaction with semicircular canal activity. Here, we tested the hypothesis that vestibular cues about self-orientation relative to gravity are most reliable when both the semicircular canals and the otolith organs are optimally activated. To compare the error behavior in estimations of the visual vertical and perceived body position, we used self-controlled passive tilts at constant velocity or acceleration.

Reciprocal error behavior in estimated body position and subjective visual vertical.

This study investigated the reciprocal relation between estimation of body tilt and visual vertical by using self-controlled passive body tilts at constant velocity (slow tilts with no semicircular canal activation) or constant acceleration (rapid tilts with canal activation). In both conditions, the visual vertical was overestimated in the luminous line setting paradigm, whereas body tilt was underestimated in the position estimation paradigm. These errors were larger after slow than rapid tilts. During slow tilts, the range of actually reached positions was on average underestimated by about 25% with respect to the desired positions. Interestingly, there were no significant differences in the estimated positions for tilts in the roll and pitch plane. Most remarkably, in the range of +/-45 degrees the resulting means of position and luminous line setting errors of the velocity and acceleration paradigms as a function of the desired roll positions were close to zero. Furthermore, the resulting means of the two paradigms showed a high correlation in the tested range of +/-90 degrees. We conclude that: (a). the otoliths provide the main information for the spatial reference for both the estimation of body positions and the luminous line settings, at least in the range of about +/-45 degrees where the resulting mean errors between the two paradigms are close to zero, and (b). coactivation of semicircular canals improves the estimations.

Combined influence of vergence and eye position on three-dimensional vestibulo-ocular reflex in the monkey.

This study examined two kinematical features of the rotational vestibulo-ocular reflex (VOR) of the monkey in near vision. First, is there an effect of eye position on the axes of eye rotation during yaw, pitch and roll head rotations when the eyes are converged to fixate near targets? Second, do the three-dimensional positions of the left and right eye during yaw and roll head rotations obey the binocular extension of Listing's law (L2), showing eye position planes that rotate temporally by a quarter as far as the angle of horizontal vergence? Animals fixated near visual targets requiring 17 or 8.5 degrees vergence and placed at straight ahead, 20 degrees up, down, left, or right during yaw, pitch, and roll head rotations at 1 Hz. The 17 degrees vergence experiments were performed both with and without a structured visual background, the 8.5 degrees vergence experiments with a visual background only. A 40 degrees horizontal change in eye position never influenced the axis of eye rotation produced by the VOR during pitch head rotation. Eye position did not affect the VOR eye rotation axes, which stayed aligned with the yaw and roll head rotation axes, when torsional gain was high. If torsional gain was low, eccentric eye positions produced yaw and roll VOR eye rotation axes that tilted somewhat in the directions predicted by Listing's law, i.e., with or opposite to gaze during yaw or roll. These findings were seen in both visual conditions and in both vergence experiments. During yaw and roll head rotations with a 40 degrees vertical change in gaze, torsional eye position followed on average the prediction of L2: the left eye showed counterclockwise (ex-) torsion in down gaze and clockwise (in-) torsion in up gaze and vice versa for the right eye. In other words, the left and right eye's position plane rotated temporally by about a quarter of the horizontal vergence angle. Our results indicate that torsional gain is the central mechanism by which the brain adjusts the retinal image stabilizing function of the VOR both in far and near vision and the three dimensional eye positions during yaw and roll head rotations in near vision follow on average the predictions of L2, a kinematic pattern that is maintained by the saccadic/quick phase system.

Geometrical properties of three-dimensional binocular eye position in light sleep.

We examined three-dimensional binocular positions in the alert and sleepy monkeys. In contrast to the tightly yoked eye movements observed in alertness, the eyes were usually converged, vertically misaligned and had a much larger torsional variability during light sleep. While in alertness eye position vectors were confined to fronto-parallel planes, the corresponding planes were rotated temporally (e.g. leftward for the left eye) in light sleep. There was no correlation between temporal rotation of the eye position planes and horizontal vergence. All these observations can be explained by randomly innervated extraocular muscles that are rotating the two eyes about anatomically determined axes.

Direction of heading and vestibular control of binocular eye movements.

To optimize visual fixation on near targets against translational disturbances, the eyes must move in compliance with geometrical constraints that are related to the distance as well as the speed and direction relative to the target. It is often assumed that the oculomotor system uses the vestibular signals during such movements mainly to stabilize the foveal image irrespective of the peripheral vision. To test this hypothesis, trained rhesus monkeys were asked to maintain fixation on isovergence targets at different horizontal eccentricities during 10 Hz oscillations along different horizontal directions. We found that the two eyes moved in compliance with the geometrical constraints of the gaze-stabilization hypothesis, although response gains were generally small ( approximately 0.5). The best agreement with the gaze stabilization hypothesis occurred for heading directions within +/-30 degrees from straight-ahead, whereas lateral movements exhibited greater variability and larger directional errors that reflected the statistical response variability inherent in the non-linear dependence on heading direction. In contrast to undercompensatory version (conjugate) components, the disjunctive part of the response (vergence) exhibited unity or higher than unity gains. The high vergence gains might reflect a strategy that aims at maintaining the binocular coordination of the gaze lines despite the low gain of the version movements.

Vestibular signals in self-orientation and eye movement control.

The central vestibular system receives afferent information about head position as well as rotation and translation. This information is used to prevent blurring of the retinal image but also to control self-orientation and motion in space. Vestibular signal processing in the brain stem appears to be linked to an internal model of head motion in space.

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.

Effect of light sleep on three-dimensional eye position in static roll and pitch.

We examined three-dimensional eye positions in alertness and light sleep when monkeys were placed in different roll and pitch body orientations. In alertness, eye positions were confined to a fronto-parallel (Listing's) plane, torsional variability was small and static roll or pitch induced a torsional shift or vertical rotation of these planes. In light sleep, the planes rotated temporally by about 10 degrees, torsional variability increased by a factor of two and the static otolith-ocular reflexes were reduced by about 70%. These data support the importance of a neural control of the thickness and orientation of Listing's plane, and suggest that part of the vestibular input underlying otolith-ocular reflexes depend on polysynaptic neural processing.

Central versus peripheral origin of vestibuloocular reflex recovery following semicircular canal plugging in rhesus monkeys.

We have previously shown that there is a slowly progressing, frequency-specific recovery of the gain and phase of the horizontal vestibuloocular reflex (VOR) in rhesus monkeys following plugging of the lateral semicircular canals. The adapted VOR response exhibited both dynamic and spatial characteristics that were distinctly different from responses in intact animals. To discriminate between adaptation or recovery of central versus peripheral origin, we have tested the recovered vestibuloocular responses in three rhesus monkeys in which either one or both coplanar pairs of vertical semicircular canals had been plugged previously by occluding the remaining semicircular canals in a second plugging operation. We measured the spatial tuning of the VOR in two or three different mutually orthogonal planes in response to sinusoidal oscillations (1.1 Hz, +/-5 degrees, +/-35 degrees /s) over a period of 2-3 and 12-14 mo after each operation. Apart from a significant recovery of the torsional/vertical VOR following the first operation we found that these recovered responses were preserved following the second operation, whereas the responses from the newly operated semicircular canals disappeared acutely as expected. In the follow-up period of up to 3 mo after the second operation, responses from the last operated canals showed recovery in two of three animals, whereas the previously recovered responses persisted. The results suggest that VOR recovery following plugging may depend on a regained residual sensitivity of the plugged semicircular canals to angular head acceleration.

Three-dimensional organization of vestibular related eye movements to rotational motion in pigeons.

During rotational motions, compensatory eye movement adjustments must continually occur in order to maintain objects of visual interest as stable images on the retina. In the present study, the three-dimensional organization of the vestibulo-ocular reflex in pigeons was quantitatively examined. Rotations about different head axes produced horizontal, vertical, and torsional eye movements, whose component magnitude was dependent upon the cosine of the stimulus axis relative to the animal's visual axis. Thus, the three-dimensional organization of the VOR in pigeons appears to be compensatory for any direction of head rotation. Frequency responses of the horizontal, vertical, and torsional slow phase components exhibited high pass filter properties with dominant time constants of approximately 3 s.

Low-frequency otolith and semicircular canal interactions after canal inactivation.

During sustained constant velocity and low-frequency off-vertical axis rotations (OVAR), otolith signals contribute significantly to slow-phase eye velocity. The adaptive plasticity of these responses was investigated here after semicircular canal plugging. Inactivation of semicircular canals results in a highly compromised and deficient vestibulo-ocular reflex (VOR). Based on the VOR enhancement hypothesis, one could expect an adaptive increase of otolith-borne angular velocity signals due to combined otolith/canal inputs after inactivation of the semicircular canals. Contrary to expectations, however, the steady-state slow-phase velocity during constant velocity OVAR decreased in amplitude over time. A similar progressive decrease in VOR gain was also observed during low-frequency off-vertical axis oscillations. This response deterioration was present in animals with either lateral or vertical semicircular canals inactivated and was limited to the plane(s) of the plugged canals. The results are consistent with the idea that the low-frequency otolith signals do not simply enhance VOR responses. Rather, the nervous system appears to correlate vestibular sensory information from the otoliths and the semicircular canals to generate an integral response to head motion.

Three-dimensional vestibuloocular reflex of the monkey: optimal retinal image stabilization versus listing's law.

If the rotational vestibuloocular reflex (VOR) were to achieve optimal retinal image stabilization during head rotations in three-dimensional space, it must turn the eye around the same axis as the head, with equal velocity but in the opposite direction. This optimal VOR strategy implies that the position of the eye in the orbit must not affect the VOR. However, if the VOR were to follow Listing's law, then the slow-phase eye rotation axis should tilt as a function of current eye position. We trained animals to fixate visual targets placed straight ahead or 20 degrees up, down, left or right while being oscillated in yaw, pitch, and roll at 0.5-4 Hz, either with or without a full-field visual background. Our main result was that the visually assisted VOR of normal monkeys invariantly rotated the eye around the same axis as the head during yaw, pitch, and roll (optimal VOR). In the absence of a visual background, eccentric eye positions evoked small axis tilts of slow phases in normal animals. Under the same visual condition, a prominent effect of eye position was found during roll but not during pitch or yaw in animals with low torsional and vertical gains following plugging of the vertical semicircular canals. This result was in accordance with a model incorporating a specific compromise between an optimal VOR and a VOR that perfectly obeys Listing's law. We conclude that the visually assisted VOR of the normal monkey optimally stabilizes foveal as well as peripheral retinal images. The finding of optimal VOR performance challenges a dominant role of plant mechanics and supports the notion of noncommutative operations in the oculomotor control system.

Canal-otolith interactions after off-vertical axis rotations I. Spatial reorientation of horizontal vestibuloocular reflex.

We examined the three-dimensional (3-D) spatial orientation of postrotatory eye velocity after horizontal off-vertical axis rotations by varying the final body orientation with respect to gravity. Three rhesus monkeys were oriented in one of two positions before the onset of rotation: pitched 24 degrees nose-up or 90 degrees nose-up (supine) relative to the earth-horizontal plane and rotated at +/-60 degrees /s around the body-longitudinal axis. After 10 turns, the animals were stopped in 1 of 12 final positions separated by 30 degrees. An empirical analysis of the postrotatory responses showed that the resultant response plane remained space-invariant, i.e., accurately represented the actual head tilt plane at rotation stop. The alignment of the response vector with the spatial vertical was less complete. A complementary analysis, based on a 3-D model that implemented the spatial transformation and dynamic interaction of otolith and lateral semicircular canal signals, confirmed the empirical description of the spatial response. In addition, it allowed an estimation of the low-pass filter time constants in central otolith and semicircular canal pathways as well as the weighting ratio between direct and inertially transformed canal signals in the output. Our results support the hypothesis that the central vestibular system represents head velocity in gravity-centered coordinates by sensory integration of otolith and semicircular canal signals.

Primate translational vestibuloocular reflexes. I. High-frequency dynamics and three-dimensional properties during lateral motion.

The dynamics and three-dimensional (3-D) properties of the primate translational vestibuloocular reflex (trVOR) for high-frequency (4-12 Hz, +/-0.3-0.4 g) lateral motion were investigated during near-target viewing at center and eccentric targets. Horizontal response gains increased with frequency and depended on target eccentricity. The larger the horizontal and vertical target eccentricity, the steeper the dependence of horizontal response gain on frequency. In addition to horizontal eye movements, robust torsional response components also were present at all frequencies. During center-target fixation, torsional response phase was opposite (anticompensatory) to that expected for an "apparent" tilt response. Instead torsional response components depended systematically on vertical-target eccentricity, increasing in amplitude when looking down and reversing phase when looking up. As a result the trVOR eye velocity vector systematically tilted away from a purely horizontal direction, through an angle that increased with vertical eccentricity with a slope of approximately 0.7. This systematic dependence of torsional eye velocity tilt on vertical eye position suggests that the trVOR might follow the 3-D kinematic requirements that have been shown to govern visually guided eye movements and near-target fixation.

Three-dimensional extraocular motoneuron innervation in the rhesus monkey. I: Muscle rotation axes and on-directions during fixation.

The rotation axis for each of the six extraocular muscles was determined in four eyes from three perfused rhesus monkeys. Measurements of the locations of muscle insertions and origins were made in the stereotaxic reference frame with the x-y plane horizontal and the x-z plane sagittal. The computed rotation axes of the horizontal recti were close to being in the x-z plane at an angle of about 15 degrees to the z axis. The rotation axes of the vertical recti and the obliques were close to being in the x-y plane at an angle of about 30 degrees to the y axis. In five alert rhesus monkeys, we simultaneously recorded extraocular motoneuron activity and eye position in three dimensions (3D). The activity of 51 motoneuron axons was obtained from the oculomotor (n=34), trochlear (n=11), and abducens nerve (n=6) during spontaneous eye movements. To extend the torsional range of eye position, the animals were also put in different static roll positions, which induced ocular counterroll without dynamic vestibular stimulation. Periods of 100 ms during fixation or slow eye movements (<10 degrees/s) were chosen for analysis. For each motoneuron, a multiple linear regression was performed between firing frequency and 3D eye position, expressed as a rotation vector, in both stereotaxic and Listing's reference frame. The direction with the highest correlation coefficient (average R=0.94+/-0.07 SD) was taken as the on-direction. Each unit's activity could be unequivocally attributed to one particular muscle. On-directions for each motoneuron were confined to a well-defined cone in 3D. Average on-directions of motoneurons differed significantly from the corresponding anatomically determined muscle rotation axes expressed in the stereotaxic reference frame (range of deviations: 11.9 degrees to 29.0 degrees). This difference was most pronounced for the vertical recti and oblique muscles. The muscle rotation axes of the vertical rectus pair and the oblique muscle pair form an angle of 58.3 degrees, whereas the corresponding angle for paired motoneuron on-directions was 105.6 degrees. On-directions of motoneurons were better aligned with the on-directions of semicircular canal afferents (range of deviation: 9.4-18.9 degrees) or with the anatomically determined sensitivity vectors of the semicircular canals (range of deviation: 3.9-15.9 degrees) than with the anatomically determined muscle rotation axes, but significant differences remain to be explained. The on-directions of motoneurons were arranged symmetrically to Listing's plane, in the sense that the torsional components for antagonistically paired muscles were almost equal, but of opposite sign. Thus, the torsional components of motoneuron on-directions cancel when eye movements are confined to Listing's plane. This arrangement simplifies the neuronal transformations for conjugate head-fixed voluntary eye movements, while the approximate alignment with the semicircular canal reference frame is optimal for generating compensatory eye movements.

Inertial processing of vestibulo-ocular signals.

New evidence for a central resolution of gravito-inertial signals has been recently obtained by analyzing the properties of the vestibulo-ocular reflex (VOR) in response to combined lateral translations and roll tilts of the head. It is found that the VOR generates robust compensatory horizontal eye movements independent of whether or not the interaural translatory acceleration component is canceled out by a gravitational acceleration component due to simultaneous roll-tilt. This response property of the VOR depends on functional semicircular canals, suggesting that the brain uses both otolith and semicircular canal signals to estimate head motion relative to inertial space. Vestibular information about dynamic head attitude relative to gravity is the basis for computing head (and body) angular velocity relative to inertial space. Available evidence suggests that the inertial vestibular system controls both head attitude and velocity with respect to a gravity-centered reference frame. The basic computational principles underlying the inertial processing of otolith and semicircular canal afferent signals are outlined.

Oculomotor control of primary eye position discriminates between translation and tilt.

We have previously shown that fast phase axis orientation and primary eye position in rhesus monkeys are dynamically controlled by otolith signals during head rotations that involve a reorientation of the head relative to gravity. Because of the inherent ambiguity associated with primary otolith afferent coding of linear accelerations during head translation and tilts, a similar organization might also underlie the vestibulo-ocular reflex (VOR) during translation. The ability of the oculomotor system to correctly distinguish translational accelerations from gravity in the dynamic control of primary eye position has been investigated here by comparing the eye movements elicited by sinusoidal lateral and fore-aft oscillations (0.5 Hz +/- 40 cm, equivalent to +/- 0.4 g) with those during yaw rotations (180 degrees/s) about a vertically tilted axis (23.6 degrees). We found a significant modulation of primary eye position as a function of linear acceleration (gravity) during rotation but not during lateral and fore-aft translation. This modulation was enhanced during the initial phase of rotation when there was concomitant semicircular canal input. These findings suggest that control of primary eye position and fast phase axis orientation in the VOR are based on central vestibular mechanisms that discriminate between gravity and translational head acceleration.