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L Borel

Publications and source records attributed to L Borel.

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Functional coupling of the stabilizing eye and head reflexes during horizontal and vertical linear motion in the cat.

The otolith contribution and otolith-visual interaction in eye and head stabilization were investigated in alert cats submitted to sinusoidal linear accelerations in three defined directions of space: up-down (Z motion), left-right (Y motion), and forward-back (X motion). Otolith stimulation alone was performed in total darkness with stimulus frequency varying from 0.05 to 1.39 Hz at a constant half peak-to-peak amplitude of 0.145 m (corresponding acceleration range 0.0014-1.13 g). Optokinetic stimuli were provided by sinusoidally moving a pseudorandom visual pattern in the Z and Y directions, using a similar half peak-to-peak amplitude (0.145 m, i.e., 16.1 degrees) in the 0.025-1.39 Hz frequency domain (corresponding velocity range 2.5 degrees-141 degrees/s). Congruent otolith-visual interaction (costimulation, CS) was produced by moving the cat in front of the earth-stationary visual pattern, while conflicting interaction was obtained by suppressing all visual motion cues during linear motion (visual stabilization method, VS, with cat and visual pattern moving together, in phase). Electromyographic (EMG) activity of antagonist neck extensor (splenius capitis) and flexor (longus capitis) muscles as well as horizontal and vertical eye movements (electro-oculography, EOG) were recorded in these different experimental conditions. Results showed that otolith-neck (ONR) and otolith-ocular (OOR) responses were produced during pure otolith stimulation with relatively weak stimuli (0.036 g) in all directions tested. Both EMG and EOG response gain slightly increased, while response phase lead decreased (with respect to stimulus velocity) as stimulus frequency increased in the range 0.25-1.39 Hz. Otolith contribution to compensatory eye and neck responses increased with stimulus frequency, leading to EMG and EOG responses, which oppose the imposed displacement more and more. But the otolith system alone remained unable to produce perfect compensatory responses, even at the highest frequency tested. In contrast, optokinetic stimuli in the Z and Y directions evoked consistent and compensatory eye movement responses (OKR) in a lower frequency range (0.025-0.25 Hz). Increasing stimulus frequency induced strong gain reduction and phase lag. Oculo-neck coupling or eye-head synergy was found during optokinetic stimulation in the Z and Y directions. It was characterized by bilateral activation of neck extensors and flexors during upward and downward eye movements, respectively, and by ipsilateral activation of neck muscles during horizontal eye movements. These visually-induced neck responses seemed related to eye velocity signals. Dynamic properties of neck and eye responses were significantly improved when both inputs were combined (CS).(ABSTRACT TRUNCATED AT 400 WORDS)

Acceleration

[Vestibular studies: current status, prospects].

Equilibrium is based on gaze stabilization. Clinical examinations currently test, individually the three components involved in equilibrium: vestibular, visual and proprioception. New investigations are being developed to test equilibrium dynamic conditions. After briefly reviewing vestibular physiology, under the authors discuss the various tests of vestibular function and equilibrium.

Eye Movements

Functional coupling of the stabilizing gaze reflexes during vertical linear motion in the alert cat.

Eye-head coordination is mainly achieved by means of stabilizing reflexes (VOR, VCR, OKR) and orienting movements (eye-neck surgery) underlying the close cooperation of the visual and vestibular systems in gaze stabilization. The functional coupling of these different sensorimotor subsystems has been principally analysed using rotatory stimulation of the whole body and/or of the visual surround. The aim of the present study was to investigate the dynamic properties of these stabilizing gaze reflexes and their coupling during linear motion in the vertical plane. These investigations were performed in the alert cat under open-loop conditions (head fixed). Otolith stimulation consisted of vertically translating the cat in total darkness using sinusoidal linear motion (0.025 Hz-1.39 Hz; 290 mm peak-to-peak amplitude). Optokinetic stimulation was provided by sinusoidally moving a pseudo-random visual pattern in front of the cat and in the vertical plane, with identical kinematic parameters. Normal visual-otolith interaction was performed by translating the cat in front of the stationary visual surround while conflicting interaction was provided by moving the animal and the visual pattern in phase and at the same velocity (visual stabilization). Results showed that the vertical otolith-neck reflex is very poorly developed or absent in the low frequency range of motion (0.025 Hz-0.25 Hz) while consistent EMG activity is found during pure optokinetic stimulation. EMG responses are in phase with the visual surround velocity in the upward direction and with the upward OKR velocity. A close correlation is observed between the EMG gain and the OKR gain, which both decrease in this low frequency range, indicating that gaze stabilization would be mainly ensured by the OKR and a functional oculo-collic coupling or eye-neck surgery in the vertical plane. On the contrary, gaze stabilization is principally achieved by way of the otolith-neck reflex in the higher frequency range of motion (above 0.25 Hz). EMG responses recorded during otolith stimulation exhibit a relatively constant gain and a phase lead with respect to motion velocity which progressively reduces as the stimulus frequency increases up to 1.39 Hz. When present, EMG responses evoked during the optokinetic stimulation show strong gain attenuation and phase lag. Normal visual-otolith interaction induces neck muscle activity which parallels the optokinetic and the otolith responses in the low and high frequency ranges, respectively. The motor responses are however improved in terms of gain and phase values in the whole frequency range when both sensory inputs are combined.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals

Neuronal coding of linear motion in the vestibular nuclei of the alert cat. II. Response characteristics to vertical optokinetic stimulation.

Extracellular activity from vestibular nuclei neurons and vertical eye movements were recorded in the alert cat during sinusoidal optokinetic stimulation in the vertical plane at frequencies varying from 0.0125 Hz to 0.75 Hz. Among a population of 96 vestibular units located in and around Deiters' nucleus, 73 neurons (76%) displayed a firing rate modulation which followed the input at the standard parameters of visual stimulation (0.05 Hz; 10.1 deg/s or 9.1 cm/s peak to peak velocity). Two different patterns of modulation were found. In 42 cells (57%) an increase in the firing rate was observed during motion of the visual scene in the downward direction, while 31 neurons (43%) showed the opposite behavior, with an enhanced firing rate during upward movement. The phase of the neuronal responses was close (+/- 45 degrees) to the velocity peaks (+90 degrees: downward and -90 degrees: upward) of visual scene motion for 65 among the 73 neurons. Mean values of phase was -6.1 +/- 19.5 degrees (SD) and -3.2 +/- 15.5 degrees (SD) with respect to the +90 degrees and -90 degrees velocity peaks, respectively. In the frequency range 0.0125-0.75 Hz, the phase of the neuronal responses remained almost stable, with only a slight lag which reaches -22 degrees at the 0.25 Hz visual stimulation. The firing rate modulation was found to be predominant at low frequencies (0.0125 Hz-0.25 Hz), with three distinct peaks of modulation occurring either at 0.025 Hz, 0.10 Hz or 0.25 Hz, depending on the recorded cells. Above 0.5 Hz, the cell modulation was very poorly developed or even absent. A gain attenuation was observed in all units, which was more important in cells showing a peak of modulation at 0.025 Hz as compared with the others (-20.7 dB vs -9.6 dB, respectively, in the 0.025 Hz-0.25 Hz decade). The gain of the optokinetic reflex (OKR) progressively decreased from mean values of 0.78 +/- 0.15 to 0.05 +/- 0.06 in the 0.025 Hz-0.5 Hz frequency range. A close correlation was observed between the OKR slow phase velocity and the modulation of the neuronal responses in the two cell populations with maximal modulations at 0.10 Hz or 0.25 Hz. No correlations were noticed in the third population characterized by a peak of modulation at 0.025 Hz. In all units, the phase of eye movement velocity and of neuronal responses were both related to the velocity of the visual surround motion. These correlations were also found when varying the amplitude of the visual stimulation at a fixed frequency.(ABSTRACT TRUNCATED AT 400 WORDS)

Action Potentials

Neuronal coding of linear motion in the vestibular nuclei of the alert cat. III. Dynamic characteristics of visual-otolith interactions.

In the present study we have investigated in the awake cat the response dynamics of vestibular nuclei neurons to visual or/and otolith stimulation elicited by vertical linear motion. Of the 53 units tested during sinusoidal motion at 0.05 Hz (9.1 cm/s), 1 (1.9%) was responsive to the otolith input only, 13 (24.5%) were influenced by the visual input only and 23 (43.4%) responded to both modalities. Neurons were excited either during upward or downward animal or visual surround movement. Most units displayed a firing rate modulation very close to motion velocity. All the neurons receiving convergent visual and otolith inputs (0.05 Hz, 9.1 cm/s) exhibited synergistic patterns of response. Motion velocity coding was improved in terms of input-output phase relationship and response sensitivity when visual and otolith signals were combined. Depending on the units, visual-otolith interactions in single neurons could follow a linear or a nonlinear mode of summation. The dynamic characteristics of visual-otolith interactions were examined in the 0.05 Hz-0.50 Hz frequency bandwidth. Visual signals seemed to predominate over otolith signals at low stimulus frequencies (up to 0.25 Hz), while the contrary was found in the higher frequency range of movement (above 0.25 Hz). The effects of visual stabilization (VS: suppression of visual motion cues) was observed in a small sample of units. As a rule, VS induced a reduction in the amplitude of unit response as compared to visual + otolith stimulation, the lower the motion frequency, the more pronounced the attenuation. VS also decreased the amplitude of the otolith-dependent component of response. The possible modes of visual-vestibular interactions in single cells are discussed. The present study supports the hypothesis that visual and vestibular motion cues are weighted according to their internal relevance.

Action Potentials

Dynamic properties of the vertical otolith neck reflexes in the alert cat.

Electromyographic activity of dorsal neck muscles elicited by sinusoidal vertical linear accelerations was studied in alert cats over a wide range of frequencies. Experiments were performed in head-fixed cats and total darkness in order to activate selectively the otolith system. The polyunitary EMG activity was recorded from splenius capitis muscles in normal and labyrinthectomized cats during vertical translations varying from 0.05-1 Hz with a fixed 290 mm peak-to-peak amplitude. The corresponding accelerations ranged from 0.003-1.2 g. In normal cats, the results showed a bilateral and sinusoidal modulation of the EMG activity characterized by two typical EMG patterns depending on the stimulus frequency. In the low-frequency range (0.05-0.25 Hz), the neck muscles responses were composed of a second harmonic (frequency double that of the input signal: H2 responses). The H2 pattern was characterized by an increase in EMG activity during both the upward and downward parts of translation. These two components of the H2 response were closely related to the two peak velocities (+90 degrees and -90 degrees) of the animal motion. Only slight decreases in amplitude and shifts in phase were observed when increasing the frequency. In the higher frequency range (0.25-1 Hz), the neck muscles response was composed of a fundamental frequency corresponding to the input signal (H1 response). The H1 pattern was in phase with the peak of downward acceleration at 0.25 Hz. A phase lag (up to 45 degrees) and a gain attenuation (16.5 dB) were observed when increasing the frequency. The two H1 and H2 EMG patterns were totally absent in bilateral vestibular neurectomized cats. In unilateral vestibular neurectomized cats, a strong drop in gain and phase advance was noted, which mainly affected the H1 pattern. The present results describe some characteristics of otolith-spinal reflexes acting on the head musculature during vertical motion. They are compared with the neuronal responses that we have recorded within the vestibular nuclei complex in the same experimental conditions. The functional role of the vertical otolith-neck reflexes in stabilizing the head in space during many real-life situations is discussed.

Acceleration

Neuronal coding of linear motion in the vestibular nuclei of the alert cat. I. Response characteristics to vertical otolith stimulation.

The aim of the present study was to investigate some aspects of the central processing of otolith information during linear motion. For this purpose, the response characteristics of 69 vestibular nuclei units to sinusoidal otolith stimulation in the vertical Z axis were analysed in the alert cat. Among this population of neurons which responded to a 0.05 Hz, 290 mm translation, 47 units (70%) displayed a firing rate modulation which followed the input frequency (H1 units). The majority of these neurons exhibited an increase in discharge rate during upward displacement, with a response phase close to the motion velocity or slightly leading downward acceleration. The acceleration related units were divided into two groups according to whether they showed clear increases or only a slight change in discharge rate when the stimulus frequency was increased. The former group was characterized by an average -16.3 dB drop in gain (from 43.9 +/- 1.8 dB, S.D. to 27.6 +/- 7 dB, S.D.) within the 0.05 Hz-0.5 Hz frequency range, while the latter group displayed an average -31.2 dB gain attenuation (from 45.1 +/- 1.1 dB, S.D. to 13.9 +/- 0 dB) within the same decade. In contrast to differences in response gain, all the units tested exhibited a relatively stable phase lead of about 20 degrees with respect to downward peak acceleration. Conversely, units whose response was close to motion velocity in the lower frequency range (0.05 Hz-0.10 Hz) displayed a strong phase lead of about 100 degrees when the stimulus frequency was increased (up to 0.50 Hz). These neurons were thus characterized by an acceleration related response in the higher frequency range. At the same time, an average -24.8 dB gain attenuation (from 47.7 +/- 3.4 dB to 22.9 +/- 3.7 dB) was found in the 0.05 Hz-0.5 Hz decade. The remaining 22 neurons (30%) were called H2 units since they displayed a response waveform double that of the input frequency, a response already described during sinusoidal rotation. Unit discharge reached a peak approximately in phase with maximum upward and downward velocity. Asymmetrical change in unit firing rate about the resting discharge level and different dynamic behavior of the upward and downward response components were usually found. These response characteristics suggest that the H2 patterns are centrally constructed and could result from convergence of otolith afferents having opposite polarization vectors.(ABSTRACT TRUNCATED AT 400 WORDS)

Animals