Synaptic linkage in the vestibulo-ocular reflex pathway of rabbit

Summary Microelectrodes were inserted into IIIrd cranial nucleus of anaesthetized rabbit. IIIrd nucleus was identified by observing the field potentials evoked antidromically by stimulation of IIIrd cranial nerve. After stimulation of VIIIth nerve extracellular field potentials, spike potentials in secondary vestibular fibers, and postsynaptic potentials in IIIrd nucleus neurones were recorded. VIIIth nerve impulses either excite or inhibit IIIrd nucleus neurones postsynaptically with disynaptic latencies around 1.7 msec. By local stimulation of the medulla, it was found that the secondary vestibular impulses inhibiting IIIrd nucleus neurones are mediated by the superior nucleus. The excitatory impulses are relayed by the rostral half of the medial nucleus as well as a certain structure(s) relevant to the brachium conjunctivum. Preliminary pharmacological investigations on the inhibition of IIIrd nucleus neurones are reported.     

The carotid sinus baroreceptor reflex in the pregnant rabbit

1. The reflex responses to baroreceptor stimulation have been compared in eight pregnant and eight non-pregnant anaesthetized female rabbits.2. The vascularly isolated, innervated carotid sinus was exposed for 30 sec to a series of non-pulsatile pressures ranging from 30 to 230 mmHg. The contralateral sinus nerve and both aortic nerves were cut. Systemic arterial pressure and heart rate were measured at each sinus pressure.3. The range of arterial pressure change which could be evoked from the isolated innervated sinus was less in the pregnant than in the non-pregnant rabbits. Mean changes were 98 and 61 mmHg respectively. On the other hand changes in heart rate were similar in the two groups (45 and 43 beats/min respectively).4. The smaller blood pressure response in the pregnant animals resulted from a lesser rise in systemic arterial pressure at low levels of sinus pressure. At high sinus pressures the blood pressure fell to a similar level in both groups of animals.5. Pressure on the great vessels by the gravid uterus was not a factor since there was no consistent difference between the responses obtained with the rabbit lying on its back or on its side.6. Mechanisms which might be responsible for the difference found are discussed.     

Carotid sinus reflex function in the alloxan diabetic rabbit.

Baroreflex function was studied in three groups of adult rabbits. Seven animals were given alloxan (100-200 mg/kg) and became diabetic (group D) with mean blood sugar values of 348 +/- 30 mg/dl. Eight animals were given alloxan, but did not develop significant hyperglycemia (135 mg/dl) (group A). Nine controls (group C) were also studied (glucose, 101 mg/dl). All animals were anesthetized with pentobarbital (30 mg/kg). Blood pressure (BP) and heart rate (HR) responses to bilateral carotid occlusion (BCO) were measured before and after depressor nerve sectioning (DNx) and sinus nerve sectioning (SNx). Before sectioning, BCO caused a rise in BP of 30 +/- 4 mmHg in group C. 35 +/- 3 in group A, and 36 +/- 4 in group D. HR increased about 13 beats/min in each group. After DNx, resting BP increased in group C from 97 to 104 mmHg (P less than 0.005), but no change occurred in the other groups. Responses to BCO were significantly but similarly enhanced in all groups after DNx. HR did not increase in group D. Resting BP increased after SNx only in the controls (group C). Differences in BP elevation with BCO before and after SNx ("pure" reflex response) were identical, averaging about 35 mmHg. Thus, no alteration of BP or HR responses to BCO was identified in early alloxan diabetes. However, resting tone in the buffer nerves may have been less.     

The reflex bradycardia during brain ischemia in the rabbit.

We examined the responses of whole aortic nerve activity, aortic baroreceptor activity, heart rate (HR), and arterial pressure (AP) to brain ischemia sustained for approximately 30 s in anesthetized spontaneously breathing rabbits. The minimum values of HR observed during brain ischemia were 76 +/- 11 beats/min (mean +/- S.E., n = 14) before sectioning the bilateral aortic nerve (BAN), and 161 +/- 12 beats/min after sectioning the left aortic nerve (LAN), and 225 +/- 11 beats/min after sectioning the LAN and right aortic nerve (RAN). Averages for reflex fall in HR during BAN, LAN, and RAN activation were 140 +/- 9, 78 +/- 7, and 62 +/- 7 beats/min, respectively, by subtracting the HR fall responses to brain ischemia in the absence of aortic baroreceptor afferents from their control values. The heights of the integrated whole left and right aortic nerve activities in systole slightly increased during brain ischemia, whereas the brain ischemia remarkably increased those activities during the diastolic phases. The brain ischemia induced a hysteresis in the mean AP-aortic barorecept or activity relationship. These results suggest that the total activity of aortic nerve fibers would determine the bradycardia evoked by brain ischemia and that the difference between the relative contributions of LAN and RAN on the brain ischemia-induced reflex bradycardia would reflect the total impulse frequency of aortic myelinated and non-myelinated fibers.     

Contribution of the maculo-ocular reflex to gaze stability in the rabbit

Summary The contribution of the maculo-ocular reflex to gaze stability was studied in 10 pigmented rabbits by rolling the animals at various angles of sagittal inclination of the rotation and/or longitudinal animal axes. At low frequencies (0.005–0.01 Hz) of sinusoidal stimulation the vestibulo-ocular reflex (VOR) was due to macular activation, while at intermediate and high frequencies it was mainly due to ampullar activation. The following results were obtained: 1) maculo-ocular reflex gain decreased as a function of the cosine of the angle between the rotation axis and the earth's horizontal plane. No change in gain was observed when longitudinal animal axis alone was inclined. 2) At 0° of rotation axis and with the animal's longitudinal axis inclination also set at 0°, the maculo-ocular reflex was oriented about 20° forward and upward with respect to the earth's vertical axis. This orientation remained constant with sagittal inclinations of the rotation and/or longitudinal animal axes ranging from approximately 5° upward to 30° downward. When the longitudinal animal axis was inclined beyond these limits, the eye trajectory tended to follow the axis inclination. In the upside down position, the maculo-ocular reflex was anticompensatory, oblique and fixed with respect to orbital coordinates. 3) Ampullo-ocular reflex gain did not change with inclinations of the rotation and/or longitudinal animal axes. The ocular responses were consistently oriented to the stimulus plane. At intermediate frequencies the eye movement trajectory was elliptic because of directional differences between the ampullo- and maculo-ocular reflexes. 4) In the upright position the coactivation of the optokinetic reflex (OKR) eliminated the eye disalignment with respect to the stimulus plane and the elliptic trajectory. 5) Combined vertical OKR and VOR gain in the prone position (VOKR + VVOR 0°) was higher than that of the combined VOKR + VVOR in the 90° nose up position. The VVOR + VOKR 90° gain was in turn higher than the VVOR + VOKR gain in the 180° upside down position. 6) We suggest that, in the dark, the maculo-ocular response tends to reduce the disalignment of both eyes with respect to the horizon rather than inducing oculocompensatory responses. In the light, this maculo-ocular reflex increases the gain of combined optokinetic and vestibular responses.     

The organization of the vestibulo-oculomotor and trochlear reflex pathways in the rabbit

Summary Intracellular and extracellular responses were recorded with glass micro-electrodes from motoneurons in the IIIrd and IVth cranial nuclei of anesthesized rabbits. Five subgroups of neurons innervating the superior rectus (SR), inferior oblique (IO), inferior rectus (IR), medial rectus (MR), and superior oblique (IVth) extraocular muscles were identified by their antidromic activation from the branches of the IIIrd and IVth cranial nerves. The relative positions of the subgroups thus determined were consistent with the histological data on the rabbit. In the SR, IO, IR, and IVth subgroups the effects of ipsilateral VIIIth nerve stimulation were inhibitory, producing disynaptic IPSPs, while the effects of contralateral VIIIth nerve stimulation were excitatory, producing disynaptic EPSPs. In the MR subgroup, however, a mixture of EPSPs and IPSPs was produced by VIIIth nerve stimulation: this was particularly clear on the ipsilateral side. Sites relaying these VIIIth nerve effects to each of the five subgroups were explored by direct stimulation of various brain stem sites. Stimulation of the superior vestibular nucleus (SV) produced IPSPs monosynaptically in all five subgroups on the ipsilateral side as well as in the contralateral MR subgroup. Stimulation of the medial vestibular nucleus (MV) produced EPSPs monosynaptically in all of the five subgroups on the contralateral side as well as in the ipsilateral MR subgroup. Stimulation of the brachium conjunctivum (BC) also produced EPSPs monosynaptically in the contralateral SR, IO, and IR subgroups. Further, while the recording electrode was placed within each of the five subgroups to observe the extracellular potentials corresponding to the intracellularly recorded IPSPs and EPSPs, the medulla and cerebellum were systematically tracked with a monopolar stimulating electrode. It was thus confirmed that the SV is the sole inhibitory relay site, while excitation is relayed by both the MV and the BC. The origin of the BC pathway was traced to the Y-Group for the IO, to the lateral nucleus of the cerebellum (LN) for the IR, and to both the Y-Group and the LN for the SR subgroup.     

Effects of brainstem lesions on the nasopharyngeal reflex in the conscious rabbit

The cardiovascular responses evoked through the nasopharyngeal reflex by the inhalation of formaldehyde vapour were studied in conscious rabbits after bilateral electrolytic lesions of the ventrolateral medulla coinciding with the A1 group of catecholamine cells. Arterial blood pressure was measured in the central ear artery, heart rate was determined from the arterial pressure trace, and iliac blood flow was determined using a Doppler ultrasonic flow meter placed around the lower abdominal aorta. There were no significant changes in the heart rate, blood pressure and iliac conductance responses elicited through the nasopharyngeal reflexes of sham operated animals in which electrodes were inserted without the passage of current. The blood pressure changes produced by inhalation of formaldehyde in control rabbits were not significantly altered in animals with ventrolateral medullary lesions. However, the magnitude of the bradycardia and of the fall in iliac conductance evoked by the inhalation of formaldehyde were approximately halved 4 h and 1 day after the lesions but were fully restored at 2 weeks. These experiments suggest that the A1 group of catecholamine neurones help mediate the bradycardia and vasoconstriction elicited through nasopharyngeal reflexes in terrestial animals exposed to noxious vapours.     

Cerebellar inhibitory control of the vestibulo-ocular reflex investigated in rabbit IIIrd nucleus

Summary In anaesthetized rabbits, the vestibulo-ocular reflex was evoked by electric stimulation of VIIIth nerve and was observed by recording postsynaptic potentials and relevant field potentials in Illrd nucleus. The electric stimulation of flocculus produced a prominent inhibition of the vestibulo-ocular reflex in both the inhibitory component relayed by the superior vestibular nucleus and the excitatory component mediated by the brachium conjunctivum. The excitatory component mediated by the medial vestibular nucleus appeared to be free of the flocculus inhibition. The flocculus inhibition was blocked very effectively by systemic injection of picrotoxin. That the flocculus inhibitory action is due to monosynaptic postsynaptic inhibition of secondary vestibular neurones was demonstrated by direct stimulation of, and also by recording from, the superior nucleus. Recording from the superior nucleus was also performed in anaesthetized cats. All of these above results indicate that Purkinje cells in flocculus projecting to vestibular and cerebellar nuclei cells have inhibitory synaptic action. Flocculus stimulation produced also an excitatory effect upon vestibular nuclei neurones. However, this effect could be attributed to intracerebellar activation of the primary vestibular fibers which pass into the flocculus.     

Kinetic and frequency-domain properties of reflex and conditioned eyelid responses in the rabbit

Eyelid position and the electromyographic activity of the orbicularis oculi muscle were recorded unilaterally in rabbits during reflex and conditioned blinks. Air-puff-evoked blinks consisted of a fast downward phase followed sometimes by successive downward sags. The reopening phase had a much longer duration and slower peak velocity. Onset latency, maximum amplitude, peak velocity, and rise time of reflex blinks depended on the intensity and duration of the air puff-evoking stimulus. A flashlight focused on the eye also evoked reflex blinks, but not flashes of light, or tones. Both delayed and trace classical conditioning paradigms were used. For delayed conditioning, animals were presented with a 350-ms, 90-dB, 600-Hz tone, as conditioned stimulus (CS). For trace conditioning, animals were presented with a 10-ms, 1-k/cm(2) air puff, as CS. The unconditioned stimulus (US) consisted of a 100-ms, 3-k/cm(2) air puff. The stimulus interval between CS and US onsets was 250 ms. Conditioned responses (CRs) to tones were composed of downward sags that increased in number through the successive conditioning sessions. The onset latency of the CR decreased across conditioning at the same time as its maximum amplitude and its peak velocity increased, but the time-to-peak of the CR remained unaltered. The topography of CRs evoked by short, weak air puffs as the CS showed three different components: the alpha response to the CS, the CR, and the reflex response to the US. Through conditioning, CRs showed a decrease in onset latency, and an increase in maximum amplitude and peak velocity. The time-to-peak of the CR remained unchanged. A power spectrum analysis of reflex and conditioned blink acceleration profiles showed a significant approximately 8-Hz oscillation within a broadband of frequencies between 4 and 15 Hz. Nose and mandible movements presented power spectrum profiles different from those characterizing reflex and conditioned blinks. It is concluded that eyelid reflex responses in the rabbit present significant differences from CRs in their profiles and metric properties, suggesting different neural origins, but that a common approximately 8-Hz neural oscillator underlies lid motor performance. According to available data, the frequency of this putative oscillator seems to be related to the species size.     

Dark threshold and steady-state reactions of the pupillary light reflex in the pigmented rabbit

Summary The light reflex of the pupil was used as an indicator of retinal activity in the awake, unanesthetized pigmented rabbit.The change of the area of the pupil in response to light flashes subtending a circular field of 22° and of 0.1 sec duration was measured continuously by reflecting infrared light from the iris to a photocell (dynamic response). The photopupillar contraction showed latencies between 0.136 sec (at 700 cd/m²) and 0.290 sec (at 0.12 cd/m²) and was maximal within 0.7 sec. The least stimulus luminance producing a just perceptible contraction of the iris muscle in the most sensitive animal was 4×10^–2 cd/m² which is nearly 10⁴ times the intensity threshold in the human eye. It is concluded that some interaction takes place in the rod-dominated rabbit's retina which results in a masking effect of the rod-mediated pupillomotor response to flashes. Evidence for this effect is the large change of threshold of the rabbit's pupillomotor response during illumination to steady lights of threshold luminance.The actual size of the pupil was measured photographically on infrared film first in the dark, then at increasingly higher levels of illumination. With illumination of white light the size of apparent pupillary diameter decreases from 6,0 mm (after dark adaptation) to 4,5 mm (during illumination with 500 cd/m²). Evidence is given that the steady state reactions of the rabbit's pupil are governed by the activity of the photopic system.     

An analysis of adrenergic influences on the sural-gastrocnemius reflex of the decerebrated rabbit

Summary The sural-gastrocnemius reflex was observed in decerebrated rabbits during intrathecal application of four -adrenoceptor antagonists. Idazoxan and yohimbine, which are antagonists at the ₂-receptor, caused facilitation of the reflex, although idazoxan was more potent and produced a larger overall increase in the reflex response. However, when given after yohimbine, idazoxan elicited no further increase in reflex responses. The differences between the two drugs may result from the interaction of yohimbine with receptors for 5-hydroxytryptamine. The selective ₁-receptor antagonist prazosin had no consistent effects when given alone, but reduced the facilitatory effects of idazoxan. The putative selective post-junctional ₂-receptor blocker SK&F 104078 had no significant effects when given alone, nor did it influence the facilitatory action of a subsequent dose of idazoxan. Section of the spinal cord in the presence of idazoxan always caused a decrease in gastrocnemius responses to sural nerve stimulation. These data show that the facilitatory effects of idazoxan are almost certainly mediated at the spinal cord and that they do not involve blockade of ₁-receptors. It appears that idazoxan acts by blockade of adrenergic descending inhibition in combination with increased descending facilitation. The inhibition is probably mediated through noradrenaline acting at ₂-receptors, and the facilitation may be the result of release of noradrenaline (acting at ₁-receptors) and 5-hydroxytryptamme in the spinal cord.     

Contributions of the amygdaloid central nucleus to the modulation of the nictitating membrane reflex in the rabbit

The contributions of the amygdaloid central nucleus (ACe) to the modulation of the amplitude of the nictitating membrane reflex (NMR) were determined. Experiment 1 demonstrated that low-level electrical stimulation of the ACe enhances the amplitude of the NMR when administered immediately preceding the elicitation of the reflex by an eyelid stimulus. In Experiment 2 the anterograde transport of wheat germ agglutinin conjugated to horseradish peroxidase determined that the ACe projects to the entire rostrocaudal extent of the lateral tegmental field (LTF), the brainstem area in which the multisynaptic component of the unconditioned NMR pathway is believed to be located. These results are consistent with the hypothesis that the ACe, via its projections to the LTF, modulates reflex sensitivity during conditioned arousal and may contribute to the associative enhancement of the unconditioned NMR that occurs early during Pavlovian nictitating membrane conditioning.     

Loss of vision-guided adaptation of the vestibulo-ocular reflex after depletion of brain serotonin in the rabbit

Brain serotonin in pigmented rabbits was depleted by intraventricular injection of 5,7-dihydroxytryptamine (5,7-DHT). Before depletion, an adaptive increase in the gain of the horizontal vestibulo-ocular reflex (HVOR) could be induced regularly by continuously rotating the animals in combination with optokinetic stimulus. After the depletion, such an increase in HVOR gain did not occur, even though dynamic characteristics of the HVOR and of the optokinetic eye movement were not altered. 5,7-DHT treatment also reduced brain norepinephrine, but depletion of norepinephrine to a similar extent by intraventricular injection of 6-hydroxydopamine in other rabbits did not affect the HVOR adaptation. These results suggest that brain serotonin plays an important role in maintaining adaptive modifiability of the HVOR.     

心-肾反射在调节兔肾水钠排出功能中的作用

目的： 研究不同浓度盐溶液所致的心房及血容量扩张通过心-肾反射对肾排钠利尿功能的影响，探讨心-肾反射在肾脏功能调节中的作用。方法： 取健康家兔分为生理盐水和高渗盐溶液组。去窦弓神经，左肾神经在近肾门处切断。导管经右颈外静脉插至右心房，以10 mL/min输入15%血容量的生理盐水和1.8%氯化钠溶液前后，观察中心静脉压（CVP）、左肾交感神经传出活动（ERSNA）、左、右肾尿量和排钠系数的变化。结果： 输液后高渗盐和生理盐水组 CVP分别升高75.00%±27.40%和64.00%±15.56%；ERSNA频率分别减慢63.00%±12.49%和44.00%±13.64%，平均群集时间分别缩短37.00%±16.49%和31.00%±10.69%，平均群集间期分别延长68.00%±27.04%和60.00%±18.38%；左肾尿量分别增加640.00%±155.39%和158.00%±28.10%，排钠系数分别增加376.00%±121.72%和132.00%±35.23%；右肾尿量分别增加 1 343.00%±429.95%和192.00%±32.26%，排钠系数分别增加856.00%±261.48%和300.00%±76.99%。结论： 在去窦弓神经家兔，不同浓度溶液所致的心房及血容量扩张可刺激心肺感受器，通过心-肾反射活动抑制ERSNA，使肾的排钠和排水增多，进而调整和维持机体血容量的相对稳定。     

Conditioning-specific reflex modification of rabbit (Oryctolagus cuniculus) heart rate

Conditioning-specific reflex modification (CRM) describes changes in rabbit (Oryctolagus cuniculus) nictitating membrane responses (NMR) to an unconditioned stimulus (US) when the US is tested by itself after pairings of tone and electrodermal stimulation. Although CRM has been replicated, it is unclear whether it occurs in response systems other than that of the NMR. The authors report that CRM of rabbit heart rate (HR) can occur following HR conditioning. A US that elicits HR acceleration before conditioning can elicit HR deceleration after conditioning. The rabbits' electrocardiograms showed both HR conditioning and HR CRM were correlated with an increased PQ interval--an index of parasympathetic function mediated by the vagus. The data suggest conditioned HR deceleration can generalize from conditioned stimulus to US as a function of conditioning.     

Chaotic components in the high-frequency EEG of rabbit cortex during formation of a conditioned defensive reflex

The chaotic component of the high-frequency EEG of the rabbit cortex was studied during the development of a conditioned defensive reflex to nonrhythmic light stimulation with electric reinforcement applied to the animal's paw. Regular changes in the dynamic parameters of the chaotic component of the high-frequency EEG were demonstrated. Characteristic changes in the asymptotic evaluations of the correlational dimensionality of the attractor of the chaotic component (DCC) of the high-frequency EEG were detected in the cortical regions studied both during the action of the conditioned stimulus and in background conditions before presentation of combinations. These changes affected both the afferent and efferent cortical regions of the conditioned reflex arc, as well as regions not directly addressed by the conditioned or unconditioned stimuli. These results allow the dynamics of changes in the morphofunctional structures of the conditioned reflex to be seen at the cortical level during learning. Significant differences in the early and late stages of conditioned reflex formation were detected. Institute of Higher Nervous Activity and Neurophysiology, Russian Academy of Sciences, Moscow. Translated from Zhurnal Vysshei Nervnoi Deyatel'nosti imeni I. P. Pavlova, Vol. 47, No. 5, pp. 858–869, September–October, 1997.