Velocity of head movements and sensory-motor adaptation during and after short spaceflight

To investigate to time course of sensory-motor adaptation to microgravity, we tested spatially-directed voluntary head movements before, during and after short spaceflight. We also tested the re-adaptation of postural responses to sensory stimulation after space flight. The cosmonaut performed in microgravity six cycles of voluntary head rotation in pitch, roll and yaw directions. During the first days of weightlessness the angular velocity of head movements increased. Over the next days of microgravity the velocity of head movements gradually decreased. On landing day a significant decrease of head rotation velocity was observed compared to the head movement velocity before spaceflight. Re-adaptation to Earth condition measured by body sway on soft support showed similar time course, but re-adaptation measured by postural responses to vestibular galvanic stimulation was prolonged. These results showed that the angular velocity of aimed head movements of cosmonauts is a good indicator of sensory-motor adaptation in altered gravity conditions.     

Vestibular Function after Repeated Space Flights

This paper presents the results from testing the vestibular function on return from repeated space flights (SF) in 32 cosmonauts of the International Space Station that were in long SFs of 125–215 days. The cosmonauts were tested twice before the flight (baseline data collection) and on days 1–2, 4–5, and 8–9 after landing. The testing was made using two methods for recording eye movements (with simultaneous recording of head movements): electro-oculography and video-oculography. It is shown that the repeated stay in the long SF leads to a considerable statistically significant reduction in the de-adaptation period. Atypical vestibular disorders and changed patterns of the otolith-semicircular canal interaction are observed mostly in the cosmonauts who have made their maiden flights to microgravity.     

Vestibular function and space motion sickness

The vestibular system plays an important role in intersensory interactions and gravitation is a natural stimulus for its receptors. Weightlessness alters the input signals of the otoliths and their effect on the pattern and dynamics of changes in the vestibular function (VF), which is accompanied by development of space adaptation syndrome (SAS) and space motion sickness (SMS). These changes occur both during the spaceflight (SF) and after returning to Earth, but the mechanisms of their development are still poorly understood and require special studies. In total, 47 Russian cosmonauts (crewmembers of long-term International Space Station (ISS) missions) have participated in the studies into VF before and after SF and nine of them, in onboard studies during SF (129–215 days) as a part of the Virtual space experiment (stage 1). Electro- and video-oculography are used to record spontaneous eye movements (SpEM), static vestibular–ocular responses during head tilts to the right or left shoulder (static otolith–cervical–ocular reflex, OCOR), and dynamic vestibular-ocular response during the head rotation around the longitudinal axis of the body. The examination is accompanied by personal and questionnaire survey on subjective responses and complaints of cosmonauts about SAS and SMS. Significant changes in SpEM (drifts of eyes, spontaneous and gaze-evoked nystagmus, and arbitrary saccades) and a decrease in OCOR (statistically significant decrease in the amplitude of ocular counter-rolling in response to head tilts up to its absence or inversion, an atypical OCOR) are observed during SF. An atypical OCOR is observed at the beginning of adaptation to weightlessness in seven of the nine cosmonauts (the first one to two weeks of SF) and repeatedly throughout the flight in all cosmonauts regardless of whether it is their first flight or not. Atypical vestibular responses after SF, similar to the responses during SF, are observed in several cosmonauts by day 9 after flight. It has been shown that atypical OCOR variants are more frequently observed in the subjects lacking any previous space experience, as well as a more pronounced decrease in this response with a concurrent increase in the response of the semicircular canals. It is also demonstrated that repeated SFs lead to a considerable shortening in the after-flight readaptation to terrestrial conditions and a considerable decrease in the degree of vestibular disorders. In the initial period of SF, the changes in VF are correlated with the complaints and manifestations of SAS and SMS; however, the complaints and the corresponding symptoms are unobservable during the further flight despite significant changes in the VF state. The patterns of the VF disorders associated with the impact of weightlessness and observed during and after SF are very similar, allowing these disorders to be regarded as SAS and SMS of different severities (intensities).     

Effects of long-term space flights on the organization of the horizontal gaze fixation reaction

The results of the Russian-Austrian space experiment Monimir, which was a part of the international space program Austromir, are presented. The characteristics of the horizontal gaze fixation reaction (hGFR) to the visual targets were studied during long-term space flights. Seven crewmembers of the space station Mir participated in our experiment. The subjects were tested four times before the flight, five times during the flight, and three to four times after landing. During the flight and after accomplishing, the characteristics of gaze fixation reaction changed regularly: the reaction time and coefficient of the gain of vestibular-ocular reflex increased; the velocities of eye-head movements increased and decreased. These changes were indicative of a disturbed control of the vestibular-ocular reflex under microgravity conditions because of variability of the vestibular input activity. The cosmonauts that had flight and non-flight professional specializations differed in strategies of their adaptation to the microgravity conditions. In the former, exposure to microgravity was accompanied by gaze hypermetry and inhibition of head movements; conversely, in the latter, the velocity of head movements increased, whereas that of saccades decreased.     

Head Movements Evoked in Alert Rhesus Monkey by Vestibular Prosthesis Stimulation: Implications for Postural and Gaze Stabilization

The vestibular system detects motion of the head in space and in turn generates reflexes that are vital for our daily activities. The eye movements produced by the vestibulo-ocular reflex (VOR) play an essential role in stabilizing the visual axis (gaze), while vestibulo-spinal reflexes ensure the maintenance of head and body posture. The neuronal pathways from the vestibular periphery to the cervical spinal cord potentially serve a dual role, since they function to stabilize the head relative to inertial space and could thus contribute to gaze (eye-in-head + head-in-space) and posture stabilization. To date, however, the functional significance of vestibular-neck pathways in alert primates remains a matter of debate. Here we used a vestibular prosthesis to 1) quantify vestibularly-driven head movements in primates, and 2) assess whether these evoked head movements make a significant contribution to gaze as well as postural stabilization. We stimulated electrodes implanted in the horizontal semicircular canal of alert rhesus monkeys, and measured the head and eye movements evoked during a 100ms time period for which the contribution of longer latency voluntary inputs to the neck would be minimal. Our results show that prosthetic stimulation evoked significant head movements with latencies consistent with known vestibulo-spinal pathways. Furthermore, while the evoked head movements were substantially smaller than the coincidently evoked eye movements, they made a significant contribution to gaze stabilization, complementing the VOR to ensure that the appropriate gaze response is achieved. We speculate that analogous compensatory head movements will be evoked when implanted prosthetic devices are transitioned to human patients.     

Sensory processing in the vestibular nuclei during active head movements

Many secondary vestibular neurons are sensitive to head on trunk rotation during reflex-induced and voluntary head movements. During passive whole body rotation the interaction of head on trunk signals related to the vestibulo-collic reflex with vestibular signals increases the rotational gain of many secondary vestibular neurons, including many that project to the spinal cord. In some units, the sensitivity to head on trunk and vestibular input is matched and the resulting interaction produces an output that is related to the trunk velocity in space. In other units the head on trunk inputs are stronger and the resulting interaction produces an output that is larger during the reflex. During voluntary head movements, inputs related to head on trunk movement combine destructively with vestibular signals, and often cancel the sensory reafferent consequences of self-generated movements. Cancellation of sensory vestibular signals was observed in all of the antidromically identified secondary vestibulospinal units, even though many of these units were not significantly affected by reflexive head on trunk movements. The results imply that the inputs to vestibular neurons related to head on trunk rotation during reflexive and voluntary movements arise from different sources. We suggest that the relative strength of reflexive head on trunk input to different vestibular neurons might reflect the different functional roles they have in controlling the posture of the neck and body.     

Effects of long-term space flights on organization of horizontal gaze fixation reaction

Results of Russian-Austrian space experiment "Monimir" which was a part of international space program "Austromir" are presented in this paper. Characteristics of horizontal gaze fixation reaction (hGFR) to visual targets were analyzed. Seven crewmembers of "Mir" space station expeditions took part in the experiment. Experiments were carried out 4 times before space flight, 5 times in flight and 3-4 times after landing. There were revealed significant alterations in characteristics of gaze fixation reaction during flight and after its accomplishing, namely: an increase of the time of gaze fixation to the target, changes of eye and head movements' velocity and increase of the gain of vestibular-ocular reflex, that pointed out to the disturbances of the control mechanisms of vestibular-ocular reflex in weightlessness caused by changes of vestibular input's activity. There was discovered also the difference in the strategies of adaptation to microgravity conditions among the cosmonauts of flight and non-flight occupation: in the first group exposure to weightlessness was accompanied by gaze hypermetry and inhibition of head movements; in the second one--on the contrary--by increase of head movement velocity and decrease of saccades' velocity.     

Integrative responses of neurons in nucleus tractus solitarius to visceral afferent stimulation and vestibular stimulation in vertical planes

Anatomical studies have demonstrated that the vestibular nuclei project to nucleus tractus solitarius (NTS), but little is known about the effects of vestibular inputs on NTS neuronal activity. Furthermore, lesions of NTS abolish vomiting elicited by a variety of different triggering mechanisms, including vestibular stimulation, suggesting that emetic inputs may converge on the same NTS neurons. As such, an emetic stimulus that activates gastrointestinal (GI) receptors could alter the responses of NTS neurons to vestibular inputs. In the present study, we examined in decerebrate cats the responses of NTS neurons to rotations of the body in vertical planes before and after the intragastric administration of the emetic compound copper sulfate. The activity of more than one-third of NTS neurons was modulated by vertical vestibular stimulation, with most of the responsive cells having their firing rate altered by rotations in the head-up or head-down directions. These responses were aligned with head position in space, as opposed to the velocity of head movements. The activity of NTS neurons with baroreceptor, pulmonary, and GI inputs could be modulated by vertical plane rotations. However, injection of copper sulfate into the stomach did not alter the responses to vestibular stimulation of NTS neurons that received GI inputs, suggesting that the stimuli did not have additive effects. These findings show that the detection and processing of visceral inputs by NTS neurons can be altered in accordance with the direction of ongoing movements.     

Head position during various locomotor executions after prolonged microgravity exposure

Head orientation in the sagittal plane was assessed before and the second and sixth days after a prolonged microgravity exposure (6 months) in 7 astronauts, which were instructed to execute various locomotor movements (stair ascent, jump down and level walk). Crewmembers systematically flexed down the head when they executed the locomotion the second day post-flight compared to the pre-flight walk. Cephalic orientation was normally positioned when movements was performed the sixth day after Earth re-entry. This head flexion may reflect a strategy used by cosmonauts to increase the vestibular sensibility, and then improve the control of the dynamic equilibrium during locomotor movements. The high rate of re-adaptation indicated that cosmonauts rapidly recovered their normal walking capacities despite the prolonged microgravity exposure.     

Comparative measurement of visual stability in Earth and cosmic space.

Three theories have been suggested as to the cause of space motion sickness: 1) eye and vestibular sensory mismatch, 2) abnormal shift of body fluids producing increased intracranial pressure and 3) pre-warning signals for unpleasant physical situations by self-produced neurotoxic substances released in the body. We are interested in the possible functional disabilities/incongruities of eye, head and body movements in 0-G. Space motion sickness might be explained from the viewpoint of lack of coordination of the movements of the eye and head. It is important to ascertain the significance of gravity in the maintenance of human visual stability. We will examine the coordination of Japanese Payload Specialist (JPS) eye and head movement by electrooculogram and neck muscle electromyogram recordings, as well as obtaining a subjective evaluation of visual stability from the PS during space flight. We hypothesize that 1) poor performance of the eye movement will be observed, 2) unusual neck muscle activity will be observed and 3) there will be decreased visual stability in micro gravity. We obtained all digital data and VCR taped image data in [TEXT MISSING]     

Effects of vestibular and support afferentation upon visual pursuit in microgravity

Evaluation of the accuracy of eye turns (saccades) to fix a jerky pointed stimulus, and smooth pursuit of slow linear and sinusoidal movements of both pointed and optokinetic stimuli was performed in 31 cosmonauts on flight days 2-3, 5-8, 30, and once in one or two months of mission. An additional investigation of the eye pursuit function involved 10 cosmonauts, who, after testing during free floating, fulfilled stimulus tracking following a cycle of active head rotation, and 14 cosmonauts who received support afferentation. It was found that at the beginning of adaptation and periodically in the course of long mission, the systems of slow pursuit tracking adopted the strategy of saccadic approximation whereby gaze fixation was achieved through a sequence of macro- or microsaccadic movements. It was demonstrated that these disturbances, practically in all investigated cosmonauts, were consequent to the vestibular deprivation developing in microgravity. Vestibular afferentation produced by active head rotation improved characteristics of visual pursuit. Support deprivation also affects pursuit tracking by cosmonauts who form the concept of space orientation based on perception of their head and leg position. With support afferentation, these cosmonauts demonstrated improved visual pursuit characteristics.     

High school and college baseball pitchers response and glove movements to line drives

The timing of glove movements used by baseball pitchers to catch fast approaching balls (i.e., line drives) was examined in two tests to determine the responses and temporal characteristics of glove movements in high school and college baseball pitchers. Balls were projected toward the head of participants at 34.8 m.s-1 (78 mph) on average in an indoor test and at speeds approaching 58.1 m.s-1 (130 mph) in a field test. Pitchers caught over 80% and 15% of the projected balls in the indoor and field tests, respectively. Analyses of glove responses indicated that all pitchers could track the line drives and produce coordinated glove movements, which were initiated 160 ms (+/-47.8), on average, after the ball was launched. College pitchers made initial glove movements sooner than high school pitchers in the field test (p=0.012). In contrast, average glove velocity for pitchers increased from 1.33 (+/-0.61) to 3.45 (+/-0.86) m.s-1 across the tests, but did not differ between experience levels. Glove movement initiation and speed were unrelated, and pitchers utilized visual information throughout the ball's flight to catch balls that approached at speeds exceeding the estimated speeds in competitive situations.     

Vergence and Standing Balance in Subjects with Idiopathic Bilateral Loss of Vestibular Function

There is a natural symbiosis between vergence and vestibular responses. Deficits in vergence can lead to vertigo, disequilibrium, and postural instability. This study examines both vergence eye movements in patients with idiopathic bilateral vestibular loss, and their standing balance in relation to vergence. Eleven patients participated in the study and 16 controls. Bilateral loss of vestibular function was objectified with many tests; only patients without significant response to caloric tests, to video head impulse tests and without vestibular evoked myogenic potentials were included in the study.

Vergence testing (from 8 patients and 15 controls)

A LED display with targets at 20, 40, and 100 cm along the median plane was used to elicit vergence eye movements, recorded with the IRIS device.

Standing balance (11 patients and 16 controls)

Four conditions were run, each lasting 1 min: fixation of a LED at 40 cm (convergence of 9°), at 150 cm (convergence of 2.3°); this last condition was repeated with eyes closed. Comparison of the eyes closed-eyes open conditions at 150 cm allowed evaluation of the Romberg Quotient. In the forth condition, two LEDS, at 20 and at 100 cm, were light on, one after the other for 1 sec, causing the eyes to converge then diverge. Standing balance was recorded with an accelerometer placed at the back near the center of mass (McRoberts, Dynaport).

Results

Vergence

Relative to controls, convergence eye movements in patients showed significantly lower accuracy, lower mean velocity, and saccade intrusions of significantly higher amplitude.

Balance

The normalized 90% area of body sway was significantly higher for patients than for controls for all conditions. Yet, similarly to controls, postural stability was better while fixating at near (sustained convergence) than at far, or while making active vergence movements. We argue that vestibular loss deteriorates convergence, but even deficient, convergence can be helpful for postural control.     

Covert Anti-Compensatory Quick Eye Movements during Head Impulses

Background

Catch-up saccades during passive head movements, which compensate for a deficient vestibulo-ocular reflex (VOR), are a well-known phenomenon. These quick eye movements are directed toward the target in the opposite direction of the head movement. Recently, quick eye movements in the direction of the head movement (covert anti-compensatory quick eye movements, CAQEM) were observed in older individuals. Here, we characterize these quick eye movements, their pathophysiology, and clinical relevance during head impulse testing (HIT).

Methods

Video head impulse test data from 266 patients of a tertiary vertigo center were retrospectively analyzed. Forty-three of these patients had been diagnosed with vestibular migraine, and 35 with Menière’s disease.

Results

CAQEM occurred in 38% of the patients. The mean CAQEM occurrence rate (per HIT trial) was 11±10% (mean±SD). Latency was 83±30 ms. CAQEM followed the saccade main sequence characteristics and were compensated by catch-up saccades in the opposite direction. Compensatory saccades did not lead to more false pathological clinical head impulse test assessments (specificity with CAQEM: 87%, and without: 85%). CAQEM on one side were associated with a lower VOR gain on the contralateral side (p<0.004) and helped distinguish Menière’s disease from vestibular migraine (p = 0.01).

Conclusion

CAQEM are a common phenomenon, most likely caused by a saccadic/quick phase mechanism due to gain asymmetries. They could help differentiate two of the most common causes of recurrent vertigo: vestibular migraine and Menière’s disease.     

Neuronal co-processing of course deviation and head movements in locusts

Summary Descending deviation detector neurons (DDNs) of Locusta migratoria are characterized physiologically by their responses to light on/off stimuli, simulated course deviation (rotation of an artificial horizon), passive rotation of the head, frontal wind, and flight activity. The investigation emphasises on the co-processing of exteroceptive input signalling course deviation (mainly movement of the retinal image, but also wind), and proprioceptive input signalling head movement and position. Stimuli were presented in combinations as expected during natural behavior. Eight DDNs are described for the first time, and 3 previously described DDNs are characterized further. Responses to horizon rotation and imposed head movements are assigned to one of 4 response types: (1) the horizon-only type codes retinal slip and/or the position of the horizon in the visual field but ignores cervical proprioception; (2) the head-only type ignores visually simulated course deviation but codes for movement or position of the head; (3) in the compensating type, head rolling causes visual input and cervical proprioceptive input of opposite signs, so that head movements themselves are ignored, whereas course deviations are recognized; (4) in the amplifying type, head rolling causes visual input and cervical proprioceptive input of the same sign, i.e. one input amplifies the other. This classification does not take the various responses to wind into account. In several DDNs, responses to phasic and tonic stimuli of the same modality, and/or responses to deviations about different axes could be assigned to different response types. Activity in DDNs has been shown previously to result in steering responses of wings, legs, abdomen and/or the head. It is proposed that different kinds of flight steering (e.g. corrective course control, intentional steering, orientation towards or away from a target) may be controlled by selective enhancement or suppression of responses or motor effects of DDN-subpopulations.Abbreviations AP action potential - DDN descending deviation detector neuron - DNI, DNC, DNM descending deviation detector neurons receiving major input from the ipsilateral, contralateral, and median ocellus respectively - PDDSMD protocerebral, descending direction-selective motion-detecting neuron - PI(2)5 descending deviation detector neuron with the cell body in the pars intercerebralis medialis - TCG tritocerebral commissure giant neuron     

Gelatinase activities in the airways of premature infants and development of bronchopulmonary dysplasia

Matrix-degrading metalloproteinases may play a role in the pathophysiology of bronchopulmonary dysplasia (BDP). We, therefore, evaluated correlations between gelatinase activities [metalloproteinase (MMP)-2 and MMP-9] or tissue inhibitor of metalloproteinase (TIMP)-1 levels present in the airways during the initial phase of hyaline membrane disease and the onset of BPD. Tracheal aspirates were obtained within 6 h of birth (day 0) from 64 intubated neonates with a gestational age < or =30 wk. Forty-five neonates were resampled on day 3 or 5. Total MMP-2 level measured by zymography fell with time, whereas total MMP-9 level and TIMP-1 levels, assayed by ELISA, increased; the MMP-9 increase correlated with the increase in airway inflammatory cell numbers. Among the parameters measured on day 0, 3, or 5, lower total MMP-2 level, lower birth weight, and higher fraction of inspired oxygen on day 0 were significantly and independently associated with the development of BPD. In conclusion, MMP-9 level and TIMP-1 levels increased after birth but are not linked to BPD outcome. In contrast, low MMP-2 level at birth is strongly associated with the development of BPD.     

Vestibular reafference shapes voluntary movement

The vestibular organs in the inner ear are commonly thought of as sensors that serve balance, gaze control, and higher spatial functions such as navigation. Here, we investigate their role in the online control of voluntary movements. The central nervous system uses sensory feedback information during movement to detect and correct errors as they develop. Vestibular organs signal three-dimensional head rotations and translations and so could provide error information for body movements that transport the head in space. To test this, we electrically stimulated human vestibular nerves during a goal-directed voluntary tilt of the trunk. The stimulating current waveform was made identical to the angular velocity profile of the head in the roll plane. With this, we could proportionally increase or decrease the rate of vestibular nerve firing, as if the head were rotating faster or slower than it actually was. In comparison to movements performed without stimulation, subjects tilted their trunk faster and further or slower and less far, depending upon the polarity of the stimulus. The response was negligible when identical stimulus waveforms were replayed to stationary subjects. We conclude that the brain uses vestibular information for online error correction of planned body-movement trajectories.     

Maintenance of pancreatic endocrine B cells of neonatal rat: Part-XIV--Effect of maternal hyperglycemia on the secretion of insulin and glucagon

Effects of gestational hyperglycemia on A and B cells were examined in pancreatic monolayer islet cell cultures of neonatal rats from mothers of normoglycemia (C) and made slightly (SH), moderately (MH) and highly hyperglycemic (HH) by streptozotocin injection. Monolayer cultures were maintained for 7 days in the medium with 5.5 mM glucose plus 1 mM 2-deoxyglucose. On day 0, B cells of the SH group were more responsive to glucose and 2-ketoisocaproate than those of other groups. On day 7, the response of B cells in the C and SH groups was remarkably enhanced, thus displaying a dose-dependent increasing pattern of insulin secretion in response to glucose, 2-ketoisocaproate and arginine, and a convex-type secretion to leucine. However, there was no response by B cells in the MH and HH groups. Further, a dose-dependent inhibition of glucagon secretion due to glucose was seen in A cells of the C and SH groups on day 0 and day 7. The responses of these A cells to other nutrients were slightly decreased or were of a low convex-type. In the MH group, however, the glucagon secretion was remarkably enhanced due to leucine and 2-ketoisocaproate on day 0 and day 7, and due to arginine on day 7, although it remained suppressed by glucose. A cells of the HH group were unresponsive through the whole culture period. These results suggest that the development of A- and B-cell responses in vitro of neonates was differently affected by the degree of maternal hyperglycemia.     

Gestational regulation of the gene expression of C-type natriuretic peptide in mouse reproductive and embryonic tissue

C-Type natriuretic peptide (CNP) is a vasoactive hormone and the endothelial component of the natriuretic peptide system. We examined the expression of CNP in mouse reproductive organs and embryos at different stages of gestation. Pregnant mice were killed and embryos were dissected on gestational days 9.5, 12.5, 15.5, 18.5 postconceptionem (pc) and at term. Nonpregnant mice were used as controls. Total RNA was isolated from placenta, ovaries, myometrium and from head and trunk of embryos and neonates. CNP-mRNA was quantified by ribonuclease-protection assay (RPA). Uterine CNP-mRNA concentrations increase during pregnancy up to the sevenfold concentration, whereas in the ovaries these levels decrease to 10% compared to nonpregnant controls. In the placenta, a peak of CNP expression has been observed around day 15.5 pc, whereby placenta showed the strongest CNP signals. CNP-mRNA concentrations in embryos are gestational age-dependent with a high level at day 9.5 pc in head and trunk. These results indicate that CNP has a regulatory function in pregnancy and embryonic development.