Postural control under clinorotation in upside-down catfish, Synodontis nigriventris.

The upside-down catfish Synodontis nigriventris has a unique habit of swimming and resting upside-down in free water. This behavior leads to the assumption that the catfish has a specific gravity information processing system. We examined the postural control behaviors in the catfish under clinorotation which is usually used for producing pseudo-microgravity. Synodontis nigriventris kept its body posture at a stable area of the rotated flask in which the catfish was kept, when it was clinorotated at the rate of 60 rpm. In contrast to Synodontis nigriventris, a related species, Corydoras paleatus, did not show such steady postural control. When the flask was rotated at a lower rate of 30 rpm or a higher rate of 100 rpm, Synodontis nigriventris as well as Corydoras paleatus showed a considerable disturbed control of body posture. In this condition, they were frequently rotated with the flask. These findings suggest that Synodontis nigriventris has a high ability to keep upside-down posture and the gravity sensation in this catfish is likely to contribute to its different postural control from that of many other fishes.     

High transmittance of X-rays in the utricular otolith of upside-down swimming catfish, Synodontis nigriventris.

The upside-down swimming catfish (Synodontis nigriventris) has unique behavior, i.e., it frequently shows a stable upside-down posture during swimming and resting. To examine whether the unique postural control in S. nigriventris results from the characteristics of the vestibular organ, we observed the morphological aspects of the otolith and the orientation of sensory hair cells in the utricle. Soft X-ray densitometry analysis showed that the transmittance of soft X-rays in the otolith of S. nigriventris was higher than that in a closely related species (Synodontis multipunctatus) belonging to Synodontis family, goldfish (Carassius auratus) or miniature catfish (Corydoras paleatus) which shows upside-up swimming. The higher transmittance of soft X-rays suggests that the density of the otolith in S. nigriventris is lower than that in S. multipunctatus, C. auratus or C. paleatus. It is possible that the low density of the otolith may have a relation to the control of the unique upside-down posture of S. nigriventris. The hair cells in S. nigriventris were present at the ventral to ventro-lateral site of the utricular epithelium, forming a single hair cell layer as in the other 3 species of fish. The orientation of the sensory hair cells does not appear to cause the unique postural control.     

Unique postural control of upside-down swimming catfish,Synodontis nigriventris,not affected by the change of gravity

In general, most fishes maintain a swimming posture with the dorsal side towards the water surface under normal gravity condition. In contrast to normal fishes, a catfish Synodontis nigriventris, shows a unique postural control. The catfish keeps its posture with the ventral side towards the water surface and the dorsal side towards water bottom under normal gravity. This evidence leads one to assume that the upside-down posture of the catfish is controlled by gravity sensation in a manner different from that of other fishes. However, it has remained unclear to date whether the gravity sensation contributes to the unique postural control of this catfish. We examined its postural control in intact and labyrinth-removed catfish using a clinostat which generates a specific gravity environment (pseudo-microgravity) on earth. In addition, we examined its postural control under microgravity during parabolic flights.     

Relationship between frequency of upside-down posture and space size around upside-down catfish, Synodontis nigriventris.

The catfish Synodontis nigriventris shows a unique habit taking a stable upside-down posture in free water regardless of an above, one-sided illumination. This upside-down posture can be observed when the catfish is apart from objects because the catfish usually orients its ventral side towards the water bottom or objects due to a so-called ventral substrate response. Thus, it is not easy to study the mechanism of the upside-down posture. To resolve this problem, the frequency of the upside-down posture was measured by using various sizes of vessel in which the catfish was kept. Video analysis showed that the frequency of the upside-down posture depended on the space size around the catfish. The smaller the size became, the higher the frequency of the upside-down posture became. Furthermore, the frequency of the upside-down posture depended on the shape of the vessel bottom. Curved-bottom vessels induced the upside-down posture more frequently than flat bottom. These findings suggest that a small, curved-bottom vessel is ideal for researching the upside-down postural control mechanism.     

Postural control mechanisms in the upside-down catfish (Synodontis nigriventris)

The catfishSynodontis nigriventris normally swims upside-down but can assume any posture in response to a substrate it swims close to. Postural reflexes of the body and the eyes, labyrinthine anatomy and passively maintained posture were investigated to obtain indications for possible mechanisms controlling the peculiar postural behavior of this fish. Saccade-like resetting movements of the eyes during counter-roll to body tilt about the longitudinal axis, and maintained tilted swimming positions in blinded fish suggest that these animals reset their vestibular CNS circuits to zero when in tilted positions.Synodontis nigriventris is thus able to maintain any posture without interference from tilt-counteracting vestibular reflexes. The normal upside-down swimming apparently results from a central bias for this position and a supporting ventral light response.We conclude that if the reafference principle applies to the phenomena investigated, the efference copy may be fed through an integrator before reaching vestibular reflex circuits.     

Comparative study of otolith organs between two species of upside-down swimming catfish Synodontis nigriventris showing converse swimming postures

To clarify whether the unique postural control of the upside‐down swimming catfish (Synodontis nigriventris, family Mochokidae) is related to the histological characteristics of the otolith organs, we performed light microscopic observation of the utricle, the saccule and the lagena. The histological aspects of the otolith organs were compared between S. nigriventris and Synodontis multipunctatus, which belong to the same genus. S. multipunctatus usually shows upside‐up swimming posture except for feeding behaviour near water surface. As controls, we additionally used a miniature catfish, Corydoras paleatus and goldfish, Carassius auratus, which shows upside‐up swimming posture. We concluded that the structural aspects of the otolith organs did not cause the unique postural control of S. nigriventris. Light microscopic observation clarified the following aspects: (1) The utricle of S. nigriventris was located at the anterior region of the otocyst and under the semicircular canals, and the saccule and the lagena were located at the posteroventral region of the otocyst like those of S. multipunctatus and the other two fishes. (2) The hair cells of the utricle were arranged on the horizontal plane of the fishes with a variation in cell size at the ventral and ventrolateral sites in S. nigriventris, S. multipunctatus and the other two fishes. (3) The hair cells of the saccule and lagena of S. nigriventris, S. multipunctatus and C. auratus presented perpendicular to the horizontal plane of the fish. (4) Region‐specific differences in the size and shape of the hair cells of S. nigriventris were observed along the three‐dimensional axes of the otolith organs like those of S. multipunctatus and the other two fishes. It is unlikely that the unique postural control of upside‐down catfish is related to the localization of the utricle, the saccule and the lagena and the distribution of the different types of hair cell of the otolith organs. Furthermore, the distribution of the hair cells suggests that the otolith organs in S. nigriventris can detect three‐dimensional postural changes like the organs of other fishes showing generally observed upside‐up swimming posture.     

Cell proliferation of Paramecium tetraurelia under clinorotation.

It has been reported that Paramecium proliferates faster under microgravity in space, and slower under hypergravity (Kato et al., 2003). Effects of gravity on cell proliferation could be discussed in terms of energetics of swimming. Because of the characteristics of 'gravikinesis' as well as 'gravitaxis', Paramecium would decrease the energy expenditure under microgravity and increase it under hypergravity. The larger stock of energy would enhance the proliferation under microgravity. In order to simulate the effect of microgravity, we investigated the proliferation under clinorotation. When cells were rotated at 2.5 rpm, the proliferation rate decreased. Similar but less pronounced decrease was also found under low speed clinorotation (0.2 rpm).     

Vestibular Lesion-Induced Developmental Plasticity in Spinal Locomotor Networks during Xenopus laevis Metamorphosis

During frog metamorphosis, the vestibular sensory system remains unchanged, while spinal motor networks undergo a massive restructuring associated with the transition from the larval to adult biomechanical system. We investigated in Xenopus laevis the impact of a pre- (tadpole stage) or post-metamorphosis (juvenile stage) unilateral labyrinthectomy (UL) on young adult swimming performance and underlying spinal locomotor circuitry. The acute disruptive effects on locomotion were similar in both tadpoles and juvenile frogs. However, animals that had metamorphosed with a preceding UL expressed restored swimming behavior at the juvenile stage, whereas animals lesioned after metamorphosis never recovered. Whilst kinematic and electrophysiological analyses of the propulsive system showed no significant differences in either juvenile group, a 3D biomechanical simulation suggested that an asymmetry in the dynamic control of posture during swimming could account for the behavioral restoration observed in animals that had been labyrinthectomized before metamorphosis. This hypothesis was subsequently supported by in vivo electromyography during free swimming and in vitro recordings from isolated brainstem/spinal cord preparations. Specifically, animals lesioned prior to metamorphosis at the larval stage exhibited an asymmetrical propulsion/posture coupling as a post-metamorphic young adult. This developmental alteration was accompanied by an ipsilesional decrease in propriospinal coordination that is normally established in strict left-right symmetry during metamorphosis in order to synchronize dorsal trunk muscle contractions with bilateral hindlimb extensions in the swimming adult. Our data thus suggest that a disequilibrium in descending vestibulospinal information during Xenopus metamorphosis leads to an altered assembly of adult spinal locomotor circuitry. This in turn enables an adaptive compensation for the dynamic postural asymmetry induced by the vestibular imbalance and the restoration of functionally-effective behavior.     

How experienced alpine-skiers cope with restrictions of ankle degrees-of-freedom when wearing ski-boots in postural exercises

The present study investigates the mechanisms underlying changes in postural strategy that occur to compensate for mechanical ankle joint restrictions induced by wearing ski-boots during postural exercises. Fourteen experienced skiers were asked to stand as still as possible in a stable (STA) posture and in 2 postures with instability in the medio/lateral and antero/posterior (ML and AP postures) direction. Postural tasks were performed with eyes open or closed and while wearing or not wearing ski-boots. The electromyographic (EMG) activity of representative lower limb muscles and positions of centre-of-foot pressure (COP) were recorded and analyzed. Our results illustrated enhanced postural performances with ski-boots in the STA posture, whereas postural performances remained unchanged when wearing ski-boots in the ML and AP postures. Analysis of COP sways in the frequency domain did not illustrate any modification in the contribution of different neuronal loops when the study subjects wore ski-boots. EMG showed that the mechanical effects of wearing ski-boots were compensated by changes in postural strategy through the reorganization of muscle coordination, made possible by inherent redundancies in the human body. The preservation of postural performances, despite restrictions of ankle degrees-of-freedom induced by ski-boots, emphasizes the subjects’ capacity to exploit the additional support provided by ski-boots by adequately adjusting muscle coordination to control posture in different balance conditions.     

N-Acetyl-L-Leucine Accelerates Vestibular Compensation after Unilateral Labyrinthectomy by Action in the Cerebellum and Thalamus

An acute unilateral vestibular lesion leads to a vestibular tone imbalance with nystagmus, head roll tilt and postural imbalance. These deficits gradually decrease over days to weeks due to central vestibular compensation (VC). This study investigated the effects of i.v. N-acetyl-DL-leucine, N-acetyl-L-leucine and N-acetyl-D-leucine on VC using behavioural testing and serial [¹⁸F]-Fluoro-desoxyglucose ([¹⁸F]-FDG)-μPET in a rat model of unilateral chemical labyrinthectomy (UL). Vestibular behavioural testing included measurements of nystagmus, head roll tilt and postural imbalance as well as sequential whole-brain [¹⁸F]-FDG-μPET was done before and on days 1,3,7 and 15 after UL. A significant reduction of postural imbalance scores was identified on day 7 in the N-acetyl-DL-leucine (p < 0.03) and the N-acetyl-L-leucine groups (p < 0.01), compared to the sham treatment group, but not in the N-acetyl-D-leucine group (comparison for applied dose of 24 mg i.v. per rat, equivalent to 60 mg/kg body weight, in each group). The course of postural compensation in the DL- and L-group was accelerated by about 6 days relative to controls. The effect of N-acetyl-L-leucine on postural compensation depended on the dose: in contrast to 60 mg/kg, doses of 15 mg/kg and 3.75 mg/kg had no significant effect. N-acetyl-L-leucine did not change the compensation of nystagmus or head roll tilt at any dose. Measurements of the regional cerebral glucose metabolism (rCGM) by means of μPET revealed that only N-acetyl-L-leucine but not N-acetyl-D-leucine caused a significant increase of rCGM in the vestibulocerebellum and a decrease in the posterolateral thalamus and subthalamic region on days 3 and 7. A similar pattern was found when comparing the effect of N-acetyl-L-leucine on rCGM in an UL-group and a sham UL-group without vestibular damage. In conclusion, N-acetyl-L-leucine improves compensation of postural symptoms after UL in a dose-dependent and specific manner, most likely by activating the vestibulocerebellum and deactivating the posterolateral thalamus.     

Altered gravity affects ventral root activity during fictive swimming and the static vestibuloocular reflex in young tadpoles (Xenopus laevis)

During early periods of life, modifications of the gravitational environment affect the development of sensory, neuronal and motor systems. The vestibular system exerts significant effects on motor networks that control eye and body posture as well as swimming. The objective of the present study was to study whether altered gravity (AG) affects vestibuloocular and spinal motor systems in a correlated manner. During the French Soyuz taxi flight Andromède to the International Space Station ISS (launch: October 21, 2001; landing: October 31, 2001) Xenopus laevis embryos were exposed for 10 days to microgravity (microg). In addition, a similar experiment with 3g-hypergravity (3g) was performed in the laboratory. At onset of AG, embryos had reached developmental stages 24 to 27. After exposure to AG, each tadpole was tested for its roll-induced vestibuloocular reflex (rVOR) and 3 hours later it was tested for the neuronal activity recorded from the ventral roots (VR) during fictive swimming. During the post-AG recording periods tadpoles had reached developmental stages 45 to 47. It was observed that microgravity affected VR activity during fictive swimming and rVOR. In particular, VR activity changes included a significant decrease of the rostrocaudal delay and a significant increase of episode duration. The rVOR-amplitude was transiently depressed. Hypergravity was less effective on the locomotor pattern; occurring effects on fictive swimming were the opposite of microg effects. As after microgravity, the rVOR was depressed after 3g-exposure. All modifications of the rVOR and VR-activity recovered to normal levels within 4 to 7 days after termination of AG. Significant correlations between the rVOR amplitude and VR activity of respective tadpoles during the recording period have been observed in both tadpoles with or without AG experience. The data are consistent with the assumptions that during this period of life which is characterized by a progressive development of vestibuloocular and vestibulospinal projections (i) microgravity retards the development of VR activity while hypergravity weakly accelerates it; (ii) that microgravity retards the rVOR development while hypergravity caused a sensitization, and that (iii) AG-induced changes of VR activity during fictive swimming have a vestibular origin.     

T cell activation responses are differentially regulated during clinorotation and in spaceflight.

Studies of T lymphocyte activation with mitogenic lectins during spaceflight have shown a dramatic inhibition of activation as measured by DNA synthesis at 72 h, but the mechanism of this inhibition is unknown. We have investigated the progression of cellular events during the first 24 h of activation using both spaceflight microgravity culture and a ground-based model system that relies on the low shear culture environment of a rotating clinostat (clinorotation). Stimulation of human peripheral blood mononuclear cells (PBMCs) with soluble anti-CD3 (Leu4) in clinorotation and in microgravity culture shows a dramatic reduction in surface expression of the receptor for IL-2 (CD25) and CD69. An absence of bulk RNA synthesis in clinorotation indicates that stimulation with soluble Leu4 does not induce transition of T cells from G0 to the G1 stage of the cell cycle. However, internalization of the TCR by T cells and normal levels of IL-1 synthesis by monocytes indicate that intercellular interactions that are required for activation occur during clinorotation. Complementation of TCR-mediated signaling by phorbol ester restores the ability of PBMCs to express CD25 in clinorotation, indicating that a PKC-associated pathway may be compromised under these conditions. Bypassing the TCR by direct activation of intracellular pathways with a combination of phorbol ester and calcium ionophore in clinorotation resulted in full expression of CD25; however, only partial expression of CD25 occurred in microgravity culture. Though stimulation of purified T cells with Bead-Leu4 in microgravity culture resulted in the engagement and internalization of the TCR, the cells still failed to express CD25. When T cells were stimulated with Bead-Leu4 in microgravity culture, they were able to partially express CD69, a receptor that is constitutively stored in intracellular pools and can be expressed in the absence of new gene expression. Our results suggest that the inhibition of T cell proliferative response in microgravity culture is a result of alterations in signaling events within the first few hours of activation, which are required for the expression of important regulatory molecules.     

Effect of spaceflight on isoflavonoid accumulation in etiolated soybean seedlings

In order to explore the potential impact of microgravity on flavonoid biosynthesis, we examined isoflavonoid levels in soybean (Glycine max) tissues generated under both spaceflight and clinorotation conditions. A 6-day Space Shuttle-based microgravity exposure resulted in enhanced accumulation of isoflavone glycosides (daidzin, 6"-O-malonyl-7-O-glucosyl daidzein, genistin, 6"-O-malonyl-7-O-glucosyl genistein) in hypocotyl and root tissues, but reduced levels in cotyledons (relative to 1g controls on Earth). Soybean seedlings grown on a horizontally rotating clinostat for 3, 4 and 5 days exhibited (relative to a vertical clinorotation control) an isoflavonoid accumulation pattern similar to the space-grown tissues. Elevated isoflavonoid levels attributable to the clinorotation treatment were transient, with the greatest increase observed in the three-day-treated tissues and smaller increases in the four- and five-day-treated tissues. Differences between stresses presented by spaceflight and clinorotation and the resulting biochemical adaptations are discussed, as is whether the increase in isoflavonoid concentrations were due to differential rates of development under the "gravity" treatments employed. Results suggest that spaceflight exposure does not impair isoflavonoid accumulation in developing soybean tissues and that isoflavonoids respond positively to microgravity as a biochemical strategy of adaptation.     

Physiological changes of premotor nonspiking interneurons in the central compensation of eyestalk posture following unilateral sensory ablation in crayfish

We investigated how the physiological characteristics and synaptic activities of nonspiking giant interneurons (NGIs), which integrate sensory inputs in the brain and send synaptic outputs to oculomotor neurons innervating eyestalk muscles, changed after unilateral ablation of the statocyst in order to clarify neuronal mechanisms underlying the central compensation process in crayfish. The input resistance and membrane time constant in recovered animals that restored the original symmetrical eyestalk posture 2 weeks after operation were significantly greater than those immediately after operation on the operated side whereas in non-recovered animals only the membrane time constant showed a significant increase. On the intact side, both recovered and non-recovered animals showed no difference. The frequency of synaptic activity showed a complex pattern of change on both sides depending on the polarity of the synaptic potential. The synaptic activity returned to the bilaterally symmetrical level in recovered animals while bilateral asymmetry remained in non-recovered ones. These results suggest that the central compensation of eyestalk posture following unilateral impairment of the statocyst is subserved by not only changes in the physiological characteristics of the NGI membrane but also the activity of neuronal circuits presynaptic to NGIs.     

Molecular mechanisms of Brainstem plasticity

Vestibular compensation is the process of behavioral recovery that occurs following unilateral deafferentation of the vestibular nerve fibers (unilateral labyrinthectomy, UL). Since UL results in a permanent loss of vestibular input from the ipsilateral vestibular (VIIIth) nerve, vestibular compensation is attributed to CNS plasticity and has been used as a general model of lesion-induced CNS plasticity. Behavioral recovery from the ocular motor and postural symptoms of UL is correlated with a partial return of resting activity to neurons in the vestibular nucleus (VN) on the deafferented side (the "deafferented VN"), and lesions to the deafferented VN prevent compensation; therefore, the regeneration of resting activity within the deafferented VN is believed to have a causal role in vestibular compensation. The biochemical mechanisms responsible for the adaptive neuronal changes within the deafferented VN are poorly understood. Neuropeptide hormone fragments, such as adrenocorticotrophic hormone (ACTH)-4-10, have been shown to accelerate vestibular compensation and can act directly on some VN neurons in vitro. Antagonists for the N-methyl-D-aspartate (NMDA) receptor have been shown to inhibit vestibular compensation if administered early in the compensation process. Biochemical studies in frog indicate marked alterations in the phosphorylation patterns of several proteins during compensation, and the in vitro phosphorylation of some of these proteins is modulated by ACTH-(1-24), calcium (Ca2+), and calmodulin or protein kinase C. It is therefore possible that ACTH fragments and NMDA antagonists (via their effects on NMDA receptor-mediated Ca2+ channels) modulate vestibular compensation through their action on Ca(2+)-dependent pathways within VN neurons. Recent studies have shown that some Ca2+ channel antagonists and the Ca(2+)-dependent enzyme inhibitor calmidazolium chloride facilitate vestibular compensation. How the regulation of Ca2+ may be related to the neuronal changes responsible for vestibular compensation is unclear at present.     

Feeding ecology and postural behaviour of the three-toed sloth (Bradypus variegatus flaccidus) in northern Venezuela

We studied the diet, activity budget, vertical ranging, and postural behaviour in relation to weather of the three-toed sloth (Bradypus variegatus flaccidus) in disturbed montane forest remnants (1150 m asl) in northern Venezuela. Sloths spent most (72.9%) of their time resting and had a nearly exclusive (99.4%) leaf diet. While resting they assumed a sitting – not hanging – posture mostly (90.2% of observations). Species of three families, Clethraceae, Cecropiaceae, and Clusiaceae accounted for 77% of feeding records. Young leaves (67.2%) accounted for most of the leaf diet. Activity and posture were dependent on weather conditions. Sloths fed more often during mid-day hours and tended to rest more at dawn and dusk. In northern Venezuela sloths tended to use more frequently the upper strata of the canopy, while in warmer lowland sites they use intermediate levels more often. They adopted postures that maximized exposure of ventral surfaces to incident solar radiation when sunny, but minimized their surface area by huddling when cloudy, foggy or rainy. We propose that sunning behaviour of sloths may speed up their fermentation rate, and ultimately, might have been an important selective factor in the evolution of derived upside-down posture of sloths.     

The development of the static vestibulo-ocular reflex in the Southern Clawed Toad,Xenopus laevis

The static vestibulo-ocular reflex was investigated in tadpoles at different times following unilateral destruction of the labyrinth during the period of early organogenesis and premetamorphosis. Balance compensation is completed after a few weeks, while gain compensation only occurs partially (Figs. 2-4). Tadpoles hemilabyrinthectomized in the age of 2.5 days (stage 38) develop no vestibular nuclei on their lesioned side, while tadpoles operated later in their life, possess these nuclei (Figs. 5, 6) even if they were not detectable at the operation day (Fig. 7). For their dorsal vestibular nucleus (DVN), the number of neurons is usually larger on the intact than on the lesioned side; while for the ventral vestibular nucleus (VVN), there is either numerical symmetry or a transient decrease of cell number on the intact side (Fig. 5). The results demonstrate that vestibular compensation occurs even if vestibular nuclei have developed only on one side, i.e. the vestibular commissure is not a prerequisite for a successful compensation process. It is discussed whether the use of extra-vestibular error signals for balance but not for gain compensation may cause the differences in time courses of both compensation processes.     

Dynamic stability of a human standing on a balance board

The neuromuscular system used to stabilize upright posture in humans is a nonlinear dynamical system with time delays. The analysis of this system is important for improving balance and for early diagnosis of neuromuscular disease. In this work, we study the dynamic coupling between the neuromuscular system and a balance board—an unstable platform often used to improve balance in young athletes, and older or neurologically impaired patients. Using a simple inverted pendulum model of human posture on a balance board, we describe a surprisingly broad range of divergent and oscillatory CoP/CoM responses associated with instabilities of the upright equilibrium. The analysis predicts that a variety of sudden changes in the stability of upright postural equilibrium occurs with slow continuous deterioration in balance board stiffness, neuromuscular gain, and time delay associated with the changes in proprioceptive/vestibular/visual-neuromuscular feedback. The analysis also provides deeper insight into changes in the control of posture that enable stable upright posture on otherwise unstable platforms.     

The vestibular cortical projection during spinal decompensation

The removal of spinal afferents in rabbits submitted to unilateral lesion of the labyrinth and fully compensated greatly modified the field potentials as well as the single unit responses recorded from the cortical vestibular area during stimulation of the intact eighth nerve. In particular, an increase of contralateral and a decrease of ipsilateral potentials were observed. These asymmetrical responses indicate that the influence of the spinal cord in compensating the effects of unilateral lesion of the labyrinth is directed to balance not only the electrical activity of the brain-stem structures, but also that of the cerebral cortex. It appears, therefore, that spinal signals intervene not only in the compensation of vestibulospinal and vestibulo-oculomotor functions but also of cortical functions, such as that related to vestibular sensation.