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Biomedical subjects

R Wassersug

Publications and source records attributed to R Wassersug.

At least 19 recordsLinked to original sources

Effects of spaceflight and cage design on abdominal muscles of male rodents.

We examined the effects of a 16-day spaceflight mission on the size of muscle fibers in the rectus abdominis, external oblique and transversus abdominis muscles of adult male Fisher rats. The rats were individually housed in orbit, in contrast to the one previous spaceflight investigation of the same muscles, where the rats were group-housed pregnant females. The cross-sectional area of the muscle fibers was used as a measure of muscle atrophy or hypertrophy. The transversus, which is presumed to be the primary expiratory muscle and consequently works against internal hydrostatic pressures that are not likely to change much between 1 G and weightlessness, did not change in size. However, both the rectus abdominis (a spinal flexor) and the external oblique (a rotator of the torso), which resist gravity in the 1 G environment, showed significant signs of atrophy after extended exposure to microgravity. The atrophy of the external oblique was diametrically opposite to hypertrophy of the same muscle observed in group-housed rodents previously exposed to spaceflight. Although the two missions differed in several factors, such as the gender of the rats and mission duration, we believe that housing of the animals was the key factor that accounted for the different responses of the external oblique. Previous research has shown that group-housed rats in spaceflight exhibited seven times more rotations of their torsos than matched ground controls. Thus unloading of the musculoskeletal system may not be achieved in weightlessness when animals have the freedom to interact with each other.

Abdominal Muscles↗

Effects of laparotomy, cage type, gestation period and spaceflight on abdominal muscles of pregnant rodents.

We studied the effects of four variables on the histological properties of three body wall muscles-rectus abdominis (RA), transversus abdominis (TA), and external oblique (EO)-from pregnant rats. The variables examined were (1) gestation period; (2) cage design; (3) the effect of a midline laparotomy, performed to determine fetus numbers; and (4) exposure to a nine-day spaceflight. We measured fiber cross-sectional area (CSA), metabolic enzyme levels (succinate dehydrogenase, glycerophosphate dehydrogenase), and myosin heavy chain (MHC) immunoreactivity in samples from each muscle. A major effect of spaceflight was an increase of 42-171% in fibers double-labeled for MHC in all three muscles. Based on fiber CSA, the TA and RA muscles showed signs of stretching with increased gestation; i.e., the CSA decreased 11-12% over a nine-day period. The EO, a torso rotator, hypertrophied by 9% in rats group-housed in cages with a complex 3-D structure, compared to controls housed singly in standard flat-bottom cages. The TA and EO, whose contractions would pull on the suture line, showed signs of atrophy in laparotomized animals, exhibiting a 12% decrease in muscle fiber CSA. Exposure to weightlessness is known to induce atrophy in most skeletal muscles. Surprisingly, the EO actually hypertrophied 11% in our flight animals; however, this can be explained by the fact that those rats actively rotated their torsos seven times more often than ground controls. The flight rats also had twice as many contractions as controls. However, they were still able to give birth on time postflight.

Abdominal Muscles↗

Swimming kinematics and respiratory behaviour of xenopus laevis larvae raised in altered gravity

We examined the respiratory behaviours and swimming kinematics of Xenopus laevis tadpoles hatched in microgravity (Space Shuttle), simulated microgravity (clinostat) and hypergravity (3 g centrifuge). All observations were made in the normal 1 g environment. Previous research has shown that X. laevis raised in microgravity exhibit abnormalities in their lungs and vestibular system upon return to 1 g. The tadpoles raised in true microgravity exhibited a significantly lower tailbeat frequency than onboard 1 g centrifuge controls on the day of landing (day0), but this behaviour normalized within 9 days. The two groups did not differ significantly in buccal pumping rates. Altered buoyancy in the space-flight microgravity tadpoles was indicated by an increased swimming angle on the day after landing (day1). Tadpoles raised in simulated microgravity differed to a greater extent in swimming behaviours from their 1 g controls. The tadpoles raised in hypergravity showed no substantive effects on the development of swimming or respiratory behaviours, except swimming angle. Together, these results show that microgravity has a transient effect on the development of locomotion in X. laevis tadpoles, most notably on swimming angle, indicative of stunted lung development. On the basis of the behaviours we studied, there is no indication of neuromuscular retardation in amphibians associated with embryogenesis in microgravity.

Journal Article↗

Swimming kinematics and respiratory behaviour of Xenopus laevis larvae raised in altered gravity.

We examined the respiratory behaviours and swimming kinematics of Xenopus laevis tadpoles hatched in microgravity (Space Shuttle), simulated microgravity (clinostat) and hypergravity (3 g centrifuge). All observations were made in the normal 1 g environment. Previous research has shown that X. laevis raised in microgravity exhibit abnormalities in their lungs and vestibular system upon return to 1 g. The tadpoles raised in true microgravity exhibited a significantly lower tailbeat frequency than onboard 1 g centrifuge controls on the day of landing (day0), but this behaviour normalized within 9 days. The two groups did not differ significantly in buccal pumping rates. Altered buoyancy in the space-flight microgravity tadpoles was indicated by an increased swimming angle on the day after landing (day1). Tadpoles raised in simulated microgravity differed to a greater extent in swimming behaviours from their 1 g controls. The tadpoles raised in hypergravity showed no substantive effects on the development of swimming or respiratory behaviours, except swimming angle. Together, these results show that microgravity has a transient effect on the development of locomotion in X. laevis tadpoles, most notably on swimming angle, indicative of stunted lung development. On the basis of the behaviours we studied, there is no indication of neuromuscular retardation in amphibians associated with embryogenesis in microgravity.

Animals↗

The three-dimensional hydrodynamics of tadpole locomotion.

Tadpoles are unusual among vertebrates in having a globose body with a laterally compressed tail abruptly appended to it. Compared with most teleost fishes, tadpoles swim awkwardly, with waves of relatively high amplitude at both the snout and tail tip. In the present study, we analyze tadpole propulsion using a three-dimensional (3D) computational fluid dynamic (CFD) model of undulatory locomotion that simulates viscous and unsteady flow around an oscillating body of arbitrary 3D geometry. We first confirm results from a previous two-dimensional (2D) study, which suggested that the characteristic shape of tadpoles was closely matched to their unusual kinematics. Specifically, our 3D results reveal that the shape and kinematics of tadpoles collectively produce a small 'dead water' zone between the head-body and tail during swimming precisely where tadpoles can and do grow hind limbs--without those limbs obstructing flow. We next use our CFD model to show that 3D hydrodynamic effects (cross flows) are largely constrained to a small region along the edge of the tail fin. Although this 3D study confirms most of the results of the 2D study, it shows that propulsive (Froude) efficiency for tadpoles is overall lower than predicted from a 2D analysis. This low efficiency is not, however, a result of the high-amplitude undulations of the tadpole. This was demonstrated by forcing our 'virtual' tadpole to swim with fish-like kinematics, i.e. with lower-amplitude propulsive waves. That particular simulation yielded a much lower Froude efficiency, confirming that the large-amplitude lateral oscillations of the tadpole do, indeed, provide positive thrust. This, we believe, is the first time that the unsteady flow generated by an undulating vertebrate has been realistically modelled in three dimensions. Our study demonstrates the feasibility of using 3D CFD methods to model the locomotion of other undulatory organisms.

Animals↗

Regulative development of Xenopus laevis in microgravity.

To test whether gravity is required for normal amphibian development, Xenopus laevis females were induced to ovulate aboard the orbiting Space Shuttle. Eggs were fertilized in vitro, and although early embryonic stages showed some abnormalities, the embryos were able to regulate and produce nearly normal larvae. These results demonstrate for the first time that a vertebrate can ovulate in the virtual absence of gravity, and that the eggs can develop to a free-living stage.

Animals↗

A computational fluid dynamics study of tadpole swimming

The hydrodynamics and undulating propulsion of tadpoles were studied using a newly developed two-dimensional computational fluid dynamics (CFD) modeling method. The mechanism of thrust generation associated with the flow patterns during swimming is discussed. Our CFD analysis shows that the kinematics of tadpoles is specifically matched to their special shape and produces a jet-stream propulsion with high propulsive efficiency, as high as that achieved by teleost fishes. Investigation of the effect of Reynolds number indicates that the Froude efficiency increases with increasing Reynolds number with no ceiling in generating the jet-stream propulsion. Further studies using tadpole- and fish-shaped models with hindlimbs added to their body profiles reveal that the tadpole shape ­ a globose head with a tapered tail and hindlimbs at the base of the tail ­ allows tadpoles, but not fish, to develop hindlimbs with very little handicap on propulsion. The shapes and kinematics of tadpoles appear to be specially adapted to the requirement of these organisms to transform into frogs.

Journal Article↗

Effects of space flight on Xenopus laevis larval development.

Fifty-three fertilized Xenopus laevis embryos at early tail bud stage were launched into orbit aboard a Biocosmos satellite and remained in microgravity for 11.5 days. During this period, the embryos hatched and continued to develop as free-living larvae. Forty-eight individuals survived the mission. Upon recovery these tadpoles had smaller heads/bodies and proportionately longer tails than ground controls. Almost all the flight animals had caudal lordosis and consequently swam in backward somersaults. Compared to ground-based controls, their notochords were significantly larger in cross-sectional area and were deformed. Caudal muscle fibers were less dense and involuted in a fashion indicative of degeneration. In contrast, cranial muscles associated with buccal pumping did not differ between the flight and control animals. Upon landing, the flight larvae were found to be negatively buoyant and lay on the bottom when they were not swimming. They had significantly smaller lungs than controls, suggesting that they had failed to inflate their lungs in microgravity. Additionally, the branchial baskets, gill filters and thymuses all showed signs of retarded development or degeneration. The caudal deformity that we observed in the flight X. laevis has been independently observed in three other space flight experiments where embryos were launched then hatched in space. In contrast, Xenopus larvae from another orbital experiment that were raised from fertilization through hatching in space did not exhibit any caudal abnormalities. These divergent results suggest that either features of the launch itself (i.e., high acceleration and vibration) or an abrupt decrease in gravity during the tail bud stage detrimentally affects musculoskeletal development in anurans.

Animals↗

Behavioral reactions of the bat Carollia perspicillata to abrupt changes in gravity.

As part of an ongoing survey of the behavioral responses of vertebrates to abrupt changes in gravity, we report here on the reactions of bats (Carollia perspicillata) exposed to altered gravity during parabolic aircraft flight. In microgravity, mammals typically behave as if they were upside-down and exhibit repetitive righting reflexes, which often lead to long axis rolling. Since bats, however, normally rest upside-down, we hypothesized that they would not roll in microgravity. Only one of three specimens attempted to fly during microgravity. None rolled or performed any righting maneuvers. During periods of microgravity the bats partially extended their forearms but kept their wings folded and parallel to the body. Between parabolas and occasionally during microgravity the bats groomed themselves. Both the extended limbs and autogrooming may be stress responses to the novel stimulus of altered gravity. This is the first behavioral record of Chiroptera in microgravity.

Animals↗

Should the Space Station be an ark?

This essay explores the pros and cons of maximizing the number of species that can be maintained on the Space Station. It reviews some of the history of comparative space biology to show that different cultures have different perspectives on the study of non-traditional research organisms (ie non-rodents) in space. Despite these differences, there are simple principles that all international partners in the Space Station endeavour should be able to uphold when deciding what facilities to build and what species to fly. As an argument for maximizing the taxonomic diversity on the Space Station, examples are given to show how very similar organisms may have different reactions to microgravity. At the same time the political pressure in the USA to make the Space Station an institution specifically servicing the 'health, well-being and economic benefits of people on earth', is acknowledged. Ultimately the justification for what species will be on the Space Station should rest with the quality of the scientific questions being asked.

Adaptation, Physiological↗

Lung use and development in Xenopus laevis tadpoles.

Shortly after hatching, Xenopus laevis tadpoles fill their lungs with air. We examined the role played by early lung use in these organisms, since they are able to respire with both their lungs and their gills. We investigated the effect on X. laevis development when the larvae were prevented from inflating their lungs, and whether early lung use influenced the size of the lungs or the tadpole's ability to metamorphose. Tadpoles that were denied access to air had lungs one-half the size of those of controls. This difference in lung size was too large to be explained merely by a stretching of the lung due to inflation. The longer tadpoles were denied access to air, the longer they took to metamorphose, and their probability of completing metamorphosis diminished. One tadpole raise throughout its larval life without access to air successfully metamorphosed but had abnormal, solidified lungs and an enlarged heart. Collectively, these experiments demonstrate that early lung use in tadpoles is important in determining both ultimate lung size and the probability of successfully metamorphosing. Lung use during early larval development in X. laevis is not absolutely necessary for survival through metamorphosis, but its absence severely handicaps growth.

Air↗

The behavioral reactions of a snake and a turtle to abrupt decreases in gravity.

We report here on the behavioral reaction of two reptiles to abrupt decreases in gravity. One striped rat snake, Elaphe quadrivirgata, and three striped-neck pond turtles, Mauremys japonica, were exposed to microgravity on parabolic flight, during the filming of a documentary for the NHK television station in Japan. The video films revealed that the snake reflexively responded to the shift from hyper- to hypogravity by taking up a defensive posture--on the first parabola, the snake struck at itself. The turtles actively extended their limbs and hyper-extended their neck in microgravity, a posture which is identical to the displayed during their contact "righting reflex", when placed upside-down in normal gravity. The aggressive display of the snake was unexpected, although the righting response of the turtles was consistent with that shown by other vertebrates, including fish and mammals, exposed to microgravity. An implication of these observations is that the afferent signal for the righting reflex of vertebrates in normal gravity must be the unloading of ventral receptors in the sensory system, rather than the loading of dorsal receptors. These are the first behavioral records for any reptiles exposed to hypogravity.

Aggression↗

Evolution of spinal nerve number in anuran larvae.

We examined the number of segmental spinal nerves in 60 premetamorphic tadpoles, representing 43 species, 33 genera and 12 families of anurans. The number of spinal nerves shows a clear phylogenetic pattern, in which the primitive condition is variable, ranging from 23 to 29 pairs. Reduction of caudal spinal nerve number has occurred independently at least 7 times during anuran phylogeny. Reduction events have occurred among species within genera, among subfamilies within families, as well as at familial and higher taxonomic levels. In general, arachaeobatrachian larvae have significantly more spinal nerves than neobatrachian larvae. While 2 of the 7 reduction events have occurred within the Archaeobatrachia, both of these have occurred at low taxonomic levels. In the Neobatrachia, 3 of the 5 reductions have occurred at subfamilial or higher taxonomic levels. Overall, reduced spinal nerve number correlates with tadpole body size, but not with relative tail length or with developmental stage, at least within the range of stages we examined. There is a positive correlation between number of spinal nerves and minimum time from hatching to metamorphosis, at least within the Archaeobatrachia. Species with more pairs of nerves take longer to reach metamorphosis, possibly because they have more neural tissue to produce and to resorb. Spinal nerve number per se does not appear to be associated with larval microhabitat ecology or swimming behavior. Tadpoles exhibit greater variation in spinal nerve number than adult frogs. This greater variation is tolerated presumably because the tail is an expendable organ, which is lost at metamorphosis.

Animals↗

Morphology of the caudal spinal cord in Rana (Ranidae) and Xenopus (Pipidae) tadpoles.

Using a variety of neuroanatomical and histological techniques, we compare the spinal cord and peripheral nerve distribution in the tails of larvae from Xenopus laevis and three species of Rana. The relatively large, postsacral spinal cord of Xenopus contains abundant motoneurons and their axons. Spinal nerves exit from the spinal cord in a regular array, one nerve per myotome, from the cervical region to near the end of the tail. Somata of motoneurons innervating caudal myotomes are found along the entire length of the tail. In contrast, the caudal cord of Rana is reduced to a filum terminale consisting of little more than an ependymal tube; spinal nerves to all caudal myotomes leave the cord in the sacral region and reach their motor targets via a cauda equina and caudal plexus. Motoneuron cell bodies innervating caudal myotomes are found only in the sacral region. The Rana larval pattern is similar to that of adult frogs and mammals, whereas the Xenopus larval pattern is more like that of salamanders and reptiles. These gross neuroanatomical differences are not due to differences in the size or developmental stage of the tadpoles, but instead are associated with differences in the swimming behavior of the larvae. The presence of motoneurons in the caudal spinal cord of Xenopus may provide local intermyotomal control within the tail; the elongated topography of the cord appears to permit finer, rostral-to-caudal regulation of neuromuscular activity. The Rana spinal cord, on the other hand--with motoneurons clustered anteriorly--may produce concurrent firing of adjacent ipsilateral myotomes, but at the expense of fine intermyotomal regulation. The fact that nerves in the tail of Xenopus enter and exit from the spinal cord locally, as opposed to far anteriorly as in Rana, means that for tadpoles of the same size, reflex arc lengths are many times shorter in Xenopus.

Animals↗