Neurophysiology of Developing Fish at Altered Gravity: Background—Facts–Perspectives

Abstract

During the entire evolution of life on Earth, the phylogenetic as well as the individual development of all organisms took place under constant gravity conditions, against which they achieved specific countermeasures for compensation and adaptation. On the one side, gravity represents a factor of physical restriction, which compelled the ancestors of all extant living beings to develop basic achievements to counter the gravitational force (e.g., elements of statics like any kind of skeleton--from actin to bone--to overcome gravity enforced size limits or to keep form). On the other side, already early forms of life possibly used gravity as an appropriate cue for orientation and postural control, since it is continuously present and has a fixed direction. Due to such a thorough adaptation to the Earthly gravity vector, both orientation behaviour as well as the ontogenetic development of animals is impaired, when they have to experience altered gravity (delta g; i.e., hyper- or microgravity). On this background, it is still an open question to which extent delta g affects the normal individual development, either on the systemic level of the whole organism or on the level of individual organs or even single cells. The present review provides information on these questions, focusing on developing fish as model systems. Special emphasis is being laid on the effect of delta g on the developing brain and vestibular system, comprising investigations on behaviour and plastic reactivities of the brain and inner ear. Moreover, clues and insights into the possible basic causes of space motion sickness-phenomena (SMS; a kinetosis) are provided. Overall, the results speak in favour of the following concept: short-term altered gravity (< or = 1 day) can induce transitional aberrant behaviour due to malfunctions of the inner ear, originating from asymmetric otoliths or, generally, from a mismatch between canal and otolith afferents. The vanishing aberrant behaviour is due to a reweighing of sensory inputs and neurovestibular compensation, probably on bioelectrical basis. During long-term altered gravity (several days and more), step by step neuroplastic reactivities on molecular basis (i.e., molecular facilitation) in the brain and inner ears obviously activate feedback mechanisms between the CNS and the vestibular organs for the regain of normal behaviour. Mainly, the following areas of research with animals at altered gravity need to be addressed in the future: (1) Maintenance of animals through two complete life cycles in the space environment (developmental deficiencies?). (2) Investigation of the peripheral and central vestibular system by ground-based studies (mutants, hypergravity experiments...), focusing on plasticity in developing animals as well as in adults. (3) Investigation of the effect of microgravity during critical developmental periods (imprinting phase for graviperception?). Answers to these questions may be of crucial interest for basic gravitational research.