Plant responses to hypergravity: a comprehensive review

Hypergravity is an effective novel stimulus to elucidate plant gravitational and mechanobiological behaviour. Here, we review the current understanding of phenotypic, physio-biochemical, and molecular plant responses to simulated hypergravity. Plants readily respond to altered gravity conditions, such as microgravity or hypergravity. Hypergravity-a gravitational force higher than that on the Earth's surface (> 1g)-can be simulated using centrifuges. Exposing seeds, seedlings, or plant cell cultures to hypergravity elicits characteristic morphological, physio-biochemical, and molecular changes. While several studies have provided insights into plant responses and underlying mechanisms, much is still elusive, including the interplay of hypergravity with gravitropism. Moreover, hypergravity is of great significance for mechano- and space/gravitational biologists to elucidate fundamental plant behaviour. In this review, we provide an overview of the phenotypic, physiological, biochemical, and molecular responses of plants to hypergravity. We then discuss the involvement of hypergravity in plant gravitropism-the directional growth along the gravity vector. Finally, we highlight future research directions to expand our understanding of hypergravity in plant biology.