Abstract

Background Investigations into the diurnal ascent and descent of leaves of beans and other species, as well as experimental interventions into these movements, such as exposures to light at different times during the movement cycle, led to the concept of an endogenous 'clock' as a regulator of these oscillations. The causal origin of leaf movement can be traced to processes that modulate cell volume in target tissues of the pulvinus and petiole. However, these elements of the leaf-movement process do not sufficiently account for the rhythms that are generated following germination in constant light or dark conditions, or when plants are transferred to similar free-running conditions. Scope To further unravel the regulation of leaf-movement rhythms, many of the published time courses of leaf movements that provided evidence for the concept of the endogenous clock were analysed in conjunction with the contemporaneous time courses of the lunisolar tidal acceleration. This was accomplished by application of the Etide program, which estimates, with high temporal resolution, local gravitational changes as a consequence of the diurnal variations of the lunisolar gravitational force due to the orbits and relative positions of Earth, Moon and Sun. To substantiate the results obtained in earthbound laboratories additional experiments were performed in the International Space Station (ISS). Tidal recurrence within the ISS exhibited a periodicity of 45 min. In all instances investigated, it was evident that a synchronism exists between the times of the turning points of both the lunisolar tide and of the leaftide when the direction of leaf movement changes. This finding of synchrony documents that the lunisolar tide is a regulator of the leaftide, and that the rhythm of leaf movement is not of endogenous origin but is an expression of an exogenous lunisolar clock impressed upon the leaf-movement apparatus. Conclusions A huge number of correlations between leaftide and Etide time courses were established for leaf movement rhythms in natural conditions of the greenhouse, in conditions of constant light or dark, and under the microgravity conditions of the International Space Station. Even the apparently spontaneous short-period, small-amplitude rhythms recorded from leaves under unusual growth conditions are consistent with the hypothesis of a lunisolar zeitgeber. Synchronism between leaftide and Etide is discussed in terms of classical and quantum mechanics.

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