The complex dynamic properties of biological timing in organisms remain a central enigma in biology despite the increasingly precise genetic characterization of oscillating units and their components. Although attempts to obtain the time constants from oscillations of gene activity and biochemical units have led to substantial progress, we are still far from a full molecular understanding of endogenous rhythmicity and the physiological manifestations of biological clocks. Applications of nonlinear dynamics have revolutionized thinking in physics and in biomedical and life sciences research, and spatiotemporal considerations are now advancing our understanding of development and rhythmicity. Here we show that the well known circadian rhythm of a metabolic cycle in a higher plant, namely the crassulacean acid metabolism mode of photosynthesis, is expressed as dynamic patterns of independently initiated variations in photosynthetic efficiency ( PSII) over a single leaf. Noninvasive highly sensitive chlorophyll fluorescence imaging reveals randomly initiated patches of varying PSII that are propagated within minutes to hours in wave fronts, forming dynamically expanding and contracting clusters and clearly dephased regions of PSII. Thus, this biological clock is a spatiotemporal product of many weakly coupled individual oscillators, defined by the metabolic constraints of crassulacean acid metabolism. The oscillators operate independently in space and time as a consequence of the dynamics of metabolic pools and limitations of CO 2 diffusion between tightly packed cells.A current trend in physics and life sciences is the investigation of spatiotemporal patterns that appear in phenomena that were formerly thought to be solely time dependent. Rhythmic phenomena are common in nature and are excellent models for investigation of spatial dynamics because of their regular structure. However, the complex dynamic properties of timing in organisms and biological systems remain a central enigma in biology despite the increasingly precise genetic characterization of oscillating units and their components (1, 2). For example, phase synchronization of predator-prey dynamics in spatially extended ecological systems emerges only when the building blocks are spatially arranged and interrelated by flows of signaling compounds (3). The challenge is to understand how the spatial organization of the components of a system can influence overall temporal development.The circadian rhythm of net CO 2 exchange (JCO 2 ) in crassulacean acid metabolism (CAM) plants in continuous light is regarded as a time-dependent generic model system for exploration of endogenous rhythmicity in a well understood metabolic pathway (4-6). During the normal day͞night cycle, CAM can be divided in four phases (7). In phase I, nocturnal CO 2 fixation by phosphoenolpyruvate-carboxylase (PEPCase) leads to the formation of malic acid that is removed from the site of its formation in the cytoplasm by active accumulation to the central cell vacuole. Malic acid exerts ...