Precise measurements were conducted in continuous flow seawater mesocosms located in full sunlight that compared metabolic response of coral, coral-macroalgae and macroalgae systems over a diurnal cycle. Irradiance controlled net photosynthesis (P net), which in turn drove net calcification (G net), and altered pH. P net exerted the dominant control on [CO3 (2-)] and aragonite saturation state (Ωarag) over the diel cycle. Dark calcification rate decreased after sunset, reaching zero near midnight followed by an increasing rate that peaked at 03:00 h. Changes in Ωarag and pH lagged behind G net throughout the daily cycle by two or more hours. The flux rate P net was the primary driver of calcification. Daytime coral metabolism rapidly removes dissolved inorganic carbon (DIC) from the bulk seawater and photosynthesis provides the energy that drives G net while increasing the bulk water pH. These relationships result in a correlation between G net and Ωarag, with Ωarag as the dependent variable. High rates of H(+) efflux continued for several hours following mid-day peak G net suggesting that corals have difficulty in shedding waste protons as described by the Proton Flux Hypothesis. DIC flux (uptake) followed P net and G net and dropped off rapidly following peak P net and peak G net indicating that corals can cope more effectively with the problem of limited DIC supply compared to the problem of eliminating H(+). Over a 24 h period the plot of total alkalinity (AT ) versus DIC as well as the plot of G net versus Ωarag revealed a circular hysteresis pattern over the diel cycle in the coral and coral-algae mesocosms, but not the macroalgae mesocosm. Presence of macroalgae did not change G net of the corals, but altered the relationship between Ωarag and G net. Predictive models of how future global changes will effect coral growth that are based on oceanic Ωarag must include the influence of future localized P net on G net and changes in rate of reef carbonate dissolution. The correlation between Ωarag and G net over the diel cycle is simply the response of the CO2-carbonate system to increased pH as photosynthesis shifts the equilibria and increases the [CO3 (2-)] relative to the other DIC components of [HCO3 (-)] and [CO2]. Therefore Ωarag closely tracked pH as an effect of changes in P net, which also drove changes in G net. Measurements of DIC flux and H(+) flux are far more useful than concentrations in describing coral metabolism dynamics. Coral reefs are systems that exist in constant disequilibrium with the water column.
Keywords: Algae; Aragnite saturation state; Boundary layers; Calcification; Coral; Coral reef; Phase lag; Photosynthesis; Proton flux.