6. Boron Isotopic Systematics in Scleractinian Corals and the Role of pH Up-regulation

The boron isotopic composition (δ¹¹B) of scleractinian corals has been used to track changes in seawater pH and more recently as a probe into the processes controlling bio-calcification. For corals that precipitate aragonite skeletons, up-regulation of pH appears to be a general characteristic, typically being ~0.3 to ~0.6 pH units higher than ambient seawater. The relationship between the pH of the corals calcifying-fluid (pH_cf) and seawater pH_T (total scale) is shown to be dependent on both physiological as well environmental factors. In laboratory experiments conducted on symbiont-bearing (zooxanthellate) corals under conditions of constant temperature and seawater pH, changes in the δ¹¹B derived calcifying fluid pH_cf is typically ¹/₃ to ¹/₂ of that of ambient seawater. Similar linear relationships are found for cold water corals that live in relatively stable, cold, deep-water environments but at significantly elevated levels of pH_cf (~0.5–1 pH units above seawater), a likely response to the lower pH of their deep-sea environments. In contrast, zooxanthellae-bearing corals living in shallow-water reef environments that experience significant natural variations in temperature, light, nutrients and seawater pH, show different types of responses. For example, over seasonal time-scales Porites corals from the Great Barrier Reef (GBR) have a large range in pH_cf of ~8.3 to ~8.5, significantly greater (~×2 to ~×3) than that of reef-water (pH_T ~8.01 to ~8.08), and an order of magnitude greater than that expected from ‘static’ laboratory experiments. Strong physiological controls, but of a different character, are found in corals grown in a Free Ocean Carbon Enrichment Experiment (FOCE) conducted in situ within the Heron Island lagoon (GBR). These corals exhibit near constant pH_cf values regardless of external changes in temperature and seawater pH. This pattern of strong physiologically controlled ‘pH-homeostasis’, with elevated but constant pH_cf has been found despite large natural seasonal variations in the pH (±0.15 pH units) of the lagoon waters, as well as the even larger super-imposed decreases in seawater pH (~0.25 pH units) designed to simulate year 2100 conditions. In natural reef environments we thus find that the processes influencing the up-regulation of pH_cf in symbiont-bearing corals are subject to strong physiological controls, behaviour that is not well simulated in the current generation of aquaria-based experiments with fixed seawater pH and temperature. Conversely, cold-water corals that lack symbionts and inhabit the relatively stable deep-sea environments hold the best prospects for providing reliable reconstructions of seawater pH. Clearly, further studies utilising the δ¹¹B-pH_cf proxy combined with other DIC/carbonate-ion proxies (e.g. B/Ca), but conducted under realistic ‘natural’ conditions, are required to elucidate the processes controlling coral bio-calcification and to better understand the vulnerability of scleractinian corals to anthropogenic driven warming and ocean acidification.