Coral bleaching : photosynthetic impacts on symbiotic dinoflagellates
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Global climate change is leading to the rise of ocean temperatures and is triggering mass coral bleaching events on reefs around the world. This involves the expulsion of the symbiotic dinoflagellate algae, known as zooxanthellae, from the coral host. Coral bleaching is believed to occur as a result of damage to the photosynthetic apparatus of these symbionts, although the specific site of initial impact is yet to be conclusively resolved. This thesis examined a number of sites within the light reactions of photosynthesis and evaluated the efficiency of photoprotective heat dissipating pathways. Upon expulsion, the capacity for long-term survivorship of expelled zooxanthellae in the water column was also assessed. A reduction in photosystem II (PSII) photochemical efficiency during exposure to elevated temperature and high light (bleaching conditions) was found to be highly dependent upon the increase in abundance of QB non-reducing PSII centres (inactive PSII centres), indicating damage to the site of the secondary electron acceptor, QB, resulting in a limited capacity for its reduction. Therefore, this reduced the rate of the reoxidation of the primary electron acceptor, QA-. Fast induction curve (FIC) analysis of the rise from minimum fluorescence to maximum fluorescence revealed a lower amplitude in the J step along this curve, which was consistent with a reduction in the rate of QA reoxidation. This photoinhibition of PSII was found to occur once the effectiveness of excess energy dissipation through energy-dependent quenching and state-transition quenching was exceeded, suggesting that these mechanisms were incapable of preventing photodamage. Antenna size heterogeneity showed little change under bleaching conditions with a significant increase in PSIIbeta only apparent in one species of coral. The thermostability of the oxygen evolving complex (OEC) and thylakoid membrane were found to increase during exposure to bleaching conditions and exceeded bleaching thresholds of corals. This rapid rise in temperature-dependent thermostability also occurred over seasons, where variation in ocean temperatures was matched by gradual shifts in OEC and thylakoid membrane thermotolerance. Variation in thermostability between species was not found to be linked to zooxanthellae genotype, and instead was related to the bleaching susceptibility of the host. Despite this capacity for resilience to bleaching conditions, the PSII reaction centres did not exhibit such a mechanism for rapid acclimatisation. Corals can only be as tolerant to bleaching conditions as their most sensitive component allows. The formation of nonfunctional PSII centres is therefore suggested to be involved in the initial photochemical damage to zooxanthellae which leads to a bleaching response. Zooxanthellae were found to be expelled irrespective of OEC function and thylakoid membrane integrity, as these sites of the photosynthetic apparatus were still intact when cells were collected from the water column. Although zooxanthellae were photosynthetically competent and morphologically intact upon expulsion, their longevity in the water column was dependent on the time of expulsion following the onset of bleaching and the ambient water temperatures. The survivorship of these zooxanthellae was restricted to a maximum of 5 days in the water column which suggests that unless expelled zooxanthellae inhabit other environs of coral reefs which may be more favourable for survival, their capacity for persistence in the environment is extremely limited. Chlorophyll a fluorescence measurements are a common tool for investigating photosynthetic impacts to in hospite zooxanthellae of corals. Pathways causing dark-reduction of the plastoquinone pool are shown to be active in corals and affect measurements which require dark-adaptation. Pre-exposure to far-red light was found to be an effective procedure to oxidise the inter-system electron transport chain and ensure determination of the true maximum quantum yield of PSII and accurate FICs. It is concluded that the trigger for coral bleaching lies in the photosynthetic apparatus of zooxanthellae and evidence is presented in support of this impact site not being the OEC or thylakoid membrane.
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