Mutualisms are often viewed as reciprocal exploitations that nonetheless provide net benefits to each partner. While the benefits of symbiosis with dinoflagellates of the genus Symbio- dinium (zooxanthellae) for corals are well understood, the costs of the association, particularly when under stress, remain a focus of much research. One of the primary impediments to exploring the costs of symbiosis in zooxanthellate corals is that it is impossible to examine the animal host in isolation. Evidence for a cost of symbiosis with zooxanthellae includes the fact that direct transmission of zoox- anthellae between generations is rare, particularly in broadcast spawning corals. Fortuitously, the absence of zooxanthellae in oocytes of many species and the ability to readily infect larvae with zoox- anthellae provide an opportunity to compare individuals with and without these symbionts. Here, we use this larval model to show that individuals with zooxanthellae have lower survival than those lack- ing zooxanthellae when exposed to high temperature. Higher activity of anti-oxidant defenses and higher levels of oxidative cellular damage in larvae with zooxanthellae suggest that oxidative stress originating in the symbionts is a cause of tissue damage in the host under heat stress. We hypothesize that this may be one reason for the absence of direct transmission of zooxanthellae in most broadcast spawning corals whose propagules must spend at least 1 d on the ocean surface.

Coral reefs are declining worldwide due to increased incidence of climate-induced coral bleaching, which will have widespread biodiversity and economic impacts. A simple method to measure the sub-bleaching level of heat-light stress experienced by corals would greatly inform reef management practices by making it possible to assess the distribution of bleaching risks among individual reef sites. Gene expression analysis based on quantitative PCR (qPCR) can be used as a diagnostic tool to determine coral condition in situ. We evaluated the expression of 13 candidate genes during heat-light stress in a common Caribbean coral Porites astreoides, and observed strong and consistent changes in gene expression in two independent experiments. Furthermore, we found that the apparent return to baseline expression levels during a recovery phase was rapid, despite visible signs of colony bleaching. We show that the response to acute heat-light stress in P. astreoides can be monitored by measuring the difference in expression of only two genes: Hsp16 and actin. We demonstrate that this assay discriminates between corals sampled from two field sites experiencing different temperatures. We also show that the assay is applicable to an Indo-Pacific congener, P. lobata, and therefore could potentially be used to diagnose acute heat-light stress on coral reefs worldwide.