The Coral Ecosystem o Corals support 25-33% of all marine species The ocean is relatively nutrient-poor; corals create

e nutrient hotspots Shelter Food Support for cyanobacteria, which create nitrogen (essential for most organisms survival) o Mutualism with algae The oxygen and sugars produced from algal photosynthesis are used by corals Algae provide corals with more than 50% of their nutrients and give them their colors Conversely, corals produce nitrogen and carbon dioxide, which both enhance algal growth o Trophic structure If some event removed corals or their algae from the ecosystem, it would decimate the populations of dependent species (removal of the main primary producer) Death of birds who eat coral fish Impacts can even reach terrestrial species Reduced protein availability for developing nations who are very dependent on fish as a food source

Coral Bleaching & Climate Change o Bleaching = death of zooxanthellae Zooxanthellae = unicellular organisms present in large quantities in marine invertebrates; for corals, most are algae Occurs naturally in seasonal phases, but if bleaching episodes are prolonged, they can result in widespread coral death o Correlation with temperature 70% of all coral bleaching reports were strongly associated with warmerthan-normal ocean waters (at least 1C), while higher temperatures consistently raised the lethality of bleaching As temperatures have increased by about 1-2C over the past century, the number of recorded mass bleaching episodes has skyrocketed The 2-5C temperature increases expected over the next century will inevitably result in much more coral bleaching

Microbes & Mediterranean Bleaching o V. shiloi & O. patagonica Mediterranean coral, i.e. temperate regions Therefore, the water experiences a 14C gradient in temperature annually Periodic episodes of bleaching, usually in the summer at the moment, short enough to allow complete or near-complete recovery of the reefs o Rosenberg et al. study Series of studies over the past 2 decades Rather than inherently being a result of temperature stress, its instead the elevated temperatures that better allow the bacteria to prosper In fact, not only was V. shiloi absent in uninfected specimens, but elevated temperatures also failed to cause significant bleaching of O. patagonica in laboratory settings if the pathogen was absent (zooxanthellae unharmed) o Patchy distribution If it was simply environmental stressors, one would expect the bleaching to occur much more uniformly However, the spread of contagious microbial disease typically occurs in such a haphazard fashion, due to some individuals immunity to that pathogen and the inherently random nature of its spread o Preceding every bleaching episode, the number of V. shiloi grew dramatically, quickly succeeded by a drop in the number of photosynthesizing zooxanthellae Suggests that the presence of the bacterium is a cause of the bleaching, rather than an effect

New Evidence: Tropical Reefs o Another member of the Vibrio genus, V. coralliilyticus, has been found to be responsible for infecting the tropical coral Pocillopora damicornis o Moreover, this another variety of this same bacteria has also infected large colonies of the Acropora millepora coral in the Great Barrier Reef off the coast of Townsville o Although so far only bleaching in only a few species of coral has been attributed to pathogens, this is simply due to the fact that the hypothesis is relatively new o Strengthening it, however, are the results in the table In corals infected with Vibrio bacteria, tissue was bleached and broken down in increasing amounts as temperature increased At 20C, the corals remained completely healthy At 24.5C, two thirds of the corals were bleached within 15 days, and almost all of them were bleached within 20 days At 29C, almost all of the corals were broken down within 10 days, with the last remnants of healthy coral succumbing within another 5 However, at 29C with no bacteria present, the coral experienced no bleaching whatsoever o This suggests that elevated temperatures do not inherently cause coral bleaching, as previous literature has suggested, but simply allow the bacteria to act more effectively

Biological Mechanisms o The marine fireworm is the primary vector for the spread of most Vibrio bacteria species, but once a few individuals in a community are infected, the bacteria can spread from coral to coral with no vector o Intracellular multiplication Attaches to the coral by chemically bonding itself to receptors in the mucus of the corals surface Once attached, the bacteria multiply within the coral, all the while producing the toxins responsible for bleaching o Superoxide dismutase Superoxide is the anion O2- and is a products of the zooxanthellaes photosynthesis (very high concentrations) Typically, it the coral converts superoxide to H2O and O2 via various enzymes, but this process typically occurs at the level of individual coral cells during respiration Superoxide is highly toxic to the infectious bacteria, so they need to produce SOD (which converts superoxide into harmless molecules) in order to reduce its high concentrations to survivable levels o Peptide toxin (Toxin P) and ammonia On its own, Toxin P had negligible effects on the zooxanthellae Meanwhile, ammonia alone stopped about 18% of photosynthesis Interrupts the pH gradient within the chloroplast (more basic on the outside than the inside) Because this gradient is an important source of energy used in photosynthesis, the more the two sides are equalized, the less energy is available for photosynthesis However, when the two were combined, the bacteria successfully shut down over 64% of algal photosynthesis Reduces the formation of further superoxide, prolonging bacterial survival and reducing energy expenditure to create SOD Stopping photosynthesis = bleaching (algae die because they dont produce energy for themselves, corals die because they dont get energy from the algae) When both ammonia and Toxin P were produced, there was a much more significant drop in the pH of the water than when just ammonia was used Suggests that the peptide toxin allows more basic ammonia to exit the water and enter the photosynthesizing algae, disrupting the pH gradient even more o Temperature dependence At temperatures below 20C, the bacteria cannot produce SOD and Toxin P, or even form the chemical bond needed to attach to the coral Optimal temperature range is above 26C Unhindered, climate change will only exacerbate the problem by increasing the length of the periods during which Vibrio species can contribute to coral bleaching, with catastrophic ecological repercussions globally

Phage Therapy o Probiotics are microbes introduced externally into an organism; they provide some sort of health benefit, from strengthening the immune system to improving digestive health o Some researchers have applied the idea of probiotics, which is prevalent in human and veterinary medicine, as well as agriculture, to coral bleaching o Specifically, they have experimented with using bacteriophages, viruses that target bacteria, in treating coral bleaching While a bacteriophage has not yet been found for V. shiloi in the Mediterranean, the virus YB2 has been found to target the tropical V. coralliilyticus o Safety and practicality of use Most of the corals that remain healthy at high temperatures in the presence of the bacteria actually contain populations of the YB2 phage Moreover, the YB2 phage only targets V. coralliilyticus These 2 factors mean that artificially introduced YB2 would not harm the ecosystem, since it does not affect other coral microbes and is natural Although it needs to be introduced about every 3 months, only about 13 gallons are needed to treat 30 miles of coral; YB2 is also cheap to produce Gene Transfer Agents o Gene transfer agents, or GTAs, are virus-like particles that transfer random strands of DNA between cells (typically unicellular organisms) Usually, this is done among bacteria in nature to improve genetic diversity o A number of researchers have proposed to use GTAs to inject pre-set genetic codes into elements of the coral community involved in bleaching One possibility is to use GTAs to inhibit superoxide dismutase and/or ammonia production in the Vibrio bacteria They need superoxide dismutase to survive Ammonia is the main reagent inhibits photosynthesis Another possible use would be to disable the receptors in the corals mucus to which Vibrio attach themselves however, other repercussions of this are unknown Finally, in the Red Sea, a number of corals incredibly resistant to bleaching have been found; this resilience can be attributed to their zooxanthellae and is almost certainly genetic. Thus, these genes could be transferred by GTAs to improve the resilience of more vulnerable corals. o Advantages Because the pathogens are not being harmed directly, they are less likely to adapt to become resistant, especially considering that they themselves use GTAs to evolve While phages may need to be introduced every year, using GTAs would likelier make more lasting changes to the ecosystem o Disadvantages Artificial GTAs could mutate to make the pathogens more virulent or transfer genes boosting pathogens resistance to corals native antibiotics