Friday, 18 November 2016

Annual bleaching on the reefs: survival by acclimation has winners and losers

As the seawater surface temperature increases due to global warming, the existance of our coral reefs is at risk. The zooxanthellae that provide the carbon needed for the animal’s respiration,  disappear with increasing temperature and leave the coral in an ill, sometimes lethal state. Since the emission of greenhouse gasses isn’t to decrease in the near future, we can expect the loss of these microbes due to heat stress is to occur more frequently. For example, a study from Van Hooidonk et al. (2013) predicts annual bleaching events in some parts of the ocean by 2025.  In this blog the paper of Grottoli et al. (2014) is discussed, in which coral acclimation and recovery is studied in the context of annual bleaching. The hypothesis predicted survival and acclimation to annual bleaching. Acclimation was here defined as “the improved performance of the physiological variables measured here from being significantly lower than the control corals after bleaching to being no different from controls after bleaching the following year”.

The zooxanthellae, in the paper of Grottoli et al. (2013) more accurately called Symbiodinium (endosymbiotic dinoflagellates), make use of photosynthesis to provide the coral’s host with fixed carbon. However after bleaching, the animal gets more dependent on other sources of carbon, such as from energy reserves or increased heterotrophy. Also, the Symbiodinium community in a certain coral colony might change after bleaching, by a shifting distribution of Symbiodinium spp. These variables (photosynthesis, heterotrophy, energy reserves and Symbiodinium type distribution) are, amongst others, measured over a time span of two years. The non-control coral fragments were treated with an artificially imposed bleaching event in 2009 and again in 2010. This was executed by raising the water temperature approximately 1 degree over seven days in a seawater filled tank, after which the increased temperature was maintained for another eight and ten days in 2009 and 2010 respectively. Immediately after the bleaching treatment, the coral fragments were used to measure respiration, photosynthesis or ‘percent Contribution of Zooxanthellae to Animal Respiration’ (CZAR) and the feeding rate or ‘percent Contribution of Heterotrophy to Animal Respiration’ (CHAR) and placed back in the reef. After six weeks on the reef the following variables were measured or identified: Symbiodinium concentration, Symbiodinium type distribution and change in energy reserves.

The experiment was done with three different coral species: Porites divaricate, Porites astreoides, Orbicella faevolata. P. divarcata appeared to have the highest acclimation capacity, since the coral only showed a significant decline in calcification and Symbiodinium cell density after the first bleaching event in 2009. After six weeks these variables fully recovered and didn’t change after the following bleaching in 2010. The endosymbiont loss and decrease in calcification compared to controls after the first bleaching was higher for P. astreoides, but fully recovered too. However, after the bleaching treatment in 2010, the decrease of these parameters was even worse and persisted troughout the following six weeks. Like both Porites species, O. faveolata showed similar bleaching effects and recovery of them in the first year, but endosymbiont density and calcification did also decrease and fully recover in the following year. Interestingly, the energy reserve of O. faveolata didn’t change after bleaching in 2009, but did decrease after the treatment in 2010. The energy reserves for the Porites species decreased after both the first and second bleaching in the first six weeks back on the reef.  As can be expected with decreases in endosymbionts, all coral species showed a declined CZAR, albeit not always significant. For P. astreoides however, this loss in fixed carbon supply was compensated for after the first bleaching event by an increase of CHAR of 147%, although this trend wasn’t proven statistically significant. These results for the fixed carbon acquisition imply that heterotrophy doesn’t play a significant role in ability of corals to acclimate to annual bleaching events. A factor that was associated with acclimation of the corals, was the shifting in the dominant Symbiodinium species in a certain coral holobiont. For example, P. divarcata’s dominant endosymbiont was C47 in 2009 and changed substantially to A4 over the next year. Of all three of the corals, P. astreaoides showed the weakest shift in dominant endosymbiont type. Together with having the highest energy reserve, the change in distribution of Symbiodinium spp. might have caused P. divarcata to be the coral best resisting and recovering from annual bleaching. With O. faveolata this appeared to be more complex and P. astreaoides just didn’t seem to acclimate.


What I find remarkable about this paper is that the results differ a lot per coral species. This is useful information to show how acclimation capacity of corals is species-specific. However, the coral species combination used here occurred to me as if it was specifically chosen to show the distinction in the results. The authors of this paper didn’t mention any reason for working with these species in particular, but if they had more knowledge about them in relation to the research topic it would have been interesting to read their expectations. Besides that, some figures show results that are not proven to be significant, but do comply with significant trends in other samples. In some cases this is mentioned, but in my opinion the authors could have been more elaborative on this. Ofcourse, these trends can be explained better with more extensive research, using different coral species and working with a longer time scale. However, this research gave an interesting insight in the abilities and inabilities of coral species to survive and acclimate to more frequent bleaching event. Some will win, some will lose.

Paper reviewed:
Grottoli, A. G., Warner, M. E., Levas, S. J., Aschaffenburg, M. D., Schoepf, V., McGinley, M., ... & Matsui, Y. (2014). The cumulative impact of annual coral bleaching can turn some coral species winners into losers. Global change biology20(12), 3823-3833.

References:
Van Hooidonk, R., Maynard, J. A., & Planes, S. (2013). Temporary refugia for coral reefs in a warming world. Nature Climate Change3(5), 508-511.

3 comments:

  1. Hi Thyrza interesting read, I like how the paper looks at the effects of blenching over a two year period many of the papers I have read usually only focus on the immediate effect of blenching over a period of a few weeks and don't look at how the coral respond/recover and the microbes associated with the corals may change following the blenching event as the corals being to recover. It’s interesting how even if the coral are able to tolerate and recover from the first year they may not be recovered enough to tolerate a second blenching event the following year, something that appears to be becoming a much bigger issue with climate change. It’s also interesting how different species of coral are more tolerant and able to recover more effectively.

    You mentioned how the researchers appeared to choose 3 coral species that would show difference responses but didn't really explain or justify why they choose those species I was wondering if you had done any research into why there was a different between the species the paper mentions that P. divarcata appeared to have the highest acclimation capacity which was likely due to it having the highest energy reserve and being able to most effectively change the distribution of Symbiodinium spp. Have you come across any research which provides any suggestions/explanation as to why it has a higher energy reserve or is able to more effectively change its Symbiodinium spp. Distribution.

    Thanks for your time
    Alisha

    ReplyDelete
  2. Hi Alisha,

    Thanks for your comment. It's true that most papers on the subject of heat-stress survival by corals only researched single bleaching events or bleaching events with a longer time period in between. However the results from the paper I reviewed often comply with some results from this earlier research. For example, several studies already indicated that Porites species had the capacity to adapt to bleaching events seperated by more years (see Smith et al. (2013) for example). Furthermore, the feature of having a high energy reserve was also expected to play an important role in reducing coral susceptibility and increasing recovery rates (Anthony et al., 2009). From what I understood, this probably has to do with the fact that the coral still needs carbon resources to recover of course, if not more than it did before. However after bleaching, most corals lost a significant amount of their Symbiodinium population, which provide the coral host with fixed carbon. Therefore, other carbon resources need
    to be used, like heterography or the energy reserves. In the paper I reviewed, it appeared that heterography doens't play as much of an important role as it was expected to do. Energy reserves did however and apparently P. divarcata has relatively more lipids, proteins and carbohydrates that can be used as energy reserves.
    Why P. divarcata is able to more effectively shift it's Symbiodinium population is not really clear to me. I think it has more to do with the heat resistancy of the microbes themselves than it has to do with the coral host, but ofcourse there could be a question of animal-endosymbiont interaction that influences this, which is also put forward by the article reviewed.

    I hope this answers your questions :)

    Thyrza

    ReplyDelete
  3. Oh the articles mentioned in my comments are:

    Anthony, K. R., Kline, D. I., Diaz-Pulido, G., Dove, S., & Hoegh-Guldberg, O. (2008). Ocean acidification causes bleaching and productivity loss in coral reef builders. Proceedings of the National Academy of Sciences, 105(45), 17442-17446.

    Smith, T. B., Brandt, M. E., Calnan, J. M., Nemeth, R. S., Blondeau, J., Kadison, E., ... & Rothenberger, P. (2013). Convergent mortality responses of Caribbean coral species to seawater warming. Ecosphere, 4(7), 1-40.

    ReplyDelete

Comments from external users are moderated before posting.

Note: only a member of this blog may post a comment.