Saturday, 28 March 2015

Is Mucus Maximising Microlayer Microbes?



The Sea-surface Microlayer (SML) is a gelatinous biofilm-like habitat with a distinct microbial community teaming with bacteria, flagellates, algae and fungi. These communities are significantly 'enriched' compared to the water below due to rising TEPs (Transparent Exopolymer Particles) that supply nutrition and attached microbes to the microlayer habitat. Although TEP is largely formed from phytoplankton DOM, coral mucus may play a similar role. Corals produce vast amounts of mucus, some of which rises to the surface as transparent filaments and strings, due to trapped air bubbles. This could potentially supply both microorganisms and nutrients to the microlayer in waters above reefs. Therefore, it may be that the communities on the SML above reefs could be more enriched with bacteria than other ecosystems and enrichment may increase as coral coverage does.

To test this, Nakajima et al. sampled a fringing coral reef off Bidong Island, Malaysia for three consecutive days in June at three sites. Two shallow sites, one with high coral cover and the other with low coral cover (dominated by Acropora nobilis and A. formosa) and also an offshore site, where I assume coral was absent. The SML was sampled using a metal mesh sampler, which collected the upper 250µm of water and subsurface water samples were taken a depth of 10cm. Water samples were immediately brought to the lab on ice. DOC and Chlorophyll a concentrations were measured. Microbes were enumerated using epi-fluorescence after filtration onto membrane filter, 0.2µm for heterotrophic bacteria and 0.8µm for cyanobacteria, heterotrophic and autotrophic nanoflagellates. Bacterial (heterotrophic) production was the measurement of bromodeoxyuridine incorporation in the dark and bacterial growth rate was calculated. Enrichment factors were calculated for all parameters and multivariate statistics carried out. Unsurprisingly, the SML was significantly enriched over every parameter bar bacterial production and growth rate. More interestingly, enrichment factors for all the microbial groups maybe higher than in other marine habitats. These higher enrichment factors could have implications for the exchange of gases, if matched to an increased metabolism. However, it is difficult to confirm this due to varying methodologies used in different studies. Remarkably, the high coral cover site had the highest enrichment of heterotrophic bacteria. This may support the hypothesis that coral mucus transports trapped pelagic and coral-associated bacteria into the SML. Furthermore, transport into the SML may be more important than in-situ production given a low bacterial growth rate. Additionally, at the high coral sites there was a very low ratio of heterotrophic flagellates to bacteria/cyanobacteria possibly suggesting that there could be a higher level of grazing above the high coral site.

I found the study very interesting, but it can be summarised as being very 'old school', given that there was no focus on taxonomy. A full phylogenetic analysis using 16S/18S rRNA sequences would have allowed direct comparison between the SML communities above the coral with other marine systems and the coral microbes themselves. It would also have been much more informative to sample a number of sites with differing coral coverage to give a clearer picture of whether the enrichment really increased with increasing coverage. Although I am cautious about some of the conclusions made, I think that the study does raise interesting questions and provides a good starting point for future studies where more powerful techniques could be used.  

Ref: Nakajima, R., Tsuchiya, K., Nakatomi, N., Yoshida, T., Tada, Y. Konno, F., Toda, T., Kuwahara, V.S., Hamasaki, K., Othman, B.H.R., Segaran, T.C., Effendy, A.W.M. (2013). Enrichment of microbial abundance in the sea-surface microlayer over a coral reef: implications for biogeochemical cycles in reef ecosystems. Marine Ecology Progress Series, 490, 11-22.

9 comments:

  1. Hi Tom,

    It would be interesting to see if different types of coral reefs produce different amounts of mucus and how the corresponding micro-layer above is affected. Did the paper mention anything about this?

    Cheers!
    Bekki

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  2. Hi Bekki,

    Great idea, no they dident! I think the best way of doing that would have been to sample all of the corals in an area. Then put the coverage data/mucus production data into a multivariate statistics package and then see if the SLM microbes varied with coral assemblage. I do think that kind of approach would definitely be benefited by using phylogenetics. However, at the sites they chose it was largely dominated by two species of Acropora, so perhaps it would be better to assess that at more diverse coral assemblages.

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    1. I think it is definitely a good idea to look at different sites with different genera of coral as here the seem to have only used Acropora. There are certainly corals which produce more mucus than the Acroporids. This would have benefited, like you said, from a phylogenetic analysis - as they appear to be assuming that the higher proportion of bacteria has directly come from the corals releasing mucus - however it could just be a nice habitat for pelagic bacteria. Maybe they could sequence the bacterial community associated with the coral to use as a comparison with the SML?

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    2. Hi Katrina,

      Yes I think sequencing the coral microbes would have been extremely beneficial and came to this conclusion in the main post. I suppose you could just take samples of the coral mucus. Presumably you can take scrapings or just whack a plastic bag over the top of the coral and then sample from the accumulated mucus?

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    3. Yep sorry I see that now, I thought you meant just different reefs.
      The results of Bourne and Munn 2005 show that the most accurate representation of coral associated bacteria is achieved by removing the coral tissue. This is done using a pressurised airgun, but it does mean killing the coral unfortunately.
      For some corals such as the Fungia you may be able to just collect some mucus from sloughing - these guys produce ridiculous amount of mucus and are free living - but im not sure about sticking a bag over it - might kill the zooxanthellae.

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    4. I read about the plastic bag method in a paper from the 70's (Richman et al. 1975) although I think they were just quantifying total amount of mucus and so were not worried about microbes etc. Thanks for the info!

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  3. Hi Tom

    Thanks for an interesting post on the links with coral and the SML. It’s a shame that they didn’t do any taxonomy as this would have given more insight into the differences in microbial communities between the SML, sub-surface waters and coral communities. What did they mean by enrichment factors? Was this a measure of bacterial diversity?

    I agree that sampling a number of sites with different coral coverage may have had an impact on the SML communities above the coral. Presumably, the coral-produced mucus floats up to the surface to aggregate with the rest of the SML? It’s great to see that there is still so much more to explore in terms of SML and factors affecting community composition.

    Cheers,
    Anita

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    1. Hi Anita,

      Enrichment factor is just how much more of a parameter the SML possess over the sub-surface water. When they use this to refer to bacteria they mean just the abundance of bacteria, there is no information on diversity included in that metric.

      With regards to the mucus yes they hypothesize that a certain proportion rises due to trapped air bubbles, its less than half as over fifty percent just dissolves straight away. So your right presumably it does aggregate with the TEP.

      With regards to the study in general its extremely basic, I was quite surprised given the current era of every study involving a molecular component. Given this I am fairly conservative about the conclusions drawn in the paper but I blogged it anyway as it was quite novel. Its more of a hypothesis generating paper than one that provides robust answers.

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    2. Hi Tom

      Thanks, that makes sense. I agree that the study is basic compared to other studies but it's nice to have another and different perspective on the SML idea. Thanks for the info.

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