Friday, 11 January 2013

Functionality of different Symbiodinium clades


Functionality of different Symbiodinium clades

Coral reefs are founded on the symbiotic relationship between the coral animals and photosynthetic algae (Symbiodinium a.k.a. zooxanthellae). Symbiodinium is a genus of dinoflagellate which contains many different edosymbiotic algae found in a diverse group of host organisms, primarily tropical cnidarians, but also other groups such as molluscs and sponges. Within the Symbiodinium genus there are numerous clades, it has been reported that single organisms can harbour one or more clades at any one time. Whilst the molecular diversity of the Symbiodinium is well documented, the functional significance of such diversity has received less attention. In the paper reviewed, Loram et al. investigated the functional significance of different zooxanthellae clades in the giant anemone Condylactis gigantean. It is important to increase our understanding on the physiological functioning of zooxanthellae in order to learn more about bleaching. This directly relates to the Adaptive Bleaching Hypothesis (ABH) proposed by Buddheimer & Fautin, (1993) which says that when environmental conditions change the loss of one or more kinds of zooxanthellae is rapidly followed by the formation of a new consortium to adapt. In other words coral, or in this case the closely related anemone, can pick and choose their zooxanthellae to help them adapt to different environmental situation. The ABH relies heavily on assumptions that different zooxanthellae clades are functionally diverse and have varying ability to cope with environmental parameters such as irradiance and temperature.

Loram et al. (2007) genetically identified Symbiodinium clades A and B in their test organisms; 32% of organisms contained clade A, 20% contained clade B and 48% contained a mixture of both A and B. An experiment was sent up to have two environmental situations, 25 °C represented a control and 30°C represented elevated temperature. As a measure of nutritional function, the photosynthetically fixed carbon in the algae was measured as well as the translocation of fixed carbon into the animal tissues. This parameter was measured by radio isotope analysis of 14C. The paper presents its results in an overcomplicated way, I feel to disguise the lack of biologically relevant statistical significance. Loram et al. make statements such as “carbon fixation rates per algal cell differed among the clades, with total fixation rates elevated at 30 °C for symbioses bearing clade A and depressed for symbioses bearing B” although close inspection of the results table reveal that this difference is not statistically significant. Overall, and most importantly, the results do indicate that clades A and B are functionally different, although the biological significance of this is perhaps over interpreted in the discussion section.

This paper fits nicely into one of our lectures with Emma “Coral Health and The Environment” and builds on work presented in the lecture by Rowan et al. (1997) and Rowan (2004). I decided to review the paper as it focuses is on  an anemone rather than the reef building corals, and although the corals are vitally important and should never be underestimated, I feel that other interesting reef species get overlooked. I think the way the paper has used genetic techniques for I.D. and more traditional experimental method to assay functionality works really well. Often too much emphasis is put onto the genes present and genomic potential without any physiological evidence of function. It is important to note that this study was done five (ish) years ago now in a time when "holobiont" research increased rapidly. It would be interesting to review more recent work to see how the field has progressed.
Loram, J. E., Trapido-Rosenthal, H. G., & Douglas, A. E. (2007). Functional significance of genetically different symbiotic algae Symbiodinium in a coral reef symbiosis. Molecular ecology, 16(22), 4849–57.
 
 
 


 

4 comments:

  1. Hi Vicky
    I think you made some excellent points in this blog. I agree that it should be of primary interest to understand the importance of bearing different clades and the physiological differences between them.

    Did the symbiosis with clade B at 25°C have high carbon fixation rates than clade A (even if it wasn't statistically significant)? otherwise I wouldn't see in what case it would be advantageous to form a symbiosis with clade A. Also do you have any ideas how the coral would control which clade colonises it?

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  2. Hi Vicky (and Anna),
    As you expressed an interest in more recent work relating to the functionality of different Symbiodinium clades, I thought of an interesting investigation by Littman et al (2010), which seems to fit the bill perfectly.

    The authors found that the photochemical efficiency of tropical coral (Acropora tenuis) juveniles which hosted different clades of Symbiodinium where found to vary significantly under elevated temperatures: Juveniles with clade D Symbiodinium underwent a 44% decline in photochemical efficiency at 32oC (in comparison to at the control temperature of 28oC), whereas juveniles associated with clade C Symbiodinium exhibited only a 10% decline in photochemical efficiency at 32oC.

    Additionally, the microbial community of the D-juveniles was observed to shift to a more Vibrio dominated state under the elevated temperature, and no such shift in microbiota structure was observed in the C-juveniles. I think that this link between the functionality of different Symbiodinium clades and the changes in anthozoan microbiota under increased temperature is, potentially, particularly important to our understanding of holobiont health and the future of coral reefs.
    I’d be interested to get your opinion.

    Littman, Raechel a, David G Bourne, and Bette L Willis. 2010. “Responses of Coral-associated Bacterial Communities to Heat Stress Differ with Symbiodinium Type on the Same Coral Host.” Molecular Ecology 19: 1978–90.

    http://www.ncbi.nlm.nih.gov/pubmed/20529072

    Also (Anna), re: clade colonisation, I believe that one prevailing opinion is that various Symbiodinium clades exist as cryptic members in anthozoan microbial communities (i.e. in very few numbers; hard to detect), and that when conditions are favourable, temperature tolerant clades outcompete (and thus replace) less tolerant ones, over time. I’m actually half way through a post on this!
    Jo

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  3. Hi Jo and Anna,
    Thanks for the great comments!
    So firstly Anna your question about the results and differences between clade A and B at 25°C; again there was no statistical difference between clade A, clade B or a mixture of A and B. At this point I agree with you, what is the advantage of clade A? However, when clade A and B are present at elevated temperature there is a statistically significant increase in fixed carbon. The results are slightly more certain for percentage of fixed carbon translocated to the animal, here we see that at 25°C, Clade A has a (statistically) higher rate than B. Whereas for 30°C A and B are the same and the organisms with both A and B have the highest. There is also an interesting two-way ANOVA for Clade*Temp. Like I said there are clearly functional differences occurring, but it is hard to pick apart the biological significance of this, particularly because of the way the results section is presented and communicated. Bearing in mind what Jo has said about the Littmann (2010) study, I think the harbouring of different clades must be linked to some sort of cost/benefit assessment. It may be useful to have a clade present which is inefficient at normal temperature but will protect you when times get tough, almost like an investment for possible change in the future.
    Moving on to the point about how coral control the symbionts which colonise them… Jo makes a good point here RE: the prevailing opinion, but I think it’s fair to say that there is still some debate on this topic. We know that corals acquire zooxanthellae from the environment at the larval stage or, they are transmitted vertically in the eggs. In this case it is easy to imagine how evolution has driven this relationship. For all animal phyla that have dinoflagellate symbionts, digestion is intracellular, certain dinoflagellates could have become resistant to digestion and thus establishing a relationship with the coral. However when you consider corals picking and choosing their symbionts in order to adapt to the changing environment, the evolutionary picture becomes somewhat complicated. I would be interested to read the blog Jo is preparing about this topic.
    Jo I agree with you on your idea about the link between Symbiodinium and microbiota, I can’t help but think that currently we are trying to compartmentalise everything too much. In Colin’s recent lecture we discussed Koch’s postules and how it is difficult to apply them to the consortium which is associated with disease. Building on this idea, I feel that more emphasis should be placed on the bigger “holobiont” picture. Whilst a deeper mechanistic understanding of individual clades physiology etc. is important, I still feel this kind of research would benefit from more integration.

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  4. Hi Vicky,

    Yes, compartmentalisation for understanding vs. knowing how something works in an actual system (the big picture): one of the trickiest aspects of biology I reckon! I suppose how this is normally addressed is to try and understand how component parts of a system work (e.g. photosystems of Symbiodinium), and then to compare how they function under different conditions in situ (e.g. when the holobiont is experiencing high temperatures).

    I guess the major problem is that it is impossible to measure everything, and if you did then how would you ever pin down causation?

    Ah, science!
    (p.s. next blog nearly there!)

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