Previously the work by Thomson et al, 2012 has been discussed during a class seminar. The
cyanobacteria UCYN-A work was concluded to be in a symbiotic relationship with a
host photosynthesising prymnesiophyte. With the symbiotic relationship believed
to be of an obligate nature due to the provision of nitrogen by the UCYN-A to
the prymnesiophyte and the provision of carbon to the UCYN-A by the
prymnesiopyte. This latest study by Thomson et
al, 2014 takes the research a step further by examining the phylogeny of
UCYN-A cyanobacteria (with its numerous clades) and the host prymnesiophyte.
Through phylogenetic analysis of UCYN-A samples possessing
the nitrogenase fixation gene (nifH) three distinct clades were identified, all
of which did not possess the Photosystem II (carbon fixation pathway) and TCA
cycle. These clades were appropriately denoted as UCYN-A1, UCYN-A2 and UCYN-A3.
The Thomson et al, 2014 study focused on the UCYN-A2 cyanobacteria from a
coastal site, examining the cell’s morphology and genetic identification and
comparing the findings with the original UCYN-A1 cyanobacteria. The nifH gene expression
of UCYN-A2 was measured over 3 diel cycles and was found to increase during
daylight hours much like the UCYN-A1 clade, reflecting the fixation of nitrogen
during the day time. This coincides with the most appropriate time for
photosynthesis to occur by photosynthesising organisms (including its symbiotic
host). As the UCYN-A clades do not possess carbon fixation pathways the production
of N2 does not have to be separated into different cells, allowing
the fixation of N2 at any time, day or night.
Analysis of the rRNA sequences revealed the two
cyanobacteria UCYN-A1 and UCYN-A2 to be of two distinct clades, this led to the
conclusion that each clade co-evolved separately with a specific prymnesipohyte
host. The analysis identified that there were differences between the clades at
nucleotide level, reflected by only 86% similarity at amino acid level yet more
than 96% similarity in genes over all. The differences between the clades may
be due to habitation within different marine environments, leading to the UCYN-A
nifH sequences being further examined to see if environmental habitats of each
clade were separate or overlapped. UCYN-A 2 nifH sequences were found in the
same geographical locations as previously recorded UCYN-A1 locations, confirming
the overlap of clades geographic locations, this was also the case with the
UCYN-A3 clade. In addition to the three known clades of UCYN-A, further
unidentified sequences were also found using flow cytometry, indicating there
may be additional UCYN-A clades in the global oceans. Additional clades within the
oceans could lead to previous levels of N2 input by nitrogenising
cyanobacteria being under estimated, meaning their ecological importance could be
greater than initially thought.
To identify whether each clade was in partnership with a
specific host prymnesiophyte cell sorting, microscopy and PCR techniques were
undertaken. From the samples of UCYN-A2 positive cells 10 phytoplankton cells
were exposed to nested PCR using 18s rRNA primers to identify the host. Seven
cells possessed the prymensiophyte sequences showing a great similarity to the
UCYN-A1 host, however, there was a 2% variation between the nucleotide
sequences. This supports the theory that the different UCYN-A clades 1 and 2
possess genetically different prymnesiophyte hosts from each other. The
presence of diverse UCYN-A clades with genetically different pymnesiophytes may
also support the theory of co-evolution. During the sequencing of the UCYN-A2
host a 100% genetic match was made to Braarudosphaera bigelowii. As B.bigelowii is a
calcareous plate forming cell it is likely that the UCYN-A2 host is also a
calcareous plate forming cell as discussed as a possibility in Thomson et al,
2012.
The research conducted in Thomson et al 2014 answered
questions from the Thomason et al, 2012 study by identifying further clades of
the UCYN-A cyanobacteria, and their close symbiotic relationship with
genetically specific B.bigelowii hosts.
This could be investigated further by completing investigations upon the
UCYN-A3 and its associated symbiotic partner to see if the same conclusions can
be ascertained through genetic sequencing. Further investigations into the
unidentified UCYN-A clades may also provide additional insight into the
symbiotic relationships between nitrogen fixing cyanobacteria and their carbon
fixing hosts.
Thompson, A.,Carter, B.J., Turk-Kubo, K., Malfatti, F.,
Azam, F and Zehr, J.P. (2014) Genetic diversity of the unicellular nitrogen-fixing
cyanobacteria UCYN-A and its prymensiophyte host. Environmental Microbiology 16
3238-3249
http://onlinelibrary.wiley.com/doi/10.1111/1462-2920.12490
Hi Emma
ReplyDeleteI think that it would be really interesting if this study was extended further by looking at the number of UCYN-A cells per host and whether there is a maximum number of UCYN-A cells that the host can accommodate at any one time. What do you think?
Anita