|Synechococcus PCC 7002 cells in DIC microscopy|
Synechococcus (from the Greek synechos (in succession) and the Greek kokkos (berry) is a unicellular cyanobacterium that is very widespread in the marine environment. Its size varies from 0.8 µm to 1.5 µm. The photosynthetic coccoid cells are preferentially found in well–lit surface waters where it can be very abundant (generally 1,000 to 200,000 cells per millilitre). Many freshwater species of Synechococcus have also been described.
Synechococcus is one of the most important components of the prokaryotic autotrophic picoplankton in the temperate to tropical oceans. The genus was first described in 1979, and was originally defined to include "small unicellular cyanobacteria with ovoid to cylindrical cells that reproduce by binary traverse fission in a single plane and lack sheaths". This definition of the genus Synechococcus contained organisms of considerable genetic diversity and was later subdivided into subgroups based on the presence of the accessory pigment phycoerythrin. The marine forms of Synechococcus are coccoid cells between 0.6 µm and 1.6 µm in size. They are gram negative cells with highly structured cell walls that may contain projections on their surface. Electron microscopy frequently reveals the presence of phosphate inclusions, glycogen granules and more importantly highly structured carboxysomes.
Cells are known to be motile by a gliding type method and a novel uncharacterized, non-phototactic swimming method that does not involve flagellar motion. While some cyanobacteria are capable of photoheterotrophic or even chemoheterotrophic growth, all marine Synechococcus strains appear to be obligate photoautotrophs that are capable of supporting their nitrogen requirements using nitrate, ammonia or in some cases urea as a sole nitrogen source. Marine Synechococcus are traditionally not thought to fix nitrogen (this perception may be changing).
The main photosynthetic pigment in Synechococcus in chlorophyll a, while its major accessory pigments are phycobilliproteins. The four commonly recognized phycobilins are phycocyanin, allophycocyanin, allophycocyanin B and phycoerythrin. In addition Synechococcus also contains zeaxanthin but no diagnostic pigment for this organism is known. Zeaxanthin is also found in Prochlorococcus, red algae and as a minor pigment in some chlorophytes and eustigmatophytes. Similarly phycoerythrin is also found in rhodophytes and some cryptomonads.
Phylogenetic description of Synechococcus is difficult. Isolates are morphologically very similar, yet exhibit a G+C content ranging from 39% to 71%, illustrating the large genetic diversity of this provisional taxon. Initially attempts were made to divide the group into three sub-clusters, each with a specific range of genomic G+C content. The observation that open-ocean isolates alone nearly span the complete G+C spectrum however indicates that Synechococcus is composed of at least several species. Bergey's Manual (Herdman et al. 2001) now divides Synechococcus into five clusters (equivalent to genera) based on morphology, physiology and genetic traits.
Cluster one includes relatively large (1–1.5 µm) non-motile obligate photoautotrophs that exhibit low salt tolerance. Reference strains for this cluster are PCC6301 (formerly Anacycstis nidulans) and PCC6312, which were isolated from freshwater in Texas and California respectively. Cluster 2 also is characterized by low salt tolerance. Cells are obligate photoautrotrophs, lack phycoerythrin and are thermophilic. The reference strain PCC6715 was isolated from a hot spring in Yellowstone National Park. Cluster 3 includes phycoerythrin lacking marine Synechococcus that are euryhaline i.e. capable of growth in both marine and fresh water environments. Several strains, including the reference strain PCC7003 are facultative heterotrophs and require vitamin B12 for growth. Cluster 4 contains a single isolate, PCC7335. This strain is obligate marine. This strain contains phycoerthrin and was first isolated from the intertidal zone in Puerto Peñasco, Mexico. The last cluster contains what had previously been referred to as ‘marine A and B clusters’ of Synechococcus. These cells are truly marine and have been isolated from both the coastal and the open ocean. All strains are obligate photoautrophs and are around 0.6–1.7 µm in diameter. This cluster is however further divided into a population that either contains (cluster 5.1) or does not contain (cluster 5.2) phycoerythrin. The reference strains are WH8103 for the phycoerythrin containing strains and WH5701 for those strains that lack this pigment (Waterbury et al. 1986b). More recently Badger et al. (2002) proposed the division of the cyanobacteria into a α- and a β-subcluster based on the type of rbcL (large subunit of ribulose 1,5-bisphosphate carboxylase/oxygenase) found in these organisms. α-cyanobacteria were defined to contain a form IA, while β-cyanobacteria were defined to contain a form IB of this gene. In support for this division Badger et al. analyze the phylogeny of carboxysomal proteins, which appear to support this division. Also, two particular bicarbonate transport systems appear to only be found in α-cyanobacteria, which lack carboxysomal carbonic anhydrases.
Ecology and distribution
Synechococcus has been observed to occur at concentrations ranging between a few cells per ml to 106 cells per ml in virtually all regions of the oceanic euphotic zone except in samples from the McMurdo Sound and Ross Ice Shelf in Antarctica. Cells are generally much more abundant in nutrient rich environments than in the oligotrophic ocean and prefer the upper well-lit portion of the euphotic zone. Synechococcus has also been observed to occur at high abundances in environments with low salinities and/or low temperatures. Synechococcus is usually far outnumbered by Prochlorococcus in all environments, where they co-occur. Exceptions to this rule are areas of permanently enriched nutrients such as upwelling areas and coastal watersheds. In the nutrient deplete areas of the oceans, such as the central gyres, Synechococcus is apparently always present, although only at low concentrations ranging from a few to 4×10³ cells per ml. Vertically Synechococcus is usually relatively equitably distributed throughout the mixed layer and exhibits an affinity for the higher light regime. Below the mixed layer, cell concentrations rapidly decline. Vertical profiles are however strongly influenced by hydrologic conditions and can be very variable both seasonally and spatially. Overall Synechococcus abundance often parallels that of Prochlorococcus in the water column. In the Pacific HNLC (High Nutrient Low Chlorophyll) zone and in temperate open seas where stratification was recently established both profiles parallel each other and exhibit abundance maxima just about the SCM.
The factors controlling the abundance of Synechococcus still remain poorly understood, especially considering that even in the most nutrient deplete regions of the central gyres, where cell abundances are often very low, population growth rates are often high and not very drastically limited. Factors such as grazing, viral mortality, genetic variability, light adaptation, temperature as well as nutrients are certainly involved, but remain to be investigated on a rigorous and global scale. Despite the uncertainties it has been suggested that there is at least a relationship between ambient nitrogen concentrations and Synechococcus abundance and an inverse relationship to Prochlorococcus in the upper euphotic zone, where light is not limiting. One environment where Synechococcus thrives particularly well are coastal plumes of major rivers. Such plumes are coastally enriched with nutrients such as nitrate and phosphate, which drives large phytoplankton blooms. High productivity in coastal river plumes is often associated with large populations of Synechococcus and elevated form IA (cyanobacterial) rbcL mRNA.
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