Seasonal dynamics of the endosymbiotic, nitrogen-fixing cyanobacterium Richelia intracellularis in the eastern Mediterranean Sea

Edo Bar Zeev, Tali Yogev, Dikla Man-Aharonovich, Nurit Kress, Barak Herut, Oded Béjà, Ilana Berman-Frank

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Biological nitrogen fixation has been suggested as an important source of nitrogen for the ultra-oligotrophic waters of the Levantine Basin of the Mediterranean Sea. In this study, we identify and characterize the spatial and temporal distribution of the N-fixing (diazotrophic) cyanobacterium Richelia intracellularis. R. intracellularis is usually found as an endosymbiont within diatoms such as Rhizosolenia spp and Hemiaulus spp. and is an important diazotroph in marine tropical oceans. In this study, two stations off the Mediterranean coast of Israel were sampled monthly during 2005-2007. R. intracellularis was identified by microscopy and by reverse transcribed-PCR which confirmed a 98.8% identity with known nifH sequences of R. intracellularis from around the world. The diatom-diazotroph associations were found throughout the year peaking during autumn (October-November) at both stations. Abundance of R. intracellularis ranged from 10 to 55 heterocysts l 1 and correlated positively with the dissolved Si(OH) 4 /(NO 3 NO 2) ratio in surface waters. Although the rates of nitrogen fixation were very low, averaging 1.1 nmol N l 1 day 1 for the R. intracellularis size fraction (10 m) from surface waters, they correlated positively with heterocyst counts during thermal stratification. The lack of large-scale diatom-diazotroph blooms and the low rates of nitrogen fixation by these diazotrophs may result from the P-starved conditions affecting the Levantine basin.

Original languageEnglish
Pages (from-to)911-923
Number of pages13
JournalISME Journal
Issue number9
StatePublished - Sep 2008

Bibliographical note

Funding Information:
We thank the Captain (Dani Sheffer) and Engineer (Etai Catzman) of the R/V Mediterranean Explorer and the EcoOcean Marine Research and Education Organization as well as the captain and crew of the R/V Shikmona for their expert assistance at sea. The R/V Shikmona cruises were funded by a donation of the Israel Oceanographic and Limnological Research North American Friends. Many thanks to R Foster for bringing to our attention the importance of the diatom–diazotroph symbioses as well as for insightful suggestions, tips, and much help throughout. Thanks to B Kimor, E Carpenter, and N Gordon for aid in identification of the diatom–diazotrophs and to E Rahav for technical help at sea. We thank I Gertman from IOLR for generating Figure 1. This research is part of the requirements for an MSc thesis for EB-Z and for a Ph D thesis for TY at Bar Ilan University. This study is supported by an Israel Science Foundation Grant (No. 458/04) to IB-F and OB.


  • Cyanobacteria
  • Diatoms-diazotroph symbiosis
  • Mediterranean sea
  • Nitrogen-fixation
  • Richelia intracellularis


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