This document summarizes heterocyst differentiation in cyanobacteria. Heterocysts are specialized cells that allow cyanobacteria to fix nitrogen in aerobic conditions. In the absence of combined nitrogen, cyanobacteria like Anabaena produce heterocysts at regular intervals along filaments. Heterocysts stop producing oxygen and develop additional envelope layers to protect the nitrogenase enzyme from oxygen. They rely on nearby vegetative cells for energy and carbon in return for fixed nitrogen. 2-oxoglutarate signals nitrogen deprivation and the transcriptional regulator NtcA induces genes for heterocyst development and nitrogen assimilation.
Cyanobacteria use two mechanisms to separate these activities: a biological circadian clock to separate them temporally, and multicellularity and cellular differentiation to separate them spatially
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For example, the unicellular
Cyanothece sp. strain ATCC 51142 stores
glycogen during the day and fixes nitrogen at
night (Toepel et al. 2008), whereas the filamentous
Trichodesmium erythraeum IMS101 fixes
nitrogen during the day in groups of specialized
cells (Sandh et al. 2009).
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Heterocysts are typically distinguishable from vegetative cells:
larger
rounder shape
diminished pigmentation
thicker cell envelopes
prominent cyanophycin granules at poles adjacent to vegetative cells
The additional envelope layers surrounding
heterocysts help to protect the enzyme nitrogenase
from oxygen
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Nitrogenase, sequestered within these cells, transforms dinitrogen into ammonium at the expense of ATP and reductant—both generated by carbohydrate metabolism, a process that is supplemented, in the light, by the activity of PS I. Carbohydrate, probably in the form of sucrose, is synthesized in vegetative cells and moves into heterocysts. In return, nitrogen fixed in heterocysts moves into the vegetative cells, at least in part in the form of amino acids. az wiki
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Photosynthetic C fixation through the Calvin cycle (CC) occurs in the vegetative cells and could lead to Suc and glycogen biosynthesis. Heterocysts act as an important sink for carbohydrates from vegetative cells and as a source of fixed N (Wolk et al., 1994). In heterocysts, which could also synthesize glycogen and Suc, the reductants for N2 and O2 reduction are generated by the activity of the oxidative pentose-P cycle (OPPC), the NADPH heterocyst-specific ferrodoxin, and respiratory electron transport (RET), as well as the ATP synthesis by cyclic phosphorylation (PSI). Suc enzymes are indicated as (1) SuS; (2) A/N-Inv; (3A) SPS-A; (3B) SPS-B; and (4) SPP. αKG, α-Ketoglutarate. az http://www.plantphysiol.org/content/143/3/1385/F8.expansion
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combined nitrogen
such as ammonium or nitrate inhibits the
differentiation of heterocysts
An artificial analog of
2-oxoglutarate, 2,2-difluoropentanoic acid, DFPA,
added to medium resulted in heterocyst development
even in the presence of ammonium
Nitrogenlimiting
conditions result in an increase in the
levels of 2-oxoglutarate showing that 2-oxoglutarate plays a key role in
controlling heterocyst development (Laurent
et al. 2005)..
NtcA is conserved in all cyanobacteria and regulates a number of genes involved in carbon and nitrogen metabolism