Not that I understand it but…………….
https://gupea.ub.gu.se/bitstream/2077/28022/1/gupea_2077_28022_1.pdf
Microcolony growth
One of the inherent properties of nitrifying bacteria in biofilms is their ability to
form microcolonies of varying size, shape and density (eg Schramm et al., 1996,
Schramm et al., 1999, Gieseke et al., 2003, Okabe et al., 2004, Hallin et al., 2005,
Papers I-IV, Fig. 6). These microcolonies are often structurally stable and difficult
to disintegrate without killing the bacteria (Larsen et al., 2008, Frank Persson,
persson-al communication) and it was recently shown that production of
extracellular DNA (eDNA) provides structural strength to the microcolonies which
contained high amounts of these molecules (Dominiak et al., 2011). Okabe et al.,
(2004) measured how nitrifier microcolony average size varied with depth and
organic carbon availability in the medium. They observed that AOB microcolony
size was rather constant throughout the biofilm when organic carbon was not
added. In contrast, size distribution was significantly stratified in the biofilm
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residing in a reactor with a C/N ratio of 1, probably an effect of heterotrophic
bacteria outcompeting autotrophic nitrifiers in the surface layers of the biofilm,
where AOB microcolony size was the smallest. Gieseke et al., (2003) also observed
hetereogenous size distribution of AOB microcolonies and reported that
Nitrosococcus mobilis microcolonies were smaller in less densely populated biofilm
regions, whereas larger ones were often found together with AOB from the
Nitrosomonas europaea/eutropha lineage (cluster 7). In addition, Okabe et al., (2004)
observed that microcolonies of two different groups of AOB, belonging to
Nitrosomonas and Nitrosospira respectively, differed in their areal cell density and it
was speculated that the looser colonies of Nitrosospira would facilitate oxygen and
ammonium diffusion which could partly compensate for a lower growth rate. Such
loose microcolony structures have also been observed in Nitrosococcus mobilis
(Gieseke et al., 2003) and Nitrospira (Daims et al., 2001a). Microcolony
disintegration in Nitrospira has been reported as an effect of nitrate accumulation in
the system (Spieck et al., 2006). Spieck and colleagues (2006) hypothesized that
switching from microcolony to planctonic growth would be a straightforward way
of escaping detrimental changes in environmental conditions. Furthermore, it was
shown for the AOB N. europaea that NO gas functions as a signal for switching
between planctonic and biofilm growth (Schmidt et al., 2004a). Thus, several
observations indicate that nitrifier microcolony size distribution and density reflects
the ecophysiology of the organisms and the conditions prevailing in their
environment. Further discussion on this topic is found in the “Conclusions and