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\section{Discussion}
\subsection{Biomass Pool Size} The goal of this study was to evaluate how
changes in available C affected the biomass pool
size,
membership size and
structure composition of
planktonic and biofilm communities. Our results suggest that C subsidies
increased heterotroph biomass in both plankton and biofilm
communities as predicted. communities.
C amendments also resulted in decreased photoautotroph biomass in the plankton
community, but there was no significant change in biofilm
photoautotroph biomass between resource treatments. Although the DOC
concentration in the highest C treatment was significantly higher than the
other treatments, the concentrations we measured were in the range of those
reported in natural marine ecosystems \citep{Mopper1980} and it is has been
noted that glucose concentrations in coastal marine ecosystems may fluctuate
over several orders of magnitude \citep{Alonso2006}. The changes in the biomass
pool size that did occur were consistent with changing relationships (commensal
to competitive) between the autotrophic and heterotrophic components of the
plankton communities but not necessarily of the biofilm communities. While we
recognize that other mechanisms may drive the shift in biomass pool size of
these two components of the microbial community (e.g. increased grazing
pressure on the algae with C additions, or production of secondary metabolites
by the bacteria that inhibit algal growth) previous studies
\cite{Stets_2008,Cotner_2002} and the data reported here suggest that altered
nutrient competition among heterotrophic and photoautotrophic members of the
plankton is the most parsimonious explanation for this shift in biomass pool
size.
\subsection{Biofilm and Plankton Alpha and Beta Diversity} Beyond changes
in the biomass pool size of each community, we explored how shifts in
resource C affected
a) the membership and structure of each community, and
b)
the recruitment of plankton during biofilm community assembly.
Intuitively, shifts in planktonic community composition should alter the
available pool of microorganisms that can be recruited into a biofilm. For
example, if planktonic diversity increases, the number of potential
...
represent a temporally integrated sample of the planktonic organisms moving
through the reactor resulting in higher apparent alpha diversity (i.e. mass
effects would be the dominant assembly mechanism). Second, the biofilm
environment may disproportionately enrich
for the
least abundant low abundance members of the
planktonic community. In this case it is probable that the biofilm would
incorporate the most abundant members from the planktonic community (i.e. mass
effects) but also select and enrich (i.e. species sorting) the least abundant
...
populations when compared to the most abundant members of the plankton
community. As noted above, in the highest C treatment (C:P = 500) the
heterotroph biofilm and plankton community membership had significant overlap
at the final timepoint (Figure~\ref{fig:pcoa}). However, heterotrophic plankton
membership community composition for the highest C treatment among timepoints (8 and
17 days) were also qualitatively as similar to each other as any other
community. Thus, variable planktonic community composition among timepoints
would not explain the higher diversity observed in the biofilm compared to the
...
treatment levels and both timepoints. While we did not (could not) specifically
measure niche diversity within the biofilm communities our results suggest that
the biofilm habitat selected for unique members of the photoautotroph and
heterotrophic community that were in low abundance in the planktonic habitat
but readily became major constituents of the biofilm community.
% Fakesubsubsection:few studies have
Few studies have simultaneously evaluated the relationship among membership
...
intermediate peak in OTU richness is consistent with a classic
productivity-diversity relationship that has been shown for many ecosystems and
communities both microbial and otherwise. However, as with other experiments
with this result our experimental design did not allow us to tell whether resources drove
productivity that
subsequently drove changes in diversity or whether resources
drove diversity which altered productivity. Rather, we note that as diversity
decreased in the highest C treatment, heterotrophic plankton and biofilm
membership became increasingly similar. This suggests that environments that
contained high amounts of labile C selected for fewer dominant taxa,
overwhelming the lifestyle species sorting mechanisms that appeared to dominate
biofilm community assembly in all other treatments. Similarly, while we did not
measure extracellular polymeric substances (EPS), direct microscopy showed
that planktonic cells in the highest C treatment (C:P = 500) were surrounded by
what appeared to be EPS. Because biofilm EPS appeared also to increase moving
from the low to high C treatments it is possible that more abundant planktonic
cells were more readily incorporated into biofilms due both to increased
"stickiness" of the planktonic cells as well as the biofilm itself. While we
did not observe flocculating DOC which has been shown to dominate high DOC
environments in nature, we did measure a substantial increase in DOC in the C:P
= 500 treatment which was more than 2-fold higher than any of the other
treatments. Thus additional adhesion of the plankton and the biofilm may also
explain the merging of the planktonic and biofilm heterotroph membership in the
highest C treatment.
\subsection{Lifestyle (biofilm or planktonic) Enriched OTUs} There are only a
few studies that attempt to compare biofilm community composition and the
...
the plankton) but are readily enriched in the biofilm lifestyle. Most of the
biofilm enriched photoautotroph OTUs were \textit{Bacillariophyta} although
there were also many \textit{Bacillariophya} OTUs enriched in the planktonic
libraries. We also found
more \textit{Cryptophyta} and \textit{Viridiplantae}
were
more uniformly enriched in the planktonic photoautotroph libraries. It appears that
these broad taxonomic groups were selected against in biofilms under our
experimental conditions. Heterotroph OTUs enriched in planktonic samples
displayed more dramatic differential abundance patterns than heterotroph OTUs
enriched in biofilm samples, but, biofilm enriched heterotroph OTUs were spread
...
in the highest C treatment. Consistent with a growing body of work our results
suggest that complex environmental biofilms are a unique microbial community
that form from taxa that are found in low abundance in the neighboring
communities. This membership was affected by C amendments for heterotrophic but
not
autotrophic photoautotrophic microbes and then only in the most extreme resource
environment. This suggests that lifestyle is a major division among
environmental microorganisms and although biofilm forming microbes must travel
in planktonic form at some point, reproductive success and metabolic
contributions to biogeochemical processes comes from those taxa primarily if