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biofilm communities. Our results suggest that C subsidies increased
bacterial biomass in both plankton and biofilm communities as predicted. Carbon
subsidies also resulted in decreased algal biomass in the plankton community,
but there was no
signifcant significant change in algal biomass of the biofilm communities
among resource treatments. The changes in the biomass pool size that did
occurr occur
were consistent with changing relationships (commensal to competitive) between
the autotrophic and heterotrophic components of the plankton communities but
not
neccessarily necessarily of the biofilm communities.
\subsection{Biofilm and Plankton Alpha and Beta Diversity}
Beyond changes in the biomass pool size of each community we explored how
...
structure of the bacterial biofilm and plankton communities were more similar
within a lifestyle (plankton vs. biofilm) than within a resource treatment.
However, for the bacteria in the highest C:P treatment (C:P = 500) both
membership and structure of biofilm and planktonic communities at day 17 were
more similar to each other than to communities from other treatments
(Figure~\ref{fig:pcoa}). Third, C subsides acted differently on the algal and
bacteria communities. Specifically while the highest level of C subsidies (C:P
= 500) resulted in a merging of membership in the bacterioplankton and
bacterial biofilm communities the same merging of membership was not observed
for the algal biofilm and plankton communities which had distinct membership in
all treatments.
We propose two potential mechanisms that could result in the increased
diversity of the biofilm communities relative to the planktonic communities.
...
community (i.e. mass effects) but also select and enrich (i.e. species sorting)
the lease abundant members of the planktonic community resulting in a higher level
of detectable alpha diversity.
The second mechanism would result if the biofilm
enivronment environment represented a more
diverse habitat including sharply delineated oxygen, nutrient and pH gradients
that are not present in the planktonic environment. In this case the more
diverse habitat would be able to support a more diverse community due to an
abundance of additional environmental habitats (i.e. niches). We evaluated the
first mechanism by comparing membership among the plankton samples taken 9 days
apart (t=8 and t=17). While bacterioplankton communities were not
indentical identical
between the time points (Figure~\ref{fig:pcoa}), communities within a treatment
were more similar to each other between timepoints than any other
bacterioplankton community (treatment or timepoint). In addition, the control
...
relative abundance of the populations within each community. Second, an
analysis of the OTU relative abundance in biofilm and planktonic libraries
where OTUs are sorted by planktonic sample rank (Figure~\ref{fig:rank_abund}) shows
that the least abundant members of the plankton community were
rountinely routinely
highly abundant within the biofilm community. This was true for both algal and
bacterial communities, at all treatment levels and both timepoints. While we
did not (could not) specifically measure niche diversity within the biofilm
...
reported that planktonic diversity was significantly higher relative to biofilm
diversity (the opposite of what we found in our study). Given the differences
in the source of the planktonic community among studies, this result is not
suprising. surprising. While biofilm communities were
establishsed established on glass beads in
\citet{22237539} and glass slides (this study) over a similar time period (~21
days, \citet{22237539} and ~17 days this study) the origin of the planktonic
community in each study was very different. The \citet{22237539} study was
...
depending on how hydrologic pathways differed among precipitation events. In
this study the source community was a marine intake located approximately
200m from the shore during July when communities are more stable over the 17
day
perios period of the incubation. A separate study conducted in alpine and
sub-alpine streams of the Rocky Mountains clearly showed that stream plankton
communities reflected localized precipitation events and could be traced
largely to sources of soil communities of drainages within the watershed
...
appears, however, that sample-wise bacterial biofilm rarefaction curves may
exceed the integrated planktonic curve upon extrapolation and most exceed the
integrated planktonic curve at sampling depths where data is present for the
biofilm and
itegrated integrated planktonic library (Figure~\ref{fig:integ_rarefaction}). This
result is consistent with our conclusion that temporal
heterogenetity heterogeneity in the
plankton was not sufficient to
explain the higher diversity in the biofilm sample but would explain the
relative differences between planktonic and biofilm diversity found in
Besemer
et al. (2012) \citet{22237539}} compared to this study.
In addition, for this study, it is important to note that biofilm community
richness peaked at the intermediate treatment (C:P = 100) and appeared to
...
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 drove
changes in diversity or whether
resoruces resources drove diversity which altered
productivity. Rather we note that, as diversity decreased in the highest C
treatment bacterioplankton and biofilm membership became increasingly
similar. This suggests that
enviornments environments that contain high amounts of labile
C selected for fewer dominant taxa that came to dominate the biofilm
community, overwhelming the species sorting mechanisms that appeared to
dominate biofilm community assembly in all other treatments. Similarly, while
...
(Figure~\ref{fig:microscope}) 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
floculating flocculating DOC which has been shown to dominate high DOC
environments in nature, we did measure a
substanial 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 bacterial membership in the
highest C treatment.
\subsection{Lifestype \subsection{Lifestyle (biofilm or planktonic) Enriched OTUs}
There are only a few studies that attempt to compare biofilm community
composition and the
overlyng overlying planktonic community \citep{Besemer_2007,
22237539, Jackson_2001, Lyautey_2005}. Those studies illustrate community
composition among the two habitats are unique with very few taxa found in both.
This is consistent with our findings in this experimental system with a natural
marine planktonic source
commmunity. community. In addition, our study also evaluated
algal community composition which showed a similar result suggesting that both
the algal and bacterial biofilm communities form from phylogenetically unique
organisms that exist in low abundance in surrounding habitat (i.e. the
...
bacterial OTUs tended to have members in that were enriched in both the
plankton and the biofilm suggesting the phylogenetic coherence of lifestyle is
not captured at the level of Order. It should be noted however that taxonomic
annotations in reference
databasees databases and therefore environmental sequence
collections show little equivalency in phylogenetic breadth between groups at
the same taxonomic rank \citep{Schloss_2011}.
Unfortunatley, Unfortunately, at higher
taxonomic resolution (e.g. Genus-level), groups did not possess a sufficient
number of OTUs to evaluate coherence between taxonomic annotation and
lifestyle. Similar to the richness results, we found the
...
\subsection{Conclusion}
In summary this study shows mechanistic links between large scale community
level dynamics and the underlying constituent populations that compose them. We
found that
autrotrophic autotrophic pools and heterotrophic pools responded differently to
ammendments amendments of labile C as hypothesized. Notably while C
ammendments amendments
altered both pool size and membership of the bacterial communities we did not
see similar dynamics within the algal communities. Planktonic algae decreased
in response to C amendments presumably in response to increased
competition from a larger bacterial community, however there was not a
similar decrease in biofilm algal community. In addition membership of the
algal communities
betweeen between the plankton and biofilm lifestyles did not become
more similar in the algae as it did for the bacteria 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 (both heterotrophs and autotrophs alike) that are found in low abundance
in the neighboring communities. This membership was affected by resource
ammendements amendments for heterotrophic but not autotrophic 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
biogeohemical biogeochemical processes comes from those taxa
primarily if not exclusively while they are part of a biofilm. Our results point to lifestyle (planktonic or biofilm) as an
important trait that explains a portion of the exceptional diversity found in
snapshots used to characterize environmental microbial communities in space and
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\textbf{Figure 7}
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Principal coordinates ordination of bray-curtis distances for 23S rRNA plastid libraries and 16S rRNA gene libraries. OTU points are weighted pricipal coordinate averages (weights are relative abundace values in each sample) and the variance along each pricipal axis is expanded to match the site variance. Point annotations denote the amended C:P ratio for the mesocosm from which each sample was derived.
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\textbf{Figure 5 }Principal Principal coordinates ordination of bray-curtis distances for 23S rRNA plastid
libraries and 16S rRNA gene libraries. OTU points are weighted
pricipal principal
coordinate averages (weights are relative
abundace abundance values in each sample) and
the variance along each
pricipal principal axis is expanded to match the site variance.
Point annotations denote the amended C:P ratio for the mesocosm from which each
sample was derived.
test
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Rarefaction curves for each biofilm versus planktonic sequence library (SSU rRNA genes for bacteria, LSU rRNA plastid genes for algae). Samples from the same time and treatment are shown in the same panel. Panel strip text at top denotes time (15 or 24 days) and whether the column is for bacterial or algal sequence libraries. Panel strip text on the right denotes the amended C:P ratio for the mesocosm.
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\textbf{Figure 4}
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\textbf{Figure 4}
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Rarefaction curves for all biofilm versus plankton libraries. Each panel
represents a single C:P
treament treatment and time point. Richness is greater for all
planktonic communities when compared to corresponding biofilm communities.
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\textbf{Figure 1} Carbon subsidies in the form of glucose alleviate the dependence of
heterotrophic
bactria bacteria on algal derived
carbon C (C) exudates. This should result in
an increase in resource space and biomass for bacteria and a decrease in
resource space and biomass for algae due to increased competition for
phosphorus (P). We hypothesized that this predicted change in biomass pool size
of these two
grouops groups will result in changes in the plankton community
composition of both groups that will
propogate propagate to to the composition of biofilm
communities for both groups.
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\textbf{Figure 8} Rarefaction plots for all samples. Planktonic libraries have been integrated
such that the count for each OTU is the sum of counts across all samples.
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\textbf{Figure 6}
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\textbf{Figure 6} $log_2$ of
environment type lifestyle OTU
proportion mean ratios
(bioflm (biofilm or plankton) and
corresponding, adjusted p-values. Each point represents one OTU
proportion mean
ratio and points are grouped along the x-axis by Order. Outlined points have
adjusted p-values below a false discovery rate of 0.10.
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\textbf{Figure 3} The structure (density of cells and thickness of biofilm) and the amount of EPS
(cloudy material) increased with increasing
carbon C subsidies from the control to the
highest
carbon C treatment (C:P=500).
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\textbf{Figure 2} Increases in carbon resulted in decreases in A) planktonic algal biomass (estiated as Chl \textit{a} but B) not algal biomass present in the biofilm \textit{a} in each. In contrast both C) biofilm total biomass and D) number of planktonic bacterial cells increased with increasing carbon subsidies. Responses in treatments separated by different letters were statistically different from one another (p<0.05) as was the highest C:P treatment for planktonic bacterial abundance compared to the the control or the C:P = 10 treatment (p<0.05). The bacterial abudance sample for the C:P = 100 treatment was lost before analysis and is therfore not reported in panel d.
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Increases in C resulted in decreases in A) planktonic algal biomass (estimated
as Chl \textit{a} but B) not algal biomass present in the biofilm \textit{a} in
each. In contrast both C) biofilm total biomass and D) number of planktonic
bacterial cells increased with increasing C subsidies. Responses in treatments
separated by different letters were statistically different from one another
(p<0.05) as was the highest C:P treatment for planktonic bacterial abundance
compared to the control or the C:P = 10 treatment (p<0.05). The bacterial
abundance sample for the C:P = 100 treatment was lost before analysis and is
therefore not reported in panel d.
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\textbf{Figure 7}
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\textbf{Figure 7} Rank abundance plots. Each panel represents a single time point and C:P. The
"rank" of each OTU is based on planktonic sample relative abundance. Each
position along the x-axis represents a single OTU. Both the x and y axes are
scaled logarithmically.
test
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\textbf{Figure 9} Rank abundance plots for all date-amendment combinations.
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\textbf{Table 1}
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