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\section{Introduction}
Biofilms are diverse and complex microbial consortia that are the rule rather
than the exception for microbial lifestyle in many environments. Large and
small-scale architectural features of biofilms play an important role in their
ecology and influence their role in localized biogeochemical cycles
\citep{17170748}. While fluid mechanics have been shown to be important drivers
of biofilm structure and assembly \citep{hoedl_2011,19571890, 14647381}, it is
less clear how other abiotic factors such as resource availability affect
biofilm assembly. Aquatic biofilms are initiated with seed propagules that
originate within the planktonic community \citep{hoedl_2011, 22120588}. Thus,
how resource amendments influence planktonic communities has the potential to
influence the formation of microbial biofilms during community assembly.
test In a crude sense, biofilm and planktonic microbial communities can be broken
into two key groups: phototrophic eukaryotes (hereafter algae) and
heterotrophic bacteria and archaea. This dichotomy, while somewhat artificial,
has been shown to be a powerful paradigm for understanding community shifts
across ecosystems of varying trophic state \citep{Cotner_2002}. Heterotrophic
bacteria meet some to all of their organic carbon (C) requirements from algal
produced C while simultaneously competing with algae for limiting nutrients
such as phosphorous (P). The
presence of external C inputs, such as terrigenous C leaching from the
watershed \citep{Jansson_2008, Karlsson_2012} or C exudates derived from
macrophytes \citep{Stets_2008, Stets_2008b}, can alleviate bacterioplankton
reliance on algal derived C and shift the relationship from commensal and
competitive to strictly competitive
\citep[see][Figure~\ref{fig:conceptual}]{Stets_2008}. Under this mechanism
increased C supply should increase the resource space available to the
bacteria and lead to increased competition for P, decreasing P available for
algae {\textendash} assuming that bacteria are superior competitors for P as
has been observed \citep[see][Figure~\ref{fig:conceptual}]{COTNER_1992}. These
dynamics should result in the increase in bacterial biomass relative to the
algal biomass along a gradient of increasing labile C inputs.
Biofilms are diverse While these gross level dynamics have been discussed conceptually
\citep{Cotner_2002} and
complex microbial consortia that are the rule rather than to some extent demonstrated empirically
\citep{Stets_2008}, the
exception for microbial lifestyle effect that these shifts in
many environments. Large the bulk biomass pool have
on membership and
small-scale architectural features structure of
biofilms play an important role in their ecology the algal and
influence their role in localized biogeochemical cycles \cite{17170748}. While fluid mechanics have bacterial community has not been
shown directly evaluated in planktonic or biofilm communities. In addition, how
dynamics in planktonic communities are propagated to
biofilms during community
assembly is not well understood. Intuitively, shifts in planktonic community
composition should alter the available pool that can be
important drivers recruited into a
biofilm. For example, if bacterioplankton diversity increases, the number of
potential bacterial taxa that can be recruited to the biofilm
structure should also
increase, potentially increasing bacterial diversity within the biofilm.
Similarly, a decrease in P available to algae should decrease algal pool size,
potentially decreasing algal diversity and
assembly \cite{hoedl_2011,19571890, 14647381}, it is less clear how other abiotic factors such as resource availability affect biofilm assembly. Aquatic biofilms are initiated with seed propagules therefore candidate algal taxa that
originate within are available for biofilm formation. In addition, C in excess of resource
requirements may increase the
production of extra cellular polysaccharides
(EPS) by planktonic
community \cite{hoedl_2011, 22120588}. Thus, how resource amendments influence cells thus increasing the probability that planktonic
communities has cells
are incorporated into a biofilm by adhesion. Each of these mechanisms suggest
that an increase in labile C to the
potential system should result in increased
alpha diversity in both bacterioplankton and bacterial biofilm communities
while decreasing alpha diversity within both planktonic and biofilm algal
communities. To evaluate these ideas we designed this study to
influence test a) if
C subsidies shifted the biomass balance between autotrophs and
heterotrophs within the
formation biofilm or its seed pool (the plankton) and b) measure
how these putative changes in pool size altered membership and structure of
microbial biofilms the
plankton communities and affected recruitment of plankton during
biofilm
community assembly.
In a crude sense, biofilm and planktonic microbial communities can be broken into two key groups: phototrophic eukaryotes (hereafter algae) and heterotrophic bacteria and archaea. This dichotomy, while somewhat artificial, has been shown to be a powerful paradigm for understanding community shifts across ecosystems of varying trophic state \cite{Cotner_2002}. Heterotrophic bacteria meet some to all of their organic C requirements from algal produced C while simultaneously competing with algae for limiting nutrients such as P. The presence of external C inputs, such as terrigenous carbon leaching from the watershed \cite{Jansson_2008, Karlsson_2012} or C exudates derived from macrophytes \cite{Stets_2008, Stets_2008b}, can alleviate bacterioplankton reliance on algal derived carbon and shift the relationship from commensal and competitive to strictly competitive (\citet{Stets_2008}, Figure 1). Under this mechanism increased carbon supply should increase the resource space available to the bacteria and lead to increased competition for P, decreasing P available for algae – assuming that bacteria are superior competitors for P as has been observed (\citet{COTNER_1992}, Figure 1). These dynamics should result in the increase in bacterial biomass relative to the algal biomass along a gradient of increasing labile carbon inputs.
While these gross level dynamics have been discussed conceptually \cite{Cotner_2002} and to some extent demonstrated empirically \cite{Stets_2008}, the effect that these shifts in the bulk biomass pool have on membership and structure of the algal and bacterial community has not been directly evaluated in planktonic or biofilm communities. In addition, how dynamics in planktonic communities are propagated to biofilms during community assembly is not well understood. Intuitively, shifts in planktonic community composition should alter the available pool that can be recruited into a biofilm. For example, if bacterioplankton diversity increases, the number of potential bacterial taxa that can be recruited to the biofilm should also increase, potentially increasing bacterial diversity within the biofilm. Similarly, a decrease in P available to algae should decrease algal pool size, potentially decreasing algal diversity and therefore candidate algal taxa that are available for biofilm formation. In addition, carbon in excess of resource requirements may increase the production of extra cellular polysaccharides (EPS) by planktonic cells thus increasing the probability that planktonic cells are incorporated into a biofilm by adhesion. Each of these mechanisms suggest that an increase in labile carbon to the system should result in increased alpha diversity in both bacterial plankton and bacterial biofilm communities while decreasing alpha diversity within both planktonic and biofilm algal communities. To evaluate these ideas we designed this study to test a) if carbon subsidies shifted the biomass balance between autotrophs and heterotrophs within the biofilm or its seed pool (the plankton) and b) measure how these putative changes in pool size altered membership and structure of the plankton communities and affected recruitment of plankton during biofilm community assembly.