Chuck Intro submitted version  over 9 years ago

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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.