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$^{13}$C-xylose assimilating organisms utilized $^{13}$C-xylose as a sole
carbon source.
\subsection{Cellulose degrader DNA
exhibits greater bouyant density shifts
further along the BD gradient upon
$^{13}$C incorporation than xylose degrader DNA}
Cellulose responders exhibited a greater shift in BD
(i.e. assimilated more
$^{13}$C per unit DNA) than xylose responders in
response to isotope incorporation (Figure~\ref{fig:shift}, p-value
1.86e$^{-06}$).
Cellulose $^{13}$C-cellulose responders
exhibited an shifted on average
shift of 0.0163 g/mL
(sd 0.0094) whereas xylose responders
exhibited an shifted on average
shift of 0.0097 (sd
0.0094).
One hundred percent For reference, 100\% $^{13}$C DNA
has a buoyant density shifts X.XX g/mL
higher than relative to the BD of its $^{12}$C counterpart. DNA
buoyant density BD increases
as
the its ratio of $^{13}$C
carbons to $^{12}$C increases. An organism that only
assimilates C into DNA from a $^{13}$C isotopically labeled source, will have a
greater $^{13}$C:$^{12}$C ratio in its DNA than an organism utilizing a mixture
of isotopically labeled and unlabeled C sources. Upon labeling, DNA from
the an
organism that
incoporates incorporates exclusively $^{13}$C will
shift its increase in buoyant density
position further relative to its original $^{12}$C-DNA position more than
the DNA
buoyant density shift from an organism that
doesn't does not exclusively utilize
isotopically labeled C. Therefore
the magnitude DNA buoyant density shifts
(labeled versus
unlabeled DNA)
indicate substrate specificity given our experimental design
(only as only one
substrate was labeled in each
amendment). amendment. We measured density shift
as the change in an OTU's density profile center of mass between corresponding
contol and labeled gradients. Density shifts, however, should not be evaluated
on an individual OTU basis as a small number of density shifts are observed for
each OTU and the variance of the density shift metric at the level of
...
utilizers for xylose (Figure~\ref{fig:shift}), and, each responder group
exhibits a range of substrate specificites (Figure~\ref{fig:shift}).
\subsection{Xylose responders at day 1 have more estimated rRNA operon copy
numbers per genome than xylose responders at days 3 and 7, and, Xylose
responders have more rRNA operon copy numbers
than cellulose responders.}
Estimated rRNA operon genome copy numbers per $^{13}$C-xylose responder OTU
genome and day of first response are correlated (p-value 2.02e$^{-15}$,
Figure~\ref{fig:copy}). $^{13}$C-xylose responder rRNA operon
geneome genome copy number is inversely related
to
time; that is, time of first response (p-value 2.02e$^{-15}$, Figure~\ref{fig:copy}). OTUs
that first respond at later time points have fewer estimated rRNA operons per
genome than OTUs that first respond earlier (Figure~\ref{fig:copy}). rRNA
operon copy number estimation is a recent advance in microbiome science
\citep{Kembel_2012}
and while the relationship of rRNA operon copy number per genome
with ecological strategy is well established \citep{Klappenbach_2000}.
Specifically, microorganisms Microorganisms with a high number of rRNA operons per genome tend
to be fast growers specialized to take advantage of boom-bust environments
whereas
a microorganisms with low rRNA operon copy
number numbers per genome
tends to occur in microorganisms that favor
slower growth under lower and more consistent nutrient input
\citep{Klappenbach_2000}. At the beginning of our incubation, OTUs with
estimated high rRNA operon copy numbers per genome or ``fast-growers''
assimilate xylose into biomass and with time slower growers (lower rRNA operon
number per genome) begin to respond to the xylose addition. Further,
$^{13}$C-xylose responders have more estimated rRNA operon copy numbers per
genome than $^{13}$C-cellulose responders (p-value 1.878e$^{-09}$) suggesting
xylose respiring
mircrobes microbes are generally faster growers than cellulose
degraders.
\subsection{Xylose responders are more abundant in the soil community than cellulose
responders}
$^{13}$C-xylose responders are generally more abundant members based on
relative abundance in bulk DNA SSU rRNA gene content than $^{13}$C-cellulose
responders (Figure~\ref{fig:shift}, p-value 0.00028). However, both
$^{13}$C-xylose
and $^{13}$C-cellulose responders were found in abundant and
rare OTUs
responded to $^{13}$C-xylose and $^{13}$C-cellulose
(Figure~\ref{fig:shift}). For instance, a \textit{Delftia} $^{13}$C-cellulose
responder is fairly abundant in the bulk samples (``OTU.5'',
Table~\ref{tab:cell}) with a mean bulk rank of 13 (\textit{i.e.} Table~\ref{tab:cell}). OTU.5 was on average the
13th most abundant
OTU) and a OTU in bulk samples. A $^{13}$C-xylose responder (``OTU.1040'',
Table~\ref{tab:xyl}) has a mean relative abundance in bulk samples of
2.85e$^{-05}$. Only one substrate responder ($^{13}$C-cellulose) was not found
in any bulk samples ("OTU.862", Table~\ref{tab:cell}). Of the
top 10
responders sorted by descending mean rank (essentially the 10 most abundant
responders
in the bulk samples), 8 are $^{13}$C-xylose responders and 5 of these 8
have mean ranks less than are consistently
among the 10
most abundant OTUs in bulk samples.
\subsection{Variation in bulk soil DNA microbial community structure is
significantly less than variation in gradient fractions}
Using a distance metric that incorporates relative abundance information
(weighted Unifrac metric, \citep{Lozupone_2005}) bulk sample beta diversity was
less than gradient fraction beta diversity
(p.value (p-value 0.003). Time was
significantly correlated to bulk sample phylogenetic profile variation (p-value
0.23, R$^{2}$ 0.63, Figure~\ref{fig:bulk_ord}) but
treatment (\textit{i.e.} the contrast between only
$^{12}$C additions with additions that included
isoptically isotopically labeled
substrates) substrates
was not (p-value 0.35).