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wat e r r e s e a r c h 4 5 ( 2 0 1 1 ) 5 2 1e5 3 4
Avai lab le a t www.sc iencedi rec t .com
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Biostabilization and erodibility of cohesive sediment depositsin wildfire-affected streams
M. Stone a,*, M.B. Emelko b, I.G. Droppo c, U. Silins d
aDepartment of Geography and Environmental Management, University of Waterloo, Waterloo, Ontario, Canada N2L3G1bDepartment of Civil and Environmental Engineering, University of Waterloo, Waterloo, Ontario, Canada N2L3G1cNational Water Research Institute, Environment Canada, Burlington, Ontario, Canada L7R4A6dDepartment of Renewable Resources, University of Alberta, Alberta, Canada T6G2H1
a r t i c l e i n f o
Article history:
Received 10 May 2010
Received in revised form
8 September 2010
Accepted 12 September 2010
Available online 21 October 2010
Keywords:
Wildfire
Sediment transport
Biostabilization
Cohesive sediment
Colmation
Critical shear stress
Treatability
Water treatment
* Corresponding author.E-mail address: [email protected]
0043-1354/$ e see front matter ª 2010 Elsevdoi:10.1016/j.watres.2010.09.016
a b s t r a c t
The erosion characteristics and bed stability of wildfire-affected stream sediment were
measured in an annular flume. Biofilms were grown in the flume on cohesive streambed
sediments collected from a wildfire affected stream and a reference undisturbed stream in
southern Alberta, Canada. Examined factors that influence sediment erosion, settling and
bed stability included applied shear stress, geochemical and physical properties of the
sediment, floc structural characteristics and consolidation period (2, 7, 14 days). Erosion
characteristics and sediment properties were strongly influenced by wildfire, consolidation
period and bed biostabilization. The fire-modified sediment was more resistant to erosion
than the referenceunburned sediment. Settling velocitieswere lower in theburnedsediment
due to higher organic content and porosity. The critical shear stresses for erosion were 1.6
and 1.8 times higher for the burn-associated sediment after 7 and 14 days of consolidation.
The differences are related to the greater degree and spatial extent (depth) of biofilm
attachment in the burned sediment. Erosion depths were 4e8 times higher in burned sedi-
ment as a result of wildfire-associated biostabilization.
ª 2010 Elsevier Ltd. All rights reserved.
1. Introduction stability of cohesive sediment deposits is governed by factors
In aquatic systems, many contaminants of concern are bound
to and transported by cohesive sediment (inorganic and
organic particles<63 mm). Depending upon the hydrodynamic
and biogeochemical characteristics of the system, fine-
grained materials flocculate in the water column (Droppo,
2001) and upon settling are the building blocks of sediment
deposits referred to as surficial fine-grained laminae (SFGL)
(Stone and Droppo, 1994). These deposits contribute to both
external and internal colmation that occurs at the interface
between ground water and surface water (Brunke, 1999). The
(M. Stone).ier Ltd. All rights reserved
such as electrochemical reactions (Mehta, 1989), consolida-
tion (Droppo and Amos, 2001), dewatering (Tolhurst et al.,
2000) and biostabilization (Dade and Norwell, 1990; Paterson,
1997). When the critical shear stress for erosion (sc) is excee-
ded (Stone et al., 2008), these deposits are remobilized within
the water column where they can adversely impact down-
stream lotic environments (Wood and Armitage, 1997).
In aquatic systems, sediment resuspension is a function
of shear stress and sediment characteristics such as grain
size, density and mineralogy (Partheniades, 1990). Benthic
flora and fauna are also being increasingly recognized as
.
wat e r r e s e a r c h 4 5 ( 2 0 1 1 ) 5 2 1e5 3 4522
important contributors to sediment stability (Paterson, 1997).
Benthic bacteria, microalgae and macrofauna secrete poly-
mers (extracellular polymeric substances; EPS) that bind
mineral grains (biostabilization) together in a mucilaginous
matrix (Dade and Norwell, 1990; Paterson, 1997). For bio-
stabilization to occur, threshold concentrations of organic
compounds that drive microbial metabolism must be avail-
able for energy maintenance and microbial enzyme induc-
tion (Schmidt et al., 1985). Biofilms develop on a variety of
interfaces by attachment processes; aquatic sediments, in
particular, provide excellent substrata for biofilm growth
(Neu, 1994).
The development of biofilms on aquatic sediments can
change the characteristics of deposited sediment (i.e. particle
structure, morphology, size, porosity, shape, degree of consol-
idation) and influence erosion rates (Droppo and Amos, 2001;
Droppo et al., 1997; Lau, 1995). Several studies have demon-
strated that biostabilization can significantly increase the
energy required to erode sediments by horizontal shear (Amos
et al., 2004; Gerbersdorf et al., 2008). While the influence of
biogenic sediment stabilization on sediment erosion and
contaminant transport has been extensively examined in
marine environments (Underwood and Paterson, 2003; Friend
et al., 2003), much less is known about the role of biofilms in
stabilizing sediment deposits in freshwater systems (Droppo
et al., 2007; Droppo, 2009; Gerbersdorf et al., 2008); particularly
those formed in streams draining forested landscapes
impacted by severe, landscape-altering disturbances such as
wildfire.
The frequency and severity of large-scale natural distur-
bances such as wildfire in many forested regions of the globe
has significantly increased in recent decades (Westerling et al.,
2006). Becauseof the severity andmagnitudeofwildfire related
landscape disturbances, sediment fluxes (Silins et al., 2008,
2009) are modified at rates and magnitudes that cause
profound and often irreversible changes in river system func-
tion (DeBanoet al., 1998; Bladon et al., 2008) anddrinkingwater
treatability (Emelko et al., 2011). For example, Blake et al. (2009)
demonstrated that post-fire sediment and nutrient transport
dynamics in streams are strongly related to coarsening of the
effective particle size distributions in burned material via the
aggregation of fines into composite particles. They reported
that burned composite particles (predominantly <63 mm) had
higher concentrations of bio-available phosphorus than
unburned sediment. The subsequent increased flux of bio-
available phosphorus and terrestrial organic matter from hill-
slopes to streamsproducedbywildfiredisturbance increase in-
streamstorageandavailability of particle-associatednutrients
(Petticrew et al., 2006), which increases biofilm formation and
growth rates, particularly in areaswhere the forest canopyhas
been lost (Minshall, 2003). Little information currently exists
regarding biofilm formation and its potential role in stabilizing
cohesive sediment deposits and associated contaminants in
streams draining fire impacted catchments. Given the
increasing severityandspatial extentof landscapedisturbance
by wildfire at the global scale and its impacts on sediment
availability, transport and storage in streams, knowledge of
processes that govern sediment transport is critical to quantify
and model sediment and associated contaminant fluxes from
fire impacted watersheds to downstream environments. Such
information is alsohighly relevant to the impacts ofwildfire on
reservoir management and drinking water treatability.
Here, wildfire and bed age were studied using an annular
flume to elucidate their impacts on the stability of cohesive
stream sediment deposits. The objectives of this work were (1)
to quantify the physical (particle size, morphology, density,
porosity, settling velocity) and geochemical (major element
composition, mineralogy, total carbon) properties of burned
and unburned river sediment (2) to characterize the microbial
communities comprising the sediment-associated biofilms and
(3) determine the effect of biofilms on the transport (sc), depo-
sition (settling behaviour) and erodibility (erosion rate) of the
two sediment types.
2. Methods
2.1. Study area and sample collection
The 2003 Lost Creek wildfire burned >21,000 ha in the
Crowsnest Pass, Rocky Mountain region of southwestern
Alberta. It was particularly severe and consumed most forest
cover and floor organic matter across much of the headwater
regions of theOldmanRiver Basin. One of thewatersheds, Lynx
Creek, was severely impacted (67.3% burned), which dramati-
cally altered the discharge and sediment and nutrient charac-
teristics of the river (Silins et al., 2008, 2009; Bladon et al., 2008).
Increased post-fire light levels and particle-associated nutrient
inputs to the stream have significantly increased the presence
and abundance of biofilms in Lynx Creek compared to neigh-
bouring unburned watersheds (Silins et al., in review). Here,
cohesive sediment and riverwaterwere collected in LynxCreek
(burned watershed) and the Castle River (reference unburned
watershed) in 2007 (4 years post-fire). Surface deposits of fine
sediment were collected with a plastic scoop, immediately
refrigerated at 4 �C then transported within one week to the
Canada Centre for Inland Waters in Hamilton, Ontario where
they were studied using an annular flume.
2.2. Experimental procedure
A stainless steel annular flumewas used tomeasure sc, erosion
rate and erosion depth of burned and unburned river sediment
(Lau, 1995). The outside diameter of the flume is 2 m and the
trough is 20cmwideand12cmhigh.A clear glass top,whichfits
inside the trough, is lowered until it touches the water surface
then rotated to generate flow. Calibration of the flume was
described by Lau and Droppo (2000). Sediment and river water
were placed in the flumeand then the coverwas rotated at high
speed. After the sediment bed was completely entrained and
well mixed, the cover rotation was reduced gradually and then
stopped. Suspended solids in the water column settled under
low shear to form a cohesive sediment bed approximately 1 cm
thick. Thebedwas thenallowed to settle for three consolidation
periods (2, 7 and 14 days) during which eight wide-spectrum
fluorescent grow lights (40Weach totalling ameasured1250 Lx)
were activated above the flume for a 12 h light and 12 h dark
scenario to promote biofilm growth.
The flume experiments were conducted in sequence to
simulate sediment transport conditions that could occur in
wat e r r e s e a r c h 4 5 ( 2 0 1 1 ) 5 2 1e5 3 4 523
stream reaches where cohesive sediments are deposited and
biofilms form and subsequently resuspend at conditions of
higher flow. Accordingly, for each sediment type (burned/
unburned), the first consolidation period studied was 2-days.
Thereafter, the flume was rotated and shear stress increased
incrementally (every 10 min) until all of the bed material had
eroded; thereby enabling the calculation of Tcrit. After the
estimated critical bed shear level had been applied for 10 min
the flume was stopped to monitor still water settling. Sus-
pended solids were collected every 2 min for 10 min. Two
subsequent samples were then collected at 5 min intervals
and four samples collected at 10min intervals thereafter. This
process was repeated for subsequent, longer consolidation
periods (i.e. the 7-day consolidation period experiment was
conducted after the 2-day experiment; the 14-day experiment
was conducted after the 7-day experiment). The same sedi-
ment remained in the flume for all three erosion experiments
conducted for a given source sediment. The river water was
not changed except when suspended solids were collected
from the sampling port and then an equal amount of river
water was added to the flume to maintain a constant water
level. This approach is non-ideal for simulating the biogeo-
chemical environment and biofilm growth in natural streams;
nonetheless, it provides a rigorous quantitative assessment of
sediment erosion under controlled hydrodynamic conditions
and a relative comparison of the effects of biofilms on the
erosion of the burned and unburned sediments.
The light conditions (intensity and duration) in forested
regions of the southern Rockies are highly variable and
dependent upon site specific conditions such as the time of
year, slope, aspect, canopy cover, forest age class and degree
of burn. In this study, fluorescent lights were not used to
simulate actual light conditions present at the stream surface
at a specific location in the complex forested environment
from which the sediment samples were obtained, but rather
to generate biofilm growth in the flume under controlled
conditions (12 h on and 12 h off) to compare and contrast
potential biofilm generation in the two systems.
Changes in sediment bed stability were observed through
windows on the side of the flume and with a video imaging
Lennox� boreoscope. The suspended sediment concentration
wasmonitoredcontinuouslyusinganoptical backscatter (OBS)
turbidimeter. Suspended solids concentrations were also
measured gravimetrically from samples collected directly
from theflume througha sampleport at 1, 5 and9minafter the
start of every 10-min shear interval.
Type I erosion is defined as an erosion rate that exponen-
tially decays to zero and is comprised of both Type 1A (erosion
of the loosely bound floc layer referred to as SFGL) and Type 1B
(erosion of the stronger bed [or armour layer] below the SFGL)
erosion (Villaret and Paulic, 1986). Here, the critical shear
stress for erosion (sc) for Type 1B erosion was determined
using visual and boreoscope observations as well as the
temporal suspended sediment concentration data for each
sediment type during each of the annular flume experiments.
2.3. Settling analysis
Total suspended solids concentrations over time were plotted
and the characteristic kinetic plots for still water settling were
examined. Still water settling was considered as the period
after the flume was stopped and the flow of water could no
longer be observed (360 s and thereafter). Mass settling during
each experiment was evaluated by calculating the appropriate
(second order in all cases as discussed below in the results and
discussion) kinetic decay constant (k), which is obtained by
regressing the inverse of the suspended solids concentration
(L mg�1) at each sampling point (360, 480, 600, 900, 1200 and
1800 s; in some cases, also 2400, 3000, and/or 3600 s) on time (s)
and determining the slope by least squares linear regression
to yield k (L mg�1 s�1). The TSSmass settling rate at each point
in time was then calculated by multiplying k by the square of
the measured suspended solids concentration. The rates of
change in mass settling rate during still water settling in the
burned and unburned systems with the various consolidation
periods were compared using an F-test for equality of slopes
(SAS/STAT� Version 9.1).
2.4. Physical and geochemical sediment properties
Sediment bulk density after the growth/consolidation periods
was used to evaluate the influence of consolidation (and
potentially biofilm formation) on bed stability. Sediment
samples were settled in 10 cm of river water in each of three
glass beakers to form beds with similar thickness to those
formed in the flume. Bulk density profiles were measured in
1 mm increments (Berkhout, 1994).
Eroded floc settling velocity was determined using
a settling column interfaced with a Nikon SMZ-2T (Nikon
Canada Inc., Mississauga, Canada) stereo-scopic microscope
and Open Lab image analysis system (Improvision, Coventry,
UK) to size and track particle settling trajectories for the
measurement of settling velocity (Droppo et al., 1997). Stokes’
law was used to obtain density and porosity estimates (Li and
Ganczarczyk, 1987).
Surface deposits of cohesive sediment were collected
from a wildfire-affected stream and a reference undisturbed
stream and immediately submitted to a commercial labora-
tory for grain size and geochemical analyses to characterize
and compare the sediment geochemistry between the two
rivers. The grain size distribution of the two sediment types
was determined using a Malvern Mastersizer Model 2000.
The concentration of major elements (Al2O3, CaO, Cr2O3,
Co3O4, CuO, Fe2O3, K2O, MgO, MnO, Na2O, NiO, P2O5, SiO2,
TiO2) was determined by X-ray fluorescence spectrometry.
The sediment loss on ignition (LOI) was determined at 475 �Cfor 12 h. Themineralogical composition of the sedimentswas
evaluated by X-ray diffraction (Philips X’pert PW3040-PRD
diffractometer with a Cu X-ray source operated at 40 kV and
50 mA). Certified reference materials USGS GXR-1, GXR-2,
GXR-4 and GXR-6 were analyzed at the beginning and end of
each batch of samples. Internal control standards were
analyzed after every 10 samples and a duplicate was
analyzed after every 10 samples. Volatile organic carbon was
determined by heating a 0.1 g sample in a pure oxygen
environment at 380 �C, releasing the volatile organic carbon
species, binding them with the oxygen to form CO and CO2,
the majority being CO2. Carbon was measured as carbon
dioxide in the IR cell.
wat e r r e s e a r c h 4 5 ( 2 0 1 1 ) 5 2 1e5 3 4524
2.5. Microscopy
Eroded floc morphology (size distribution and structure) was
examined by light microscopy (Zeiss Axiovert 100 fitted with
a flow cell interfaced directly to the flume) (Droppo et al.,
2007). Environmental scanning electron microscopy (ESEM;
ElectroScan model 2020 operated at 20 kV, ElectroScan
Corporation, Wilmington, MA, USA) was used to observe fine-
scale structural characteristics of the eroded flocs using
a Peltier stage cooled to 1 �C (Leppard, 1986). Transmission
electron microscopy (TEM) was used to determine the micro-
structure and internal composition of the eroded flocs.
Ultrathin sectionswere imaged in transmissionmode (TEM) at
an accelerated voltage of 80 kV (JEOL 1200 Ex II TEMSCANTEM)
(Leppard, 1986).
2.6. Biofilm mass
Biofilm is a heterogeneous andamorphousmaterial comprised
of living organisms and other solid materials (e.g., organic
substancessuchasEPS, inorganicsolids suchasclays). Liuetal.
(1994) suggested determination of total attached poly-
saccharide (carbohydrate) as an indirect measure of biofilm
mass because biofilms contain relatively large amounts of
polysaccharide. Here, water samples from the flume were
collected in a polyethylene bottle and filtered onto Whatman
GF/C glass-fiber filters and frozen (220 �C) for subsequent
analysis (performed on the filters within one week). The filter
residue was extracted with acetone and assessed at a wave-
length of 663 nm (Pacepavicuis et al., 1997). This method is
particularly suitable for the determination of total poly-
saccharide content in suspended biofilm because it is sensitive
(i.e. a few mg is sufficient) and specific (i.e. there is little inter-
ference from environmental factors such as salt and water
hardness). The amount of eroded biofilmmass is expressed as
mass of total polysaccharide as glucose per liter of water
(mg L�1) and is determined directly from a glucose calibration
curve. The contribution of biofilm mass (as glucose) to sus-
pended solids concentration in the burned and unburned
sediment was evaluated using least squares linear regression
and an F-test for equality of slopes (SAS/STAT� Version 9.1).
2.7. Microbial community analysis
Phospholipid fatty acids (PLFA) are the primary constituents of
cell membranes of all living cells. Different groups of micro-
organisms synthesize varieties of PLFA through different
biochemical pathways. Some PLFAs can be used as “signa-
tures” to analyze changes in viable microbial biomass and
community structure (Tunlid and White, 1992); accordingly,
changes in PLFA profiles are indicative of changes in the total
viable microbial community, thereby making PLFA an effec-
tive tool for monitoring microbial responses to environmental
conditions. Here, lipid extraction and PLFA analyses were
performed using the modified Bligh and Dyer-method (1959)
as described by White et al. (1979).
Themicrobial communities comprising the biofilm formed
on the two sediment types (burned and unburned) after 2, 7
and 14 days of consolidation were determined using a phos-
pholipid fatty acid (PLFA) technique (White et al., 1979).
Biofilmwas formed using a plastic bin containing a 1 cm layer
of bed sediment covered by a water layer of 2 cm. The bins
were placed under the grow lights and frosted glass slides
were placed on the surface of the sediment to allow biofilm
growth for the designated consolidation periods of 2-, 7-, and
14-days.While biofilm growth on slides is likely different from
that formed in streams where flow conditions, light and
nutrient levels are variable over short temporal scales (days to
weeks), this method provides a comparison of relative differ-
ences in biofilm growth that may form on various sediment
types (Lau, 1995; Lau and Liu, 1993). Twenty-four slides were
added to the plastic bins (8 for each consolidation period). Of
these eight slides: the biofilm from four was combined and
analyzed by PLFA (Microbial Insights; Rockford, Tennessee) to
yield a composite determination of viable microbial biomass
and a PLFA profile reflecting the proportions of organisms
present in the sample, according to six PLFA structural groups
(monoeonic, terminally branched saturated, branched mon-
oeonic, mid-chain branched saturated, normal saturated, and
polyeonic). Biofilm from the remaining four slides was
analyzed by TEM and ESEM.
3. Results and discussion
3.1. Physical and geochemical sediment properties
The geochemical composition and claymineral assemblage of
sediment in aquatic systems are largely dependent upon the
general characteristics of the source area which include
geology, weathering, vegetation, soils, mass wasting
processes and land use (Griggs and Hein, 1980). Resulting
differences in the sediment properties (density, grain size,
geochemistry) will influence sediment transport characteris-
tics. Themajor element composition of the study sediments is
presented in Table 1 and indicates that Lynx Creek sediment
(burned) has higher concentrations of CaO (9.16%) and carbon
(LOI ¼ 16.7%) but lower concentrations of SiO2 (53.74%) and
Al2O3 (8.79%) than Castle River (unburned) sediment. Lynx
Creek sediment contains less quartz than Castle River sedi-
ment, but elevated levels of dolomite andmuscovite (Table 2).
The measured volatile organic carbon content of Lynx Creek
was 4.73% compared to <0.05% in Castle River.
The median density, porosity and settling velocity of the
burned and unburned sediments after consolidation are pre-
sented in Table 3. While the geochemical composition of both
sediments varied slightly, their densities were similar. The
median settling velocity and porosity of the unburned sedi-
ment were higher than of burned sediment. In contrast to the
results presented herein, Blake et al. (2007) examined the
impact of wildfires on the effective size distribution of burned
and unburned soils and reported that burned soils had signif-
icantly higher settling velocities than unburned particles of
equivalent diameter; they attributed this to increased density
and decreased organic matter and pore space in burned soils.
Wildfires can influence the structure and stability of soil
aggregates but the degree of aggregationwill vary as a function
burn temperature, soil depthaswell as themineral andorganic
properties of the soil (Andreu et al., 2001; Fox et al., 2007).
However, once soil aggregates enter receiving streams they are
Table 1 e Major element composition of fine sedimentdeposits in Burned and Unburned Watersheds (%).
Major element Burned Unburned
Al2O3 8.79 10.49
CaO 9.16 2.02
Cr2O3 0.01 0.01
Co3O4 0.005 0.005
CuO 0.005 0.005
Fe2O3 2.92 3.59
K2O 3.05 3.24
MgO 4.87 2.88
MnO 0.055 0.052
Na2O 0.86 0.99
NiO 0.003 0.003
P2O5 0.12 0.11
SiO2 53.74 71.26
TiO2 0.33 0.51
LOI 16.66 4.44
Table 3 e Physical characteristics of eroded sediment.
Consolidationtime(days)
Median settlingvelocity(mm s�1)
Medianporosity
(%)
Mediandensity(g cm�3)
Burned 2 2.22 84.7 1.09
7 2.81 85.6 1.09
14 2.96 93.3 1.05
Unburned 2 3.18 89.3 1.08
7 3.26 91.3 1.07
14 3.82 93.4 1.04
wat e r r e s e a r c h 4 5 ( 2 0 1 1 ) 5 2 1e5 3 4 525
influenced by a range of physical, chemical and biological
processes that alter the grain size distribution and biogeo-
chemical nature of the particles (Blake et al., 2009). Here, flocs
were formed within the flume (i.e. not eroded soil aggregates)
with a significant microbial component (biofilm interactions).
The higher organic content (Table 1) for Lynx Creek is likely
responsible for the larger floc size and increased floc porosity
resulting in a slower settling velocity (although the density
analysis showed no difference between rivers). Moreover, the
river sediments assessed in this study are four years post-fire
whereas Blake et al. (2009) investigated recently burned soils.
3.2. Morphology and physical characteristics oferoded flocs
Eroded flocs were collected during the flume experiments to
assess changes in floc structure between rivers and over
different consolidation times. Within these experiments it is
difficult to compare eroded floc morphology, size, settling
velocity, porosity and density for a given shear, because for
a given shear value, flocs will originate from different depths
within the bed where different physicochemical and biological
properties may exist. Further, a comparison of floc character-
istics at sc also provides little insight as these comparisons are
different for each consolidation period and bed type (burned
and unburned) resulting in a wide range of properties given the
different forces affecting the floc. Accordingly, differences in
floc properties can only be viewed on an average trend basis.
Table 2 eMineralogy of fine sediment deposits in Burnedand Unburned Watersheds (%).
Burned Unburned
Quartz 23 42
Calcite 1 1
Dolomite 8 2
Albite 7 12
Microcline 15 14
Muscovite 41 22
Chlorite 5 7
In general, fire modified eroded flocs were larger than
unburned flocs and both floc types decreased in size with
increasing shear levels.Using threedifferentmicroscopeswith
a range of resolution, it was observed (e.g., Fig. 1) that Castle
Creek exhibited more inorganic networks relative to Lynx
Creek, which possessed substantially more organic networks
encompassing both cellular material and extracellular poly-
meric substances in the form of fibrils (see TEM image Fig. 1c
and f). COM images (Fig. 1a and d) show the eroded Lynx Creek
flocs to be more diffuse and irregular in shape while the ESEM
images (Fig. 1b and e) showmore organic coatings on the Lynx
Creek flocs relative to the inorganic dominated Castle Creek
flocs. The above differences relative to floc structure were
consistent over time, however, the level of organic dominance
increased with time of consolidation/growth.
3.3. Biofilm analysis
Visual inspection of streambeds draining unburned (Fig. 2a)
and burned (Fig. 2b) catchments even 5 years post-fire clearly
indicated the possibility of significant differences in
streambed-associated biomass amount, composition and
activity. Biomass can attach to sediment in the form of biofilm
and several studies have demonstrated that biostabilization
can significantly increase the energy required to erode sedi-
ments by horizontal shear (Amos et al., 2004; Gerbersdorf et al.,
2008; Droppo, 2009). A simple estimate of biofilmaccumulation
on the sediment bed within the flume expressed as mass of
total polysaccharide as glucose per liter of water indicated that
burned sediments had significantly higher biomass (glucose
mass) per suspended solids mass than unburned sediments
( p < 0.0001; Fig. 3). PLFA analyses of the same biofilms did not
yield a clear difference in accumulated biomass associated
with the burned and unburned sediments (Fig. 4). However,
only a very limited number of samples could be processed
thereby precluding a more thorough assessment of intra- and
inter-sample variability.
Themicrobial community structure comprising the biofilms
formed on the unburned and burned sediments with various
consolidation periods is organized according to the PLFA
structural groups and presented in Fig. 5. As observed with the
total accumulated biomass evaluated using PLFA (Fig. 4), few
differences in microbial community structure between the
unburned and burned catchments and the various consolida-
tion periods were observed. The only notable difference
appeared tobean increase in themid-chainbranchedsaturated
PLFA structural group indicative of Actinobacteria (e.g.,
Fig. 1 e Eroded Castle River (a) and Lynx Creek (b) flocs using conventional optical microscopy. Representative ESEM images
of 14-day Castle River (c) and 14-day Lynx Creek sediment (d). TEM images of 14-day Castle River (e) and 14-day Lynx Creek
sediment (f).
wat e r r e s e a r c h 4 5 ( 2 0 1 1 ) 5 2 1e5 3 4526
Actinomycetes spp.) and sulfate-reducing bacteria in the biofilms
formed on burned relative to unburned sediments (Fig. 6).
Increasedamounts ofbacteria likeActinomycetes spp. inbiofilms
obtained on burned relative to unburned sediments are not
surprising given that Actinomycetes spp. form conidia (asexual
spores) that may help them better survive harsh soil environ-
ments such as those exposed to dessication and heat
(Alexander, 1998) that result fromwildfire.
Increases in Actinomycetes spp. may be of concern when
impacted waters are used as drinking water sources because
thesebacteria are associatedwithearthy/mustyodors inwater
and may contribute to formation of geosmin and 2-
methylisoborneol (MIB), which are major sources of taste and
odor causing compounds in drinking water (Zaitlin and
Watson, 2006). Biofilm growth in plastic bins occurred under
quiescent (no flow) conditions on a glass substratum which
likely resulted in differences observed between the biomass
amount, composition andactivity in the lab compared to those
found in nature. Glass is a relatively smooth substratum;
however, in many circumstances biofilms accumulate more
readily on rougher substrata (e.g., Percival et al., 1999).
Accordingly, biomass growth in the source watersheds was
likely underpredicted by the simple approaches used herein to
estimate biomass accumulation. Nonetheless, caution should
Fig. 2 e Typical biofilm accumulation in riverbeds draining (a) reference and (b) burned catchments five years post-fire.
wat e r r e s e a r c h 4 5 ( 2 0 1 1 ) 5 2 1e5 3 4 527
be exercised when considering increases (or lack thereof) in
amounts of total estimated biomass (Figs. 3 and 4) or in
biomass associated with specific structural groups such as
those indicative of Actinomycetes spp. (Fig. 6) because differ-
ences in biomass are not necessarily indicative of biomass
activity. For example, even when substantial differences in
substratum roughness exist and may contribute to significant
differences in biomass accumulation, differences in biomass
activity may still be insignificant (e.g., Emelko et al., 2006).
Regardless of how representative biofilm mass formed on
a glass substratum is of the biofilm formed in the source
watersheds, it is essential to note that secondary metabolite
(geosmin and MIB) production by Actinomycetes spp. in a given
situation is also not necessarily linked to the amount of
biomass present (Zaitlin and Watson, 2006). Accordingly,
further investigation of the potential risks (e.g., increased
propensity for taste and odor events as related to sediment-
associated microorganism transport) resulting from land
disturbances to downstream drinking water supply and
further in situ analysis and characterization of biofilms in fire
impacted streams is warranted.
Fig. 3 e Estimated contribution of biofilm mass (as glucose)
to suspended solids concentration arising from the Castle
River (unburned) and Lynx Creek (burned) sediments with
various bed consolidation periods.
3.4. Critical erosion values
Flume experiments were conducted to determine sc of cohe-
sive sediment deposits in a wildfire impacted and reference
stream. Suspended solids concentrations increased exponen-
tially with applied bed shear stress, but the degree of erosion
was influenced by bed age and sediment type (Fig. 7). The scvalues for Type 1A and 1B erosion varied both within (i.e.
different consolidation/biostabilization periods) and between
sediment types (Table 4). The level of shear stress required for
Type 1A erosion increased with consolidation time for the
burned and unburned sediment, but higher shear stresses
were required to erode the SFGL of the burned sediments
compared to unburned (Fig. 7). During the 2-day consolidation
experiments, the level of shear stress required to produceType
1B erosion was comparable for both sediment types; however,
the measured sc was 1.6 and 1.8 times higher for the burn-
associated sediment after 7 and 14 days of consolidation,
respectively than for unburned sediment (Table 4). Accord-
ingly, sediment resistance to erosion generally increased with
increased consolidation time regardless of sediment type;
however, this increase was more pronounced for the burn-
associated sediment.
The observed increase in shear stress required for Type 1A
and 1B erosion to occur with increasing bed age is strongly
related to the nature of the biofilm and its associationwith the
underlying sediments. Boreoscope observations of sediment
movement as a function of applied shear stress show that
surface biofilm began to erode before the underlying sedi-
ments. The process of biofilm erosion typically began at low
shear with the creation of small fractures on the biofilm
surface. With increasing shear, segments of biofilm would
partially dislodge and the biofilm would roll up upon itself in
long narrow segments up to a few cm in length until it
detached completely from the sediment bed. As it was being
dislodged from the bed, the eroded Lynx Creek biofilm
appeared to contain more sediment than Castle River biofilm.
These visual observations coupledwith the flumedata suggest
that the degree and spatial extent (depth) of biofilm attach-
mentwere greater on the burned (Lynx Creek) sediment. Table
4 shows that burned sediment was approximately twice as
Fig. 4 e Estimated amount of bacterial and eukaryotic biomass on Castle River (reference/unburned) and Lynx Creek
(burned) sediment with various bed consolidation periods as determined by phospholipid fatty acid (PLFA) analysis.
wat e r r e s e a r c h 4 5 ( 2 0 1 1 ) 5 2 1e5 3 4528
resilient to the applied bed stress for the 7-day (1.6 times) and
14-day (1.9 times) deposit compared to the unburned sedi-
ment. The higher sc values required to erode Lynx Creek sedi-
ment suggests that the sediment-pore biofilmcomplex ismore
integrated and the biological community associated with the
SFGL and eroded flocs in these deposits is more active than in
unburned materials. Networks of EPS can permeate void
spaces and promote inter-particle linkages (Gerbersdorf et al.,
2008, 2009) and the resultant increased level of attachment
within and between deposited flocs can lead to increased bed
stability (Droppo, 2009). Recent sediment biostabilization
studies in freshwaterwavedominated (Droppoet al., 2007) and
Fig. 5 e Relative percentages of total PLFA structural groups
in Castle River (reference/unburned) and Lynx Creek
(burned) sediments with various bed consolidation
periods. Structural groups are assigned according to PLFA
chemical structure, which is related to fatty acid
biosynthesis.
riverine (Gerbersdorf et al., 2008, 2009) environments show
that bacterial EPS production correlateswellwith bed stability.
Droppo (2009) examined biofilm structure and bed stability of
five contrasting freshwater sediments and demonstrated that
Fig. 6 e Biomass composition of mid-chain branched
saturated structures of PLFA (indicative of Actinobacteria
[e.g., Actinomycetes spp.] and sulfate-reducing bacteria)
comprising the biofilms formed on Castle River (reference/
unburned) and Lynx Creek (burned) sediments with
various bed consolidation periods.
Fig. 7 e Changes in suspended solids concentration as a function of applied bed shear stress for the 2-, 7-, and 14-day
consolidation (Lynx Creek e A, B, C; Castle River e D, E, F).
wat e r r e s e a r c h 4 5 ( 2 0 1 1 ) 5 2 1e5 3 4 529
Table 4 e Comparison of erosion depths at Type 1A and 1B erosion for Castle River (unburned) and Lynx Creek (burned)sediment.
Consolidationtime (days)
1A Shear(Pa)
1A Erosiondepth (mm)
1B Shear(Pa)
1B Erosion depth(mm)
Burned 2 0.08 0.0295 0.12 0.336
7 0.16 0.041 0.23 0.426
14 0.18 0.095 0.31 1.54
Unburned 2 0.039 0.0024 0.105 0.0126
7 0.097 0.0009 0.141 0.008
14 0.094 0.0009 0.165 0.014
wat e r r e s e a r c h 4 5 ( 2 0 1 1 ) 5 2 1e5 3 4530
the resultant increased level of attachment within and
between deposited flocs increased bed stability.
Wetweight bulkdensitymeasurementsof the twosediment
typesweremade toevaluate the influenceof consolidation (and
potentially biofilm formation) on bed stability. The wet density
of sediment from the unburned watershed (Castle River) was
approximately 1.28 g cm�3 and changed very little with depth.
In contrast, some compaction of the Lynx Creek sediment
deposit is suggested by the increase in density of deposited
sediment with depth (Fig. 8). For example, at a depth of 5 mm,
the density of the burned sediment was 0.19, 0.16 and
0.14 g cm�3 after 2, 7 and 14 days of consolidation, respectively.
However,at adepthof11mm, thedensity increasedto1.16, 1.05
and 0.89 g cm�3 after the respective consolidation periods.
Droppo andAmos (2001) developed a general three-layermodel
to describe the formation processes and characteristics of SFGL
deposits. Themodel describesa surfaceorganicfloc layer (Layer
1), amiddle collapse zone (Layer2) anda lowerconsolidatedbed
(Layer 3). They describe SFGL as a porous, low density and high
water content deposit with high yield resistance due to bio-
stabilization. Here, the bulk density (Fig. 8), the sc (Table 4) and
floc morphology analyses indicate that cohesive sediment
deposits in wildfire-impacted streams consist primarily of
porous, low density flocs that are associated with an active
biological community. Accordingly, the formational processes
Fig. 8 e Bulk density (g mL3) of Ly
and erosion dynamics in streams draining wildfire-impacted
landscapes are consistentwith the conceptualmodel proposed
by Droppo and Amos (2001).
3.5. Erosion rates
The erosion rates and depths for each flume experiment are
shown in time series plots in Fig. 9. For all runs, the initial
erosion event occurred at a lower applied shear than for
subsequent runs with longer consolidation periods (Table 4).
The peak in erosion rate occurred at the beginning of each
shear increment but then decreased as the underlying more
stable bed sediment was exposed. The maximum erosion
rates for the 2, 7 and 14-day runs occurred at 0.22, 0.34 and
0.39 Pa for unburned sediment (Castle River) and 0.25, 0.33 and
0.30 Pa for burned sediment (Lynx Creek), respectively. The
erosion depths at 0.25 Pa for the 2, 7 and 14-day consolidation
periods were 0.084, 0.051 and 0.033 mm for Castle River and
2.78, 0.426 and 0.181 mm for Lynx Creek sediment, respec-
tively. The time series plots show that increasing shear stress
was required to erode the wildfire-affected Lynx Creek sedi-
ment over time with increasing bed age. However, this effect
was less pronounced in the Castle River sediment.
Erosion depths for Castle River sediments never exceeded
0.5mmwhilemaximumerosiondepths in thewildfire-affected
nx Creek (burned) sediment.
Fig. 9 e Changes in erosion rate and depth as a function of applied bed shear stress for the 2-, 7-, and 14-day consolidation
(Lynx Creek e A, B, C; Castle River e D, E, F).
wat e r r e s e a r c h 4 5 ( 2 0 1 1 ) 5 2 1e5 3 4 531
wat e r r e s e a r c h 4 5 ( 2 0 1 1 ) 5 2 1e5 3 4532
LynxCreek sedimentwere 1.4, 4 and 3.7mm for 2, 7 and 14-day
runs respectively.
3.6. Still water settling
Still water settling rates have been described using zero (Amos
et al., 2003; Droppo, 2009) and first order kinetic models else-
where (Amos et al., 2004 based on Einstein and Krone, 1962). In
contrast to single point-based approaches that have been
utilized in the literature, the approach used herein is based on
all of the collected still water settling data (5e8 points per land
disturbanceeconsolidation period combination). Character-
istic zero, first, and second order kinetic plots were made for
still water mass settling of total suspended solids and indi-
cated that still water settling (the change in TSS concentration
over time) was best described as a second order reaction. For
this type of reaction, the rate of reaction decreases rapidly
(faster than linearly) as the suspended solids concentration
decreases. All combinations of land disturbance (unburned/
burned) and bed consolidation period (2, 7, and 14 days)
resulted in high (i.e. >98%) coefficients of determination for
second order kinetic models.
The still water TSS mass settling rates (mg TSS L�1 s�1)
derived from the Castle River (unburned) and Lynx Creek
(burned) eroded sediments after the various consolidation
periodswerecalculatedandevaluatedover time.TheTSSmass
settling rates associated with the burned eroded sediments
duringstillwater settlingwerehigher than thoseobservedwith
unburned eroded sediments (Fig. 10). The temporal changes in
TSS mass settling rates of the unburned and burned eroded
sediments in still water (Fig. 10) were significantly different
( p ¼ 0.0053, p ¼ 0.0295, p ¼ 0.0054 for the 2, 7, and 14-day
consolidation periods respectively). These analyses suggest
that TSSmass settling rates of the unburned eroded sediments
decreasedmore rapidly than those associated with the burned
sediments, possibly implying that settling velocities of the
unburned sediment were higher than those of the burned
sediments, a result consistent with the settling velocities
reported in Table 3. Differences in mass settling rates for the
two sediment types are likely explained by differences in the
Fig. 10 e Temporal changes in still water TSS mass settling
rates of Castle River (unburned) and Lynx Creek (burned)
sediments after various bed consolidation periods.
floc size distribution and characteristics (i.e. porosity, settling
velocity, floc strength and composition). Here, it is hypoth-
esised that burned sediment and attached biofilm eroded into
the water column formed organic rich (Table 1; Fig. 3), low
porosity (Table 2) flocs that settled more slowly than those
formed in the unburned system. From a water management
perspective, mass settling rate data imply that suspended
solids in wildfire-affected streams will remain in suspension
longer than in streams draining unburned landscapes.
3.7. Implications for water management
Results of the present study suggest that biogenic sediment
stabilization will have a significant effect on the rates and
magnitudes of sediment erosion and associated contaminant
transport in wildfire-affected streams. The initial influx of fire
modified sediment and associated nutrients from hillslopes
(Blake et al., 2009) combined with the loss of forest canopy in
wildfire-affected watersheds can create conditions which
promote biostabilized in-channel deposits of cohesive sedi-
ment. Silins et al. (in review) deployed fixed area ceramic tiles
in the wildfire-affected streams and reported that the mean
algal biomass in these streamswas 10e14 times higher than in
reference streams. The flume experiments show that the
erosion depth of wildfire-influenced sediments is higher,
suggesting that more material will likely be eroded once the
biofilm is removed compared to streams in unburned water-
sheds. Accordingly, once sc in the wildfire-affected streams is
exceeded, the eroded cohesive materials, which have low
settling velocities, will remain in the water column for pro-
longed periods of time and will more likely be transported to
downstream reservoirs, potentially contributing to significant
drinking water treatment challenges (Emelko et al., 2011).
4. Conclusions
1. Burned sediment had lower settling velocities, increased C
and carbohydrate levels compared to unburned sediment.
2. Biofilm formation on burned sediment increased sc for 7
and 14-day deposits by a factor of 1.6 and 1.8, respectively.
3. Erodibility (erosion depth) of burned sediment after 2, 7,
and 14 days of consolidationwas respectively 26, 53 and 110
times greater than unburned sediment.
4. Lower settling velocities of burned sediment are related to
increased organic content and higher porosity compared to
unburned sediment.
5. Changes in suspended solids concentrations as related to
shear stress indicated that sc increased with consolidation
period andas a result ofwildfire-associatedbiostabilization.
6. Erosion depths significantly decreased with bed age;
however, these depths were significantly greater as a result
of wildfire-associated biostabilization.
7. As a result of upstream land disturbance, fine sediments
(and their associated-contaminants) will stay in riverbeds
for longer periods of time due to disturbance-associated
biostabilization.
8. Increases in biofilm communities of Actinomycetes spp. (and
other species) may be associated with wildfire and may be
indicative of an increased propensity for taste and odor
wat e r r e s e a r c h 4 5 ( 2 0 1 1 ) 5 2 1e5 3 4 533
events related to sediment-associated microorganism
transport to downstream water suppliers.
9. Temporal changes in TSS mass settling rates of the
unburned and burned eroded sediments in still water were
significantly different for the 2, 7, and 14-day consolidation
periods, respectively.
Acknowledgements
The assistance of S. Deignan, B. Trapp and C. Jaskot in con-
ducting the flume experiments and K. Bladon in collecting
sediment and water samples is greatly appreciated.
r e f e r e n c e s
Alexander, D.B., 1998. Bacteria and archaea. In: Sylvia, D.M.,Fuhrmann, J.J., Hartel, P.G., Zuberer, D.A. (Eds.), Principles andApplications of Soil Microbiology. Prentice Hall, NJ, pp. 65e66.
Amos, C.L., Bergamasco, A., Umgiesser, G., Cappucci, S.,Clouthier, D., DeNat, C., 2004. The stability of tidal flats in theVenice Lagoon e the results of in situ measurements usingtwo benthic, annular flumes. J. Marine Syst. 51, 211e241.
Amos, C.L., Droppo, I.G., Gomez, E.A., Murphy, T., 2003. Thestability of a remediated bed in Hamilton Harbour, LakeOntario, Canada. Sedimentology 50 (1), 149e165.
Andreu, V., Imeson, A.C., Rubio, J.L., 2001. Temporal changes insoil aggregates and water erosion after a wildfire ina Mediterranean pine forest. Catena 44 (1), 69e84.
Berkhout, N., 1994. Manual “UHCM” Ultra High ConcentrationMeter. Delft Hydraulics, The Netherlands.
Blake, W.H., Droppo, I.G., Wallbrink, P.J., 2009. Sedimentaggregation and water quality in wildfire affected river basins.Mar. Freshw. Res. 60, 653e659.
Blake, W.H., Droppo, I.G., Humphreys, G.S., Doerr, S.H.,Shakesby, R.A., Wallbrink, P.J., 2007. Structural characteristicsand behaviours of fire-modified soil aggregates. J. Geophys.Res. Earth Surf. 112 (F2) article number: F02020.
Bladon, K.D., Silins, U., Wagner, M.J., Stone, M., Emelko, M.B.,Mendoza, C.A., Devito, K.J., Boon, S., 2008. Wildfire impacts onnitrogen concentration and production from headwaterstreams in southern Alberta’s Rocky Mountains. Can. J. for For.Res. 38, 2359e2371.
Bligh, E.G., Dyer, W.J., 1959. A rapid method of total lipidextraction and purification. Can. J. Biochem. Physiol. 37,911e917.
Brunke, M., 1999. Colmation and depth filtration: retention ofparticles in streambeds. Int. Rev. Hydrobiol. 84, 99e117.
Dade, W.B., Norwell, A., 1990. Moving Muds in the MarineEnvironment. In: Proceedings of Coastal Sediments 91, vol. 1.American Society of Civil Engineers. 54e71.
Droppo, I.G., 2001. Rethinking what constitutes suspendedsediment. Hydrolog. Process. 15, 1551e1564.
DeBano, L.F., Neary, D.G., Folliott, P.F., 1998. Fire’s effects onecosystems. John Wiley and Sons Inc, New York, NY, 333 pp.
Droppo, I.G., 2009. Biofilm structure and bed stability of fivecontrasting freshwater sediments. Marine Freshwater Res. 60,690e699.
Droppo, I.G., Amos, C.L., 2001. Structure, stability andtransformation of contaminated lacustrine surface fine-grained laminae. J. Sediment. Res. 71, 717e726.
Droppo, I.G., Leppard, G.G., Flannigan, D.T., Liss, S.N., 1997. Thefreshwater floc: A functional relationship of water and organic
and inorganic floc constituents affecting suspended sedimentproperties. Water Air Soil Pollut 99, 43e53.
Droppo, I.G., Jaskot, C., Nelson, T., Milne, J., Charlton, M., 2007.Aquaculture waste sediment stability: implications for wastemigration. Water Air Soil Pollut. 183 (1e4), 59e68.
Einstein, H.A., Krone, R.B.J., 1962. Experiments to determinemodes of cohesive sediment transport in salt water.J. Geophys. Res. 67, 1451e1461.
Emelko, M.B., Huck, P.M., Smith, E.F., 2006. Full-scaleoptimization of single-stage biological filtration. J. AWWA 98(12), 61e73.
Emelko, M.B., Silins, U., Bladon, K.D., Stone, M., 2011.Implications of land disturbance on drinking watertreatability in a changing climate: demonstrating the needfor “source water supply and protection” strategies. WaterRes 45 (2), 461e472.
Fox, D.M., Darboux, F., Carrega, P., 2007. Effects of fire inducedwater repellency on soil aggregate stability, splash erosionand saturated hydraulic conductivity for different sizefractions. Hydrolog. Process. 21, 2377e2384.
Friend, P.L., Ciavola, P., Cappucci, S., Santos, R., 2003. Bio-dependent bed parameters as a proxy tool for sedimentstability in mixed habitat intertidal areas. Coastal Shelf Res.23, 1899e1917.
Gerbersdorf, S.U., Jancke, T., Westrich, B., Paterson, D.M., 2008.Microbial stabilization of riverine sediments by extracellularpolymeric substances. Geobiol. 6, 57e69.
Gerbersdorf, S.U., Westrich, B., Paterson, D.M., 2009. Microbialextracellular polymeric substances (EPS) in fresh watersediments. Microb. Ecol. 58, 334e349.
Griggs, G.B., Hein, J.R., 1980. Sources, dispersal and clay mineralcomposition of fine-grained sediment off central and northernCalifornia. J. Geol. 88 (5), 541e566.
Lau, Y.L., 1995. Relative importance of mean velocity and bedshear on biofilm accumulation in open-channel flows. WaterSci. Technol. 32 (8), 193e198.
Lau, Y.L., Liu, D., 1993. Effect of flow rate on biofilm accumulationin open channels. Water Res. 27 (3), 355e360.
Lau, Y.L., Droppo, I.G., 2000. Influence of antecedent conditions onthecritical shearstressofbedsediments.WaterRes.34, 663e667.
Leppard, G.G., 1986. The fibrillar matrix component of lacustrinebiofilms. Water Res. 20 (6), 697e702.
Li, D.H., Ganczarczyk, J., 1987. Stroboscopic determination ofsettling velocity, size and porosity of activated sludge flocs.Water Res. 21, 257e262.
Liu, D., Lau, Y.L., Chau, Y.K., Pacepavicius, G., 1994. Simpletechnique for estimation of biofilm accumulation. Bull.Environ. Contam. Toxicol 53, 913e918.
Mehta, A.J., 1989. On estuarine cohesive sediment suspensionbehaviour. J. Geophy. Res. 94, 14303e14314.
Minshall, G.W., 2003. Responses of stream benthicmacroinvertebrates to fire. Forest Ecol. Manage. 178 (1e2),155e161.
Neu, T.R., 1994. Biofilms and microbial mats. In: Krumbein, W.E.,Paterson, D.M., Stal, L.J. (Eds.), Biostabilization. OldenburgUniversity Press, Oldenburg, pp. 9e16.
Pacepavicuis, G., Lam, Y., Liu, D., Okamura, J., Aoyama, I., 1997. Arapid method for estimating biofilm mass. Environ. Toxicol.Water Qual. 12 (1), 97e100.
Partheniades, E., 1990. Microstructure of fine-grainedsediments: from mud to shale. In: Bennet, R.H., et al. (Eds.),Frontiers in Sedimentary Geology. Springer-Verlag, Berlin,pp. 175e183.
Paterson, D.M., 1997. Biological mediation of sediment erodibility:ecology and physical dynamics. In: Burt, N., Parker, R.,Watts, J. (Eds.), Cohesive Sediments.Wiley andSons, pp. 215e229.
Petticrew, E.L., Owens, P.N., Giles, T.R., 2006. Wildfire effectson the quantity and composition of suspended and
wat e r r e s e a r c h 4 5 ( 2 0 1 1 ) 5 2 1e5 3 4534
gravel-stored sediments. Water Air Soil Pollut. Focus 6,647e656.
Percival, S.L., Knapp, J.S., Wales, D.S., Edyvean, R.G.J., 1999. Theeffect of turbulent flow and surface roughness on biofilmformation in drinking water. J. Indust. Microbiol. Biotechnol.22 (3), 152e159.
Schmidt, S.K., Alexander, M., Shuler, M.L., 1985. Predictingthreshold concentrations of organic substrates for bacterialgrowth. J. Theor. Biol. 114, 1e8.
Silins, U., Stone, M., Emelko, M.B., Bladon, B., 2008. Impacts ofWildfire and Post-fire Salvage Logging on Sediment Transfer inthe Oldman Watershed, Alberta, Canada. In: IAHS Workshopon the Effects of Landuse Change on Sediment Erosion, NewZealand, vol. 325. IAHS Publication, pp. 509e515.
Silins, U., Stone, M., Emelko, M.B., Bladon, K.D., 2009. Sedimentproduction following severe wildfire and post-fire salvagelogging in the Rocky Mountain headwaters of the OldmanRiver Basin, Alberta. Catena 79 (3), 189e197.
Silins, U., Bladon, K.D., Esch, E., Spence, J.R., Kelly E.N., Stone, M.,Emelko, M.B., Boon, S., Wagner, M.J., Williams, C.H.S.,Tichkowsky, I. After the fire’s out: the legacy of wildfire andsalvage logging on nutrient runoff and aquatic plant,invertebrate, and fish productivity. Global Change Biol.(in review).
Stone, M., Droppo, I.G., 1994. In-channel surficial fine-grainedsediment laminae (Part II): chemical characteristics andimplications for contaminant transport in fluvial systems.Hydrolog. Process. 8 (2), 113e124.
Stone, M., Krishnappan, B.G., Emelko, M.B., 2008. The effect of bedage and shear stress on the size distribution and particle
morphology of eroded cohesive sediment deposits in anannular flume. Water Res. 42 (15), 4179e4187.
Tolhurst, T.J., Reithmueller, P., Paterson, D.M., 2000. In situversus laboratory analysis of sediment stability fromintertidal mudflats. Continental Shelf Res. 20 (10/11),1317e1334.
Tunlid, A., White, D.C., 1992. Biochemical analysis of biomass,community structure, nutritional status and metabolicactivity of microbial communities in soil. In: Stotzky, G.,Bollag, J.M. (Eds.), Soil Biochemistry, vol. 6. Marcel Dekker,New York, pp. 229e262.
Underwood, G.J.C., Paterson, D.M., 2003. The importance ofextracellular carbohydrate production by marine epipelicdiatoms. Adv. Bot. Res. 40, 184e240.
Villaret, C., Paulic, M., 1986. Experiments on the Erosion ofDeposited and Placed Cohesive Sediments in an AnnularFlume. Report to Coastal and Oceanographic. EngineeringDepartment, University of Florida, Gainesville.
White, D.C., Davis, W.M., Nickels, J.S., King, J.D., Bobbie, R.J., 1979.Determination of the sedimentary microbial biomass byextractable lipid phosphate. Oecologia 40, 51e62.
Westerling, A.L., Hidalgo, H.G., Cayan, D.R., Swetnam, T.W., 2006.Warming and earlier spring increase western U.S. forestwildfire activity. Science 313, 940e943.
Wood, P.J., Armitage, P.D., 1997. Biological effects of finesediment in the lotic environment. Environ. Manage. 21 (2),203e217.
Zaitlin, B., Watson, S.B., 2006. Actinomycetes in relation to tasteand odour in drinking water: myths, tenets, and truths. WaterRes. 40 (9), 1741e1753.