The effect of biofilm colonization on the stability of non-cohesive sediments

In the past decades, engineers have started to realize the importance of the interaction between vegetation, biota and water flow, in riverine and marine environments; a discipline that has been named “Eco-Hydraulics”. Scientists have valued this coupled phenomenon for much longer than their engineering colleagues. As early as 1970, marine researchers presented the evidence that colonies of micro-organisms might alter the stability of fine cohesive sediments (Neuman et al., 1970). However traditional models of sediments transport (e.g. Shields, 1936) have been derived using abiotic sediments and did not consider that most wet surfaces would soon be colonized by micro-organisms and their extracellular polymeric substances (EPS), a combination called “biofilm” (Lock, 1993). Scientists during the 1990s, after observing this phenomenon in the field, coined the term “biostabilization”. During this period they showed that colonies of cyanobacteria and diatoms coating fine sand or cohesive sediments can increase their stability by up to 960% compared to abiotic sediments (Grant and Gust, 1987; Dade et al, 1990; Paterson 1997). Only recently have engineers started to take into consideration the effect of such increased cohesion and adhesion due to biogenic forces within the sediment transport model (Righetti and Lucarelli, 2007); yet all of those studies have low applicability because they are linked to specific environmental conditions. Moreover no data are available on the effect of biofilm on larger sediments (e.g. coarse sand and gravel). The present thesis provides experimental data carried out in a flume laboratory pertaining to biostabilization of non-cohesive coarse sand and gravels at a scale representation of a real river system (from 0.2m to 1m). Four sediment substratum (glass spheres of D50 = 1.09mm and 2.00mm; sand of D50 = 1.20mm and gravel of D50 = 2.20mm) were colonized under unidirectional flow by a cyanobacterium (Phormidium sp.) for between 1 and 10 weeks. The increase in erosion threshold for biotic sediment is then investigated using a series of different methods ranging from traditional sediment transport techniques (e.g. Yalin, 1972), to image thresholding and particle image velocimetry (PIV) assessments of flow modification due to biofilm presence. Moreover, tensile strength analysis of ex-situ biofilm/substratum specimens will be presented to understand better the mechanical property of this composite material. Data indicates that: i) biostabilization of sediments in the range of coarse sand and gravel occurs (9%-150% more shear stress required to induce entrainment compared to abiotic sediments) but to a lower extent compared to critical entrainment thresholds for fine sand and cohesive sediments (Paterson, 1997); ii) flume experimentation can be employed to control specific variables affecting biostabilization and could help to unfold the complicated interactions between environmental variables, and the affect of flow on the growth and strength of biofilm colonization over sediments; iii) strong biofilm growth generated a more uniform velocity field, with reduction in shear stress (up to 82% compared with abiotic sediments) and decreases in roughness length of the bed (up to 94% compared to abiotic sediments); iv) Composite biofilm/substratum specimens presented a clear elastic behaviour when tensile tested; v) Conventional models of sediment transport (e.g. Wiberg and Smith, 1987) do not consider the presence of biofilm and will not work in the case of bio-mats smoothing the surface of the bed; hence the need for new models which include the biofilm elasticity and the bio-mat smoothing process. This thesis suggests two theoretical examples where the biofilm action is considered at a grain to grain and bio-mat scale.

[1]  W. S. Chepil,et al.  Equilibrium of Soil Grains at the Threshold of Movement by Wind , 1959 .

[2]  Claude Michel Discussion of "The Legend of A. F. Shields" , 2000 .

[3]  R. Walters,et al.  Effects of bed material properties on cohesive sediment erosion , 2004 .

[4]  David M. Paterson,et al.  Measuring the in situ Erosion Shear Stress of Intertidal Sediments with the Cohesive Strength Meter (CSM) , 1999 .

[5]  T. Tolhurst,et al.  The influence of an extracellular polymeric substance (EPS) on cohesive sediment stability , 2002 .

[6]  M. Schultz,et al.  The effect of biofilms on turbulent boundary layers , 1999 .

[7]  H. A. Einstein,et al.  Formulas for the Transportation of Bed Load , 1942 .

[8]  R. Adrian Particle-Imaging Techniques for Experimental Fluid Mechanics , 1991 .

[9]  S. Sauvage,et al.  Effect of near‐bed turbulence on chronic detachment of epilithic biofilm: Experimental and modeling approaches , 2010 .

[10]  K. Sundbäck,et al.  The influence of benthic microalgae on the stability of a subtidal sediment , 1993 .

[11]  R. Soulsby,et al.  Threshold of Sediment Motion in Coastal Environments , 1997 .

[12]  J. Grant,et al.  The interaction between benthic diatom films and sediment transport , 1986 .

[13]  H. Li,et al.  Distribution and Fractions of Phosphorus and Nitrogen in Surface Sediments from Dianchi Lake, China , 2012 .

[14]  D. Paterson,et al.  Microalgal sediment biostabilisation along a salinity gradient in the Eden Estuary, Scotland: unravelling a paradox , 2008 .

[15]  Vladimir Nikora,et al.  Hydrodynamics of aquatic ecosystems: An interface between ecology, biomechanics and environmental fluid mechanics , 2010 .

[16]  D. Vericat,et al.  The relative contributions of ecology and hydraulics to ecohydraulics , 2010 .

[17]  B. Little,et al.  Spatial relationships between bacteria and mineral surfaces , 1997 .

[18]  Peter A. Wilderer,et al.  Structure and function of biofilms. , 1989 .

[19]  L. Stal Microphytobenthos, their Extracellular Polymeric Substances, and the Morphogenesis of Intertidal Sediments , 2003 .

[20]  J. Stanford,et al.  The hyporheic habitat of river ecosystems , 1988, Nature.

[21]  V. Körstgens,et al.  Uniaxial compression measurement device for investigation of the mechanical stability of biofilms. , 2001, Journal of microbiological methods.

[22]  Pierre Francus,et al.  An image-analysis technique to measure grain-size variation in thin sections of soft clastic sediments , 1998 .

[23]  Variation in the mechanical properties of a porous multi-phase biofilm under compression due to void closure. , 2007, Water science and technology : a journal of the International Association on Water Pollution Research.

[24]  M. Righetti,et al.  May the Shields theory be extended to cohesive and adhesive benthic sediments , 2007 .

[25]  A. S. Paintal,et al.  A Stochastic Model Of Bed Load Transport , 1971 .

[26]  Erosion threshold of saturated natural cohesive sediments: Modeling and experiments , 2008 .

[27]  Heather Haynes,et al.  The impact of stress history on bed structure , 2013 .

[28]  C. Laspidou,et al.  On the calculation of the elastic modulus of a biofilm streamer. , 2008, Biotechnology and bioengineering.

[29]  T. Walker,et al.  Quantifying erosion rates and stability of bottom sediments at mussel aquaculture sites in Prince Edward Island, Canada , 2009 .

[30]  J W Costerton,et al.  How bacteria stick. , 1978, Scientific American.

[31]  N. Bury,et al.  Metal contamination in aquatic environments: science and lateral management , 2009 .

[32]  Vladimir Nikora,et al.  Some observations of the effects of micro-organisms growing on the bed of an open channel on the turbulence properties , 2002, Journal of Fluid Mechanics.

[33]  John M. Buffington,et al.  The Legend of A. F. Shields , 1999 .

[34]  M. Yalin,et al.  Inception of Sediment Transport , 1979 .

[35]  A. Grass Structural features of turbulent flow over smooth and rough boundaries , 1971, Journal of Fluid Mechanics.

[36]  B. Caussade,et al.  The relationship between stream periphyton dynamics and near-bed turbulence in rough open-channel flow , 2007 .

[37]  J. Waterbury,et al.  Generic assignments, strain histories, and properties of pure cultures of cyanobacteria , 1979 .

[38]  J. Finnigan Turbulence in plant canopies , 2000 .

[39]  Bernhard Westrich,et al.  Processes governing river water quality identified by principal component analysis , 2002 .

[40]  R. Hozalski,et al.  Micro-cantilever method for measuring the tensile strength of biofilms and microbial flocs. , 2003, Journal of microbiological methods.

[41]  Y. Monbet,et al.  Sediment erodability in sediment transport modelling: Can we account for biota effects? , 2007 .

[42]  Gerben J. de Boer,et al.  Modeling large-scale cohesive sediment transport affected by small-scale biological activity , 2008 .

[43]  Hilary M. Lappin-Scott,et al.  Growth and Detachment of Cell Clusters from Mature Mixed-Species Biofilms , 2001, Applied and Environmental Microbiology.

[44]  W. Lick,et al.  Initiation of movement of quartz particles , 2004 .

[45]  I. V. Egiazaroff,et al.  Calculation of Nonuniform Sediment Concentrations , 1965 .

[46]  Emily H. Stanley,et al.  THE FUNCTIONAL SIGNIFICANCE OF THE HYPORHEIC ZONE IN STREAMS AND RIVERS , 1998 .

[47]  S. Rice,et al.  A transferable method for the automated grain sizing of river gravels , 2005 .

[48]  R. Ananthanarayan Textbook of Microbiology , 1996 .

[49]  David M. Paterson,et al.  Muddy Sediment Erosion: Insights from Field Studies , 2009 .

[50]  T. P. Scoffin,et al.  The composition, structure and erodability of subtidal mats, Abaco, Bahamas , 1970 .

[51]  Lucas J. Stal,et al.  Interplay between biology and sedimentology in a mudflat (Biezelingse Ham, Westerschelde, The Netherlands) , 2000 .

[52]  Vladimir Nikora,et al.  A Simple Model of Stream Periphyton-Flow Interactions , 1998 .

[53]  H. Harms,et al.  Adhesion of the positively charged bacterium Stenotrophomonas (Xanthomonas) maltophilia 70401 to glass and Teflon , 1996, Journal of bacteriology.

[54]  I. Droppo,et al.  Dynamic existence of waterborne pathogens within river sediment compartments. Implications for water quality regulatory affairs. , 2009, Environmental science & technology.

[55]  V. Vanoni,et al.  Temperature Effects in Low-Transport, Flat-Bed Flows , 1972 .

[56]  P. Carling Particle over-passing on depth-limited gravel bars , 1990 .

[57]  S. J. Hall Physical disturbance and marine benthic communities: life in unconsolidated sediments , 1994 .

[58]  Y. Dufrêne,et al.  Probing microbial cell surface charges by atomic force microscopy , 2002 .

[59]  Jerome P.-Y. Maa,et al.  Critical Bed-Shear Stress for Cohesive Sediment Deposition under Steady Flows , 2008 .

[60]  J. Smith,et al.  Calculations of the critical shear stress for motion of uniform and heterogeneous sediments , 1987 .

[61]  Martin C. Miller,et al.  Threshold of sediment motion under unidirectional currents , 1977 .

[62]  A. Jopling,et al.  Mechanics of Sediment Movement (SEPM Short Course No. 3) , 1979 .

[63]  T. Tolhurst,et al.  Changes in cohesive sediment properties associated with the growth of a diatom biofilm , 2007, Hydrobiologia.

[64]  C. Ling,et al.  Criteria for incipient motion of spherical sediment particles , 1995 .

[65]  S. B. Surman,et al.  CONSIDERATION OF SOME IMPLICATIONS OF THE RESISTANCE OF BIOFILMS TO BIOCIDES , 1998 .

[66]  B. Biggs,et al.  On stream periphyton‐turbulence interactions , 1997 .

[67]  R. Hozalski,et al.  Development and testing of a novel microcantilever technique for measuring the cohesive strength of intact biofilms , 2010, Biotechnology and bioengineering.

[68]  J. Costerton,et al.  Optical sectioning of microbial biofilms , 1991, Journal of bacteriology.

[69]  P. Owens Background and summary of this issue on sediment linkages , 2007 .

[70]  Markus Raffel,et al.  Particle Image Velocimetry: A Practical Guide , 2002 .

[71]  Daniel Prats,et al.  Influence of temperature variations on the cake resistance and EPS of MBR mixed liquor fractions , 2010 .

[72]  W. S. Chepil,et al.  The use of evenly spaced hemispheres to evaluate aerodynamic forces on a soil surface , 1958 .

[73]  David M. Paterson,et al.  Working with Natural Cohesive Sediments , 2002 .

[74]  Hideki Harada,et al.  A novel method for evaluation of biofilm tensile strength resisting erosion , 1999 .

[75]  Zhi-jun Liu,et al.  River Processes: An Introduction to Fluvial Dynamics , 2005, The Professional Geographer.

[76]  T. Battin Hydrodynamics is a major determinant of streambed biofilm activity: From the sediment to the reach scale , 2000 .

[77]  P. Wilcock Estimating Local Bed Shear Stress from Velocity Observations , 1996 .

[78]  S. Gerbersdorf,et al.  Physico-chemical and biological sediment properties determining erosion resistance of contaminated riverine sediments – Temporal and vertical pattern at the Lauffen reservoir/River Neckar, Germany , 2005 .

[79]  N. Cheng,et al.  Pickup probability for sediment entrainment. , 1998 .

[80]  Nicholas Kouwen,et al.  FLEXIBLE ROUGHNESS IN OPEN CHANNELS , 1973 .

[81]  Gerard Middleton,et al.  Mechanics of sediment movement , 1978 .

[82]  J. Newbold,et al.  Contributions of microbial biofilms to ecosystem processes in stream mesocosms , 2003, Nature.

[83]  J. Best On the interactions between turbulent flow structure, sediment transport and bedform development: some considerations from recent experimental research , 1993 .

[84]  Vladimir Nikora,et al.  Spatially Averaged Open-Channel Flow over Rough Bed , 2001 .

[85]  D. Montgomery,et al.  Effects of sediment supply on surface textures of gravel‐bed rivers , 1999 .

[86]  C. R. Smith,et al.  Use of flow visualization data to examine spatial-temporal velocity and burst-type characteristics in a turbulent boundary layer , 1991, Journal of Fluid Mechanics.

[87]  P. Komar Selective grain entrainment by a current from a bed of mixed sizes; a reanalysis , 1987 .

[88]  J. F. McNabb,et al.  Quantitative characterization of microbial biomass and community structure in subsurface material: a prokaryotic consortium responsive to organic contamination , 1986 .

[89]  V. Bakkestuen,et al.  Spatial and temporal variability in epilithic biofilm bacterial communities along an upland river gradient. , 2008, FEMS microbiology ecology.

[90]  P.K Sahoo,et al.  A survey of thresholding techniques , 1988, Comput. Vis. Graph. Image Process..

[91]  K. C. Marshall,et al.  Microbial Adhesion and Aggregation , 1985, Life Sciences Research Reports.

[92]  N. Salant ‘Sticky business’: The influence of streambed periphyton on particle deposition and infiltration , 2011 .

[93]  Robert C. Grabowski,et al.  Erodibility of cohesive sediment: The importance of sediment properties , 2011 .

[94]  Stephan Scholl,et al.  Structure and shear strength of microbial biofilms as determined with confocal laser scanning microscopy and fluid dynamic gauging using a novel rotating disc biofilm reactor , 2007, Biotechnology and bioengineering.

[95]  Panayiotis Diplas,et al.  Surface Sampling in Gravel Streams , 1993 .

[96]  C. Leboulanger,et al.  Spatial heterogeneity of periphytic microbial communities in a small pesticide-polluted river. , 2009, FEMS microbiology ecology.

[97]  Hideki Harada,et al.  Adhesion strength of biofilm developed in an attached-growth reactor , 1994 .

[98]  W. Sloan,et al.  Quantifying the tensile strength of microbial mats grown over noncohesive sediments , 2012, Biotechnology and bioengineering.

[99]  P. Davies,et al.  Calibration of the high-pressure cohesive strength meter (CSM) , 2007 .

[100]  P. Klingeman,et al.  On why gravel bed streams are paved , 1982 .

[101]  Yi-Ju Chou,et al.  Rolling and Lifting Probabilities for Sediment Entrainment , 2003 .

[102]  Stefan Winkler,et al.  Color Space Conversions , 2013 .

[103]  W. Manz,et al.  Microbial assemblages as ecosystem engineers of sediment stability , 2009 .

[104]  R. Rathbun,et al.  Measurement of hydraulic and sediment transport variables in a small recirculating flume , 1967 .

[105]  G. Pender,et al.  Critical shear stress for incipient motion of sand/gravel streambeds , 2001 .

[106]  J. Costerton,et al.  Bacterial biofilms in nature and disease. , 1987, Annual review of microbiology.

[107]  L. Dendooven,et al.  Microbial communities to mitigate contamination of PAHs in soil—possibilities and challenges: a review , 2011, Environmental science and pollution research international.

[108]  D. Montgomery,et al.  A systematic analysis of eight decades of incipient motion studies, with special reference to gravel‐bedded rivers , 1997 .

[109]  Chih Ted Yang,et al.  Sediment transport : theory and practice / Chih Ted Yang , 1995 .

[110]  H. Cyr,et al.  Environmental variability influences the structure of benthic algal communities in an oligotrophic lake , 2006 .

[111]  J. Feijen,et al.  Adhesion of Escherichia coli on to a series of poly(methacrylates) differing in charge and hydrophobicity. , 1991, Biomaterials.

[112]  W. Verstraete,et al.  Distribution of extracellular polysaccharides and flotation of anaerobic sludge , 1996, Applied Microbiology and Biotechnology.

[113]  C. Doscher,et al.  Straight Benthic Flow-Through Flume for In Situ Measurement of Cohesive Sediment Dynamics , 2003 .

[114]  Peter A. Mantz,et al.  INCIPIENT TRANSPORT OF FINE GRAINS AND FLAKES BY FLUIDS-EXTENDED SHIELD DIAGRAM , 1977 .

[115]  P. Abreu,et al.  Rheological properties of marine organic aggregates: Importance for vertical flux, turbulence and microzones , 1991 .

[116]  S. R. McLean,et al.  Spatially averaged flow over a wavy surface , 1977 .

[117]  J. Newbold,et al.  Effects of Current Velocity on the Nascent Architecture of Stream Microbial Biofilms , 2003, Applied and Environmental Microbiology.

[118]  I. Klapper,et al.  Description of mechanical response including detachment using a novel particle model of biofilm/flow interaction. , 2007, Water science and technology : a journal of the International Association on Water Pollution Research.

[119]  A. R. Nowell,et al.  Effects of bacterial exopolymer adhesion on the entrainment of sand , 1990 .

[120]  D. Paterson Microbiological mediation of sediment structure and behaviour , 1994 .

[121]  Hubert Chanson,et al.  The hydraulics of open channel flow , 2004 .

[122]  B. Biggs,et al.  Periphyton responses to a hydraulic gradient in a regulated river in New Zealand , 1994 .

[123]  Hubert Chanson,et al.  The Hydraulics of Open Channel Flow: An Introduction , 1999 .

[124]  Hideki Harada,et al.  A novel concept for evaluation of biofilm adhesion strength by applying tensile force and shear force , 1996 .

[125]  Cory J. Rupp,et al.  Viscoelastic fluid description of bacterial biofilm material properties. , 2002, Biotechnology and bioengineering.

[126]  S. Gerbersdorf,et al.  Surface adhesion measurements in aquatic biofilms using magnetic particle induction: MagPI , 2009 .

[127]  Jane E. Sargison,et al.  The turbulence structure of flows over rough surfaces and freshwater biofilms , 2010 .

[128]  D. Paterson,et al.  Microbial Extracellular Polymeric Substances (EPS) in Fresh Water Sediments , 2009, Microbial Ecology.

[129]  Michael D. Abràmoff,et al.  Image processing with ImageJ , 2004 .

[130]  G. Pender,et al.  Stress history effects on graded bed stability , 2007 .

[131]  M. C. Powers A New Roundness Scale for Sedimentary Particles , 1953 .

[132]  R. Zare,et al.  Microscale Location, Characterization, and Association of Polycyclic Aromatic Hydrocarbons on Harbor Sediment Particles , 2000 .

[133]  Charles R. Neill,et al.  Quantitative Definition of Beginning of Bed Movement , 1969 .

[134]  Stephen P. Rice,et al.  Sampling surficial fluvial gravels; the precision of size distribution percentile sediments , 1996 .

[135]  W. Graf Hydraulics of Sediment Transport , 1984 .

[136]  J. Costerton,et al.  River epilithon: toward a structural-functional model , 1984 .

[137]  J. Southard,et al.  Erosion of fine-grained marine sediments: Sea-floor and laboratory experiments , 1978 .

[138]  V. Phoenix,et al.  Magnetic Resonance Imaging of Structure, Diffusivity, and Copper Immobilization in a Phototrophic Biofilm , 2008, Applied and Environmental Microbiology.

[139]  T. Tolhurst,et al.  In situ versus laboratory analysis of sediment stability from intertidal mudflats , 2000 .

[140]  M. Righetti,et al.  Resuspension phenomena of benthic sediments: The role of cohesion and biological adhesion , 2010 .

[141]  Paul S. Kemp,et al.  Experimentation at the interface of fluvial geomorphology, stream ecology and hydraulic engineering and the development of an effective, interdisciplinary river science , 2010 .

[142]  Vladimir Nikora,et al.  Hydraulic Resistance due to Aquatic Vegetation in Small Streams: Field Study , 2008 .

[143]  G. K. Gilbert,et al.  The Transportation of Débris by Running Water , 2010 .

[144]  Waleed M. Zahid,et al.  Suspended solids in biological filter effluents , 1990 .

[145]  Cory J. Rupp,et al.  Biofilm material properties as related to shear-induced deformation and detachment phenomena , 2002, Journal of Industrial Microbiology and Biotechnology.

[146]  John Pitlick,et al.  Observations of Flow and Sediment Entrainment on a Large Gravel-Bed River , 1996 .

[147]  W. Manz,et al.  Anthropogenic pollutants affect ecosystem services of freshwater sediments: the need for a “triad plus x” approach , 2011 .

[148]  J. Pandey,et al.  Accessing microbial diversity for bioremediation and environmental restoration. , 2005, Trends in biotechnology.

[149]  M. Callow A review of fouling in freshwaters , 1993 .

[150]  F. Hjulström Studies of the morphological activity of rivers as illustrated by the River Fyris , 1935 .

[151]  Pelton,et al.  Micromechanics: A New Approach to Studying the Strength and Breakup of Flocs , 1996, Journal of colloid and interface science.

[152]  C. Yang Incipient Motion and Sediment Transport , 1973 .

[153]  Application of Paramagnetically Tagged Molecules for Magnetic Resonance Imaging of Biofilm Mass Transport Processes , 2010, Applied and Environmental Microbiology.

[154]  Emmanuel Partheniades,et al.  Erosion and Deposition of Cohesive Soils , 1965 .

[155]  S. Lane,et al.  Coherent flow structures in a depth-limited flow over a gravel surface : The role of near-bed turbulence and influence of Reynolds number , 2009 .

[156]  André Robert,et al.  Characteristics of velocity profiles along riffle-pool sequences and estimates of bed shear stress , 1997 .

[157]  T. Day A Study of Initial Motion Characteristics of Particles in Graded Bed Materials , 1980 .

[158]  Anthony J. Grass,et al.  Initial Instability of Fine Bed Sand , 1970 .

[159]  D. Sigee Freshwater Microbiology: Biodiversity and Dynamic Interactions of Microorganisms in the Aquatic Environment , 2005 .

[160]  Gordon Rittenhouse Transportation and deposition of heavy mineral , 1943 .

[161]  D. S. Smith,et al.  Characterizing Heterogeneous Bacterial Surface Functional Groups Using Discrete Affinity Spectra for Proton Binding , 1999 .

[162]  Hans Kramer,et al.  Closure of "Sand Mixtures and Sand Movement in Fluvial Model" , 1935 .

[163]  J W Costerton,et al.  The bacterial glycocalyx in nature and disease. , 1981, Annual review of microbiology.

[164]  Carl L. Amos,et al.  The stability of a mudflat in the Humber estuary, South Yorkshire, UK , 1998, Geological Society, London, Special Publications.

[165]  E. Bouwer,et al.  Biofilms in porous media , 2000 .

[166]  P. P. Brown,et al.  Sphere Drag and Settling Velocity Revisited , 2003 .

[167]  W. C. Krumbein,et al.  Size frequency distributions of sediments , 1934 .

[168]  Lawrence P. Sanford,et al.  A unified erosion formulation for fine sediments , 2001 .

[169]  S. Findlay Importance of surface‐subsurface exchange in stream ecosystems: The hyporheic zone , 1995 .

[170]  F. Garcia-Pichel,et al.  The Evolution of a Capacity to Build Supra-Cellular Ropes Enabled Filamentous Cyanobacteria to Colonize Highly Erodible Substrates , 2009, PloS one.

[171]  Giselher Gust,et al.  Prediction of coastal sediment stability from photopigment content of mats of purple sulphur bacteria , 1987, Nature.

[172]  Stuart N. Lane,et al.  Assessment of Dem Quality for Characterizing Surface Roughness Using Close Range Digital Photogrammetry , 1998 .

[173]  Jerome P.-Y. Maa,et al.  Sediment resuspension characteristics in Baltimore Harbor, Maryland , 1998 .

[174]  C. Wentworth A Scale of Grade and Class Terms for Clastic Sediments , 1922, The Journal of Geology.

[175]  T. P. Scoffin,et al.  The Trapping and Binding of Subtidal Carbonate Sediments by Marine Vegetation in Bimini Lagoon, Bahamas , 1970 .

[176]  T. Battin,et al.  Linking Sediment Biofilms, Hydrodynamics, and River Bed Clogging: Evidence from a Large River , 1999, Microbial Ecology.

[177]  Paige J. Novak,et al.  Biofilm Cohesiveness Measurement Using a Novel Atomic Force Microscopy Methodology , 2007, Applied and Environmental Microbiology.

[178]  E. Andrews Entrainment of gravel from naturally sorted riverbed material , 1983 .

[179]  Wilbert Lick The Transport of Contaminants in the Great Lakes , 1982 .

[180]  S. Gerbersdorf,et al.  Microbial stabilization of riverine sediments by extracellular polymeric substances , 2007, Geobiology.

[181]  Y. Guan,et al.  Effect of biofilm on cast iron pipe corrosion in drinking water distribution system: Corrosion scales characterization and microbial community structure investigation , 2008 .

[182]  David M. Paterson,et al.  Short‐term changes in the erodibility of intertidal cohesive sediments related to the migratory behavior of epipelic diatoms , 1989 .

[183]  H. Christian,et al.  In situ erosion measurements on fine-grained sediments from the Bay of Fundy , 1992 .

[184]  S. Tait,et al.  Water‐worked gravel beds in laboratory flumes – a natural analogue? , 2009 .

[185]  Rosário Oliveira,et al.  The Role of Exopolymers Produced by Sphingomonas paucimobilis in Biofilm Formation and Composition , 2000 .

[186]  I. Droppo,et al.  The effect of depositional history on contaminated bed sediment stability. , 2001, The Science of the total environment.

[187]  J. Best On the entrainment of sediment and initiation of bed defects: insights from recent developments within turbulent boundary layer research , 1992 .

[188]  P. Klingeman,et al.  Bedload and Size Distribution in Paved Gravel-Bed Streams , 1983 .

[189]  Wolfgang E. Krumbein,et al.  Microbially Induced Sedimentary Structures: A New Category within the Classification of Primary Sedimentary Structures , 2001 .

[190]  B. Caussade,et al.  Interplay between turbulence and periphyton in rough open-channel flow , 2001 .

[191]  Mark G. Macklin,et al.  The significance of pollution from historic metal mining in the Pennine orefields on river sediment contaminant fluxes to the North Sea , 1997 .

[192]  John S. Halow,et al.  INCIPIENT ROLLING, SLIDING AND SUSPENSION OF PARTICLES IN HORIZONTAL AND INCLINED TURBULENT FLOW , 1973 .

[193]  B. Biggs,et al.  SUBSIDY AND STRESS RESPONSES OF STREAM PERIPHYTON TO GRADIENTS IN WATER VELOCITY AS A FUNCTION OF COMMUNITY GROWTH FORM , 1998 .

[194]  D. Paphitis,et al.  Sand grain threshold, in relation to bed ‘stress history’: an experimental study , 2005 .

[195]  Yarko Niño,et al.  Experiments on particle—turbulence interactions in the near–wall region of an open channel flow: implications for sediment transport , 1996, Journal of Fluid Mechanics.

[196]  Raymond M. Hozalski,et al.  Determination of biofilm mechanical properties from tensile tests performed using a micro-cantilever method , 2010, Biofouling.

[197]  P. Komar,et al.  Laboratory measurements of pivoting angles for applications to selective entrainment of gravel in a current , 1986 .

[198]  Z Lewandowski,et al.  Structural deformation of bacterial biofilms caused by short-term fluctuations in fluid shear: an in situ investigation of biofilm rheology. , 1999, Biotechnology and bioengineering.

[199]  A. Shields,et al.  Application of similarity principles and turbulence research to bed-load movement , 1936 .

[200]  C. Pilditch,et al.  Variation in sediment stability and relation to indicators of microbial abundance in the Okura Estuary, New Zealand , 2003 .

[201]  I. Droppo,et al.  Erosion characteristics and floc strength of Athabasca River cohesive sediments: towards managing sediment-related issues , 2011 .