Biogeosciences, 4, 269–282, 2007 www.biogeosciences.net/4/269/2007/ © Author(s) 2007. This work is licensed under a Creative Commons License. Biogeosciences Structure of mass and momentum fields over a model aggregation of benthic filter feeders J. P. Crimaldi1 , J. R. Koseff2 , and S. G. Monismith2 1 Department 2 Department of Civil and Environmental Engineering, University of Colorado, Boulder, CO, 80309-0428, USA of Civil and Environmental Engineering, Stanford University, Stanford, CA, 94305-4020, USA Received: 18 January 2007 – Published in Biogeosciences Discuss.: 9 February 2007 Revised: 19 April 2007 – Accepted: 19 April 2007 – Published: 16 May 2007 Abstract. The structure of momentum and concentration boundary layers developing over a bed of Potamocorbula amurensis clam mimics was studied. Laser Doppler velocimetry (LDV) and laser-induced fluorescence (LIF) probes were used to quantify velocity and concentration profiles in a laboratory flume containing 3969 model clams. Model clams incorporated passive roughness, active siphon pumping, and the ability to filter a phytoplankton surrogate from the flow. Measurements were made for two crossflow velocities, four clam pumping rates, and two siphon heights. Simultaneous use of LDV and LIF probes permited direct calculation of scalar flux of phytoplankton to the bed. Results show that clam pumping rates have a pronounced effect on a wide range of turbulent quantities in the boundary layer. In particular, the vertical turbulent flux of scalar mass to the bed was approximately proportional to the rate of clam pumping. 1 Introduction Shallow estuarine communities are commonly dominated by suspension feeders that filter phytoplankton and other particles from the overlaying flow. The extent to which these feeders can effectively filter the bulk phytoplankton biomass depends on the vertical distribution and flux of phytoplankton in the water column. Phytoplankton generally reproduce near the surface where there are high levels of incident light. Density stratification or low levels of vertical mixing can isolate phytoplankton in the upper layers of the water column (Koseff et al., 1993). On the other hand, turbulent mixing can distribute phytoplankton throughout the water column, where they become accessible to benthic grazers. But grazing acts as a near-bed sink of phytoplankton, Correspondence to: J. P. Crimaldi ([email protected]) and, in the absence of sufficient phytoplankton replenishment from above through mixing, can produce a phytoplanktondepleted, near-bed region called a concentration boundary layer. The severity of this concentration boundary layer depends on a balance between the rate at which grazers remove phytoplankton and the rate at which phytoplankton is delivered to the bed by turbulent mixing processes. The nature of the turbulence, and hence of the vertical mixing, depends on tidal energy, waves, bed geometry, and the presence of the benthic feeders themselves, whose geometric roughness and siphonal currents can alter the flow. In the present study, we investigate how aggregations of benthic suspension feeders can alter the structure of both the overlaying momentum and concentration fields. The physical roughness associated with benthic communities alters the turbulent velocity field above them (Butman et al., 1994; van Duren et al., 2006), significantly enhancing both turbulence intensities and Reynolds stresses. Active siphonal currents associated with filter feeding also impact the overlaying flow structure (Ertman and Jumars, 1988; Larsen and Riisgard, 1997), and have been shown to enhance turbulence intensities (Monismith et al., 1990; van Duren et al., 2006). The presence of suspension feeders also changes the structure of the concentration field above them (O’Riordan et al., 1993; Widdows and Navarro , 2007). Near-bed concentrations are reduced by the filtering action of the community, but this effect can be mitigated by additional mixing due to both physical roughness and siphonal pumping. We used model aggregations of the Asian clam Potamocorbula amurensis for this study. San Francisco Bay in California, USA is abundantly populated by this invasive species (Carlton et al., 1990), and it has become clear that the grazing pressure exerted by these clams alters the dynamics of phytoplankton blooms in the San Francisco Estuary. The goal of the study is to quantify changes to the momentum and concentration fields produced by the passive siphon roughness and active siphonal pumping of the clam aggregations. Published by Copernicus Publications on behalf of the European Geosciences Union. 270 J. P. Crimaldi et al.: Mass and momentum fields over filter feeders Crimaldi et. al: Mass and momentum fields over filter feeders 3 LDV/LIF optics LIF PMT To LIF amplifier Glass flume sidewall False Plexiglas sidewall Laser Beams 60 cm Measuring Volume 20 cm Model Clam Plates Flow False Plexiglas sidewall x Glass flume sidewall LDV PMT To LDV tracker Fig. 1.1.Top sidewalls, and and the theLDV/LIF LDV/LIFoptical opticalmeasurement measurementsystem. system. Fig. Topview viewofofthe theflume flumetest testsection sectionshowing showingthe themodel model clam clam plates, plates, false false sidewalls, Normalized mean velocity 2 1.2 8.75 Methods 1.0 2.10.8 Flume 0.6 U/U∞ The experiments were performed in an V/U open-channel, recir∞ 0.4 W/U∞Mechanics Labculating flume in the Environmental Fluid 0.2 oratory at Stanford University. The flume is constructed of 0.0 stainless steel and Plexiglas, with glass sidewalls in the test section to reduce laser refraction at the glass/water interface. -0.2 -10 -5 in normal0 operation 5is approximately 10 The flume capacity distance from flume centerline (cm)by a digital 8000 liters. Lateral A centrifugal pump commanded frequency controller draws water from a downstream reservoir charges a constant-head theofflume. Fig. 2. and Lateral profiles over a smoothtank bed upstream at z = 15ofcm mean An overspill excess water the constantstreamwise (U ), pipe lateralreturns (V ), and vertical (W )from velocities, normalized by tank the mean stream velocity Ureservoir. sidewalls are head backfree to the downstream Water from the ∞ . The false located at 10 and -10 fromthe the flume centerline. constant-head tankcm enters through a full-width diffuser and then passes through three stilling screens with decreasing coarseness to remove any large-scale structure in the geous smaller amounts of dye are flow because and homogenize the turbulence. Theused. flow then passes Excurrent flows consisted of ambient flume fluid witha through a 6.25:1, two-dimensional contraction andwith enters a rectangular dose of dyechannel. added toAit3-mm (Fig.rod 5). spanning This dosing process was the flume floor at the beginning channelas section trips the boundary layer done continuallyofinthe real-time the experiment is conducted. 2m upstream ofcontrolled the test section. The pump flow passes through An electronically centrifugal drew ambient the from test section, then through an exit section, over fluid the flume’s constant-head tank intoand thefinally excurrent an adjustable weirfluid backininto downstream reservoir. supply line. The thethe constant-head tank was Freewell streamand velocities the test section of 10 cmconcentration s−1 to 40 cm s−1 mixed had theinsame background dye as arefluid usedthat forwas this about study. to enter the flume. An electronically the The test section 3 m pumped long andconcentrated 0.6 m wide, with a rectcontrolled gear pumpisthen dye from a angular cross section and a nominal flow depth of 25 cm 20-liter reservoir into to the excurrent stream. The concentrated dye entered the excurrent stream via the dye injector, and was then mixed into the excurrent supply first by passBiogeosciences, 4, 269–282, 2007 www.biogeosciences.net/bg/0000/0001/ (Fig. 1). In the middle of the test section floor is a 1.8-m long by 20-cm wide, removable section that can accommodate either a set model clam smooth plate Topof View 6.40 plates or a single 3.20 for baseline flow measurements. A pair of thin Plexiglas sidewalls border the lateral edges of the model clam plates. 1.60 These false walls extend vertically from the bed through the free surface and act as symmetry planes3.20 to effectively model a wide bed of clams. The boundary layers developing on the false vertical walls grow to only about 1 cm 3.20thick at the Side View downstream edge, and thus have little effect on flow over the plates (Fig. 2). The flow is quite uniform across the inner test section, with no significant variation or secondary flow Fig. 3. Top and side views of a single model clam siphon pair in structure. the raised position, with the rubber siphon material shown in gray. for this siphons study were on theofflume cenTheMeasurements exhalant and inhalant (withmade diameters 1.6 mm and terline the model plates (or over smooth 3.2 mm,over respectively) areclam shown as white circlesa in the topplate view, for baseline The streamwise location thethe and as dashedmeasurements). lines in the side view. The side view shows of that measurements is denonted which the raised siphon models protrudeby 3.2x,mm into is themeasured flow. The from overlaying upstream layer edge flow of the platesleft (Fig. 1). The plates extend from boundary is from to right in the figure. Dimensions are mm. x=0in at the upstream edge to x=180 cm at the downstream edge. ing centrifugal excurrent supply pump, and then 2.2 through Model the Clams through an in-line static mixer. A flow meter measured total excurrent flow rate, and a manifold the supply Models of supply clam aggregations were placed in split the removable floor nine section of the flume. The models mimicked threethat clam into streams. Individual flow meters ensured the feautures: (1) the physical roughness associated with siphons nine streams had equal flow rates. Each of the nine streams that areinto raised flow, (2) clam the incurrent and excurpassed one into of thethe nine model plates, where a series rent siphonal flows associated with filter feeding, andof(3) of internal channels routed excurrent fluid to each one the the ability of the to plate. filter mass fromportion the flow. Two 441 excurrent jetsclams in each A small of the excurrent stream was diverted to the calibration jet for use as a reference for calibrating the LIF probe. www.biogeosciences.net/4/269/2007/ Biogeosciences, 0000, 0001–14, 2007 To LDV tracker est section showing the model clam false sidewalls, and the LDV/LIF optical measurement system 1. Top view of the flume test section plates, showing model plates, false sidewalls, and the LDV/LIF optical measurement system. J.Fig. P. Crimaldi et al.: Mass and momentum fieldsthe over filterclam feeders 271 Normalized mean velocity 1.2 8.75 8.75 1.0 0.8 Top View 0.6 U/U∞ V/U∞ W/U∞ 0.4 Top View 6.40 0.2 U/U∞ V/U∞ -0.2 W/U ∞ -10 6.40 3.20 1.60 3.20 3.20 0.0 -5 0 5 10 Lateral distance from flume centerline (cm) Side View 3.20 1.60 3.20 Fig. 2. Lateral profiles over a smooth bed at z = 15 cm of mean Fig. 2. Lateral over a smooth z = 15 cm of mean streamwise (U ), profiles lateral (V ), and verticalbed (Wat ) velocities, normalstreamwise (U ),free lateral (V ),velocity and vertical (W )false velocities, normalized by the mean stream U∞ . The sidewalls are ized by the mean free stream velocity U . The false sidewalls are ∞ located at 10 and −10 cm from the centerline. located at 10 and -10 cm from the centerline. 5 10 Fig. 3. Top and side views of a single model clam siphon pair in the raised position, with the rubber siphon material shown in gray. The exhalant and inhalant siphons (with diameters of 1.6 mm and 3.20 3.2 mm, respectively) are shown as white circles in the top view, Side View 0 and as dashed lines in the side view. The side view shows that the raised siphon models protrude 3.2 mm into the flow. The overlaying types of clam models from flume centerline (cm) were built, one with raised siphons boundary layer flow is from left to right in the figure. Dimensions (that were therefore rough) and one with flush siphons (that geous because smaller amounts of dye are used. mm. Fig.are 3. inTop and side views of a single model clam siphon pair in presented a smooth surface except for the presence of the Excurrent flows consisted of ambient flume fluid with with Fig. 3. Top and side views ofrubber a single modelshown clam siphon pai the raised position, with the siphon material in gray. siphon orifices). Tests were run with only one type of clam smooth model bed z = 15 cm of mean a doseat of dye added to it (Fig. 5). This dosing process was The exhalant and inhalant siphons (with diameters of 1.6 mm and (siphons raised or siphons flush) present the in theraised flume position, with the rubber siphon material shown in g continually in real-time as the experiment is conducted. the centrifugal excurrent supply pump, and then 3.2 ing mm,through respectively) are shown as white circles in the top view, and vertical ) Each velocities, atdone one(W time. model typenormalconsisted of nine identical 20The exhalant and inhalant siphons (with diameters ofthe1.6 and as dashed lines in the side view. The side view shows that An electronically controlled centrifugal drew ambient through an in-line static mixer. A flow meter measured total mm cm×20-cm square plates that fill theare 1.8 mpump long cutout in the velocity U . The false sidewalls ∞ raised siphon models protrude 3.2 mm into the flow. The overlaying fluid from flume’s tankarray into3.2 the excurrent excurrent supply flow rate,as andwhite a manifold split the mm, respectively) are shown circles in supply the top vi flume floor. the Each plate constant-head contains a 21×21 of individlayerstreams. flow is from left to right in the figure.ensured Dimensions supply fluid in the plus constant-head was in well boundary into nine Individual flow meters that the the centerline. ual clamline. siphonThe pairs (inhalant exhalant) , tank resulting in mm. in the side view. The side view shows that and as dashed lines mixed and had the same background dye concentration as are nine streams had equal flow rates. Each of the nine streams 441 clams per plate, and a total of 3969 clams in4the strip of the fluid thatThe wasclam aboutmodels, to enterwhile the flume. Anraised electronically passed into one of the 3.2 nine model clam the plates, whereThe a series siphon models protrude mm into flow. overlay nine plates. idealized, were fullcontrolled gear pumpofthen concentrated dye with from a of internal channels routed excurrent fluid to each one of the scale representations theirpumped biological counterparts, boundary layer441flow is from left to right in the figure. Dimensi 20-literboth reservoir into todimension the excurrent stream. The excurrent jets in each plate. A small portion of the exregard to siphonal and pumping rates.concenDiunts of dye are used. are mm. current stream was diverted to the calibration jet for use as a trated dyeand entered the excurrent stream via the dyein mensions flow rates were based on observations ofinjector, real and was then mixed into the excurrent supply first by passreference for calibrating the LIF probe. Potamocorbula amurensis clams made by Cole et al. (1992) d of ambient flume fluid with with and Thompson (personal comm.). 9.6 9.53 Fig. 5). This dosing wassiphons had the same Clam models withprocess raised and flush www.biogeosciences.net/bg/0000/0001/ Biogeosciences, 0000, 0001–14, 2007 and exhalant geometry (Fig. 3), but latter e as the inhalant experiment isorifice conducted. ingthethrough the centrifugal excurrent supply pump, and th did not protrude into the flow. Thus, the flush siphon model d centrifugal pump clam drew ambient for an individual consisted simply of a pairthrough of holes in an in-line static mixer. A flow meter measured to a smooth plate through which siphonal currents flowed. tant-headModels tankofinto the excurrent excurrent supply flow rate, and a manifold split the sup 9.53 9.6 individual clams were arrayed on plates placed the constant-head well intowasnine in the flume testtank section was (Fig. 4). This geometry re- streams. Individual flow meters ensured that peated across the entire array of 3969 clam models for both Fig.had 4. Top equal 2×2 arrayrates. of model Each clam siphon showackground dye concentration as nine streams flow ofpairs the nine strea the raised and flush siphon model types. Fig. 4. Top view of view aspacing 2 of×a 2used array model clam siphon ing the clam for theof study. The resulting clam array pairs sh Model clam plates were cast in a mold using apassed firm rubber;intoconsisted enter the flume. An electronically ofof 3969 the nine model clam plates, where a ser siphon pairs in a 21×189 pattern. Theresulting overlaying ing the clamone spacing used for the study. The clam a details of the construction process are given by O’Riordan boundary-layer flow is from left to right in the figure. Dimensions pumped(1993). concentrated dye from a of internal channels routed excurrent fluid to each one of consisted 3969 siphon pairs in a 21×189 pattern. The overla The interior of the plates included a series of chan- of are in mm. nels that linked all of the incurrent siphons together and all e excurrent stream. The concen441 excurrent jets each left plate. A small of the boundary-layer flow in is from to right in theportion figure. Dimens of the excurrent siphons together. The back of each plate rrent stream via the dye injector, current stream was diverted to the calibration jet for use a areand in one mm. had a pair of outlets: one for all incurrent siphons, Filter feeders inhale phytoplankton-laden fluid through for all excurrent siphons. Irregularities in the casting process their incurrent siphons and then exhale fluid with some frache excurrent supply first by passreference for calibrating the LIF probe. led to variations in the flow rates of individual clams. The tion of the phytoplankton removed through their excurrent siphons. Thus, while the volume of water entering and leavmeasured average deviation of the vertical excurrent velociIncurrent generated a similar ties across a row of individual clams was approximately 25% ingflows the clamwere is constant, the amountby of suspended scalarsystem, of the mean. Although this was not an intentional feature of mass (e.g. phytoplankton) is not. We model this concentrag/0000/0001/ Biogeosciences, 0000, 0001–14, 20 without dyetioninjection. An electronically controlled centr the design, it is likely more representative of the real-world change by labeling the excurrent flows with a fluorescent situation than uniform flow. dye. Thefluid model in clams inhale ambient from the flume, gal pump drew through thewater incurrent siphon orifi www.biogeosciences.net/4/269/2007/ while nine flowmeters ensured that each of the nine mo Biogeosciences, 4, 269–282, 2007 clam plates had the same incurrent flow rate. A mani then combined the nine incurrent streams, and the resul 272 J. P. Crimaldi et al.: Mass and momentum fields over filter feeders 5 Crimaldi et. al: Mass and momentum fields over filter feeders Incurrent Pump Return to Flume Reservoir Flume Constant-Head Tank Flowmeter Dye Pump Controller Excurrent Pump Controller 123.45 123.45 From Incurrents 9 Clam Plates Dye Reservoir Dye Pump To Excurrents Incurrent Pump Controller Dye To Flume From Flume Pump 123.45 Excurrent Supply Dye Injector Excurrent Pump Passive Mixer Flowmeter Manifold Fig. the incurrent incurrent and and excurrent excurrentsiphon siphonflows, flows,and andfor fordosing dosingthe theexcurrent excurrentflows flows Fig.5.5.Schematic Schematicofofthe theplumbing plumbingsystem systemresponsible responsible for for driving driving the with system, and and the the top top half halfisisthe theincurrent incurrentsystem systemAAtotal totalofof3969 3969model model withfluorescent fluorescentdye. dye.The Thebottom bottomhalf half of of the the figure figure is is the excurrent system, clam driven byby this system. clamsiphon siphonpairs pairs(grouped (groupedininnine nineplates) plates) are were driven this system. but exhale water that has a known concentration of dye. 1000they 8 Thus,6 in our system, clear fluid respresents phytonplanktonu 2 /u 2τ 4 laden water in the real system, and dyed fluid respresents wa2 2 w 2/u ter in the real system that has had phytoplankton filtered from τ 8 it.100 This inverse system, which was first used by Monismith 6 z +et al.4 (1990) and then later by O’Riordan (1993) is advantageous2 because smaller amounts of dye are used. 108 Excurrent flows consisted of ambient flume fluid with a dose64of dye added to it (Fig. 5). This dosing process was done continually in real-time as the experiment was conducted. 0 2 4 6 drew ambient 8 An electronically controlled centrifugal pump Normalized turbulence intensities fluid from the flume’s constant-head tank into the excurrent supply line. The fluid in the constant-head tank was well Fig. 6. Normalized streamwise (u) and vertical turbulence inmixed and had the same background dye (w) concentration as tensities. LDV Reflume. and lines are θ = 1320, the fluidSymbols that was are about to data enteratthe An electronically corresponding DNSpump results at Re from Spalartdye (1988). θ = 1410 controlled gear then pumped concentrated from a 20-liter reservoir into to the excurrent stream. The concentrated dye entered the excurrent stream via the dye injector, and was then mixed into the excurrent supply first by passBecause the dye fluoresces in an omni-directional pating through the centrifugal excurrent supply pump, and then tern, we were able to place the LIF receiving optics in the through an in-line static mixer. A flow meter measured total backscatter configuration without any loss of signal (as opexcurrent supply flow rate, and a manifold split the supply posed to thestreams. LDV receiving optics, which were placedthat preferinto nine Individual flow meters ensured the entially in the strong forward-scatter lobes). The LIF receivnine streams had equal flow rates. Each of the nine streams ing opticsinto andone theofLIF directly within the passed thePMT nine were modelmounted clam plates, where a series LDV front optics (using a backscatter module intended for of internal channels routed excurrent fluid to each one of the making backscatter LDV measurements). Thus, the receiv441 excurrent jets in each plate. A small portion of the exing opticsstream for thewas LIFdiverted automatically moved with current to the calibration jet the for measuruse as a ing volumefor as the LDV/LIF was traversed throughout reference calibrating thesystem LIF probe. the test section of the flume, maintaining consistent alignment. The PMT for the LIF had a pinhole section that Biogeosciences, 4, 269–282, 2007 www.biogeosciences.net/bg/0000/0001/ Incurrent flows were generated by a similar system, but 1000 8 6 dye injection. An electronically controlled centrifuwithout 4 gal pump drew fluid in through the incurrent siphon orifices 2 while nine flowmeters ensured uw / u 2τthat each of the nine model 1008 clam plates had the same incurrent flow rate. A manifold 6 ∂U 2 z + combined 4 /u streams, and the resulting ν then the nine incurrent ∂z τ stream2 returned into the flume reservoir downstream of the flume 108test section. There was a great deal of turbulence in 6 the flume reservoir (generated by the plunging action of the 4 flume flow spilling over the weir), and this turbulence con0.2 fluid in0.4 0.6 1.0 therestantly0.0 mixed the the reservoir. The0.8reservoir fore served as a well Normalized mixed source of fluid for the flume stress pump and the excurrent supply pump. The of background dye stresses. in the flume greware Fig. 7. concentration Normalized Reynolds and viscous Symbols LDV time data atdue Reto 1320, and lines are corresponding DNSflows. results with the constant dosing of the excurrent θ = at Resystem Spalartsuch (1988). θ = 1410 The wasfrom designed that this concentration growth was extremely linear in time (Crimaldi, 1998). Excurrent flows contained a fixed amount of dye added on top of the masked stray light from anywhere other than section. existing background flume concentration, so the the test difference More details on construction and operation ofthe theexcurrent LIF probe between the background concentration and the are given by Crimaldi concentration remained(1998). constant with time. The smallest scalar fluctuations in a flow occur at the scale at which viscous diffusion acts to smooth any remaining con2.3 Instrumentation centration gradients. Batchelor (1959) defines this scale as 2.3.1 Velocity measurements ηB = ηK Pr−1/2 (2) A Dantec, two-component, laser-Doppler velocimeter (LDV) where Pr to is the Prandtl number. According to Barrett was used measure velocities. This instrument was (1989), operthe Prandtl number Rhodaminefluorescence 6G, the dye(LIF) usedprobe in the ated in tandem with afor laser-induced study, is 1250. Therefore, the smallest concentration scales ηB = 3 µm were about 35 times smaller than the smallest www.biogeosciences.net/4/269/2007/ Biogeosciences, 0000, 0001–14, 2007 Fig. 5. 5. Schematic Schematic of of the the plumbing plumbingsystem systemresponsible responsiblefor fordriving drivingthe theincurrent incurrentand andexcurrent excurrentsiphon siphonflows, flows,and andfor fordosing dosingthe theexcurrent excurrentflows flows Fig. with fluorescent dye. The bottom half of the figure is the excurrent system, and the top half is the incurrent system A total of 3969 model with fluorescent dye. The bottom half of the figure is the excurrent system, and the top half is the incurrent system A total of 3969 model clam siphon pairs pairs (grouped in nine nine plates)are arefields drivenover bythis this system. J.clam P. Crimaldi et al.: Mass in and momentum filter feeders 273 siphon (grouped plates) driven by system. 10008 1000 8 10008 1000 8 6 6 4 4 22 + zz+ 6 6 4 4 2 /u22 uu2 /u ττ 22 2/u22 ww2/u ττ 1008 100 8 66 44 + uw/ u/ 2u 2τ uw τ ∂U 2 ∂U 2 ν τ ν ∂z/u/u ∂z τ 1008 100 8 z z+ 66 44 22 22 108 10 8 1010 88 66 44 66 44 00 22 44 66 88 0.0 0.0 Normalizedturbulence turbulenceintensities intensities Normalized 0.2 0.2 0.4 0.4 0.6 0.6 0.80.8 1.01.0 Normalizedstress stress Normalized Fig.6.6. 6.Normalized Normalizedstreamwise streamwise(u) (u)and andvertical vertical (w) turbulence inFig. (w) turbulence inNormalized streamwise (u) and vertical (w) turbulence intensities.Symbols Symbols are LDV data Reθθ = = and 1320, and lines are tensities. areare LDV datadata at Re =1320, lines arelines corretensities. Symbols LDV atatθ Re 1320, and are sponding DNS results at Reθ at =1410 from Spalart corresponding DNS results results at Re Reθθ = = 1410 1410 from(1988). Spalart(1988). (1988). corresponding DNS from Spalart Fig. Normalized Reynolds and viscous stresses. Symbols are Fig. Normalized Reynolds and viscous stresses. Symbols Fig. 7.7.7.Normalized Reynolds and viscous stresses. Symbols areare LDV data atRe 1320, andlines lines arecorresponding corresponding DNSresults results LDV data are LDV data atatRe =1320, and and lines are corresponding DNSDNS results at θ θ==1320, θRe Reθ θ==from 1410Spalart fromSpalart Spalart (1988). atat 1410 from (1988). Re =1410 (1988). θ Re to measure mass fluxes. The LDV was driven with an argonBecause the dye dye fluoresces in an an omni-directional patthe in omni-directional patIonBecause laser operated in fluoresces the 514.5 nm single-line mode, with we were were able to place place themeasuring LIF receiving receiving optics the we able the LIF optics the atern, nominal output of to 1.0 W. The volume wasininelbackscatter configuration without any loss of ofin signal (asopopbackscatter configuration any loss signal (as liptical in shape, with the without long axis oriented the crossposed to the optics, which were prefertodirection. the LDV LDV receiving receiving optics,of which wereplaced placed preferchannel The dimensions the measuring volume entially in the forward-scatter lobes). LIF receiventially in intensity the strong strong forward-scatter lobes). The The0.1 LIFmm receiv(to the e−2 contour) were approximately in ingvertical optics LIF were within the optics and and the the LIF PMT PMT weremounted mounted directly within the the streamwise directions, and 1directly mm in the crossLDV front optics (using aa backscatter intended for front opticsThe (using backscatter modulein intended for channel direction. smallest scales of module motion the flows making LDV the making backscatter backscatter LDV measurements). Thus, thereceivreceivmeasured in this study can measurements). be determined byThus, estimating the ing optics the automatically moved optics for for the LIF LIF(Tennekes automatically movedwith withthe themeasurmeasurKolmogorov scale as and Lumley, 1972) ing volume volume as as the the LDV/LIF LDV/LIFsystem systemwas wastraversed traversedthroughout throughout !1 the test section testκzν section of the the flume, flume, maintaining maintaining consistent consistent alignalign3 4 of ηment. (1) The PMT for the LIF had a pinhole section that K ≈ The PMT for the LIF had a pinhole section that u3τ masked stray lightfrom fromanywhere anywhere other thanthe the test section. masked light test section. station, atstray a sample rate of 80 Hz. other The than mean free stream −1and Moredetails details on construction and operation of the LIF probe More construction operation of the LIF probe velocity was Uon =11.6 cm s , which resulted in a calcu∞ aregiven givenby byCrimaldi Crimaldi(1998). (1998). are lated momentum-thickness Reynolds number of Reθ =1320. fluctuations inina aflow atatthe Thesmallest smallest scalar fluctuations flowoccur occur thescale scale We The compared thescalar data to Spalart’s DNS results simulated at atat which viscous diffusion acts to smooth any remaining conwhich viscous diffusion acts to smooth any remaining conRe =1410. Although the Reynolds numbers differ by 6%, θ centration Batchelor (1959) defines scale centration gradients. Batchelor (1959) definesthis this scale the variationgradients. of normalized turbulence parameters with Reasθas is quite weak, enabling a valid comparison. −1/2 −1/2 ηηBB(Fig. ==ηηK6) Pr Pr (2) K Turbulence intensities were normalized by the(2) square of the shear velocity uτ that was obtained by fitwhere Pr Prandtl number. According totoBarrett (1989), where Prisisthe the Prandtl number. According Barrett ting the mean velocity profile to the law of the wall. (1989), The the Prandtl number for Rhodamine 6G, the dye used ininthe the Prandtl number for Rhodamine 6G, the dye used the agreement between the LDV results and DNS calculations study, isis1250. Therefore, the smallest concentration scales study, 1250. Therefore, the smallest concentration scales by Spalart (1988) was excellent for both the turbulence inηηBB == 33µm were about 35 times than smallest about timessmaller smaller thanthe the smallest tensities andµm thewere viscous and 35 turbulent stress profiles (Fig. 7). where κ is the Kolmogorov constant, z is distance from the www.biogeosciences.net/bg/0000/0001/ www.biogeosciences.net/bg/0000/0001/ bed, ν is the viscosity, and uτ is the shear velocity. The smallest value of ηK for the flows in this study (corresponding to uτ =1.7 cm s−1 and z=0.1 cm) was approximately 0.1 mm. Although this value of ηK is comparable to the dimension of the measuring volume in the vertical and streamwise directions, it is smaller than the dimension in the cross-flow direction. Nonetheless, the LDV easily captures the larger scales responsible for the transport of mass and momentum. The LDV laser and optics were mounted on a motorized, computer-controlled three-axis traverse which permitted the LDV measuring volume to be positioned anywhere within the test section. The traverse system was accurate to within approximately 200 µm. For validation purposes, LDV measurements of boundary layer turbulence were taken in the flume over a smooth plate that was installed in the same location where the model clam plates were later installed. The LDV results were then compared with direct numerical simulations (DNS) of boundary layer turbulence over a smooth bed by Spalart (1988). Data were recorded at 23 logarithmically-spaced vertical stations between z=0.7 mm and z=180 mm, measured from the bed. Approximately 20 min of velocity data were recorded at each 2.3.2 www.biogeosciences.net/4/269/2007/ Concentration measurements Biogeosciences, Biogeosciences,0000, 0000,0001–14, 0001–14,2007 2007 We developed a laser-induced fluorescence (LIF) probe to make non-intrusive measurements of dye concentrations in the flow above model clam beds. The LIF probe used the same measuring volume as the LDV, ensuring that the velocity and concentration measurements were made in the same location. This is particlarly important for the scalar flux measurements, which resulted from correlations of velocity and concentration measurements. The laser light in the combined LDV/LIF measuring volume was absorbed by fluorescent dye in the flow and re-emitted at a different wavelength. The fluoresced light was optically filtered and converted to a electrical current with a photomultiplier tube (PMT). Finally, the current is converted to a voltage using an ideal currentto-voltage converter. Because the dye fluoresces in an omni-directional pattern, we were able to place the LIF receiving optics in the backscatter configuration without any loss of signal (as opposed to the LDV receiving optics, which were placed preferentially in the strong forward-scatter lobes). The LIF receiving optics and the LIF PMT were mounted directly within the LDV front optics (using a backscatter module intended for Biogeosciences, 4, 269–282, 2007 274 J. P. Crimaldi et al.: Mass and momentum fields over filter feeders 6 Crimaldi et. al: Mass and momentum fields over filter feeders probe. The time response of the LIF probe is extremely fast; without dyeassociated (other than background dye)with represented the time constants withthe the dye fluorescence, phytoplankton-laden fluid. Using the LIF probe, 0.8 the PMT, and with the LIF signal amplifier are extremelywe calcua nondimensional in thisin“inverse” syssmall lated relative to the time scalesconcentration of turbulent motion this 0.6 flow. tem as − CB ∗ modelC During experiments over the Cinv = clams, the, LIF calibra(3) 0.4 C CB the measureE − tion jet remained in the flume, positioned above 0.2 ment where region in freeoutput streamofofthe theLIF flow.probe A small portion C the is the at the measurement of thelocation, dyed flowCmixed for the excurrent supply was diverted B is the output due to the background dye (mea0.0 through the in calibration jet. The and LIF C probe was periodically sured the free stream), E is the output of pure excur0.0 0.2 0.4 0.6 0.8 1.0 positioned in the free stream and behind the calibration jet this “inrent fluid (measured using the calibration jet). To put Normalized calibration jet concentration during experiments to maintain the probe calibration as backverse” measurement in a more intuitive framework, we then ground and excurrent concentrations rose during the experia complementary nondimensional concentration Fig.8.8.Normalized Normalized calibration curve for LIF the LIF showing Fig. calibration curve for the probeprobe showing the thementsdefined due to dye accumulation in the flume. linear system response. linear system response. Normalized LIF output 1.0 2.3.3 ∗ C ∗ = 1 − Cinv Concentration normalization (4) ∗ such that C = 1 now corresponded (in the real system) to making backscatter measurements). Thus, the receivscales of motion ηLDV was conK . Thus, the LIF probe could not measureAs discussed earlier, a known concentration of dye fluid with full phytoplankton load, and C ∗ = 0 corresponded ing optics for the LIF automatically moved with the measurthe smallest scales of motion present in the studied flows.tinuously added to the excurrent jets. The dyed fluid repto fluid that had its entire phytoplankton concentration reing volume as the LDV/LIF system was traversed throughout However, the LIF probe could easily measure the larger con-resented filtered fluid devoid of phytoplankton, and fluid the test section of the flume, maintaining consistent alignmoved by filtration. These nondimensional concentrations centration scales that are responsible for the vast majority ofwithout dye (other than the background dye) represented ment. The PMT for the LIF had a pinhole section that could be converted dimensional concentrations phytoplankton-laden fluid. toUsing the LIF probe, we calcu-for a real the scalar flux. masked stray light from anywhere other than the measureby considering the ambient phytoplankton concentralated system a nondimensional concentration in this “inverse” sysTovolume. validateMore the LIF probe, we measured dye ment details on construction and known operation of con-tem astion and the filtering efficiency of the bivalve. centrations potential core(1998). of a jet flowing from a the LIF probe in arethe given by Crimaldi C − CB 7.5 mm diameter the at flume test sec- ∗ The smallest scalarcalibration fluctuationstube in a within flow occur the scale , (3) Cinv = − CB measurements oftothe jet fluid were made 1 mm attion. whichLIF viscous diffusion acts smooth any remaining con3 CEResults downstream of the Batchelor jet orifice(1959) (x/Ddefines = 0.13), centration gradients. thiswhich scale asensuredwhere C is the output of the LIF probe at the measurement Vertical ofdue velocity and concentration that the measuring volume was well within the potential corelocation, CB is profiles the output to the background dye (mea-data were ηof Pr−1/2 (2)from B = taken for two crossflow velocities and four clam pumping theηKjet. Eight different dye concentrations ranging sured in the free stream), and CE is the output of pure excur- rates, for eachusing of the clam model ((Table 0 to 100 were pumped theto jet, and(1989), the result-rent fluid (measured thetwo calibration jet). Totypes put this “in- 1). For where Pr isppb the Prandtl number. through According Barrett vertical profile, simultaneous LDV and ingPrandtl LIF data are linear (Fig 8). Calibration jetused concentrations verse”each measurement in a more intuitive framework, weLIF thendata were the number for Rhodamine 6G, the dye in the acquired at approximately 20 vertical stations. The stations were normalized by the maximum value used in the test (100defined a complementary nondimensional concentration study, is 1250. Therefore, the smallest concentration scales were logarithmically spaced, usually starting at z = 0.5, and LIF output was normalized so that the output from the ηppb). =3 µm were about 35 times smaller than the smallest B ∗ C ∗ =ending 1 − Cinv (4) of data at z = 120 mm. Typically, 100,000 samples 40 ppb was η0.4. A least-squares estimate of the slope of scales of jet motion . Thus, the LIF probe could not resolve K the in thedye studied flows. wereCacquired approximately from each LDV chanthesmallest line wasscales 1.006of±motion 0.003.present The actual concentrations ∗ =1 nowatcorresponded such that (in80 theHz real system) to However, the LIF probe could easily measure the larger con- ex-fluid with ∗ nel and from the LIF probe for each vertical station, although used in the measurements over the model clams rarely full phytoplankton load, and C =0 corresponded centration that are responsible for the vast range majority shorter records were phytoplankton used near the concentration upper edge ofrethe boundceeded 5 scales ppb, well within the demonstrated of of linear-to fluid that had its entire the layer where the variance of the signals was small. ityscalar of theflux. LIF probe. The time response of the LIF probe ismovedary by filtration. These nondimensional concentrations To validate thethe LIFtime probe, we measured known dye presented in thisconcentrations paper focus for ona perturbations extremely fast; constants associated with thecondye flu-could beResults converted to dimensional real centrations in the potential core of a jet flowing from a by considering the ambient concentrato the momentum andphytoplankton scalar concentration fields by the orescence, with the PMT, and with the LIF signal amplifiersystemmade 7.5 mm diameter calibration tube within the flume test secthe filtering the bivalve. presence of theefficiency clams. of These perturbations come from two are extremely small relative to the time scales of turbulenttion and tion. LIF measurements of the jet fluid were made 1 mm sources: the presence of roughness (in the case of the model motion in this flow. downstream of the jet orifice (x/D=0.13), which ensured clams with raised siphons), and the presence of siphonal curDuring experiments over the model clams, the LIF calibrathat the measuring volume was well within the potential core 3 Results rents and filtering. The flush siphonal orifices without clam in the flume, positioned above the from measureoftion thejet jet.remained Eight different dye concentrations ranging pumping alter the (Figs. 9 and region in the free stream A small portionVertical 0ment to 100 ppb were pumped throughofthethe jet,flow. and the resulting profilesdid of not velocity and flow concentration data10). were The data these twovelocities figures compares boundary-layer flow of the mixed LIF datadyed wereflow linear (Fig. for 8). the excurrent supply was divertedtaken shown for twoincrossflow and four the clam pumping a smooth withmodel the flow over the flush clam modthrough the calibration jet. Thewere LIF probe was periodically Calibration jet concentrations normalized by the rates, over for each of the plate two clam types (Table 1). For els, withprofile, no siphonal currents. experiments maximum the test ppb).theLIF output jeteach vertical simultaneous LDVBoth and LIF data werewere perpositionedvalue in theused freeinstream and(100 behind calibration was normalized so that the output from the 40 ppb jet was acquired at approximately 20 vertical stations. The formed with a free stream velocity of U 10 cm s−1 , corduring experiments to maintain the probe calibration as back∞ =stations 0.4. A least-squares estimate of the slope the line logarithmically spaced, usually startingshow at z=0.5, andflow over to Re Results that the ground and excurrent concentrations roseofduring thewas experi-were responding θ = 560. 1.006±0.003. The actual dye concentrations used in the ending at z=120 mm. Typically, 100 000 samples of data the flush siphon orifices was indistinguishable from flow over ments due to dye accumulation in the flume. measurements over the model clams rarely exceeded 5 ppb, were acquired at approximately 80 Hz from each LDV the smooth plate. Thus, the perturbations to thechanflow demonwell within the demonstrated range of linearity of the LIF nel and from the LIF probe for each vertical station, strated later are due only to the presence of although siphon roughness 2.3.3 Concentration normalization As discussed earlier, a known Biogeosciences, 4, 269–282, 2007concentration of dye was continuously added to the excurrent jets. The dyed fluid represented filtered fluid devoid of phytoplankton, and fluid and/or siphonal pumping. The effects ofwww.biogeosciences.net/4/269/2007/ siphon roughness and pumping on the shear velocity are shown in Fig. 11. Shear velocity uτ is a measure of the bed shear stress τw , where uτ = (τw /ρ)1/2 . As is Crimaldi et. al: Mass and momentum fields over filter feeders Crimaldi al: Mass momentum fields fields over filter feeders J.Crimaldi P.et.Crimaldi et and al.: Mass momentum fields over filter feeders et. al: Mass andand momentum over filter feeders 7 275 7 7 Table 1. Parameters varied in the experiments 1.1.Parameters Table 1.Table Parameters variedvaried in theinexperiments Table Parameters varied inthe theexperiments. experiments Parameter Symbol Values Comments Parameter Symbol Values Comments Parameter Symbol Comments Parameter Symbol Comments Free stream velocity U∞ValuesValues 10, 40 cm s−1 −1 −1 Clam pumping rate Q 0, 0.030, 0.045, 0.060 ml s rate per clam Free velocity 10, 4040 cm s s−1 Free stream velocity cm s−1 Freestream stream velocityU∞ U∞ U∞10, 40 10, cm −1 Clam siphon height h 0, 3.2 mm “flush” and “raised” in text s 0,0, 0.030, 0.045, 0.060 s s−1rate raterate perclam clam Clampumping pumping rate Q QQ 0, 0.030, 0.030, 0.045, 0.060 ml per clam Clam Clam pumping rate rate 0.045, 0.060 mlmls−1 per Clam siphon height h 0, 3.2 mm “flush” and “raised” in text sh Clam siphon height 0, 3.2 mm “flush” and “raised” in text Clam siphon height hs s 0, 3.2 mm “flush” and “raised” in text 1000 22/u22 ττ u 2 /u 2τ uw /u 2 2 w 2/u 2τ w /u τ 100 2 z +100 100 6 4 26 4 62 4 2 10 10 U∞ (cm/s) z + 10 26 4 62 4 6 4 40 1 30 20 2 0 1 2 0 0 2 2 4 6 20 20 4 6 10 8 10 4 6 8 Normalized turbulence intensities 1.2 1.1 1.0 0.9 0.8 8 Normalized turbulence intensities 1.5 1.3 30 40 1.5 1.3 1.4 1.2 40 40 (a) siphons (a) siphons flushflush 1.4 30 1.4 10 2 1 40 (b) siphons raised 1.5 0.7 00.6 U∞ (cm/s) U∞ (cm/s) U∞ (cm/s) 62 4 40 U∞ (cm/s) U∞ (cm/s) 6 4 z+ u 2 /u 2τ 1000 1000 (a) siphons flush 6 4 1.3 1.1 1.2 1.0 30 1.1 0.9 1.0 0.8 0.9 0.7 0.8 0.6 20 0.7 0.5 0.6 0.5 0.5 0 .030 .045 Q (ml/s) .030 .045 .060 .060 10 1.7 siphons raised (b) (b) siphons raised 1.6 1.7 1.6 1.5 1.4 30 1.3 20 1.2 1.1 20 1.0 0.9 10 0.8 0 0.7 0.6 10 0.5 30 0 1.5 1.7 1.4 1.6 1.3 1.5 1.2 1.4 1.1 1.3 1.0 1.2 0.9 1.1 0.8 1.0 0.7 0.9 0.6 0.8 0.5 0.7 0.6 0.5 .030 .045 Q (ml/s) .030 .045 .060 .060 0 .030 .045 .060 0 .030 .045 .060 Q (ml/s) Q (ml/s) Fig. 9. Comparison of streamwise and vertical turbulence intensiNormalized turbulence intensities Fig. 11. Contours of shear velocity uτ as a function of free stream Q (ml/s) Q (ml/s) ties for flow over non-pumping flush siphon orifices (closed symvelocity U∞ and clam pumping rate Q for flow over (a) flush Fig. 9. Comparison of streamwise and vertical turbulence intensiFig. Comparison streamwise vertical turbulence intensibols)9.with flow over of a smooth plate and (open symbols). Fig. 11. and Contours of shear velocity uτforasua function of siphons (b) of raised siphons. Units arefree cmstream s−1 . Fig. 9. ties Comparison of streamwise and vertical turbulence intensifor flow over non-pumping flush siphon orifices (closed symτ contours ties for flow over non-pumping flush siphon orifices (closed sym- Fig. 11. Contours shear velocity uττrate as function ofoffree Fig. 11. Contours of shear velocity u as aaQ function freestream stream velocity U clam pumping for flow over (a) flush ∞ and over plate (open symbols). ties for bols) flow over non-pumping flush siphon orifices (closed symbols)with withflow flow overa asmooth smooth plate (open symbols). −1 velocity U∞ and clam pumping pumping rate Q over (a) flush velocity U∞ and Qforfor for flow overare (a) flush siphons and (b)clam raised siphons. rate Units uτ flow contours cm s . −1−1 bols) with flow over a smooth plate (open symbols). siphons and (b) raised siphons. Units for u contours are cm s . τ siphons and (b) raised siphons. Units for u contours are cm s . 3.1 1000 6 4 uw/u 2τ 62 4 uw/u 2τ Profiles τ 3.1 Profiles present vertical profilesflow of velocity and contwoFigures figures12-18 compares the boundary-layer over a smooth 3.1 Profiles 1000 centration data in a common figure format. Because the in100 plate with 12-18 the flow over vertical the flush clam of models, with Figures present profiles velocity andnocon6 2 26 uw/u fluence of the clams is greatest in the near-bed region, dis4 τ 4 siphonal currents. experiments performed with centration data inBoth a vertical common figure were format. Because the Figures 12-18 present profiles ofinvelocity and con-inz + 100 2 tance from the bed (the vertical axis the plots) is shown −1 2 6 a free stream velocity ofisUgreatest =10 cm sthe ,near-bed corresponding ∞figure fluence of the informat. region, discentration data in aclams common Because theto in100 z + 104 on a logarithmic scale. Each figure contains four plots Re =560. Results show that the flow over the flush siphon θ tance from the bed (the vertical axis in the plots) is shown 6 26 fluence of the clams is greatest in the near-bed region, velocdisrepresenting different combinations free the stream 4 4 orifices indistinguishable fromfigure flowof over smooth z+ onfrom a was logarithmic scale. Each contains four plots 10 tance the bed (the vertical axis in the plots) is shown 2 ity U positionto (flush or demonstrated raised). For later profiles 62 ∞ and plate. Thus, thesiphon perturbations the flow representing different combinations of free stream veloc4 1 onare aoflogarithmic scale. Each figure contains plots 10 concentration-related quantities, there is no four data for the tosiphon the presence siphon roughness and/or 2 6 itydue U∞only and position of(flush or free raised). For profiles 0.0 0.2 0.4 0.6 0.8 1.0 representing different combinations of stream velocQ = 0 case since clam pumping is required for concentra4 siphonal pumping. 1 of concentration-related quantities, there is no data for the Normalized Reynolds stress 2 ity UShear siphonu position (flushof or raised). For profiles tion measurements. ∞ and velocity a measure the bed shearfor stress τw , 0.0 0.2 0.4 0.6 0.8 1.0 τ isclam Q = 0 case since pumping is required concentra1 1/2 of concentration-related quantities, there is no data for The influence of clam pumping on U was small andthe limwhere u =(τ /ρ) . As is typical in turbulent boundary Normalized Reynolds stress τ w tion measurements. Fig.10. 10.Comparison Comparison ofReynolds Reynolds stress correlations flow over Fig. of stress correlations forfor flow over 0.0 0.2 0.4 0.6 0.8 1.0 to thesince near-bed region (Fig. 12). The effect was largest Q layer =ited 0 flows, case clam pumping is required for concentrauτ increases with the freeon stream velocity, U∞ lim. The influence of clam pumping U was small and non-pumpingflush flush siphon orifices (closed symbols) with flow over non-pumping siphon orifices (closed with flow over Normalized Reynolds stresssymbols) forpresence the slowof(Uthe = 10 siphons cm s−1 produced ) flow over raised siphons, tion measurements. ∞ raised Fig. 10. Comparison of Reynolds stress correlations for flow over The an increase in smoothplate plate (open symbols). The lines DNS results Spalart ited to the near-bed region (Fig. 12). The effect was largest a asmooth (open symbols). The lines areare DNS results byby Spalart where near-bed values of U were as Qsiphons increased. non-pumping flush siphon orifices (closed symbols) with flow over u The of clam U to was and limat influence any of Upumping thesmall flush −1on retarded τ for ∞ as compared (1988)atatReRe (1988) =670. θ = 670. the given slow value (U ) flow over raised siphons, θ ∞ = 10 cm s Fig. 10. aComparison of Reynolds stress correlations for flow over This was consistent with the u contours in Fig. 11, where smooth plate (open symbols). The lines are DNS results by Spalart ited τ only (Fig. 11). Siphonal pumping caused modest increase in to the near-bed region (Fig. 12). The effect was largest where near-bed values of U were retarded as Q increased. non-pumping orifices (closed symbols) with flow over −1 (1988)flush at Resiphon the greatest sensitivity toswas changes inpronounced wall raised stress(i.e., weretheseen θ = 670. ,This although the the increase more foruτthe slow (U = 10 cm ) flow over siphons, ∞ was consistent with the uτ contours in Fig. 11, where a smooth plate (open symbols). The lines are DNS results by Spalart fornear-bed slow overwas raised siphons. For flush siphons or faster slope of theflows contours for the raised siphon clam shorter records were used near the upper edge of the bound- where values of greater) U retarded asstress Q increased. the greatest sensitivity to were changes in wall were seen (1988) at Reθ = 670. flows, the effect of Q on U uwascontours negligible milmodels at low free stream velocities. ary layer where the variance of the signals was small. This was consistent with the in beyond Fig. 11,a few where τ for slow flows over raised siphons. For flush siphons or faster limeters from the bed. typical turbulent boundary layerfocus flows,onuτ perturbations increases with Resultsinpresented in this paper the greatest in wall stress were seen flows, thesensitivity effect of Qtoonchanges U was negligible beyond a few milthe free stream velocity, U∞ . concentration The presencefields of the For the slow flow with flush siphons, clam pumping attenProfiles made to the momentum and scalar byraised the slow flows overthe raised limeters from bed. siphons. For flush siphons or faster typical in turbulent boundary layer flows, uτ increases with for3.1 siphons produced an increase in uτ at any given of U∞ uated the streamwise turbulence intensities (expressed as the presence the clams. These comeofvalue from two flows, the of Q onwith U was negligible a few milFor effect the slow flow flush siphons, beyond clam pumping attenthe free of stream velocity, U∞perturbations . The presence the raised uu)present in a narrow region centered aroundand z =con4 mm as compared to the flush siphons (Fig. 11). Siphonal pump- Figures variance sources: the presence of roughness (in the case of the model 12-18 vertical profiles of velocity the bed. turbulence produced an increase in uτ atuany given value of U∞ limeters uatedfrom the streamwise intensities (expressed as the typical siphons in boundary flows, with ingturbulent caused only modestlayer increase in uττ, increases although the the (Fig. 13a). This to theformat. height atBecause which the verclams with raised siphons), and the presence of siphonal cur-in- centration data in aacorresponds common figure the uu) in narrow region centered z =atten4being mm compared to the flush siphons (Fig. 11). Siphonal pumpvariance the freeas stream velocity, U . The presence of the raised For the slow flow with flush siphons, clamaround pumping ∞ tical excurrent achieve a horizontal crease more pronounced (i.e., the slope of the contours influence rents andwas filtering. of the jets clams is greatest in the trajectory near-bed after region, caused an only modest in given uτ , although thefree in- uated(Fig. 13a).by This corresponds to the height at which as theet versiphonsing produced increase inincrease uτsiphon at any value ofthe Ulow the over streamwise turbulence[see intensities the ∞ bent Fig.in 4 (expressed inplots) O’Riordan al. was for theorifices raised distance from thethe bedcrossflow (the vertical axis the is shown Thegreater) flush siphonal withoutclam clam models pumpingatdid not crease was more pronounced (i.e., the slope of the contours tical excurrent jets achieve a horizontal trajectory after being as compared to the flush siphons (Fig. 11). Siphonal pumpuu) in a narrow region centered around z = 4 mm variance stream Since the excurrent were laminar, streamwise alter the velocities. flow (Figs. 9 and 10). The data shown in these on(1995)]. a logarithmic scale. Each jets figure contains four plots was only greater) for the raised in siphon clam models at low by corresponds the crossflowto[see in which O’Riordan et al. 13a).over This the Fig. height4 at the vering caused modest increase uτ , although the the in- free (Fig.bent (1995)]. Since the excurrent jets were laminar, streamwise stream velocities. tical excurrent jets achieve a horizontal trajectory after being crease was more pronounced (i.e., the slope of the contours www.biogeosciences.net/bg/0000/0001/ Biogeosciences, 0001–14, www.biogeosciences.net/4/269/2007/ Biogeosciences, 4,O’Riordan 269–282, 2007 bent over by the crossflow [see Fig. 4 in0000, et 2007 al. was greater) for the raised siphon clam models at low free stream velocities. (1995)]. Since the excurrent jets were laminar, streamwise www.biogeosciences.net/bg/0000/0001/ Biogeosciences, 0000, 0001–14, 2007 1000 276 J. P. Crimaldi et al.: Mass and momentum fields over filter feeders 8 Crimaldi et. al: Mass and momentum fields over filter feeders (a) Siphons flush, U∞ = 10 cm/s (b) Siphons flush, U∞ = 40 cm/s z (mm) 100 100 Q = 0.000 Q = 0.030 Q = 0.045 Q = 0.060 10 10 1 1 0 2 4 6 8 10 0 z (mm) (c) Siphons raised, U∞ = 10 cm/s 100 10 10 1 1 2 4 6 8 20 30 40 (d) Siphons raised, U∞ = 40 cm/s 100 0 10 10 U (cm/s) 0 10 20 30 40 U (cm/s) Fig. 12. Effect of clam Q on Q vertical profiles of mean streamwise velocity U for combinations of of free stream Fig. 12. Effect ofpumping clam pumping on vertical profiles of mean streamwise velocity U different for different combinations free streamvelocity velocityU∞ −1 and −1= U∞ and siphon roughness. Theright left and right correspond columns correspond to Ucm cm s−1=40 and 40 cm s−1 , respectively. The and siphon roughness. The left and columns to U∞ =10 U∞ cmUs∞ , respectively. The top and bottom ∞ s= 10 top and bottom rows correspond to siphons flush (h (hss=3.2 = 0) and respectively. raised (hs = The 3.2 mm), respectively. The bottom row plotsdotted containline a at rows correspond to siphons flush (hs =0) and raised mm), bottom row plots contain a horizontal horizontal linethe at location z = hs of = the 3.2raised mm tosiphon denotetops the (below locationwhich of the data raised siphon (below which couldocclusion not be acquired z=hs =3.2 mm todotted denote could nottops be acquired due todata optical of the indue toEach optical the instruments). plot of contains color-coded each of(Q=0, the different clam pumping struments). plotocclusion containsof color-coded profilesEach for each the different valuesprofiles of clamfor pumping 0.030,values 0.045,ofand 0.060 ml s−1 ). −1 (Q = 0, 0.030, 0.045, and 0.060 ml s ). representing different combinations of free stream velocity turbulence intensities were locally reduced. The effect is U∞ and siphon position (flush raised). Fortheprofiles of similar for the fast flow case or (Fig. 13b), but attenuation concentration-related there is no data formore the Q=0 was closer to the quantities, bed as the jets were bent over rapidly case since pumping is required for slow concentration by theclam stronger crossflow. For the flow overmearaised surements. siphons (c), there was a similar near-bed attenuation by the pumping, but it pumping was now accompanied by a strong enTheclam influence of clam on mean velocity U was the bed. for the flow small hancement and limitedfurther to the from near-bed regionFinally, (Fig. 12). Thefast effect over the siphons (d), thecm effect clam over pumping was was largest forraised the slow (U∞ =10 s−1of) flow raised minimal as the turbulence intensities were dominated by the siphons, where near-bed values of U were retarded as Q inflow and roughness. creased. This was consistent with the uτ contours in Fig. 11, where the to changes in wall stress intensiwere Thegreatest effect ofsensitivity clam pumping on vertical turbulence tiesslow is opposite fromraised that seen for theFor horizontal streamwise seen for flows over siphons. flush siphons or 14).ofThe vertical energy imparted by the ainfaster intensities flows, the(Fig. effect Q on U was negligible beyond current andfrom excurrent flows enhanced the vertical turbulence few millimeters the bed. intensities. The effect was strongest for the slow flows (a, c) For the slow flow with flush siphons, clam pumping atwhere the relative strength of the clam pumping was stronger. tenuated the streamwise turbulence intensities (expressed as In these cases, the influence of the pumping extended deep the variance in a narrow around into theuu) boundary layer. region For thecentered faster flows (b, d)z=4 the mm effect (Fig. 13a). This corresponds to the height at which the verwas minimal except close to the bed. tical excurrent jets achieve a horizontal trajectory after being The of roughness due to theO’Riordan raised siphons probent over bypresence the crossflow (see Fig. 4 in et al., duced increased Reynolds stress relative to the flush siphon 1995). Since the excurrent jets were laminar, streamwise turcase, especially for the fast flow cases (Fig. 15). Clam pumpbulence intensities were locally reduced. The effect is similar for the fast flow case (Fig. 13b), but the attenuation was 0000, 2007 closerBiogeosciences, to the bed as the jets0001–14, were bent over more rapidly by the stronger crossflow. For the slow flow over raised siphons Biogeosciences, 4, 269–282, 2007 (c), there was a similar near-bed attenuation by the clam ing also produced an increase in Reynolds stress, with the pumping, butdramatic it was now accompanied by aover strong effect more at slower flows and the enhanceraised ment further from the bed. Finally, for the fast flow over the siphons. raised siphons (d), the effect of clam pumping was minimal In all cases, theintensities mean nondimensional concentration, C ∗ and , as the turbulence were dominated by the flow was reduced near the bed (relative to the free stream value roughness. of C ∗ = 1) due to the filtering action of the clams (Fig. 16). The effect of clam pumping on vertical turbulence intensiThe reduction was significantly more pronounced for slower ties is opposite that more seen so forfor thethe horizontal streamwise flows (a, c), andfrom slightly raised siphons (c, intensities (Fig. 14). The vertical energy imparted by the ind). Increased clam pumping also enhanced the concentration current andbut excurrent flows enhanced vertical turbulence reduction, the effect on the near-bedthe concentrations was intensities. The effect was pumping strongestproduced for the slow flows (a, c) relatively weak. Stronger a larger conwhere the relative strength of thethe clam pumping stronger. centration reduction throughout depth of thewas boundary Inlayer. these cases, the influence of the pumping extended deep into the boundary layer. For the faster flows (b, d) the∗ effect The nondimensional concentration fluctuations, c c∗ , was minimal exceptlarger closefor to the the slow bed. flow cases relative to were significantly The presence of roughness due to thetoraised siphonstheprothe fast flows since there was less mixing homogenize duced increased Reynolds stress relative to the flush(a,c), siphon concentration field (Fig. 17). For the slow flow cases case, especially for the fast flow cases (Fig. 15). Clam where the excurrent jets penetrated farther into the flow,pumpthe ing also produced an increase in was Reynolds stress, peak in the concentration variance above the wall.with Thethe effect more dramatic at the slower and overin the raised peak moved farther from wall flows and decreased magnitude as pumping increased. For the fast flow cases (b,d), the siphons. peak concentration variance was at the tops of the siphons, www.biogeosciences.net/bg/0000/0001/ www.biogeosciences.net/4/269/2007/ J. P. Crimaldi et al.: Mass and momentum fields over filter feeders 277 (a) Siphons flush, U∞ = 10 cm/s z (mm) 100 (b) Siphons flush, U∞ = 40 cm/s 100 Q = 0.000 Q = 0.030 Q = 0.045 Q = 0.060 10 10 1 1 0 1 2 3 4 5 0 z (mm) (c) Siphons raised, U∞ = 10 cm/s 100 10 10 1 1 1 2 3 uu (cm/s) 4 10 15 20 (d) Siphons raised, U∞ = 40 cm/s 100 0 5 5 0 5 2 10 15 uu (cm/s) 20 2 Fig. 13. Effect of clam pumping Q on vertical profiles of streamwise turbulence intensity uu for different combinations of free stream velocity U∞ and siphon roughness. (a) Siphons flush, U∞ = 10 cm/s z (mm) 100 (b) Siphons flush, U∞ = 40 cm/s 100 Q = 0.000 Q = 0.030 Q = 0.045 Q = 0.060 10 10 1 1 0.0 0.2 0.4 0.6 0.8 1.0 0 z (mm) (c) Siphons raised, U∞ = 10 cm/s 100 10 10 1 1 0.2 0.4 0.6 ww (cm/s) 0.8 2 2 3 4 5 (d) Siphons raised, U∞ = 40 cm/s 100 0.0 1 1.0 0 1 2 3 ww (cm/s) 4 5 2 Fig. 14. Effect of clam pumping Q on vertical profiles of vertical turbulence intensity ww for different combinations of free stream velocity U∞ and siphon roughness. www.biogeosciences.net/4/269/2007/ Biogeosciences, 4, 269–282, 2007 278 J. P. Crimaldi et al.: Mass and momentum fields over filter feeders Crimaldi et. al: Mass and momentum fieldsUover filter (a) Siphons flush, cm/sfeeders ∞ = 10 100 z (mm) z (mm) 100 Q = 0.000 Siphons flush, U∞ = 10 cm/s Q(a) = 0.030 Q = 0.045 = 0.000 QQ = 0.060 Q = 0.030 Q = 0.045 Q = 0.060 100 10 (b) Siphons flush, U∞ = 40 cm/s 100 10 10 10 1 1 1-0.5 -0.4 -0.3 -0.2 -0.1 (c) Siphons ∞ = 10 cm/s -0.5 -0.4 raised, -0.3 U-0.2 -0.1 100 0.0 -4 1 100 (c) Siphons raised, U∞ = 10 cm/s -3 -2 -1 0 (d) -4 Siphons -3 raised,-2U∞ = 40-1cm/s 0 0.0 100 z (mm) z (mm) 11 (b) Siphons flush, U∞ = 40 cm/s (d) Siphons raised, U∞ = 40 cm/s 100 10 10 10 10 1 1 1 -0.5 -0.4 -0.5 -0.4 -0.3 -0.2 -0.1 2 uw -0.3(cm/s) -0.2 -0.1 0.0 1 -4 0.0 -3 -2 -3uw (cm/s) -2 -4 2 2 -1 0 -1 0 2 uw (cm/s) uw (cm/s) Fig. 15. Effect of clam pumping Q on vertical profiles of Reynolds stress uw for different combinations of free stream velocity U∞ and siphon roughness. Fig. 15. Effect of clam pumping Q on vertical profiles of Reynolds stress uw for different combinations of free stream velocity U∞ and siphon roughness. (a) Siphons flush, U∞ = 10 cm/s z (mm) 100 (b) Siphons flush, U∞ = 40 cm/s 100 Q = 0.030 Q = 0.045 Q = 0.060 10 10 1 1 0.5 0.6 0.7 0.8 0.9 1.0 0.5 z (mm) (c) Siphons raised, U∞ = 10 cm/s 100 10 10 1 1 0.6 0.7 0.8 * C 0.9 0.7 0.8 0.9 1.0 (d) Siphons raised, U∞ = 40 cm/s 100 0.5 0.6 1.0 0.5 0.6 0.7 0.8 0.9 1.0 * C 16. Effect of clam pumping on vertical profiles meannondimensional nondimensional concentration combinations of free stream Fig. 16.Fig. Effect of clam pumping Q onQvertical profiles ofof mean concentrationCC∗ ∗forfordifferent different combinations of free stream velocity U and siphon roughness. velocity U∞ and∞siphon roughness. www.biogeosciences.net/bg/0000/0001/ Biogeosciences, 4, 269–282, 2007 Biogeosciences, 0000, 0001–14, 2007 www.biogeosciences.net/4/269/2007/ J. P. Crimaldi et al.: Mass and momentum fields over filter feeders 279 (a) Siphons flush, U∞ = 10 cm/s z (mm) 100 (b) Siphons flush, U∞ = 40 cm/s 100 Q = 0.030 Q = 0.045 Q = 0.060 10 10 1 1 0.000 0.005 0.010 0.015 0.020 0.000 z (mm) (c) Siphons raised, U∞ = 10 cm/s 100 10 10 1 1 0.005 0.010 0.015 0.010 0.015 0.020 (d) Siphons raised, U∞ = 40 cm/s 100 0.000 0.005 0.020 0.000 0.005 c* c * 0.010 0.015 0.020 c* c * Fig. 17. Effect of clam pumping Q on vertical profiles of nondimensional concentration variance c∗ c∗ for different combinations of free stream velocity U∞ and siphon roughness. (a) Siphons flush, U∞ = 10 cm/s z (mm) 100 (b) Siphons flush, U∞ = 40 cm/s Q = 0.030 Q = 0.045 Q = 0.060 100 10 10 1 1 -0.030 -0.020 -0.010 0.000 -0.030 z (mm) (c) Siphons raised, U∞ = 10 cm/s 100 10 10 1 1 -0.020 -0.010 wc* (cm/s) -0.010 0.000 (d) Siphons raised, U∞ = 40 cm/s 100 -0.030 -0.020 0.000 -0.030 -0.020 -0.010 0.000 wc* (cm/s) Fig. 18. Effect of clam pumping Q on vertical profiles of vertical scalar flux wc∗ for different combinations of free stream velocity U∞ and siphon roughness. www.biogeosciences.net/4/269/2007/ Biogeosciences, 4, 269–282, 2007 Fig. 18. Effect of clam pumping Q on vertical profiles of vertical scalar flux wc∗ for different combinations of free stream velo siphon roughness. 280 J. P. Crimaldi et al.: Mass and momentum fields over filter feeders rates fluxes by infaunal Biology, 113, 219 Turbulent scalar can be bivalves, expressed Marine as a nondimensional correlation coefficient, defined asstructure of velocity and scal Crimaldi, J.: Turbulence 100 6 a bed of model bivalves, Ph.d. thesis, Stanford Univ 4 wc∗ ρw,c = √ Ertman, √ S. C. and Jumars, P. A.: Effects of(5)bivalve siph wwonc∗the c∗ settlement of inert particles and larvae, Journ z (mm) 2 10 6 4 2 1 6 -0.5 -0.4 -0.3 rw,c -0.2 -0.1 0.0 Research, 797–813, 1988. formulation where −1≤ρw,c ≤1. The 46, correlation coefficient removes the effect ofP.the variances, reJonsson, R.,individual Petersen,w J.and K.,c Karlsson, O., Loo, L. O sulting in a true measure the correlation between two son, S. F.: of Particle depletion abovethe experimental biv (Fig. 19). Thesitu profiles collapse into a relatively tight band, measurements and numerical modeling of biva with a common peak correlation coefficient of approximately in the boundary layer, Limnology and Oceanograph -0.38. Note that the vertical location of the peak correlation 1998, 2005.higher in the flow than the correcoefficient is significantly Koseff, sponding peak of wcJ.,∗ . Holen, J., Monismith, S., and Cloern, J.: Co of vertical mixing and benthic grazing on phytopla lations in shallow, turbid estuaries, Journal of Mari 4 Discussion51, 843–868, 1993. Fig. 19. Vertical profiles of the correlation coefficient ρw,c for all Fig. 19. Vertical profiles of the correlation coefficient ρw,c for all experimental conditions listed in Table 1. Flush siphon cases are P. S. and Riisgard, H. U.: Biomixing generate experimental conditions in Table 1. Flush siphon cases The are resultsLarsen, shown in red, and raisedlisted siphon cases in black. of this study add to a growing body of literafilter feeders: A diffusion modelalter forcharnear-bottom ph shown in red, and raised siphon cases in black. ture that demonstrates how benthic filter feeders of concentration Sea Research, 37, in81–90, 1997 acteristics of depletion, momentum Journal and scalar fields In all cases, the mean nondimensional concentration, C ∗ , Monismith, S., Koseff, Thompson, O’Riordan, C the water column. The results share J., some qualitative J., simiwas reduced near the bed (relative to the free stream value larities with previous studies, despite the fact that different H.: A study of model bivalve siphonal currents, Lim of C ∗ =1) due to the filtering action of the clams (Fig. 16). species were involved. Our results that streamwise Oceanography, 35,show 680–696, 1990. turButman, A., Frechette, M., Geyer, W. R., andfor Starczak, R.: TheC. reduction was significantly more pronounced slower V.bulence intensities are relatively insensitive to clam pumping O’Riordan, C.: The effects of near-bed hydrodynamic Flume on food-supply bluesiphons mussel flowsExperiments (a, c), and slightly more so for to thethe raised (c,Mytilusand siphon roughness, whereas vertical turbulence intensibivalve filtration rates, Ph.d. thesis, Stanford Univer Edulis-L as a function of boundary-layer flow, Limnology and d). Increased clam pumping also enhanced the concentration ties increase with pumping rate and roughness. This result is reduction, but 39, the 1755–1768, effect on the near-bed O’Riordan, C.,inMonismith, S., and Koseff, Oceanography, 1994. concentrations was consistent with the increase turbulent kinetic energy (TKE, J.: A stud relatively weak. Stronger pumping L., produced a larger conboundary-layer over a bed of mo turbulence intensities in formation all three directions) Carlton, J., Thompson, J., Schemel, and Nichols, F.: Thethe re-sum of thetration centration reduction throughout the Bay, depthCalifornia of the boundary over beds ofand shutOceanography, and open mussels van Limnology 38,by 1712–1729, 199 markable invasion of San Francisco (USA) bydemonstrated the layer. Duren et al. (2006). Our measured increases in Reynolds Sobral, P., Widdows, J.: Effects of increasing current AsianThe clamnondimensional Potamocorbula amurensis, Marine Ecology Progress concentration fluctuations, c∗ c∗ , stress due to siphon roughness are qualitatively similar to bidity and particle size selection on the feeding activ Series, 66, 81–94, 1990. were significantly larger for the slow flow cases relative to measurements over mussels by Butman et al. (1994) and van for growth of Ruditapes from Cole, the B., fast Thompson, J., and Cloern, J.: Measurement of filtration flows since there was less mixing to homogenize the Duren et al. (2006). However, van Durendecussatus et al. (2006) did Ria Formo concentration field (Fig. 17). For the slow flow cases (a, c), where the excurrent jets penetrated farther into the flow, the peak in the concentration variance was above the wall. The www.biogeosciences.net/bg/0000/0001/ peak moved farther from the wall and decreased in magnitude as pumping increased. For the fast flow cases (b, d), the peak concentration variance was at the tops of the siphons, as the excurrent jets were bent over almost immediately by the crossflow. Turbulent fluxes of scalar concentration in the vertical direction, wc∗ , were always negative, meaning that mass (i.e., phytoplankton) had a net flux towards the bed as a result of near-bed turbulent processes (Fig. 18). The fluxes tended towards zero at the bed and in the free stream, with a peak near z=10 mm. The magnitude of the peak flux increased approximately linearly with clam pumping, Q, and was also significantly larger when the raised siphons were present. A surprising result is that the fluxes were largely insensitive to the mean free stream velocity U∞ . Biogeosciences, 4, 269–282, 2007 not see any significant change in Reynolds stress for inactive versus actively feeding mussels; our results over model Biogeosciences, clams show an increase in Reynolds stress as pumping ac-0000, tivity increases, especially at slow crossflow velocities. This disparity is likely due to functional differences in the feeding mechanisms between the two species. In a study over an array of artificial siphon mimics in a natural channel, Jonsson et al. (2005) found that near-bed Chl a concentration depletion increased with shear velocity. This finding went counter to the expectation (shared by the authors of the study) that increased mixing at higher values of uτ would reduce Chl a depletion through enhanced vertical mixing. The results of our study indicate that concentration depletion decreases dramatically with flow speed (and thus uτ – see Fig. 11). However, increases in uτ due to bed roughness had little effect on concentration depletion, and we did indeed find situations where the concentration depletion was larger for the raised-siphon case as compared to the flush-siphon case, even though uτ was larger with the siphons raised. It appears that uτ by itself may not be a reliable metric for concentration depletion. www.biogeosciences.net/4/269/2007/ 000 J. P. Crimaldi et al.: Mass and momentum fields over filter feeders When mixing rates are low and phytoplankton replenishment to the bed is poor, the maximum concentration depletion can be located some distance above the bed, due to vertical momentum of the exhalant jets that moves phytoplankton-depleted fluid away from the bed (Sobral and Widdows , 2000; Widdows and Navarro , 2007). It has been suggested that this scalar stratification is beneficial to the filter feeders since it would reduce the degree of re-filtrartation of phytoplankton-depleted water. In the present study, the maximum concentration depletion was always at the lowest measured location. Presumably, this is due to the fact that our flow conditions were always relatively energetic compared to the previous studies. Our study is the first of its kind to directly measure turbulent vertical mixing of mass above a bed of bivalves. The profiles of wc∗ in Fig. 18 show that the peak turbulent flux of mass to the bed is approximately 50% larger when the clam siphons are raised. This increase in flux might be expected to decrease the near-bed concentration depletion, but it does not (Fig. 16). One explanation is that the bivalves are able to access higher concentrations by raising their siphons (thus depleting more mass), and this effect overwhelms the roughness-induced increase in turbulent mass flux to the bed. This idea is supported by O’Riordan (1993), who found that incurrent flows had a lower percentage of previously filtered fluid when siphons were raised. The turbulent mass flux measurements present a detailed picture of how and where mass is transfered to the bed. For our study, the turbulent mass flux goes to zero at approximately 100 mm. This height indicates the extent of the water column that is directly impacted by the presence of the filter feeders. Supply of mass to the bed from regions above this distance would need to rely on large-scale turbulent structures that are not present in our flume. Below 100 mm, mass is actively transported to the bed by organized momentum structures in the presence of the concentration boundary layer. This flux is largest near z=10 mm. The magnitude of this flux increases with siphonal pumping rate and roughness due to raised siphons. Closer to the bed, the turbulent mass flux wc∗ goes back to zero due to the hydrodynamic requirement that ww go to zero at the bed. However, this does not mean that the overall mass flux is being reduced. Instead, the mass flux in the near-bed region is accomplished through mean processes associated with the steady (although spatially inhomogenous) siphonal currents. These fluxes are not captured in wc∗ , but have been demonstrated by O’Riordan (1993) and others. On a final note, there are experimental simplifications in this study that are not representative of real bivalve communities. These include (1) the uniformity of the model bivalve array spacing and geometry, (2) spatially consistent siphon heights, and (3) steady siphonal pumping rates. The effect of these approximations is not known, and future research would be valuable in these areas. www.biogeosciences.net/4/269/2007/ 5 281 Conclusions In this paper we presented a set of measurements in a laboratory flume over a bed of model bivalves. The model bivalves incorporated the effect of roughness in the form of raised clam siphons, incurrent and excurrent flows, and siphonal filtering of ambient scalar mass in the overlaying flow. We measured profiles of velocity and mass concentration for different free stream velocities, clam pumping rates, and siphon positions (flush or raised). The results show that clam pumping rates had a pronounced effect on a wide range of turbulent quantities in the boundary layer. In particular, the vertical turbulent flux of scalar mass to the bed was approximately proportional to the rate of clam pumping. However, the formation of a concentration boundary layer above the clams was only weakly sensitive to the pumping rate. When the bivalves pump more vigorously, the increased turbulent scalar flux of phytoplankton towards the bed mitigates the decrease in concentration of available food. The results demonstrate an important mechanism whereby bivalves are able to effectively filter a broad depth range of the water column rather than just re-filtering the layer of water adjacent to the bed. Acknowledgements. This work was supported by the National Science Foundation under Grant No. OCE950408. The authors wish to acknowledge work done by C. A. O’Riordan on design and construction of the model clam plates. The manuscript benefited greatly from detailed editing by P. Jumars, and from a review by an anonymous referee. Edited by: E. Boss References Barrett, T.: Nonintrusive optical measurements of turbulence and mixing in a stably-stratified fluid, Ph.D. thesis, University of California, San Diego, 1989. Batchelor, G.: Small-scale variation of convected quantities like temperature in turbulent fluid, J. Fluid Mech., 5, 113–133, 1959. Butman, C. A., Frechette, M., Geyer, W. R., and Starczak, V. R.: Flume Experiments on food-supply to the blue mussel Mytilus-Edulis-L as a function of boundary-layer flow, Limnol. Oceanogr., 39, 1755–1768, 1994. Carlton, J., Thompson, J., Schemel, L., and Nichols, F.: The remarkable invasion of San Francisco Bay, California (USA) by the Asian clam Potamocorbula amurensis, Mar. Ecol. Prog. Ser., 66, 81–94, 1990. Cole, B., Thompson, J., and Cloern, J.: Measurement of filtration rates by infaunal bivalves, Mar. Biol., 113, 219–225, 1992. Crimaldi, J.: Turbulence structure of velocity and scalar fields over a bed of model bivalves, Ph.d. thesis, Stanford University, 1998. Ertman, S. C. and Jumars, P. A.: Effects of bivalve siphonal currents on the settlement of inert particles and larvae, J. Mar. Res., 46, 797–813, 1988. Jonsson, P. R., Petersen, J. K., Karlsson, O., Loo, L. O., and Nilsson, S. F.: Particle depletion above experimental bivalve beds: In Biogeosciences, 4, 269–282, 2007 282 J. P. Crimaldi et al.: Mass and momentum fields over filter feeders situ measurements and numerical modeling of bivalve filtration in the boundary layer, Limnol. Oceanogr., 50, 1989–1998, 2005. Koseff, J., Holen, J., Monismith, S., and Cloern, J.: Coupled effects of vertical mixing and benthic grazing on phytoplankton populations in shallow, turbid estuaries, J. Mar. Res., 51, 843–868, 1993. Larsen, P. S. and Riisgard, H. U.: Biomixing generated by benthic filter feeders: A diffusion model for near-bottom phytoplankton depletion, J. Sea Res., 37, 81–90, 1997. Monismith, S., Koseff, J., Thompson, J., O’Riordan, C., and Nepf, H.: A study of model bivalve siphonal currents, Limnol. Oceanogr., 35, 680–696, 1990. O’Riordan, C.: The effects of near-bed hydrodynamics on benthic bivalve filtration rates, Ph.d. thesis, Stanford University, 1993. O’Riordan, C., Monismith, S., and Koseff, J.: A study of concentration boundary-layer formation over a bed of model bivalves, Limnol. Oceanogr., 38, 1712–1729, 1993. Biogeosciences, 4, 269–282, 2007 Sobral, P., Widdows, J.: Effects of increasing current velocity, turbidity and particle size selection on the feeding activity and scope for growth of Ruditapes decussatus from Ria Formosa, southern Portugal, J. Exp. Mar. Biol. Ecol., 245, 111–125, 2000. O’Riordan, C., Monismith, S., and Koseff, J.: The effect of bivalve excurrent jet dynamics on mass transfer in a benthic boundary layer, Limnol. Oceanogr., 40, 330–344, 1995. Spalart, P.: Direct simulation of a turbulent boundary layer up to Reδ2 = 1410, J. Fluid Mech., 187, 61–98, 1988. Tennekes, H. and Lumley, J.: A First Course in Turbulence, The MIT Press, Cambridge, 1972. van Duren, L. A., Herman, P. M. J., Sandee, A. J. J., and Heip, C. H. R.: Effects of mussel filtering activity on boundary layer structure, J. Sea Res., 55, 3–14, 2006. Widdows, J. and Navarro, J. M.: Influence of current speed on clearance rate, algal cell depletion in the water column and resuspension of biodeposits of cockles (Cerastoderma edule), J. Exp. Mar. Biol. Ecol., 343, 44–51, 2007. www.biogeosciences.net/4/269/2007/
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