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Dynamics of Dissolved and Biogenic Silica in the Freshwater Reaches of A Biogeochemistry(2009) 96:49-72 DOI 10.1007/sl0533-009-9344-6 Dynamicsof dissolvedand biogenicsilica in the freshwater reachesof a macrotidalestuary (The Scheldt,Belgium) VincentCarbonnel • Marie Lionard • Koenraad M uylaert* Lei Chou Received: 22 December2008 /Accepted: 1 July2009 /Publishedonline: 31 July2009 © SpringerScience+Business Media B.V. 2009 Abstract Temporal evolution of dissolved and Nevertheless,the detritalBSi was a significant biogenicsilica concentrationsalong the Scheldt tidal fractionof the total BSi pool, of whichless than riverand in its tributarieswas investigatedduring 1 10% couldbe attributedto phytoliths.The tidalriver yearin 2003. In thetributaries, dissolved silica (DSi) was dividedinto two zones forbudgeting purposes. concentrationsremained high and biogenic silica The highestproductivity was observedin the zone (BSi) concentrationswere low throughoutthe year. In thatreceived the highestwater discharge, as higher the tidalriver during summer, DSi was completely riverineDSi inputfluxes induced presumably a less consumedand BSi concentrationsincreased. Overall, restrictiveDSi limitation,but the dischargepattern mostof theBSi was associatedwith living diatoms could notexplain all by itselfthe variationsin DSi duringthe productiveperiod in the tidal river. consumption.Silica uptakeand retentionin thetidal riverwere important at theseasonal time-scale: from to 48% of the riverineDSi was V. Carbonnel(El) • L. Chou (El) May September, Laboratoired'Oceanographie Chimique et Geochimiedes consumed and 65% of the produced BSi was Eaux, Facultedes Sciences,Universite Libre de Bruxelles, deposited,leading to a silica (DSi + BSi) retention Campus de la Plaine, CP 208, Boulevarddu Triomphe, in the tidal riverof 30%. However,when annual 1050 Brussels,Belgium were DSi in thetidal river e-mail: [email protected] fluxes considered, uptake amountedto 14% of theDSi inputsand only6% of L. Chou silica was retainedin the e-mail: [email protected] the riverine (DSi + BSi) tidalriver. M. Lionard • K. Muylaert Protistologyand Aquatic Ecology, Biology Department, Keywords Biogenicsilica • Diatoms• GhentUniversity, Krijgslaan 281-S8, 9000 Ghent, Dissolvedsilica • Scheldtestuary • Belgium Silica budget• Tidal freshwater PresentAddress: M. Lionard Abbreviations des Sciences de la Mer de Universitedu Institut Rimouski, BSi silica Quebec a Rimouski,310 Allee des Ursulines, Biogenic CP 3300 Rimouski,QC G5L 3A1, Canada BSidet BSi notassociated with living diatoms BSinv BSi associatedwith living diatom PresentAddress: DiatChla Chlorophylla ascribedto diatoms K. Muylaert DSi Dissolvedsilica K.U. Leuven,Campus Kortrijk,Biology Department, E. Sabbelaan 53, 8500 Kortrijk,Belgium POC Paniculateorganic carbon 4^ Springer This content downloaded from 193.191.134.1 on Thu, 16 Jul 2015 12:07:32 UTC All use subject to JSTOR Terms and Conditions 50 Biogeochemistry(2009) 96:49-72 POCnid POC notcorresponding to livingdiatoms et al. 2005). On its way to the coastal zone, DSi SPM Suspendedpaniculate matter participatesin aquatic biogeochemicalcycles in SPMnbid SPM notcorresponding to BSi norto riversand estuaries(Conley et al. 1993), involving livingdiatoms a numberof biological, chemical and physical processes.Diatom bloomsoccurring in spring/sum- mer in rivers and estuariesresult in significant decreasesin DSi and concomitantincreases in BSi concentrations,which can accountfor 50-70% of the Introduction total riverinesilica load (Admiraalet al. 1990; Conley1997). As particulatematerial, BSi can settle Dissolvedsilica (DSi) is a keynutrient for the aquatic and accumulatein thesediments, inducing retention ecosystemas diatomshave an essentialrequirement of silica in theecosystem. It can also be transported forDSi to build up theirfrustules, rigid outer cell downstreamas suspendedmaterial (Admiraal et al. walls made of amorphousbiogenic silica (BSi) 1990) and/orbe dissolvedat biologicaltimescales (Ragueneauet al. 2000). Diatoms oftendominate (Roubeix et al. 2008; Loucaides et al. 2008). BSi the primaryproduction in turbidaquatic environ- dissolutionis howevernot expectedto occurat the mentssuch as rivers,estuaries and coastal zones same ratealong the salinity gradient: BSi dissolution (Ragueneauet al. 2000), and accountfor 75% of the rate increaseswith salinity (Loucaides et al. 2008) primaryproduction in the world's coastal areas and the bacterial communitycan significantly (Nelson et al. 1995). They can supportan efficient enhance the remineralisation(Bidle and Azam aquaticfood chain (Turner et al. 1998) and mayplay 1999; Roubeixet al. 2008). In addition,phytoliths a significantrole in theexport of carbon from surface (BSi materialproduced by terrestrialhigher plants) waters(Treguer and Pondaven2000; Ploug et al. can add a significant,ifnot major, contribution to the 2008). The availabilityof DSi and its relative BSi pool carriedby rivers(Cary et al. 2005). They abundance compared to the other nutrientscan can originatefrom top soil erosion(Cary et al. 2005) influencethe compositionof the phytoplankton as well as directlyfrom the vegetationof the river community,which can subsequentlyaffect the eco- banksor tidalmarshes (Struyf et al. 2006). Marshes logical functioningof the ecosystem(Officer and can retain significantamounts of silica in the Ryther1980; Conley et al. 1993; Lancelot 1995; abovegroundbiomass and in sediments(Struyf Turneret al. 1998). However,riverine DSi fluxes, et al. 2005) and DSi and BSi exchangebetween the which are the main source of DSi to the oceans riverchannel and the marshes can affectthe estuarine (Wollast 1974; Tregueret al. 1995), have been silica cycle(Struyf et al. 2006). Thus,because of the alteredby humanactivities during the last decades. possiblyimportant riverine and estuarineBSi con- Land-useactivities, such as deforestation,may affect centrations,its specificbehaviour and its interaction theDSi inputsto rivers(Conley et al. 2008) and the withDSi, the studyof BSi dynamicsshould not be use of silica in washingpowders (Verbanck et al. omittedwhen silica fluxesand mass-balancesin 1994) and fertilizers(Datnoff et al. 2001) may also riversand estuariesare assessed. constitutean additionalDSi source to rivers.In In some estuaries,the tidal influencepropagates contrast,eutrophication results in enhancedsilica furtherinland than salt intrusion,leading to the retentionin lakes,rivers and estuaries(Conley et al. existenceof tidalfreshwater reaches which may host 1993). Decreased silica concentrationsare also importantchemical and biologicalprocesses (McLu- induced by river dammingand river regulation, sky 1993; Schuchardtet al. 1993; Muylaertet al. which cause particletrapping and reduce contact 2005). In particular,tidal freshwater reaches exhibit betweenthe waterand the riparianvegetated zone specificfeatures which can influencethe phytoplank- containingDSi-rich interstitialwaters (Humborg tondynamics (Schuchardt et al. 1993; Lionard2006) et al. 2006). and thusthe biogeochemical cycle of nutrients.They Mostof theDSi is originallyproduced during rock differfrom the adjacent riversby longer water weatheringon landwhere it generallygoes througha residencetimes and the presenceof tides, which terrestrialcycle beforereaching the rivers(Derry induceshigher turbulence and turbidity(Schuchardt £i Springer This content downloaded from 193.191.134.1 on Thu, 16 Jul 2015 12:07:32 UTC All use subject to JSTOR Terms and Conditions Biogeochemistry(2009) 96:49-72 51 et al. 1993; McLusky1993). The resultinglow light shouldstill be performedand comparedto themodel conditionsmay limit phytoplankton growth but may results.Being a part of the suspendedpaniculate also favourdiatoms due to theirlower light require- matter(SPM), BSi in the Scheldttidal freshwater mentscompared to other algae (Reynolds 1988; reachesis expectedto followa complexbehaviour Cushing 1989; Lionard 2006). In contrastto the due to spatial and temporalvariability in SPM downstreamareas where freshwaterand seawater dynamics(Chen et al. 2005a). Due to diatom mix, thereis no phytoplanktonmortality due to mortality,resuspension of dead diatomfrustules and salinitystress in tidalfreshwater reaches (Schuchardt the possible presenceof phytoliths,a significant et al. 1993; Muylaertet al. 2000). Thus, if the fractionof theBSi maynot be associatedwith living residencetime is sufficientlylong, a maximumin the diatoms.In addition,marshes along the Scheldt tidal diatomproduction and theassociated DSi consump- freshwaterreaches may act as DSi recyclersor sinks tion may occur in the tidal freshwaterreaches forestuarine BSi, andmay play a rolein theestuarine (Anderson1986; Schuchardtand Schirmer1991; De silica cycle(Struyf et al. 2006, 2007b). As theyalso Seve 1993; Schuchardtet al. 1993; Muylaertet al. contain significantamounts of phytolithin the 2005). abovegroundbiomass (Struyfet al. 2005), BSi as In the presentstudy, we focus on the Scheldt phytolithsmay be providedto theestuarine waters by estuarywhich comprises an extensivetidal freshwater tidalexchange of thisbiogenic material between the area.The biogeochemistryof silica and thefluxes of riverchannel and marshes. thisnutrient in the continuumof the Scheldtare of In thisstudy, we presenthigh resolution DSi and primaryimportance as theycan affectthe ecosystem BSi temporalprofiles during one full annualcycle in the adjacentcoastal zone. Earlier works have alongthe tidal freshwater reaches of theScheldt and shownthat DSi drivesthe extent of theearly spring itstributaries. BSi concentrationsare comparedwith diatombloom
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