‘Muddy Coast ofthe World: Processes, Deposits and Function ‘T. Healy, Y. Wang and JA, Healy (Editors) 203, © 2002 Elsevier Science B.V. All rights reserved. Chapter Seven Biochemical factors influencing deposition and erosion of fine grained sediment PIETER G.EF. AUGUSTINUS Utrecht University ‘The Netherlands INTRODUCTION Deposition and erosion of fine cohesive particles are strongly influenced by biochemical factors, especially in the intertidal environment (Van Straaten and Kuenen 11957). Organic matter released by algae and bacteria acts as ‘biological glue’ and causes the formation of aggregates. Filter feeders like cockles and mussels filter fine particles, from the water and produce comparatively coherent faecal pellets. Diatoms and bacteria produce slime which binds sediment particles. ‘The aim of this chapter is to present a state-of-the-art review of the role that biochemical processes play in the deposition and erosion of fine-grained sediments. To achieve this purpose, the following three important influences will be discussed. ~ Flocculation of fine-grained particles by bio-substances giving rise to the formation of aggregates; ~ Filter feeding and subsequent pellet production by benthic organisms and zooplankton; and - Effect of bed fauna and flora on the strength of the bed material. In 1993, two Dutch authors contributed greatly to this subject (Eisma 1998; Ten Brinke 1993). The content of this chapter is mainly based on these sources.2 FLOCCULATION 2.1 Introduction An important property of fine cohesive particles in suspension is that they are sensitive to flocculation. Flocculation results in aggregates which consist of many small particles formed by collision and adhesion of material in suspension (Kranck 1981). The flocs or aggregates are coarser than the primary particles, and therefore sink faster. The settling velocity of aggregates is considered to be an important parameter to describe the behaviour of cohesive sediments in coastal and estuarine environments (Ten Brinke 1998; Dyer 1995). The dynamic behaviour of aggregates differs from that of non-cohesive solitary particles. The settling velocity of particles smaller than 50 pm is approximately described by the Stokes’ equation, which is based on a positive relation to the square of the grainsize (Schlichting 1968), For coarser particles, the settling velocity increasingly approaches towards the root of the grainsize, which is in accordance to the Oseen modification of Stokes’ equation (Figure 1). However, both equations are only valid for spherical grains which, especially for the smaller grainsizes, hardly exist in natural environments. Water 20 degrees Celcius 10, velocity (em / sec) 001 001 op on 7 70 10 ‘grain size (mm) Figure 1 The settling velocity in water in relation to grainsize. Up to a size of 0.05 mm. the settling velocity can be calculated using the Stokes’ equation. At higher settling velocities the Oseen modification of the Stokes’ equation gives better results.205 Chase (1977, 1979) has measured settling velocities in nature, in lacustrine as well as in marine environments. For aggregates smaller than 100 um, using the Stokes’ equation, he found the settling velocities in these environments to be underestimated by more than an order of magnitude. The same tendency has been shown for aggregates coarser than 100 um up to 1000 um, however with increasing grainsize the setiling velocity more and more tends to behave in accordance with the Stokes’ equation (Kajihara 1971). Most research on the settling of mud aggregates refers to relatively high concentrations (50-1000 mg/1). An exponential increase of the settling velocity with sediment concentration up to the point of ‘hindered settling’ has been indicated by many researchers (e.g. Delft Hydraulics Laboratory 1962; Migniot 1968; Owen 1971; Puls and Kuehl 1986; Pejrup 1988; Van Leussen 1988, 1994; Dyer 1995). This is demonstrated in Figure 2. The relation is explained by an increase of the collision frequency at a higher concentration of suspended particles, whick favours aggregation (Van Leussen 1988, 1994). settling velocity lems) on oot p01 100 "1000 wa000 00.000 ‘concentration mgt) Figure 2 The settling velocity in water in relation to suspended sediment concentration (after Dyer 1995),206 According to this relationship between settling velocity and sediment concentration, and under the assumption that this relationship also holds for concentrations <100 mg/l, the settling velocity of suspended aggregates at concentrations of about 10mg/I should be extremely low (<0.1 mm/s). In Dutch tidal basins with concentrations in the order of 10 mg/I, however, the accumulation of cohesive sediment can only be explained by assuming settling velocities on the order of 1 mm/s (Ten Brinke 1993). The discrepancy between the measured and the calculated settling velocities of aggregates may be ascribed to the following three causes (Ten Brinke 1993): (i) the changes in aggregate density as aggregates grow (e.g. Kajihara 1971; Hawley 1982), (é viscous drag reduction (Chase 1977, 1979; Hawley 1982) and (iii) disturbance of the delicate aggregate structures during sampling (Kranck 1975; Gibbs 1981; Amos and Mosher 1985; Eisma 1986). The last mentioned cause has lost much of its importance since, recently, techniques for in situ observation of flocs have become available (Honjo et al. 1984; Johnson and Wangersky 1985; Bale and Morris 1987; Eisma et al. 1990; Ten Brinke 1993). A comparison of modern in situ techniques has been carried out by Dyer et al. (1996). One of the most modern techniques for observing flocs in situ is by using the VIS (Video In Situ), by which a video camera is used to collect images of the flocs (Van Leussen 1994; Van Leussen and Cornelisse 1996). 2.2 Collision and aggregation of cohesive particles In tidal seas and estuaries, a majority of the fine mineral particles are part of flocs (Kranck 1973; Dobereiner and McManus 1983; Eisma 1986, 1993). Most flocs consist of both organic (living and dead cells surrounded by gelatinous masses of waterbearing, bio-compounds) and inorganic material (Ten Brinke 1993). The suspension population consists of macro-flocs, micro-flocs and mineral parts (Van Leussen 1994): = Micro-flocs are smaller than 100 um and strong; adhesives released by algae ete., keep them together, = Mineral particles form, together with micro-flocs, the components for the macro- flocs. = Macro-flocs vary in size from several hundreds of um to more than one mm. They are very fragile. The formation of flocs can be ascribed to a considerable number of processes (Eisma 1993). Flocculation depends on the collisions between the particles and on the cohesive properties of the material. Physico-chemical processes of flocculation ‘Suspended clayey particles are usually negatively charged (Neihof and Loeb 1972; Hunter and Liss 1979; Hunter 1982). Under these conditions the particles repel each other. However, when the intensity of the negative surface charges is controlled, the inter-particle attraction from Van der Waal's forces becomes dominant. Due to these overall attractive forces, particles agglomerate when they reach close proximity or collide, resulting in the formation of aggregates, or flocs. The flocculation potential of207 particles increases with increasing concentration because, under these circumstances the chance of colliding is enhanced. According to McCave (1984), Smetacek (1985), Van Leussen (1988, 1994), Alldredge and Gotschalk (1989), the most important mechanisms which cause collisions are: 1) The Brownian motion, which is a general property of fine dispersed particles, due to the thermal movement of (in this case) the water molecules. This mechanism is only effective at very high particle concentrations and for particle size less than about 0.5 jum (Dyer 1986). The resulting aggregates have a ragged structure and are weak 2) Velocity gradients within the suspending liquid, a mechanism which is especially important in the mixing zone of estuaries (Van Leussen 1988, 1994). The related flocs are usually more or less spherical (Krone 1962). Their strength is high compared to that of other flocs. 3) Differential settling of the suspended particles which results in the capture of small, slowly settling particles by larger, faster settling particles. In calm hydrodynamic conditions, e.g, around slack water, this mechanism efficiently increases aggregate size and settling velocity. The resulting flocs are weak and have a ragged structure. However, not all collisions will lead to flocculation. In a natural environment, particularly at higher turbulence, flocculated particles continuously vary in size (Terwindt 1971). Coarser components will dislodge and settle more easily than finer ones. So with increasing turbulence not only the size of the flocs will be reduced, but also the size of the composing particles. In this process of alternating flocculation and disaggregation, the narrow zone of high velocity gradient close to the bed, plays an important role (Van Leussen 1994), ‘The water movement in a channel can be divided into an inner region, which is directly influenced by the condition of the channel bed, and an outer region, in which the flow is only indirectly affected (Figure 3). In the case of a smooth channel bed, the A B peas penne Cece ¥ ti ae | TT iogartnmic 9, ‘turbulent one O See Ste gps, C—O Figure 3 A: The current velocity profile in a channel with zones in turbulent flow above a smooth bottom. B: A schematic picture of the disruption and aggregation of mud flocs. (After Van Leussen 1994).208 inner region can be subdivided into a viscous bottom layer and a logarithmic turbulent zone, divided by a buffer zone, where the viscous and turbulent transfers of momentum are comparable. In the lower inner region, the shear and lift forces are strongest. According to Mehta and Partheniades (1975), these control the size and shear strength distributions of the suspended aggregates. The flocs which are strong enough to resist the shear forces will settle on the bed. If the strength of the flocs is insufficient in this respect, the aggregates are broken up into smaller pieces, become uplifted by turbulent flow, and will participate in the flocculation process again (Figure 3B). Bubbles of gas or air, produced by breaking waves or escaping from organisms or bottom sediments, may contribute to the formation of aggregates as well (Eisma 1993). When a gas bubble eventually bursts or dissolves, the adsorbed organic particles will aggregate. It is not yet clear if gas bubbles significantly contribute to flocculation, compared to other flocculation processes. Cationic flocculation (often called salt flocculation) of fine mineral particles when riverine water enters sea water, as described by Van Olphen (1963) is now regarded as a less dominant mechanism. Although demonstrated in laboratory experiments (among others Whitehouse et al. 1960; Migniot 1968), the process appears to be inoperative in nature because of organic or metallic surface coatings, which change the surface characteristics of the particles (among others Neihof and Loeb 1972; Gibbs 1977, 1983; Hunter 1980; Hunter and Liss 1982; Loder and Liss 1985). Generally, surface- active organic matter is sufficiently available and the formation of oxide films such as iron oxide is a common feature. Such coatings, therefore, are expected to be present on suspended particles in almost all estuaries (Van Leussen 1994). The main exception might be in situations of short rivers with a steep gradient, draining an area characterised by episodic rain storms, and transporting vast quantities of clay particles in a very short time. Then, possibly, there is no time to get a metallic or an organic coating on the individual flocs. However, coating appears to be a very fast process. Loeb and Neihof (1977) have demonstrated that the surface of a platinum plate of one square millimetre in seawater showed a completely organic coating in only one hour. This suggests that particles in estuaries usually will be coated. Biochemical components in the flocculation process For the formation of flocs due to biochemical influences the following processes are operative (Eisma 1986, 1993; Van Leussen 1988; Ten Brinke 1993): 1) organic coatings on suspended particles which influence the electrostatic stability of the particles (Neihof and Loeb 1972; Gibbs 1977, 1983; Hunter 1980; Hunter and Liss 1982; Loder and Liss 1985); 2) polymer bridging (Harris and Mitchell 1973; Gregory 1978; Alldredge and Silver 1988); and 3) faecal pellet production by filter feeding organisms (Haven and Morales-Alamo 1972; Schubel and Kana 1972; Smetacek 1980a; Martens and Krause 1990). The effects of organic coatings and of polymer bridging will be discussed in section 3. Pellet production will be treated in section 4.209 3 AGGREGATION OF ORGANIC AND INORGANIC PARTICLES Organic matter influences both the size of the flocs as well as their settling velocity (Kranck and Milligan 1980; Kranck 1981, 1984). The way it influences the aggregation of particles is by the formation of coatings and polymer bridging. Experiments by Kranck (1984) have demonstrated that a mixture of organic and inorganic matter produces larger aggregates which settle faster than organic or inorganic particles alone (Figure 4). Organic matter forms coatings on mineral particles creating uniform electrical surface properties of these particles (Neihof and Loeb 1972; Hunter and Liss 1979; Hunter 1982; Loder and Liss 1985). The organic coatings retain or inhibit salt flocculation (Bisma 1993). Organic coatings are assumed to be present on suspended particles in almost all estuaries (Van Leussen 1988, 1994). According to Eisma (1993), there are several possibilities for the formation of coatings. They can be the result of direct adsorption of dissolved compounds onto the particle surface, or by flocculation or polymerisation of dissolved compounds in the water followed by adsorption. Coatings can also be produced by micro-organisms. cone, ees 0 Nt 50 eran sadivont CSiBt omar =] Organic matter Time (days) Figure 4 The decrease in suspended material concentration with time due to flocculation and settling for inorganic particles and organic matter (after Kranck 1984),210 3” Em Eo 3 3 27°] = : 3 vl Eo 5 Jad oct an ape of ‘ost Jen Apr 1988 Fine-grained sediment ZS] Chlorophyll a Figure 5 The (seasonal) variation in the depth-averaged concentrations of chlorophyll a and fine-grained suspended sediment over a period from October 1989 until May 1990 in the western part of the Eastern Scheldt, The Netherlands (after Ten Brinke 1993). Bridges of organic polymers are produced by algae and bacteria and bind fine grained particles. According to Ten Brinke (1993), algae release sticky biopolymers under environmental stress. This release may be due to an increased production of extracellular mucus (Myklestad 1974; Avnimelech et al. 1982; Degens and Ittckot 1984), and cell lysis (Pavoni et al. 1972; Harris and Mitchel 1973; Alldredge and Silver 1988), During his research in the Eastern Scheldt (The Netherlands), Ten Brinke (1993) found the distinct seasonal variation in suspended sediment concentration to be related to the occurrence of phytoplankton blooms. During the development of a phytoplankton bloom the background suspended sediment concentration rapidly decreases, The lowest values are reached after the disappearance of the bloom (Figure 5) Mucus is produced by bacteria as well. The rapid increase of bacterial biomass which is observed during the decline of a phytoplankton bloom therefore also makes bacteria very important producers of polysaccharides and other polymers (Pavoni et al. 1972; Harris and Mitchell 1973; Calleja 1984; Biddanda 1986). Suitable nuclei for the aggregation of particles may also be provided, by mucus produced by zooplankton (Jorgensen et al. 1984) and mucopolysaccharide remains of appendicularian houses - a species of the class Tunicata - (Alldredge and Cox 1982). The composition of large flocs shows an unequal division of organic and inorganic constituents. According to Eisma (1986, 1993), Wells and Shanks (1987) and Wells (1989) large aggregates in shallow water generally contain much more inorganic than organic constituents. This strongly depends on the time of the year. The organic fraction will predominate during phytoplankton blooms especially at low suspended sediment concentrations. Therefore, primary produced organic matter may greatly affect aggregation of suspended matter at relatively low suspended matter concentrations (Ten Brinke 1993).21 4 INFLUENCE OF FILTER FEEDING AND PELLET PRODUCTION BY BENTHIC ORGANISMS AND ZOOPLANKTON: ON DEPOSITION OF FINE-GRAINED SEDIMENT 4. Introduction Deposition of fine-grained sediment is not only accelerated by flocculation but also through filter feeding by bivalves, various hard-substratum filter feeders (tunicates, sponges) and zooplankton. Generally, benthic filter feeders are thought to contribute substantially to the settling of fine-grained sediment (Lund 1957; Haven and Morales- Alamo 1972; Rhoads 1974; Risk and Moffat 1977). The importance of zooplankton filter feeding as a mechanism for the settling of organic and inorganic matter is questioned (Alldredge 1979; Smetacek 1980a, 1980b; Peinert et al. 1982; Beers et al. 1986; Bloesch and Biirgi 1989). In this paragraph both the benthic filter feeding effect and the zooplankton grazing effect will be discussed. 42 Benthic filter feeding, faecal pellet and pseudofaeces formation and biodeposition Filter feeders versus deposit feeders Deposit feeders and filter feeders are both benthic organisms that produce faecal pellets near the sediment surface. Filter feeders feed on sediment suspended in the water column. They are thought to contribute substantially to the settling flux of cohesive sediment (Lund 1957; Haven and Morales-Alamo 1972; Rhoads 1974; Risk and Moffat 1977). Plate 1 gives a picture of an unconsolidated clay deposit, including a istinet contribution of transported faecal pellets. Deposit feeders, however, have no effect on the flux of suspended sediment settling cn the sea bed, because they feed on bottom sediment. According to Rhoads (1974), their feeding activity is generally confined to the uppermost 10 cm of the sediment, including the sediment-water interface, Their bioturbation results in a faecal rich floceulent bottom sediment with a high water content, which can easily be eroded (@hoads 1974; Aller and Dodge 1974). The resulting high turbidity will lead to clogging cf the filter mechanisms of the filter feeders (Risk and Moffat 1977). It explains why, generally, dense populations of filter feeders and deposit feeders are spatially separated (Carney 1981). Biodeposits, produced by filter feeders are more compact and cohesive, giving the bed more resistance against erosion (Rhoads 1974). This stabilizing effect is caused by microbial and algal mucus, binding the individual aggregates and pellets (Rhoads and Foyer 1982). Eenthic filter feeders and sedimentation Filter feeders like mussels, cockles and tunicates filter suspended particles from the water column and transform them into pellets. An important aspect of this activity is that they favour the deposition of individual particles that would not have settled by Fhysical processes (Ten Brinke 1993). Conditions like age, density and species determine the degree of biodeposition.212 Plate 1 A thin section of an undisturbed clay deposit of the Solo delta, Indonesia, showing faecal pellets as part of a fining upward sequence, which has settled down from a buoyant plume in front of the delta. A number of pellets have been indicated by an arrow. The width of the photo is 38 millimetre,213 Filter feeders produce true faecal pellets and pseudofaeces. The difference is that pseudofaeces are formed from suspended material which is not ingested but rejected at the maximum ingestion rate. Pseudofaeces are ejected from the shell cavity in a loosely compacted mass and dropped beside the animal together with the true faeces (Verwey 1952). Due to their loose composition, the pseudofaeces are easily resuspended. Therefore they hardly contribute to the settling of fine-grained sediment (Risk and Moffat 1977). Faecal pellets are more resistant against erosion than aggregates. This is due to the compaction of the material during its passing through the alimentary canal of the filter feeder, as well as to a number of biological (bacterial degradation of the binding mucus) and physical (water turbulence, interpelletal collisions) causes. The resistance against erosion will therefore vary seasonally, ‘The seasonal variation in biodeposition is probably also influenced by the quality of the available food. Yet there is considerable uncertainty concerning the rate of biodeposition. Tenore et al. (1982) found high biodeposition during peaks in primary production. Other researchers like Foster-Smith (1975) and Smaal et al. (1986) found no relation between particle/ (food) concentration and clearance rate. According to their observations, mussels clear the same volume of water independent of particle concentration. Therefore, the amount of biodeposition is doubled when concentration is doubled. In extensive mussel farming areas, faeces production can be an important mechanism for the deposition of fine-grained sediment (Ten Brinke 1993). In The Netherlands this subject has been’ studied extensively, especially in the Costerschelde, an estuary in the southwest of The Netherlands, Here the biodeposition is dominated by the blue mussel (Mytilus edulis (L.)) and the cockle (Cerastoderma edule (L)) (Smal et al. 1986; Prins and Smaal 1989). The bioproduction of the blue mussel and the cockle is in the same order of magnitude (Smaal et al. 1986). However, the net effect of cockle filter feeding on fine-grained sediment accumulation is considered to be minor. According to Verwey (1952) this is due to: 1) cockle faecal pellets being much smaller and thus more easily resuspended than mussel pellets; 2) a higher mucus production by mussels as compared with cockles which decreases erodibility of the faeces; and 3) a more regular distribution of the cockle biomass over the cockle banks and therefore a pellet production over wide sandy areas which facilitates resuspension. Other benthic filter feeders like tunicates, sponges and hydroids have a much lower biomass (Lewis and Waardenburg 1990). The rate at which mussels clear water can be approached in different ways. ‘According to Bayne (1976); Widdows et al. (1979) and ‘Thompson (1984), adults filter the water at an average rate of a few litres per animal per hour. Smaal et al. (1986) relate the clearance rate of the blue mussel Mytilus edulis (L.) to their body size. They calculate an average clearance rate for adults of 1.6 litre/hour per animal, whereas for juveniles an average clearance rate of 08 litre/hour is calculated. Recently, Smaal and Van Stralen (1992) mention a filtration rate of approximately 3 litres per hour for adult blue mussels of about 5 em in length.214 ‘There is a clear seasonality in filter feeding activity in the Oosterschelde. In autumn and winter the activity is low. In spring and summer, the biodeposition figures are much higher (Kautsky and Evans 1987; Jorgensen et al. 1990). This might be due to a low metabolic activity of benthic filter feeders at low water temperature (Haven and Morales-Alamo 1972; Risk and Moffat 1977; Newell 1979; Kautsky and Evans 1987). Waves and currents can influence the effect of biodeposition. In the Oosterschelde, Ten Brinke (1993) and Ten Brinke et al. (1995) demonstrated that despite high biomass, biodeposition by mussels hardly leads to a decrease in suspended sediment concentration. This is thought to be due to resuspension of most of the biodeposits by (tidal) currents and waves. 4.3 The effect of zooplankton grazing on biodeposition Zooplankton grazing Zooplankton feeds on phytoplankton and consecutively produces faecal pellets, consisting (mainly) of organic matter surrounded by a membrane. The importance of the contribution of these zooplankton pellets to biodeposition has been the subject of many studies during the last two decades (Smetacek 1980a, 1980b; Lorenzen and Welschmeyer 1983; Davies and Payne 1984; Lampitt 1985; Skjoldal and Wassmann. 1986; Riebesell 1989; Martens and Krause 1990; Ten Brinke 1993). The results of these investigations show large differences, which to a certain extent can be attributed to the degradation of zooplankton pellets during deposition. The vulnerability of zooplankton pellets during settling appears to depend on the water temperature, the coherence of the pellets and on coprophagy: the feeding activity of zooplankton on their own pellets (Smetacek 1980a, 1980b; Haney and Trout 1990; Martens and Krause 1990). Moreover, the importance of grazing for the transformation of the phytoplankton to detritus is reported to vary both regionally (Tremblay et al. 1989) and seasonally (Smetacek 1980). According to Schubel and Kana (1972) the average filtration rate for zooplankton is 5-10 ml/day per individual. Concentration, composition, taste and age of the (phytoplankton) food largely determine the ingestion rate (Dagg and Walser 1986; Burns and Xu 1990; Haney and Trout 1990; Martens and Krause 1990; Vanderploeg 11990). Light conditions are important for the production of faecal pellets as well (Small et al. 1989), due to assimilation of the ingested particles. Research on copepods has demonstrated that the faecal pellet production of adult copepods may vary between some hundred pellets per day (Smetacek 1980a) and five to eight pellets per day (Bathmann et al. 1990), probably depending on food concentration (Ten Brinke 1993). According to Paffenhifer and Van Sant (1985) copepods select their food in the sequence: living phytoplankton cells > dead phytoplankton cells > faecal pellets > polystyrene spheres. Zooplankton and sedimentation Zooplankton grazing results in the transformation of phytoplankton and detritus into compact faecal pellets that sink down rapidly. Ten Brinke (1993), by comparing measured settling velocity data of zooplankton pellets from the literature, however, shows a variation between approximately (0.2 mm/s (Turner 1977) and approximately215 20 mm/s (Dunbar and Berger 1981). Several arguments have been put forward to explain this variation: differences in species and age of the zooplankton (Smetacek 1980a), food concentration and diet (Bienfang 1980; Smetacek 1980a; Deibel 1990; Voss 1991; Cadée et al. 1992) as well as water temperature (Smetacek 1980a). The importance of the contribution of zooplankton pellets for the sedimentation flux of fine-grained material is therefore questioned in literature. The number of studies that stress the significance of zooplankton pellets in this respect (e.g. Schubel and Kana 1972; Steele 1974; Honjo 1978; Knauer et al. 1979; Sasaki and Nishizawa 1981; Pickrill 1987; Hedges et al. 1988) is in the same order of magnitude of those opposing this view (eg. Alldredge 1979; Smetacek 1980a, 1980b; Peinert et al. 1982; Cadée 1985; Beers et al. 1986; Skjoldal and Wassmann 1986; Asper 1987; Bloesch and Biirgi 1989; Wassmann et al, 1990). The overestimation of the importance of zooplankton faecal pellets as a mechanism for the settling of cohesive sediment is attributed to the easy break-down of the pellets. Therefore, the number of undisturbed pellets reaching the bottom will be small as compared to aggregates of non-faecal origin. Angel (1984) and Cadée (1988) have argued that a number of the aggregates may in fact be disintegrated faecal pellets. An important cause for the degradation of zooplankton pellets is the temperature of the sea water, which determines the microbial activity. Honjo and Roman (1978) have shown that the degradation of the protective outer membrane of the pellets by microbial activity in sea water at 20°C takes only 3 hours, whereas the membranes stayed intact for 20 days in sea water of 5°C. Martens and Krause (1990) studied the degree of disintegration in relation to water depth. They found a positive relation and could not prove any pellets at a depth of 100 metres. Other reasons for the breakdown of zooplankton pellets are differences in the coherence of the pellets (Deibel 1990) and coprophagy (Smetacek 1980a; Skjoldal and Wassmann 1986; Haney and Trout 1990). The coherence of the faecal pellets is influenced by both the concentration (Burns and Xu 1990) and the size (Haney and Trout 1990) of the available food. At lower food concentrations the pellets have a looser consistency. Larger food particles produce looser faecal pellets. Te will be clear that the importance of zooplankton for the deposition of fine-grained sediment depends on specific physical and biological processes in the area of research, which determine the degree of disintegration of the suspended pellets. During a six week period of measurements in the Oosterschelde, Ten Brinke (1993) found the larger volume of the aggregates in comparison to the zooplankton pellets to be dominant over the larger density of the pellets. Despite an abundancy of suspended pellets relative to aggregates of up to 10%, the contribution of pellets to the vertical sediment flux is relatively small (less than 2% by weight). Zooplankton grazing activity in response to a phytoplankton bloom The development of a zooplankton population in response to a phytoplankton bloom shows a lag of about one month (Legendre 1990). This means that the primary organic matter of the phytoplankton biomass settles down and largely contributes to the aggregates before the grazing activity of the zooplankton population is large enough to produce a substantial amount of faecal pellets (Wassmann et al. 1990). The importance of grazing decreases further when the phytoplankton bloom develops more rapidly (Legendre 1990).216 This slow response of zooplankton grazing on the growing phytoplankton biomass during a bloom results in a typical seasonal character in the settling of particulate organic matter. During the spring bloom, when the zooplankton population is still underdeveloped, the settling rates of primary produced organic matter shows a peak. This period of high sedimentation usually lasts no longer than a week (Smetacck 1980b). Large downward fluxes of organic aggregates, descended from the phytoplankton bloom, have been reported by several authors (Forsskahl et al. 1982; Lampitt 1985; Riesebell 1989), After the spring bloom, when the zooplankton population has fully developed, the phytoplankton biomass is low due to grazing. At that time, sedimentation of organic aggregates is at a minimum (Smetacek 1980b; Peinert et al. 1982). In these situations, organic matter may settle mainly through zooplankton faecal pellets (Tremblay et ai. 1989). In the Oosterschelde study of Ten Brinke (1993) the contribution of zooplankton pellets to the total sediment accumulation was 13% by weight. This indicates that in this area the contribution of zooplankton filter feeding is insignificant for the vertical sediment flux 5 EFFECT OF BED FLORA AND FAUNA ON THE STRENGTH OF THE BED MATERIAL 5.1 Introduction ‘An increase in the resistance of a sediment bed to erosion caused by the direct or indirect influence of biota can be termed. ‘biogenic stabilization’, according to Ten Brinke et al. 1994. In contrast to larger organisms, microbes are found on almost all submerged surfaces. Large organisms may have a more pronounced impact on sediment stabilization but because of the abundance of the microbes, their influence may be more impressive. Organisms can stabilize sediment in different ways, e.g. the building of tubes and burrows, by the excretion of cohesive compounds and by varying the surface roughness (Montague 1986). Several groups of benthic organisms have a biological impact on the strength and erodibility of the sediment. In the following sections the impact of macro-, meio- and microfauna is treated. The influence of benthic diatoms (microfauna) is emphasized in this chapter, because experiments have shown that they play an important role in stabilizing intertidal sediment. The actual developments of measuring the effects of biochemical factors on the strength of the bed material are strongly into the direction of in situ measurements (Amos et al. 1992). Measuring sediment stability in the laboratory to be representative for the natural environment, makes high demands upon the complexities arising from chemical, biological and hydrodynamical influences (e.g. Amos and Mosher 1985; Grant et al, 1986; Grant and Gust 1987). The number of in situ measurements, however, is still relatively limited.217 5.2 Macro- and meiofauna The most important effect of macrofauna on the erodibility of the bed is their presence. Their existence already modifies the sediment because, in general, the size of the macrofauna is greater than the size of a single sediment particle. By pushing and excavating, these burrowing animals move through the sediment. Smaller burrowers are part of the meiofauna and live in the upper centimetres of the sediment. Their productivity is high in comparison with the macrofauna. Polycheate worms (Gray 1974) with tubes, exhalant and inhalant siphons, may create a micro-relief, increasing the erodibility of the sediment (Nowell et al. 1981). The bed shear strength is also enhanced by an increase in stability due to the copious mucus production, which may be left in the tracks and trails formed by the burrowers (Frankel and Mead 1973). 5.3 Micro-organisms Abundant populations of phototrophic micro-organisms (phytobenthos) occur in intertidal muddy sediments. All living organisms up to 0.1 mm in size are considered micro-organisms (Yingst and Rhoads 1978). These phytobenthos can be subdivided (on the basis of their shape) into squat (coccal), single-celled organisms and organisms with a filamentous structure. The first group consists mainly of diatoms, microfauna and some species of bacteria. The latter group also includes certain species of bacteria, blue- green algae (cyanobacteria) and fungi. Most coccal micro-organisms are characterised by their excretion of mucus. Among other things, this mucus is responsible for adhesion to the sediment (Montague 1986; Rhoads and Boyer 1982). The coccal micro-organisms use the mucus to envelop sediment particles. When the population of coccal micro-organisms increases, the interstices are filled more and more (Frankel and Mead 1973; Holland et al. 1974). This results in membraneous films on the sediment surface. The strength of the elements of this film and the surface coverage of the topmost particles are responsible for an increase of the strength of the elastic, organically bounded surface (Bathurst 1967; Frankel and Mead 1973; Holland et al. 1974; Fahrbiter et al. 1983; Grant et al. 1986; Grant and Gust 1987; Paulic et al. 1986; Schrider and Heinzelmann 1987). ‘The micro-organisms with a filamentous structure form a network, in contrast to the unstructured settlement of the coccal micro-organisms. The network formed by blue- green algae and bacteria consists of separate, mutually entangled filaments, When blue-green algae are surrounded by a mucilaginous sheath, these filaments are mutually agglutinated (Ginsburg and Lowenstam 1958; Hommeril and Rioult 1965; Gebelein 1969; Neuman et al. 1970; Scoffin 1970). Benthic diatoms Benthic diatoms and their influence on the strength of the bed will be discussed extensively in this section because of their important role in stabilizing the sediment. The density and mucilage secretion of the various species determine the magnitude of the sediment stabilization. Benthic diatoms can be subdivided into epipsammic diatoms and epipelic diatoms. Epipsammic diatoms favour a sandy environment and are therefore not treated. Epipelic diatoms, however, favour a fine-grained (silt/mud)218 environment (Admiraal 1984; Ten Brinke et al. 1994) and prevail throughout the year at locations where wave and current energy are low. These diatoms are mobile micro- organisms that migrate up and down through the sediment in response to the tidal and day/night cycles (Palmer and Round 1965; Round 1979). They leave a trail of mucilage threads behind, which join separate sediment particles, Biofilms of epipelic diatoms often appear daily in response to the upward migration of the dominant diatoms as the result of the influence of light and emergence. Paterson (1989) studied the stability of sediment beds with different diatom coverage. He concluded that the length of the period, after the turn of the tide, during which the sediment bed retained its stability depended on the diatom coverage. ‘There are three effects of benthic diatoms on sediment stability 1) cohesive effect: Traditionally the cohesive strength of estuarine muds was ascribed to the electrostatic interactions between charged clay aggregates. However, more and more studies indicate the importance of the influence of organic factors on the cohesion (De Boer 1981; Montague 1986; Vos et al. 1988; Westall and Rincé 1994; Black 1997; Paterson 1997). For instance, diatoms excrete extracellular polymer substances (EPS), which enhance cohesion. Therefore, the stability of the bed will be increased because the EPS binds sediment particles at the sediment-water interface (Paterson 1997; Westall and Rincé 1994). It is still uncertain in what way EPS is bound to the sediment. A possibility may be ion-dipole interactions involving bridging by cations between the negatively charged bed and inorganic polymers by Van der Waals forces. 2) network effect: An extensive network of mutual attachment and mucus threads can develop when large amounts of epipelic diatoms migrate to the sediment surface. This network will strengthen the sediment bed against current and wave attack (Vos et al. 1988). 3) smoothed beds: The hydraulic roughness of the bed decreases when the inter- particle voids are filled by microbes and their secretions (Manzenrieder 1983) Therefore the strength of the bed increases and a larger shear velocity will be needed to entrain particles. The combined effects of biological stabilization of muddy sediments have been demonstrated by Black (1997) measuring in situ the erosion rate and the critical erosion stress (CES) of muds, which have been treated with chemical biocides and comparing the results with measurements on non-treated muds. He observed that the superficial flocs of treated muds had undergone gross disaggregation. Using copper sulphate and mercuric chloride as biocides, he found the median floc diameter to have changed from 135 um in the non-treated mud to values around 17 um in the muds treated with biocides. Comparable changes have been observed with regard to the 90% percentile of the granular composition. The critical erosion stress appeared to be depressed by a single order of magnitude. A mean value for the CES of undisturbed mud in the research area appeared to be in the order of 0.13 Nnv. For mud treated with mercuric chloride a CES value of 0.016 Nm was measured and for mud treated with copper sulphate a value of 0.029 Nmv®, The peak erosion rates appeared to be increased by up 10 650%219 During the same field experiments mentioned above, Black (1997) investigated the elation between diatoms and sediment stability. Sediment properties such as diatom biomass (chlorophyll a) and ATP content are considered to be biochemical indicators of micro-biological stabilization (Black 1997; Paterson 1997). Chlorophyll a is regarded to be the index of extracellular polysaccharides produced by diatoms (carbohydrate). The ATP (Adenosine Triphosphate) content is seen as the indicator of total living biomass in the sediment. Black (1997) found a strong co-variation between the critical erosion stress and chlorophyll a and carbohydrate (Figure 6). This indicates that benthic micro- organisms are indeed important regarding erosion resistance. The correlation between ATP content and critical erosion stress was not significant and therefore ATP content is, considered to be a poor biochemical indicator of adhesion by micro-organisms (Black 1997), 020 e Sew é : § om]? 3 i 2 som 4 a a a rr a er Selle Coots ay dy end) horn tu 3 = © +Chomical biocides E owe . = Natural sediments r ‘4 Laboratory value 5 ow] = ° ne ee ATP Concontation (ual dry sed) Figure 6 The relation between the Critical Erosion Stress and Soluble Carbohydrate (upper diagram), Chlorophyll a (middle diagram) and Adenosine Triphosphate (ATP) concentration (lower diagram) (after Black 1994).220 6 DISCUSSION ‘The impact of biochemical factors on the deposition and erosion of fine-grained sediments has received much attention in the last two to three decades. Most of the research has been directed towards the effect of biochemical activity on flocculation and (faecal) pellet production, Attention has been focussed on the role of the benthic filter feeders and on the grazing activity of zooplankton. However, it is obvious that other organisms may also have significant effects, from a cohesive sedimentary point of view. Bacteria for instance appear to have great influence on the aggregation of organic matter, especially phytoplankton. Therefore they may play an important role in the dynamics of suspended clayey sediments. More research is needed to fully understand the impact of bacteria and other micro-organisms on the sedimentation process of fine- grained material. There is a noticeable increase in scientific interest concerning the influence of the bed flora and fauna on the strength of the bed material. Much still has to be discovered concerning this complicated subject. 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