Fossil epithelial cell imprints as indicators of conodont biology

von Bitter, P.H. & Norby, R.D. 1994 10 15: Fossil epithelial cell imprints as indicators of conodont biology.
Size and growth characteristics of microsculpture polygons on the cup nodes of the bladelike Pa elements of the conodont Lochriea commutata support the hypothesis that they are epithelial cell imprints. They are ˜2–6 pn wide and ˜4–10 pn long and are the same size throughout growth of the element. The epithelial imprints increased in number with growth, either linearly or periodically; the latter possibility may be important for defining specific growth stages of conodont elements. The imprint location on the top of nodes suggests that the latter were tissue-covered and that the scissor-model of function applies to these bladelike Pa elements. The location of the imprints also reflects evolutionary history: a probable ancestor, L. cracoviensis (Belka), also lacks a platform and possesses well-developed microsculpture polygons on broad cup nodes. Finally, correlation between internal white matter and external microsculpture suggests that internal osteocytes may have supplied the external secreting cells with calcium phosphate from the inner storage bank. Conodont biology, fossil epithelial imprints, polygonal microsculpture, Carbonferous, Lochriea commutata .     

Traces of microboring organisms in Palaeozoic conodont elements

Conodont elements from Palaeozoic strata (Ordovician, Devonian and Carboniferous) contain abundant evidence for presumed post-mortem colonisation by endolithic organisms. It is possible to distinguish several morphotypes of microcavities in conodont apatite, which show some morphological similarities to microborings in calcareous substrates. Some of them are believed to be produced by microendoliths, some others are yet unknown from calcareous substrates. The present report deals with the various kinds of microendolithic traces in conodont apatite. The significance of microborings in conodont elements is discussed.     

New Middle Devonian conodont data from the Dong Van area,NE Vietnam (South China Terrane)

Middle Devonian conodonts from the Si Phai section in NE Vietnam are described. The section ranges from the Middle Devonian ensensis to timorensis conodont zones to the Late Devonian rhomboidea conodont Zone. A rich overall assemblage is described, including 27 taxa of species or subspecies rank and 11 taxa described in an open nomenclature. Among the dominant Polygnathus forms, four new taxa are described: Polygnathus linguiformis saharicus subsp. nov., Polygnathus linguiformis vietnamicus subsp. nov., Polygnathus rhenanus siphai subsp. nov., and Polygnathus xylus bacbo subsp. nov. Conodont assemblages are attributed to polygnathid, polygnathid-klapperinid, and klapperinid conodont biofacies representing hemipelagic to pelagic environments. The klapperinid biofacies, unreported in the previous literature, are here attributed to offshore areas of the external shelf. The taxonomic compositions of the studied conodont assemblages, as well as their CAI characteristics (CAI 4–5), suggest a palaeogeographic affinity of the studied strata to the Chinese Devonian Guangxi Basin, and the South China Terrane in general. Furthermore, the conodont biofacies and the palaeogeographic distribution of the fauna are discussed.     

Growth and patterning in the conodont skeleton

Recent advances in our understanding of conodont palaeobiology and functional morphology have rendered established hypotheses of element growth untenable. In order to address this problem, hard tissue histology is reviewed paying particular attention to the relationships during growth of the component hard tissues comprising conodont elements, and ignoring a priori assumptions of the homologies of these tissues. Conodont element growth is considered further in terms of the pattern of formation, of which four distinct types are described, all possibly derived from a primitive condition after heterochronic changes in the timing of various developmental stages. It is hoped that this may provide further means of unravelling conodont phylogeny. The manner in which the tissues grew is considered homologous with other vertebrate hard tissues, and the elements appear to have grown in a way similar to the growing scales and growing dentition of other vertebrates.     

Skeletal ontogeny and feeding mechanisms in conodonts

The growth and function of the conodont skeletal apparatus have important implications for early vertebrate relationships and the evolution of vertebrate hard tissues, yet they are poorly understood. Analysis of element length, platform linear dimensions, and platform area in discrete Pa elements of Carboniferous Idiognathodus and Gnathodus bilineatus reveals that the platform increased in size at a rate significantly above that required to maintain geometric similarity. Measurements of P, M and S elements in bedding-plane assemblages of Idiognathodus and G. bilineatus indicate that relative to Pa element length, Pb and S element growth was isometric, whereas M elements grew with negative allometry. There is no evidence to support loss or resorption of S and M elements in later growth stages, or to indicate periodic shedding and replacement of elements. These results are important for understanding apparatus and element Function. The positive allometry of the Pa element platform supports interpretations of a mashing or grinding tooth-like Function for platformed Pa elements. If conodonts were active suspension-feeders, the increasing food requirements of a growing conodont would require the filter array formed by the S and M elements to have grown at a rate significantly above isometry. The lack of positive allometry of S and M elements indicates that conodonts were not suspension-feeders and supports hypotheses that conodonts fed with a raptorial apparatus and teeth. □ Conodonts, vertebrates, skeletal apparatus, ontogeny, allometry, function, suspension-feeding, teeth.     

Adaptations to increasing hydraulic stress: morphology, hydrodynamics and fitness of two higher aquatic plant species

Sessile organisms often exhibit morphological changes in response to permanent exposure to mechanical stimulation (wind or water movements). The adaptive value of these morphological changes (hydrodynamic performance and consequences on fitness) has not been studied extensively, particularly for higher plants submitted to flow stress. The aim was to determine the adaptive value of morphological patterns observed within two higher aquatic plant species, Berula erecta and Mentha aquatica, growing along a natural flow stress gradient. The hydrodynamic ability of each ramet was investigated through quantitative variables (drag coefficient and E-value). Fitness-related traits based on vegetative growth and clonal multiplication were assessed for each individual. For both species, the drag coefficient and the E-value were explained only to a limited extent by the morphological traits used. B. erecta exhibited a reduction in size and low overall plant drag at higher flow velocities, despite high drag values relative to leaf area, due to a low flexibility. The plants maintained their fitness, at least in part, through biomass reallocation: one tall ramet at low velocity, but shorter individuals with many interconnected stolons when flow velocity increased. For M. aquatica, morphological differences along the velocity gradient did not lead to greater hydrodynamic performance. Plant size increased with increasing velocities, suggesting the indirect effects of current favouring growth in high velocities. The fitness-related traits did not demonstrate lower plant fitness for high velocities. Different developmental constraints linked to plant morphology and trade-offs between major plant functions probably lead to different plant responses to flow stress.     

Compostional variations and patterns of condont reworking in Late Devonian and early carboniferous calciturbidites (Moravia, Czech Republic)

Summary Compositional variations and grain-size properties of both carbonate constituents and conodonts as an alternative component group were used for interpreting the processes governing the deposition of upper Famennian and middle Tournaisian calciturbidites in Moravia, Czech Republic. Both the composition and grain-size properties of conodont element associations showed to be markedly dependant on facies type of their host sediment. Upper Devonian calciturbidite successions deposited on flanks of wide, Moravian-Silesian carbonate platform are composed mainly of echinoderm-and peloid-rich wacke/packstones and intraclastic float/rudstones (fine-grained calciturbidites, “normal” calciturbidites with Tab Bouma sequences, debris-flow breccias) with abundance of shelf-and shelf margin conodont taxa and epipelagic and “mesopelagic” conodonts. Upper Devonian calciturbidites deposited on slopes of volcanic sea-mounts are composed of echinoderm-and peloid-rich wacke/packstones and float/rudstones with increased proportion of intraclasts and volcanigenic lithoclasts (fine-grained calciturbidites, normal calciturbidites), yeilding abundant conodont associations with higher proportion of “mesopelagic” taxa compared to the platform-flank examples. Middle Tournaisian calciturbidite succession composed of crinoid-, peloid-, intraclast-and lithoclast-rich lime mudstones, wacke/packstones and float/rudstones (normal calciturbidites and debris-flow breccias) yielded conodont element associations rich in shelt-and shelf-margin taxa, “mesopelagic” conodonts and reworked Middle-and Upper Devonian conodonts. In general, the ratio of shelf-and shelf margin conodont taxa to “mesopelagic” taxa is distinctly lower in finegrained calciturbidites than it is in normal calciturbidites and debris-flow breccias. Grain-size properties (mean grain size and sorting) and percentage of fragmented conodont elements, too, are markedly dependant on the facies type: in fine-grained calciturbidites the values of mean grain-size and fragmentation are low and the sorting is good to very good whereas in normal calciturbidites and debris-flow breccias the values of mean grain-size and fragmentation are distinctly higher and the sorting is poorer. The interdependence of facies type and composition and grain-size properties of conodont element associations in gravity-flow deposits is explained as resultant from hydrodynamic sorting during turbidity current flow and final deposition of the bed. Compositional variations observed in our sections may thus be attributed to facies variability (coarsening-and thickening-upward trends) rather than to sea-level fluctuations (highstand shedding of carbonate platforms). On the other hand, significant enrichment in reworked conodont taxa in middle Tournaisian normal calciturbidites compared to scarcity and/or absence of such conodonts in essentially identical facies of upper Famennian age indicate sea-level to be the major control governing such compositional variations, with low relative sea-level stand in middle Tournaisian and high relative sea-level stand in upper Famennian. Thorough analysis of conodont evolution, palaeoecology and taphonomy, with emphasis on understanding the processes of deposition of their host rock, are recommended for any biostratigraphic and biofacies study to be done in carbonate sediments deposited under strong hydrodynamic regimes, such as calciturbidites, temperstites, debris-flow deposits, shelf-edge oolitic sands, tidal-channel facies etc.     

Hydrodynamic study of biogranules obtained from an anaerobic hybrid reactor

The bed expansion characteristics of a fluidized bed containing bacterial granules have been studied. These biogranules were obtained from an anaerobic hybrid reactor, which uses biogranules (without carrier particle) in fluidized condition. The settling velocity study of biogranules has shown that the drag coefficient of biogranule is greater than that of the rigid particle at the same Reynolds number. A new correlation based on this finding has been developed. The bed expansion study has demonstrated that a linear relationship exists between the natural logarithm of bed porosity and the natural logarithm of upflow superficial liquid velocity for the bed containing either a particular fraction of biogranule size or biogranules with wide size distribution. For a fluidized bed having a particular granule size, the bed porosity, and liquid superficial velocity could be related by the classic equation suggested by Richardson and Zaki (1954). The characteristic parameter of this correlation, the slope of the line n, has been related with Reynolds number. The intercept of the line gave a smaller value than the unhindered settling velocity of the particle. For fluidized bed having wide size distribution, the characteristic parameter n could not be related to Reynolds number. But the correlation suggested for single biogranule size has been found to predict n value with an average error of 2.3%.     

Zur Biologie, Taxonomie und Chronologie der Conodonten

The actual state of knowledge concerning the conodont animal is critically reviewed. All soft tissue documented by the specimens of the conodont animal were interpreted by the British collectors as Craniota (Vertebrata) features. Opinions on the precise affinities, however, remain controverse: The alleged chorda could have been a gut, bilateral symmetry, myomery, apatitic biomineralisation occur also in Invertebrata, caudal fins do in Chaetognatha. Even the lobes in the capital area interpreted as sensory organs could have served as aiding organs for food intake. Therefore, the originally sensational findings are reduced to the core issue that the conodont animal had an eel-like body and at the blunt end of it, behind peculiar lobes, the mineralized conodont elements were arranged. As a consequence, a relationship to the Protochordata cannot be excluded. A systematic position of the conodont animal as possible Vertebrata is likely only if organized below the Ostracodermata which is particularly so because conodonts occur so early in the Cambrian. In the second part of the article conodonts are discussed as guide fossils. The comprehensive, empirically grown taxonomy of the single elements is the prerequisite for recognizing outstanding, high resolution stratigraphy. In comparison, a multielement taxonomy remains as long problematical, as it is based on assumed, mostly statistically reconstructed rather than on natural multielement apparatuses. Taxonomical and nomenclatorial operations built on assumed apparatus affinity can bear heavily on nomenclature and stratigraphy of platform elements. It can be shown at the example of the Devonian that the platform elements have provided particularly in the pelagic facies a Standard Conodont Zonation that covers time almost without any gaps. The basis for this is that platform single element species and their phylogenetic development are extra-well known. In a special paragraph, history and applicability of the Standard Zonation is discussed in details. Different conodont biofacies are developed between the pelagic realm where the Standard Zonation originated, and the coastal areas so that conodonts can indicate temporal and bathymetric relationships of the mother rocks. In addition alternative conodont zonations were introduced mainly for the shallow water. Because of their conspicuous stratigraphic significance in the Devonian, conodonts were employed by International Committees to redefine Series and Stage Boundaries. For the boundary selection species were utilized from phylognetic lineages as only from tight evolution their point of origination could be well identified and recognized in worldwide correlation. The GSSPs (Global Stratotype Section and Point) within the Devonian are discussed in detail. In recent times, attempts are made at calibrating the duration of conodont zones on the basis of their well known rate of evolution and high temporal resolution. These analyses have to be continued in order to demonstrate the absolute time relationship of condont zones in more and continuous detail. Currently the phyletically controlled Standard Conodont Zonation is the authoritative stratigraphic method in the Devonian. It has provided together with the GSSPs precise and valuable fix points for a Global Time Scale (GTS). In combination with additional stratigraphie methods the Standard Conodont Zonation may provide further refinements in the future.     

Experimental Measurement of Settling Velocity of Spherical Particles in Unconfined and Confined Surfactant-based Shear Thinning Viscoelastic Fluids

An experimental study is performed to measure the terminal settling velocities of spherical particles in surfactant based shear thinning viscoelastic (VES) fluids. The measurements are made for particles settling in unbounded fluids and fluids between parallel walls. VES fluids over a wide range of rheological properties are prepared and rheologically characterized. The rheological characterization involves steady shear-viscosity and dynamic oscillatory-shear measurements to quantify the viscous and elastic properties respectively. The settling velocities under unbounded conditions are measured in beakers having diameters at least 25x the diameter of particles. For measuring settling velocities between parallel walls, two experimental cells with different wall spacing are constructed. Spherical particles of varying sizes are gently dropped in the fluids and allowed to settle. The process is recorded with a high resolution video camera and the trajectory of the particle is recorded using image analysis software. Terminal settling velocities are calculated from the data.The impact of elasticity on settling velocity in unbounded fluids is quantified by comparing the experimental settling velocity to the settling velocity calculated by the inelastic drag predictions of Renaud et al.¹ Results show that elasticity of fluids can increase or decrease the settling velocity. The magnitude of reduction/increase is a function of the rheological properties of the fluids and properties of particles. Confining walls are observed to cause a retardation effect on settling and the retardation is measured in terms of wall factors.     

Physiological Performance and Stream Microhabitat use by the Centrarchids Lepomis megalotis and Lepomis macrochirus

Two centrarchids, Lepomis megalotis and L. macrochirus, were compared in laboratory studies of prey capture success, swimming endurance, morphology, hydrodynamic drag, and thermal tolerance, as well as field observations of focal point velocity and depth. For both species, capture of planktonic prey declined as current velocity increased, but L. megalotis was more efficient than L. macrochirus at higher current velocities. Capture of floating prey was not influenced by current velocity, but L. megalotis was more efficient overall at all velocities. Of the two species, L. megalotis was significantly more streamlined, had relatively lower hydrodynamic drag, and had higher swimming endurance in current. Both species had significantly higher critical thermal maxima (CTMax) in summer than in winter, but variance in CTMax was greater for L. megalotis than for L. macrochirus in both summer and winter. Differences between L. megalotis and L. macrochirus in performance and morphology may have direct influence on their relative abundances in small streams. Field observations showed L. megalotis was more common than L. macrochirus in faster, shallower microhabitats.     

Optimizing the rotor design for controlled-shear affinity filtration using computational fluid dynamics

Controlled shear affinity filtration (CSAF) is a novel integrated processing technology that positions a rotor directly above an affinity membrane chromatography column to permit protein capture and purification directly from cell culture. The conical rotor is intended to provide a uniform and tunable shear stress at the membrane surface that inhibits membrane fouling and cell cake formation by providing a hydrodynamic force away from and a drag force parallel to the membrane surface. Computational fluid dynamics (CFD) simulations are used to show that the rotor in the original CSAF device (Vogel et al., 2002) does not provide uniform shear stress at the membrane surface. This results in the need to operate the system at unnecessarily high rotor speeds to reach a required shear stress of at least 0.17 Pa at every radial position of the membrane surface, compromising the scale-up of the technology. Results from CFD simulations are compared with particle image velocimetry (PIV) experiments and a numerical solution for low Reynolds number conditions to confirm that our CFD model accurately describes the hydrodynamics in the rotor chamber of the CSAF device over a range of rotor velocities, filtrate fluxes, and (both laminar and turbulent) retentate flows. CFD simulations were then carried out in combination with a root-finding method to optimize the shape of the CSAF rotor. The optimized rotor geometry produces a nearly constant shear stress of 0.17 Pa at a rotational velocity of 250 rpm, 60% lower than the original CSAF design. This permits the optimized CSAF device to be scaled up to a maximum rotor diameter 2.5 times larger than is permissible in the original device, thereby providing more than a sixfold increase in volumetric throughput.     

Shifts in drag and swimming potential during grayling ontogenesis: relations with habitat use

The drag coefficient (C_d) of grayling Thymallus thymallus was dependent on body surface conditions and rigidity. At comparable flow conditions, C_d values of a fish preserved in formalin (high body rigidity) were 15–30% lower than those obtained on a freshly-killed fish (medium rigidity); the presence of skin mucus on fish could reduce C_d by 10%. The hydrodynamic potential of grayling increased during ontogenesis, because C_d values decreased (except for yolk sac larvae, which had a particular morphology) and the swimming capacities (in terms of relative muscular mass) increased. Grayling morphology evolves towards hydrodynamically efficient shape at high velocities, and there is a relationship between these shifts in hydrodynamic abilities and the different habitats (in terms of current velocity) used by five morphological groups. Therefore, the concept of hydrodynamic potential (i.e. hydrodynamics of shape and swimming capacities) could be a useful tool in fish ecomorphology and predictions of habitat use.     

In situ measurements of hydrodynamic forces imposed on Chondrus crispus Stackhouse

Among the hydrodynamic forces experienced by intertidal organisms, drag and the impingement force are thought to have the greatest effect on macroalgae. These forces are modified by biotic factors such as algal morphology, reconfiguration, and the presence of a canopy. However, much of what is known about the hydrodynamics of macroalgae has been garnered from low-velocity laboratory flume studies. Few field studies have measured drag and none have directly measured the effects of the canopy on force. To examine in situ hydrodynamic forces imposed on the turf forming macroalga Chondrus crispus, compact digital force sensors were developed that measure and record the 3-dimensional force imposed on a macroalga without disturbing the surrounding canopy. Sensors were positioned within natural Chondrus beds and the effects of the canopy, algal morphology, and sea state on in situ hydrodynamic force were examined. Additionally, the predictions of a new model for drag on flexible macroalgae were tested by simultaneously measuring force and water velocity. Digital force recordings indicated that Chondrus only experience drag; lift and impingement force were negligible in all combinations of factors. Canopies significantly reduced drag by 15-65%. Morphology and size also influenced drag, such that lower forces were imposed on small planar algae than large arborescent individuals. Further, planar algae experienced low drag in all combinations of sea and canopy state, indicating that these individuals may not be as susceptible to wave disturbance as arborescent individuals. Overall, these data indicate that the ability for Chondrus to grow large, arborescent individuals is dependent on the drag reducing properties of the canopy, while more hydrodynamically harsh habitats may be accessible to planar morphologies. Additionally, these data suggest that drag models for canopy forming macroalgae must incorporate the effects of the canopy to predict drag accurately in situ.     

Development of net energy intake models for drift-feeding juvenile coho salmon and steelhead

We developed models to predict the effect of water velocity on prey capture rates and on optimal foraging velocities of two sympatric juvenile salmonids, coho salmon and steelhead. Mean fish size was ~80 mm, the size of age I+ coho and steelhead during their second summer in Southeast Alaska streams, when size overlap suggests that competition might be strongest. We used experimentally determined prey capture probabilities to estimate the effect of water velocity on gross energy intake rates, and we modeled prey capture costs using experimental data for search and handling times and published models of swimming costs. We used the difference between gross energy intake and prey capture costs to predict velocities at which each species maximized net energy intake rate. Predicted prey capture rates for both species declined from ~75 to 30–40 prey/h with a velocity increase from 0.30 to 0.60 m·s⁻¹. We found little difference between coho and steelhead in predicted optimum foraging velocities (0.29 m·s⁻¹ for coho and 0.30 m·s⁻¹ for steelhead). Although prey capture ability appears to be more important than are prey capture costs in determining optimum foraging velocities, capture costs may be important for models that predict fish growth. Because coho are assumed to pay a greater swimming cost due to a less hydrodynamic body form, we also modeled 10 and 25% increases in hydrodynamic drag to assess the effect of increased prey capture costs. This reduced optimum velocity by 0 and 0.01 m∙s⁻¹, respectively. Habitat segregation among equal-sized coho and steelhead does not appear to be related to the effects of water velocity on their respective foraging abilities.     

Patterns of bilateral asymmetry and allometry in Late Devonian Polygnathus conodonts

Conodont animals were early jawless vertebrates equipped with a feeding apparatus composed of several tooth‐like elements. The P1 elements, at the rear of the apparatus, were characterized by a robust shape and rapid morphological evolution. Occlusion occurred between paired right and left P1 elements, occasioning some bilateral asymmetry, which, together with allometric growth, may partially obliterate the temporal differences. The present study aims to disentangle these different components of morphological variation in Late Devonian Polygnathus P1 conodont elements. An extensive 2D geometric morphometric analysis of the platform shape was performed through the Famennian record of two outcrops. This analysis was completed by a 3D study on a subset of conodont elements. The 2D and 3D morphometric quantifications provided highly congruent results, showing that the 2D shape constitutes a good approximation of the element geometry. The 3D analysis delivered further insights into the relationship between the geometry of the elements and the constraints related to occlusion. The 2D analysis allowed a quantitative assessment of the variation among species and through time. Allometry and bilateral asymmetry were differently expressed depending on the species considered, suggesting that constraints imposed on pairing by the morphology of the elements varied even among related species. The within‐species variation was so important that it largely obliterated temporal trends; a relationship of Polygnathus shape and conodont biofacies variations through the Famennian nevertheless suggested an evolution driven by ecological interactions between conodont genera.     

Conodont affinity of the enigmatic Carboniferous chordate Conopiscius

Conopiscius shares V-shaped myomeres with the co-occurring conodont Clydagnathus but instead of a complex oral apparatus it has only a single pair of conical elements, and structures resembling scales are associated with its myomeres. Moreover, the coarsely crystalline crown tissue typical for conodonts has not been identified in the Conopiscius elements, which show only a finely lamellar skeletal tissue. The gap between conodonts and Conopiscius may be filled by isolated elements of similar morphology and structure occurring in the Late Devonian. They reveal a very thin external layer developed mostly at the tooth tip and resembling conodont crown tissue. The pulp cavity is partially filled with layered or spherulitic phosphatic tissue of the kind known also in conodonts (basal filling tissue) and early vertebrates (lamellin). Conodont elements of similar morphology and representing uni-membrate oral apparatuses have not been previously reported from the Devonian or Carboniferous but occur near the Cambrian–Ordovician transition ( Proconodontus ) and in the Late Permian ( Caenodontus ). It is proposed that Conopiscius represents a mostly cryptic conodont lineage extending from the Early Ordovician to the Permian, instead of being directly related to the agnathans.     

A spathognathodont lineage of Mississippian conodonts

Evolutionary lineages of conodont elements and of some apparatuses are gradually being documented to increase the value of these fossils in stratigraphic interpretation and to further our understanding of conodont taxonomy. The well-documented faunas from Mississippian strata of the Illinois Basin and type Mississippian beds provide a great deal of information for the recognition of phylogenies. The disjunct element genus Spathognathodus is part of a basic and long-ranging type of conodont apparatus from which a number of other forms have been derived, and some of its Mississippian disjunct element species can now be placed in multielement species. A lineage involving spathognathodont Pa elements recognized in the Mississippian, then, is the basis of the present study. The phylogeny recognized started with Synprioniodina? crassidentata, and includes S.? regularis, S.? pulchra, S.? coalescens, S.? spicula, Hinduodus cristula, and H. minutus in the Mississippian. Aputiignathus scitulus and A. penrscitulus apparently represent a separate line, which also might include disjunct element Spathognathodus rexroadi.     

Tidal variation in the settling diameters of suspended matter on a tidal mud flat

During one tidal period, measurements of the variation of current velocities, suspended sediment concentration and settling velocities of the suspended matter were carried out on a tidal mud flat 1.8 km west of Ballum Sluse in the northern part of the Lister Dyb tidal basin. The settling velocities have been converted to equivalent settling diameters. Current velocities follow a variation pattern well-known from other parts of the Wadden Sea. Variations in current velocities are responsible for the variation in suspended sediment concentration and thereby indirectly control the equivalent particle sizes, because high suspended sediment concentration favours the formation of flocs. Maximum concentrations of up to 532 mg/l were recorded at the beginning of the flood period and at the end of the ebb period, when current velocities are high. This is reflected in the median equivalent settling diameters, which show corresponding high values of 83 μm at the beginning of the flood period and 96 μm at the end of the ebb period.     

Hydrodynamic theory of swimming of flagellated microorganisms.

A theory of the type commonly used in polymer hydrodynamics is developed to calculate swimming properties of flagellated microorganisms. The overall shape of the particle is modeled as an array of spherical beads which act, at the same time, as frictional elements. The fluid velocity field is obtained as a function of the forces acting at each bead through Oseen-type, hydrodynamic interaction tensors. From the force and torque equilibrium conditions, such quantities as swimming velocity, angular velocity, and efficiency can be calculated. Application is made to a spherical body propelled by a helical flagellum. A recent theory by Lighthill, and earlier formulations based on tangential and normal frictional coefficients of a curved cylinder, CT and CN, are analyzed along with our theory. Although all the theories predict similar qualitative characteristics, such as optimal efficiency and the effect of fluid viscosity, they lead to rather different numerical values. In agreement with Lighthill, we found the formalisms based on CN and CT coefficients to be somewhat inaccurate, and head-flagellum interactions are shown to play an important role.