A hydromechanical principle for particle retention in Mytilus edulis and other ciliary suspension feeders

It is generally thought that the laterofrontal cirri of the bivalve gill act as filters that retain suspended particles in the through current and transfer the particles onto the frontal surface of the gill filaments. In Mytilus edulis calculations indicated that if water passed between the branching cilia of the cirri that are assumed to constitute the filter the pressure drop needed would amount to about 10 times the actual pressure drop across the whole gill. Thus, instead of acting as filters the laterofrontal cirri seem to move water. Presumably, the cirri together with the frontal cilia produce the water currents along the frontal surface of the gill filaments. Particle retention in the bivalve gill implies the transfer of suspended particles from the current of water about to enter an interfilamentar space into a neighbouring frontal surface current. The complex three-dimensional pattern of flow that arises where the 2 systems of current meet is characterized by steep velocity gradients. Particles that enter such steep, steady velocity gradients become exposed to transverse forces that cause the particles to migrate perpendicularly to the direction of flow. Whether particles enter the surface current, i.e. are retained, or they stay within the through current andescape, depends primarily upon particle size, and upon the steepness and height of the gradients within the boundary zone between the surface current and through current. Further studies are needed to evaluate the capacities and relative importance of this hydromechanical particle-trapping mechanism in suspension feeding bivalves. It is suggested that in downstream particle-retaining systems, e.g. on the tentacles of polychaetes and entoprocts, velocity gradients between through currents and surface currents also act as the particle-collecting mechanism.     

Methods of ciliary filter feeding in adult Phoronis muelleri (phylum Phoronida) and in its free-swimming actinotroch larva

Microscope-video recordings were used to describe basic characteristics of particle capture in adult Phoronis muelleri and in its actinotroch larva. The larva captures food particles by means of close interplay between the tentacles and the oral hood. Suspended particles in the incoming water flow set up by the lateral "pump" cilia on the tentacles approach the tentacles with a velocity of about 1.5 mm s^-1. Near the tentacles, the particles are stopped by the stiff sensory laterofrontal cilia acting as a mechanical sieve. Simultaneous with stoppage of a particle, the oral hood is rapidly lifted so that water from the surrounding area flows in to occupy the volume created under the hood. Due to the proximity of the tentacle with the arrested particle, this suction will draw the particle away from the laterofrontal cilia into the increased space under the oral hood with a velocity of about 0.6 mm s^-1 before it is subsequently carried to the mouth. If a particle is stopped near the tip of a tentacle this may trigger a tentacle flick, with a tip velocity of about 5-7 mm s^-1, which brings the particle down toward the lifting edge of the hood. A stopped particle may cause a local disruption of the metachronism of the lateral cilia for about 0.14 s. Likewise in the adult, when an incoming particle with a velocity of about 1.6 mm s^-1 is stopped near the tip of a tentacle this triggers a flick, which brings the particle down towards the mouth. The duration of the active flick phase is about one-tenth of the flick cycle. Only when a particle is stopped on the outer part of a tentacle is a flick triggered. Otherwise the particle is either transported down along the frontal surface of the tentacle by means of the frontal cilia, or transferred into the downward current created by the lophophore. The metachronal wave velocity is about 0.25 mm s^-1, the wavelength about 12 µm, and hence the ciliary beat frequency about 21 Hz (at ~16°C). Essential features of the filter-feeding process in both actinotroch larvae and adult P. muelleri reported here resemble the ciliary-sieving process described for bryozoans in recent years. It should be pointed out that the stiff, mechanosensory laterofrontal cilia may be a decisive factor for understanding ciliary upstream collecting, also in pterobranchs and brachiopods in which sensory laterofrontal cilia are present.     

Functional morphology and feeding behavior of Paraprionospio pinnata (Polychaeta: Spionidae)

The functional morphology and feeding behavior of Paraprionospio pinnata (Ehlers) were studied. The tentacular feeding palps of P. pinnata possess four groups of cilia: laterals, latero-frontal cirri, frontals and basal transverse rows. The lateral cilia beat in metachronal waves creating in current that flows toward the frontal surface of the palp. The latero-frontal cirri deflect suspended particles onto the frontal surface and potentially resuspend deposited particles. The frontal cilia line the groove of the palp and transport particles to the ciliated pharynx. The basal transverse cilia also beat in metachronal waves and together with the frontal cilia of the branchiae produce a U-shaped current that removes particles rejected by the pharynx from the burrow. The significance of particle selection at the site of the pharynx in spionid polychaetes is hypothesized to be constrained by the interaction of specific morphologies and behaviors that reduce the effects of epifaunal predation and browsing. The relationship between sediment permeability and type of respiratory or ventilation current is hypothesized to be a potentially important factor in understanding the distribution of spionid polychaetes.     

Fluid mechanics of the mussel gill: The lateral cilia

Quantitative data on the water currents produced by the ciliary tracts of the gill filaments are needed to understand the fluid mechanics of suspension feeding in bivalves, as well as in other ciliary suspension feeders. This paper investigates the water currents produced by the bands of lateral cilia, as studied on isolated gill filaments, gill fragments, and intact gills of the mussel Mytilus edulis L with severed adductor muscles. The metachronally beating cilia produce oscillatory currents near the oscillating enveloping surface of the ciliary bands and rectilinear currents, the interfilamentary through-currents, farther from the surface. It is suggested that the oscillatory currents play an important role in the fluid mechanical capture of suspended particles. In the intact gill the interfilamentary currents pass the bands of lateral cilia at velocities that are two or more times higher than those generated by the bands of isolated filaments. The mussel gill is compared with an optimized peristaltic pump.     

Ciliary feeding of tornaria larvae of Ptychodera flava (Hemichordata: Enteropneusta)

Ciliary feeding by tornariae of Ptychodera flava (Eschholtz) and other tornariae from the plankton is compared with Garstang's (1939) account of feeding by these larvae, which account contains errors, and with ciliary feeding by echinoderm larvae. Some details of ciliation are also described. As in echinoderm larvae, band cilia beat away from the food grooves and retain particles on the upstream side of the ciliated band, but tornariae use muscles less in ingestion and rejection. In early-stage and most late-stage echinoderm larvae, the ciliated band functions in both swimming and feeding, but in tornariae the ciliated band is arranged meridionally so that few portions of the ciliated band produce a posteriorly-directed current and a locomotory teletroch is needed for swimming. Faster-swimming tornariae observed in bowls achieved higher ingestion and clearance rates. These observations raise questions about form and function in the giant Planctosphaera pelagica. Cilia of the locomotory telotroch increase in length as tornariae increase in size, but there is little increase in length of cilia which capture food. Instead, the length of the ciliated band increases relative to other larval tissues by means of increasing convolution of the band. Hence, the volume of water processed for food probably increases relative to energy expended by larvae during development to the tentaculate stage. However, the length of the ciliated band may decrease relative to other larval surfaces with continued increase in size beyond this stage. These interpretations of growth and feeding efficiency are consistent with the reported geographic distribution of tornariae with and without tentacles.     

Form and feeding mechanism of a living Planctosphaera pelagica (phylum Hemichordata)

We describe aspects of the anatomy and suspension-feeding mechanism of a single Planctosphaera pelagica captured from the plankton in June 1992 off Bermuda in the western Atlantic. We also describe several unusual features of the larva, including its occurrence in surface waters, unusually large size, and limited swimming ability. Our account of the form and feeding behavior of P. pelagica is the first based on observations of a specimen captured and observed alive. Our limited observations suggest that the planctosphaera may use a suspension-feeding mechanism much like that of the other feeding deuterostome larvae (the pluteus and bipinnaria larvae of echinoderms and the tornaria larva of enteropneust hemichordates) known to capture food particles using a single ciliated band. Although we could not observe cilia directly, the movement of dye streams and food particles and the structure of the ciliated band suggest that some particles may be captured at the ciliated band by the reversal of ciliary beat. The planctosphaera possesses many prominent mucous glands near the food grooves. This suggests an important role of mucus in the biology of the larva, but we were not able to observe directly any role of mucus in particle capture.     

Functional morphology of the larval tentacles of Phragmatopoma californica (Polychaeta: Sabellariidae): composite larval and adult organs of multifunctional significance

Larvae of the sabellariid polychaete Phragmatopoma californica (Fewkes), which were collected in San Diego, California and were competent to metamorphose after 18 to 30 d of development were observed in vivo by videoequipped light microscopy, and the fine structure of the larval tentacles was examined by transmission and scanning electron microscopy. Each tentacle has tufts of at least two types of immotile cilia arranged in dorsolateral and ventrolateral rows, and a ventral groove covered by two types of motile cilia that beat independently of each other. The epidermis is regionalized into glandular, sensory, locomotory, and support cell types and contains four longitudinal bundles of basiepithelial nerve fibers. A layer of connective tissue separates the epidermis and the nerve tracts from obliquely striated muscles that occur within the peritoneum that lines the central coelomic cavities. The peritoneum forms an intact coelomic epithelium that overlies and interdigitates with the muscle cells, with no intervening basal lamina. The muscle cells are considered to be intraperitoneal because they are located above the basal lamina and they lack intercellular junctions with the peritoneal cells. Specialized peritoneal cells form a striated myoepithelial blood vessel that partitions the coelom into medial and lateral cavities. No neuromuscular junctions were found, but both muscular and ciliary movement seem to be under neuronal control. The basiepithelial nerve terminals appear to synapse into the connective tissue layer toward the intraperitoneal muscle. Several similarities in tissue organization are noted between the larval tentacles of P. californica and the tube feet of echinoderms. Observations on the ontogeny, morphology, and behavior of the tentacles suggest that they are multifunctional organs involved in feeding, construction of the juvenile sand tube, locomotion, attachment, and sensory perception during larval and adult lives.     

Particle capture in the musselMytilus edulis: The role of latero-frontal cirri

Microscope video graphs of particle paths near one-filament-thick mussel gill preparations, stimulated with a nerve transmitter (10^–6 M serotonin which restores normal ciliary activity), were used to disclose the capture of 6 m algal cells. Suspended algal cells carried with the water were stopped for a while at the entrance to the interfilament gap by the action of the latero-frontal cirri (Ifc), and transferred to the frontal side of the filament to be transported towards the marginal food groove. The event of transfer took place during approximately a time interval of 1150 to 1/25 s. To gain a better understanding of the capture mechanism and retention efficiency versus particle size, the flow through and around the Ifc was theoretically estimated. Normally beating Ifc create periodic, unsteady, three-dimensional flows at the entrance to the interfilament canal. During the active beat most of the water is deflected to flow around the branching cilia of the Ifc while some of the water is strained by these. Large particles (> 4 m) are stopped and transferred to the frontal current, whereas smaller particles either follow the flow around the Ifc and escape or they are stopped by the branching cilia.     

Ultrastructure and behavior of the larva of Phragmatopoma californica (Polychaeta: Sabellariidae): identification of sensory organs potentially involved in substrate selection

The tentacles of the larvae of Phragmatopoma californica (Fewkes) a tubicolous, reef-building polychaete, were examined by video-equipped light microscopy and transmission and scanning electron microscopy. The surface of the tentacles has a unique ciliation pattern, consisting of dorsal tufts of short immotile cilia, ventrolateral tufts of short and long immotile cilia, and ventral motile cilia. Cells bearing immotile cilia are primary sensory cells with long basal processes that form synapses with basiepithelial nerve fibers. The sensory cell cytoplasm is similar to that of nervous tissue, and contains microtubules, neurofilaments, and synaptic vesicles. Sensory cell synapses with basiepithelial nerves appear to be both axodendritic and axoaxonic. The structure of the immotile cilia is compared to that of motile cilia. Unlike motile cilia, immotile cilia are short, rigid, end in a blunt tip and possess and axoneme with typically arranged mictotubules that terminate in an electron-dense end plate. The basal feet of immotile cilia do not anastomose with adjacent basal bodies, and the ciliary membrane is loosely applied to the axoneme and is covered by a surface coat of filamentous material. The use of the larval tentacles during substrate exploration, and the location and ultrastructure of sensory cilia, indicate that they may be involved in the perception of substrateassociated chemical signals and/or mechanical cues of significance in substrate selection.     

Analysis of capture efficiency in suspension feeding: Application of nonparametric binary regression

Suspension feeding is an important mechanism by which many marine organisms obtain food. A nonparametric method for analyzing experimental results on capture efficiency in suspension feeding is described. This approach can be used to estimate capture efficiency along the length of a cilium and to compare the capture efficiencies of two different cilia. The method is applied to experimental data on feeding by larvae of the clamMerenaria merenaria (Linné), to examine changes in capture efficiency through time, and to assess the effect of particle electrostatic charge on capture efficiency.     

Activity patterns in the terebellid polychaete <Emphasis Type="Italic">Eupolymnia nebulosa</Emphasis> assessed using a new image analysis system

The activity and surface sediment displacement by the terebellid polychaete Eupolymnia nebulosa were monitored, using new image acquisition and image analysis procedures. We used a video sensor mounted on a motorized table, to acquire adjacent images of the whole studied aquarium within about 2 s. These images were then grouped within a composite image acquired every 15 s. Consecutive composite images were compared to infer activity and surface sediment displacement. This procedure proved efficient for E. nebulosa as indicated by (1) the continuity of the tentacles within composite images, and (2) the direct comparison of images and detected activities. There were important temporal changes in the relative importance of the three main types of activity: feeding, tube-building and (both partial and total) emergence from the tube, accounting, respectively, for 75, 15 and 10% of the entire experiment duration. Activity intensity (cm² min^-1) was assessed through the surface of pixels with grey levels differing by more than 20 (on a 0–255 scale) between two consecutive images. Feeding was associated with low activity intensity, whereas tube-building and emergence from the tubes were associated with high activity intensities. Surface sediment displacement mostly resulted from tentacle activity both during feeding and tube-building and was almost zero when worms emerged from their tube. We used our experimental set up to assess spatial changes in activity intensity and particle displacements along the tentacles. Most of the activity occurred within the first 4–6 cm from the tube aperture. Particle loss during their transfer along the tentacles preferentially affected larger particles as postulated by the model of particle selection and transport in tentaculate deposit-feeders. Moreover, the speed of the particles along the tentacles correlated negatively with their size. This contributed to increase in the residence time of larger particles on the tentacles and thus, to further enhance their probability of being lost. The size distributions of particles during feeding and tube-building did not differ significantly although visual observations confirmed the preferential use of larger particles for tube-building. This suggests the existence of a sorting step occurring elsewhere than on the tentacles as already demonstrated for spionid polychaetes.     

Particle capture in the crown of the ciliary suspension feeding polychaete Sabella penicillus: videotape recordings and interpretations

An experimental setup was designed for in situ videotape recording of the particle capture process in the crown of the polychaete Sabella penicillus. Intact individuals of S. penicillus (collected in the Gullmornfjord, Sweden in 1992) were exposed to either 6 m Latex spheres or Rhodomonas sp. flagellate cells (6 m). The capture of the added particles was recorded on video. From frameby-frame analyses particle velocities were estimated and the shape of the three-dimensional particle paths was inferred. The mean velocity of particles approaching the crown was estimated to be ca. 1 mm s^-1, increasing to ca. 1.7 mm s^-1 in the interpinnule channel. At the moment of capture the particles were seen to follow a curved, near circular path close to the tips of the latero-frontal cilia. The transport velocities on the frontal side of the pinnules and filaments were estimated to be up to 0.15 and 0.5 mm s^-1, respectively. Counting captured particles relative to particles arriving within the area of the pinnules gave a rough, direct estimate of nearly 100% retention rate when the polychaete was feeding undisturbed. Together with results from clearance measurements in the literature this implies that the worm is able to capture particles down to 3 m entering the interpinnule channel almost 100% effectively. In view of the 80-m wide interpinnule channel and 40-m spacing between the tips of the latero-frontal cilia on both sides of the channel, this result cannot be explained by mechanisms based solely on direct mechanical contact between cilia and particles but must involve fluid mechanical mechanisms. The present work is the experimental basis for ongoing numerical simulations of the particle motion in the interpinnule channel.     

Morphology and orientation of tube extensions on aggregations of the polychaete annelid Phragmatopoma californica

The morphology of organisms can influence fluid flow near their surfaces which, in turn, can influence the rates of mass exchange as well as the ability of the animals to capture and retain particles. In this paper, the morphologies and orientations of extensions on the tubes of aggregated individuals of the sabellariid polychaete Phragmatopoma californica from different habitats are described. The effects of these structures on fluid exchange and feeding-tentacle deflection are also assessed. The tube shape of aggregated P. californica varies among habitats. In exposed habitats, extensions on tubes (flares) encircle the entire circumference of the worms' apertures. In more protected habitats, extensions (hoods) partially encircle the circumference of apertures and are oriented so that the opening of the hood faces into the predominant direction of water flow over the aggregation. After the surface structures are damaged, hoods are repaired more quickly than flares. In this study, all damaged hodds were repaired within 24 h while flares were fully repaired only after 240 h. The repaired hoods are oriented in the same direction as the original structures Both hoods and flares decrease the rate of fluid exchange between the aggregation surface and the mainstream water flow. These structures also decrease the deflection of the feeding tentacles by the water flow. The orientation of the hoods relative to the direction of water flow affects both the rates of fluid exchange and the deflection of the tentacles. Present address: Hawaii Institute of Marine Biology, P.O. Box 1346, Kaneohe, Hawaii 96744, USA     

Effects of particle concentration on clearance rate and feeding current velocity in the marine bryozoan Flustrellidra hispida

Colonies of the marine bryozoan Flustrellidra hispida (Fabricius) were collected intertidally from Port St Mary, Isle of Man (UK) during the autumn and winter 1981/1982. Suspension feeding in F. hispida was examined in some detail, using as food the alge Tetraselmis suecica. Previous studies had not examined variation in the speed of suspended food particles travelling through the lophophore. It is clear that feeding rate, as measured by particle velocity, varies over periods of time, even when all external factors are kept constant. Moreover, particle speed within the lophophore can be shown to be positively correlated with the concentration of suspended food particles in the surrounding environment. It is also apparent that feeding current velocity varies at different positions within the lophophore. In addition observations are made on the involvement of the tentacles in particle capture.     

Suspension feeding in the brittle-star Ophiothrix fragilis : efficiency of particle retention and implications for the use of encounter-rate models

Dense beds of the suspension-feeding brittle-star Ophiothrix fragilis are common in European waters. Their potential importance in benthic–pelagic coupling has been highlighted, but little is known about the feeding dynamics of this species. Encounter-rate models provide a potential mechanism for the estimation of feeding rates on suspended material of varying sizes. This work investigates factors essential to the application of such models. Particle-retention efficiency (RE) converts encounter rate into capture, or clearance rate. Laboratory studies demonstrated that RE varied with the interactive effects of flow velocity and particle size. RE was lowest for large particles, particularly at high flow velocity where RE as low as 59% was observed. This indicates that if RE is not accounted for in encounter-rate models, significant overestimates of feeding rates on large particles may occur. Flow around feeding arms and tube feet was characterised by intermediate Reynolds numbers, precluding application of the most simple encounter-rate models. Complex secondary-flow patterns were observed, which carried particles along the downstream side of the feeding arms, but these did not appear to increase the area available for particle capture. Previously reported particle capture by arm spines was not observed. Evidence of active rejection of large particles by tube feet was recorded. Difficulties in the application of encounter-rate models for prediction of seston-removal rates are highlighted by these results. Predicted encounter rate may deviate from actual clearance rate due to the effects of retention dynamics, localised flow patterns and differential particle handling. Other methods of estimation of seston-removal rates are equally problematic however, so that encounter-rate models are likely to remain a useful tool for such estimates. Received: 23 January 1998 / Accepted: 24 June 1998     

Effects of exposure to petroleum hydrocarbons on the gill functions and ciliary activities of a marine bivalve

The effects of long-term exposure to low levels of water-accommodated fractions of Kuwait Crude oil, or to shortterm exposure to chemically dispersed oil, on the gill performance of the clam Venus verrucosa were investigated. Reduced pumping activities of the lateral cilia as well as interference with the normal beating activities of the eulaterofrontal cirri led to reduced clearance rates and retention efficiencies of food particles less than 6 m in diameter. On the other hand, frontal ciliary activities were significantly accelerated, while any retained oil droplets were conducted to the mouth region as food particles. The activities of terminal and sensory cilia were also enhanced and mucus production increased. The significance of these responses to the clam's energy budget is discussed.     

Acquisition of particle processing capability in juvenile oyster <Emphasis Type="Italic">Crassostrea gigas</Emphasis>: ontogeny of gill mucocytes

Acquisition of particle processing capability in postlarval oysters depends upon the structural development of the pallial organs, as well as the development of cilia and mucocytes used (either directly or indirectly) in particle capture and transport. Mucocyte mapping was therefore used to identify mucocyte types and distributions throughout gill development in juvenile oyster Crassostrea gigas (Thunberg 1793) specimens from 2.9 mm to 2.4 cm in shell length. Three categories of gill filaments were identified: apical, lateral and principal filaments, corresponding to filament location or future location in gill plicae. Mucocyte densities were recorded per linear μm (l μm) of frontal surface, and converted to potential total volumes, using the mean volumes of each of the two major mucocyte types: acid mucopolysaccharide (AMPS)-mucocytes and mixed mucopolysaccharide (MMPS)-mucocytes. While AMPS secretions were dominant up to 1.0 cm (flat homorhabdic gill, to semi-heterorhabdic differentiation and plication), MMPS secretions increased progressively, dominating in 2.4 cm and adult specimens (fully heterorhabdic and plicated). Mucus composition, and hence mucus viscosity, thus appears to evolve in relation to the degree of enclosure of the gill frontal surfaces. Total (AMPS + MMPS) potential mucus secretion increased allometrically with juvenile growth, characterized by a sharp increase between 10 and 24 mm shell length, suggesting a marked improvement in particle processing capability. Mucocyte distributions on the gill were heterogeneous from the onset of heterorhabdic differentiation (7.5 mm): the apical filaments of the plicae contained much greater mucocyte total volumes, compared to the lateral and principal filaments. In addition to mucus composition, total potential mucus volume thus also evolved in relation to the degree of enclosure of the gill frontal surfaces. These results show that functional specialization in mucocyte distribution precedes the complete anatomical heterorhabdic differentiation. The completely functional adult gill system is thus attained in 2.4 cm juveniles. This information should be of use in understanding the dynamics of juvenile feeding, growth, and mortality, both in natural systems and in rearing operations.     

Ammonium and free amino acid uptake by a deep-sea mussel (Bathymodiolus sp., undescribed) containing methanotrophic bacterial symbionts

The morphology of the lophophore and mouth was measured for freshly collected colonies of seven common bryozoan reef species at three sites along the central Caribbean coast of Panamá during the dry season, from the end of December 1988 through May 1989. Characters measured or scored include the number of tentacles, circumference of the lophophore, abanal and anal tentacle length, abanal and anal tentacle spacing, and perimeter of the mouth. Taxa studied include three species each of Stylophoma Levinsen, and Steginoporella (Smitt), and one species of Trematooecia Osburn. Individuals were collected from both continental and offshore fringing reefs from depths of 1 to>30 m. Terrestrial runoff from heavy tropical rains, sedimentation, and sea exposure vary greatly between sites. Congeneric species showed small but consistent differences in lophophore size, but little or no difference in spacing of tentacles or perimeter of the mouth. This suggests that closely related species feed on similarly sized particles despite their overall differences in zooidal size. Lopophore morphology also varied intraspecifically between reefs, particularly in the length of the tentacles and spacing between tentacles. Patterns of variation were similar for all species, which implies that species shift their diets in similar directions between the different reefs. The combination of these patterns suggests that niche diversification in feeding does not occur among congeneric species in the cheilostomes examined.     

Dietary study of the midwater polychaete Poeobius meseres in Monterey Bay,California

This study presents the first quantification of the diet of a gelatinous midwater organism on a temporal basis. Using the Monterey Bay Aquarium Research Institute's remotely operated vehicle Ventana, regular collections of the polychaete Poeobius meseres (Heath, 1930) over a 1 yr period (October 1990 to November 1991) in Monterey Bay yielded intact organisms for the study of feeding behavior and quantitative analysis of stomach contents. In situ observations showed P. meseres feeding in two different ways: (1) by deployment of a mucus web in the water column that passively collects particles for consumption; and/or (2) by grasping detrital material in the water column with its ciliated tentacles. Stomach-content analyses showed that P. meseres is primarily coprophagic, its diet being dominated by fecal pellets from euphausiids and copepods. These fecal pellets appear to provide P. meseres with essentially all its carbon. Although fecal pellets were the most important food item volumetrically, P. meseres also consumed large numbers of diatoms and small numbers of dinoflagellates, chrysophytes, radiolarians, foraminiferans and eggs. The diet of P. meseres appears to reflect primary productivity in the surface waters, with different food items predominant in the diet at different times of the year. Pennate diatoms were most abundant in the diet during the fall, centric diatoms were most abundant during the sumnier, and fecal pellets during the winter. The composition of P. meseres diet suggests that this and other midwater gelatinous organisms have a significant role in the remineralization of particles as they sink from the surface to the deep sea.     

Fluid motion and particle retention in the gill of Mytilus edulis: Video recordings and numerical modelling

Particle trajectories of 6.4m Latex spheres were recorded by video, both near an isolated blue mussel, Mytilus edulis, gill filament and, in place of an intact interfilamentary canal, in a model canal of width 200, 100 or 70m, formed by a transparent plate positioned next to a gill filament. Each arrangement was placed in a 2 x 10 x 10 cm test vessel filled with seawater. Serotonin (nerve-transmitter) stimulation was used to activate lateral cilia and to either lock latero-frontal cirri at the end of an active stroke (10^-5 M), or to activate them (10^-6 M), yielding lateral cilia beat frequencies of 19 and 16 Hz, respectively. With latero-frontal cirri locked, image analysis of particle tracks gave maximum velocities of ca. 2.9±0.2mm s^-1 close to the tips of lateral cilia, for both isolated filament and model canal cases. Experimental velocity profiles along the 200-m wide model canal were recorded and used as good approximations to the fluid velocity because of the low Reynolds number. A two-dimensional steady model was proposed for the gill pump, assumed to only comprise lateral cilia. This model was solved numerically for the experimental model, canal in the vessel and the results showed satisfactory agreement with experimental volocity profiles from particle tracks. The numerical approach was also applied to a model of a single interfilamentary canal in the vessel. The resulting mean velocity in the canal was 1.70 mm s^-1, but the resistance to flow in the model was less than that in an intact mussel gill. Video graphs of particle tracks indicated that active latero-frontal cirri play a role in the transfer of particles from through current to frontal current, probably by means of a strong interaction through the motion of intervening fluid rather than through a direct physical contact. M. edulis specimens used in the present study were collected in 1990 at Helsingør and in 1991 at Kerteminde, Denmark.