Filter feeding mechanism in the phoronid <Emphasis Type="Italic">Phoronopsis harmeri</Emphasis> (Phoronida,Lophophorata)

Phoronids, like other Lophophorata (Bryozoa and Brachiopoda) are filter feeders. The lophophore performs various functions, the most important of which is the collection and sorting of food particles. The mechanism of sorting has been well studied for many other groups of invertebrate, but until now it has remained obscure for phoronids. With the help of functional morphology data we are proposing a possible scheme of sorting in phoronids on the example of Phoronopsis harmeri. The lower limit of the particle size is defined by the distance between laterofrontal cilia of tentacles and equals 1.2 μm. Larger particles are transferred by frontal cilia to the basis of the tentacles, where they pass into the lophophoral groove. The distance between the epistome and the external row of tentacles regulates the upper limit of the particle size that are suitable for food. Only particles whose size does not exceed 12 μm get into the lophophoral groove and further into the mouth. Larger particles collect in the space above the epistome and are removed from the lophophore. The size of the food particles that phoronids consume by filtration lies in a range 1.2–12 μm. These are bacteria and small phytoplankton organisms. At the same time the significant individual mobility of the phoronid tentacles plays an important role in the expansion of the pabular spectrum to large inactive zooplankton and phytoplankton organisms reaching a size of 50–100 μm.     

The feeding behaviour of a deep-sea holothurian, Stichopus tremulus (Gunnerus) based on in situ observations and experiments using a Remotely Operated Vehicle

Using a Remotely Operated Vehicle (ROV) to deploy an in situ cage experiment incorporating fluorescent Luminophore particle tracers, the gut throughput time of the deposit feeding holothurian, Stichopus tremulus (Gunnerus) was determined as 23.73 h (S.D.±2.3). For a range of individuals examined at different depths (350-500 m) and locations, throughput times varied between 19 and 26 h irrespective of animal size or gut tract length. In situ video observations of feeding behaviour showed that this species uses fine oral papillae in a ‘sweeping’ motion to target particles on the seafloor. Following detection of a food source fine-branched digitate tentacles collect a large range of sediment fragments from the seabed. The main types of particles ingested include silica fragments (<20 >500 μm), pelagic foraminifera, benthic foraminifera, fine phytodetrital remains and occasional larger rock fragments (∼1 cm). Ingested sediment consisted mainly of very fine silica fragments (∼50 μm) accounting for over 50% of the total gut contents. Frame-by-frame video analysis revealed that the particle handling time (i.e. the time taken for a tentacle insertion and the subsequent collection of food) was found to be ∼54 s. Only 10 of the 20 feeding tentacles were simultaneously employed during feeding. Use of tentacles appeared to be in sequence, alternating between the reserve and active tentacles. Estimating the rate of movement over the seabed and the total effective capture area of each tentacle, the impact of this animal on the turnover and quality of surface sediment at this deepwater site is potentially substantial. The in situ experiments provided a significant improvement over previous methods used to investigate deep-sea deposit feeders and represent a useful concept for further in situ deep-sea research using an industrial ROV.     

Ciliary feeding structures and particle capture mechanism in the freshwater bryozoan Plumatella repens (Phylactolaemata)

Abstract. In contrast to marine bryozoans, the lophophore structure and the ciliary filter‐feeding mechanism in freshwater bryozoans have so far been only poorly described. Specimens of the phylactolaemate bryozoan Plumatella repens were studied to clarify the tentacular ciliary structures and the particle capture mechanism. Scanning electron microscopy revealed that the tentacles of the lophophore have a frontal band of densely packed cilia, and on each side a zigzag row of laterofrontal cilia and a band of lateral cilia. Phalloidin‐linked fluorescent dye showed no sign of muscular tissue within the tentacles. Video microscopy was used to describe basic characteristics of particle capture. Suspended particles in the incoming water flow, set up by the lateral ‘pump’ cilia on the tentacles, approach the tentacles with a velocity of 1–2 mm s^‐1. Near the tentacles, the particles are stopped by the stiff sensory laterofrontal cilia acting as a mechanical sieve, as previously seen in marine bryozoans. The particle capture mechanism suggested is based on the assumed ability of the sensory stiff laterofrontal cilia to be triggered by the deflection caused by the drag force of the through‐flowing water on a captured food particle. Thus, when a particle is stopped by the laterofrontal cilia, the otherwise stiff cilia are presumably triggered to make an inward flick which brings the restrained particle back into the downward directed main current, possibly to be captured again further down in the lophophore before being carried to the mouth via the food groove. No tentacle flicks and no transport of captured particles on the frontal side of the tentacles were observed. The velocity of the metachronal wave of the water‐pumping lateral cilia was measured to be ～0.2 mm s^‐1, the wavelength was ～7 μm, and hence the ciliary beat frequency estimated to be ～30 Hz (～20 °C). The filter feeding process in P. repens reported here resembles the ciliary sieving process described for marine bryozoans in recent years, although no tentacle flicks were observed in P. repens. The phylogenetic position of the phylactolaemates is discussed in the light of these findings.     

Numerical simulation of flow fields and particle trajectories in ciliary suspension feeding

A model describing the ciliary driven flow and motion of suspended particles in downstream suspension feeders is developed. The quasi-steady Stokes equations for creeping flow are solved numerically in an unbounded fluid domain around cylindrical bodies using a boundary integral formulation. The time-dependent flow is approximated with a continuous sequence of steady state creeping flow fields, where metachronously beating ciliary bands are modelled by linear combinations of singularity solutions to the Stokes equations. Generally, the computed flow fields can be divided into an unsteady region close to the driving ciliary bands and a steady region covering the remaining fluid domain. The size of the unsteady region appears to be comparable to the metachronal wavelength of the ciliary band. A systematic investigation is performed of trajectories of infinitely small (fluid) particles in the simulated unsteady ciliary driven flow. A fraction of particles appear to follow trajectories, that resemble experimentally observed particle capture events in the downstream feeding system of the polycheate Sabella penicillus, indicating that particles can be captured by ciliary systems without mechanical contact between particle and cilia. A local capture efficiency is defined and its value computed for various values of beat frequencies and other parameters. The results indicate that the simulated particle capture process is most effective when the flow field oscillates within timescales comparable to transit timescales of suspended particles passing the unsteady region near the ciliary bands. However, the computed retention efficiencies are found to be much lower than those obtained experimentally.     

Rejection Mechanism of Plectolophous Lophophore of Brachiopod <Emphasis Type="Italic">Coptothyris grayi</Emphasis> (Terebratulida,Rhynchonelliformea)

Brachiopoda is a relict group of invertebrate filter feeders that used a tentacle organ, lophophore, for capturing food particles from the water column. Brachiopod extinction apparently occurred due to low productivity of their filtering organ in comparison with more advanced filter-feeders. Investigation of the filtering mechanism of modern brachiopods is essential to understanding their evolutionary fate. This study is devoted to the rejection mechanism of large waste particles from the plectolophous lophophore of brachiopod Coptothyris grayi. The waste particles gather inside of the lophophore on the outer side of the brachial fold. The particles form rows along frontal grooves of outer tentacles and are carried successively to the tentacle tips and move along them, slimed by mucus. One portion of the particles comes off the lophophore and falls down the mantle, while another part is carried to the abfrontal surface of the tentacles. Due to repeated reversals of abfrontal cilia, the particles wavily move along the abfrontal surface of tentacles. Such movement contributes to the secretion of mucus and the formation of particle clots. The clots come off the lophophore and fall down the mantle. The particles are transported along the mantle by cilia to the anterior part of the mantle margin. Here the ciliary reversals that facilitate secretion of mucus and formation of pseudofeces also take place. The latter takes away from the mantle cavity. Thus, only outer tentacles participate in the rejection of large waste particles from the lophophore. Ciliary reversals of the abfrontal surface of tentacles and the mantle are discovered in brachiopods for the first time. This facilitates the additional secretion of mucus and formation of pseudofeces, easing their exit from the mantle cavity. The results contribute to the knowledge of lophophore function and evolution of tentacle organs in Bilateria.     

Feeding mechanisms and feeding strategies of Atlantic reef corals

The feeding behaviour of 35 species of Atlantic reef corals was examined in the laboratory and in the field. Observations were made during the day and at night, using freshly hatched brine shrimp nauplii and finely ground, filtered fresh fish as food sources. Three feeding strategies were observed: Group I–feeding by tentacle capture only; Group II–feeding by entanglement with a mucus net or mucus filaments; Group III–feeding by a combination of tentacle capture and mucus filament entanglement. Group I included corals of the families Poritidae and Pocilloporidae which were normally expanded during both day and night. Group II included corals of the family Agaricidae which were normally expanded at night and contracted during the day. Group III included corals of the other families examined which, with the exception of Dendrogyra cylindrus , were normally expanded only at night.
Feeding responses were elicited by both chemical and tactile stimuli. A preparatory feeding posture was assumed in response to chemical stimuli and consisted of horizontal positioning of the tentacles, elevation of the oral disk to form a cone-like mouth, a wide mouth opening and secretion of mucus by the epidermis of the oral disk. Following the assumption of the preparatory feeding posture, food capture and ingestive movements were elicited by tactile stimuli. However, food capture and ingestive movements were also elicited by chemical stimuli alone in those species which were normally contracted during the day.
While expanded corals captured food with their tentacles or with mucus filaments, contracted corals were able to feed by capturing fine particulate matter with mucus filaments only and thus acted as suspension feeders. By a combination of feeding strategies, reef corals were able to feed both day and night and a wide range of potential food ranging from fine particulate matter to large zooplankton was available to them.     

The sorting of food particles by Abra sp. (bivalvia: tellinacea)

The selection of food particles by Abra tenuis (Montagu), A. alba (Wood) and A. nitida (Müller) has been investigated.Material for ingestion smaller than 30μm is not selected according to size by A. tenius. Particles smaller than 0.5 μm are retained by the palliai organs and ingested but only particles larger than 1 μm appear to be retained with an efficiency approaching 100 %. The mesh size of the gill filter is found to be ≈ 3.0 × 0.5 μm.A. tenuis does not appear able to discriminate between particles smaller than 20 μm by their food value; however, relatively large silica particles which are devoid of food are partially rejected by the labial palps in favour of particles of similar size but having a bacterial coating. To a lesser extent the physical nature of particles seems to influence their selection by A. tenuis; clean angular particles are rejected in favour of clean rounded ones.Small, light particles appear to be transported on the gills directly to the mouth without coming into contact with the palps. Larger, heavier particles tend to drop from the gill to be caught by the palps which extend posteriorly to cup the entire ventral margin of the inner demibranch when the animal is feeding.The material ingested by A. alba is significantly finer than that taken into the mantle cavity indicating that the pallial organs actively select food by size. Selection of material for ingestion by size in A. nitida appears to be optional since only some of the animals examined had stomach contents significantly finer than material from the mantle cavity.     

Benthic suspension feeders: their paramount role in littoral marine food webs

In recent years, particular attention has been paid to coupling and energy transfer between benthos and plankton. Because of their abundance, certain benthic suspension feeders have been shown to have a major impact in marine ecosystems. They capture large quantities of particles and might directly regulate primary production and indirectly regulate secondary production in littoral food chains. Suspension feeders develop dense, three-dimensional communities whose structural complexity depends on flow speed. It has been postulated that these communities can self-organize to enhance food capture and thus establish boundary systems capable of successfully exploiting a less structured system, namely, the plankton.     

Organic coatings and ontogenetic particle selection in Streblospio benedicti Webster (Spionidae: Polychaeta)

Surface deposit feeders select food particles based upon characteristics including size, texture, specific gravity, and organic coatings. Spionid polychaetes feed at the sediment-water interface using a pair of ciliated palps and switch between surface deposit feeding and suspension feeding primarily as a function of water flow. Juveniles and adults of some spionid species have different stable isotopic carbon signals, indicating the ingestion of different food sources and potentially the ability to differentiate organic cues ontogenetically. In the present study, the feeding responses of juvenile and adult Streblospio benedicti Webster to seven organic coatings bound to glass microbeads were tested using five amino acids and two carbohydrates. Coated versus uncoated particles were presented in equal proportions based upon surface area. Juveniles and adults were highly selective for all seven types of organically coated beads—87.1% of all beads ingested were organically coated beads. For two of the organic coatings, there were ontogenetic differences; juveniles were more selective of threonine and adults were more selective of proline. These differences may result in ontogenetic diet shifts that allow maximization of energy and/or essential nutrients during critical early life-history stages. Particle selection in tentaculate surface deposit feeders is generally thought to occur primarily during particle contact and transport to the mouth, and is typically characterized as a passive process. Active particle selection at the site of the everted pharynx was observed and quantified for S. benedicti. Organically coated particles represented 50% of the ambient experimental treatment, 64.4% of the particles transported along the palp after contact, and 81.8% of the particles ingested after pharyngeal rejection behavior. Of the beads reaching the pharynx, 26.9% were rejected by ciliary sorting on the pharynx before ingestion, and 81.8% of the rejected beads at the site of the pharynx were uncoated. Our study demonstrates that microphagous feeders that generally handle food particles in bulk are capable of significant levels of active selection for organically coated particles.     

Time and extent of ciliary response to particles in a non-filtering feeding mechanism

Mechanisms of suspension feeding are usually described by the physics of inanimate filters. High-speed videorecordings in this study demonstrated that sea urchin larvae concentrate particles without filtration. They actively captured individual particles. At most times and places, the effective strokes of the swimming/feeding ciliary band were away from the circumoral field. Cilia of this band responded to particles by a reversal of beat that redirected the particle toward the circumoral field. A change of beat occurred along approximately 80 micro m of ciliary band during particle capture. Cilia responded 0.02 to 0.06 s after the particle was within reach of effective strokes and reversed beat, usually for about 0.1 to 0.2 s. The whole event (disruption of forward beat) generally lasted between 0.13 and 0.5 s. These observations imply reversed movement of a parcel of water much larger than the included captured particle, but particles are nevertheless greatly concentrated because water is directed toward the circumoral field only when and where a particle is sensed. Thus most of the concentration of particles occurs by a temporarily and locally redirected current, without filtration, and size and quality of particles captured depends on sensory capabilities, not the mechanics of filtration.     

Are Antarctic suspension-feeding communities different from those elsewhere in the world?

This paper reviews the trophic ecology of benthic suspension feeders in Antarctic shelf communities, studied within SCAR's EASIZ Programme, in comparison with published information from other seas. Dense benthic suspension-feeder communities capture large quantities of particles and may directly regulate primary, and indirectly, secondary production in littoral food chains. Most work has been performed in temperate and tropical seas; however, little is known about suspension feeders in cold environments. Recent studies on Antarctic littoral benthic suspension feeders suggest the period of winter inactivity may last only a few weeks. This contrasts with the hypothesis that in Antarctic communities there is a prolonged period of minimal activity lasting at least 6 months during the austral winter. Results from other oceans may explain how dense benthic communities could develop under such conditions. Alternative food sources, i.e. the "fine fraction", sediment resuspension, lateral advection and efficient food assimilation may play a significant role in the development of suspension-feeder dominated, very diversified, high biomass and three-dimensionally structured communities on the Antarctic shelf.     

The effect of size of polystyrene particles on their retention within the rat peritoneal compartment,and on their interaction with rat peritoneal macrophages in vitro

Polystyrene particles (size range 300nm-3μm diameter) were radioiodinated and their capture by rat peritoneal macrophages measured in vitro. For unmodified particles, most efficient accumulation was observed using a diameter of 600nm (Endocytic Index (E.I.) = 16.4 ± 2.9μl/10⁶cells/h). Particles (3μm diameter) which had been modified to become more hydrophilic by hydroxymethylation showed an increased rate of capture (E.I. = 136.6 ± 91.2μl/10⁶cells/h). Following intraperitoneal administration to rats, unmodified 3μm particles showed selective accumulation in the omentum (18.4% injected dose/g), and this was increased for the hydroxymethylated bead (35.3% dose/g). The smaller (800 nm) particles were better able to leave the peritoneal compartment. Radiolabelled particles isolated from a peritoneal wash after 5h were mostly cell-associated (72–86%, depending on the type of particle).     

Feeding behavior in freshwater bryozoans: function,form, and flow

Bryozoans are impressively active suspension feeders, with diverse feeding behaviors. These have been studied extensively in marine bryozoans, but less so in their freshwater counterparts. Here we identified 16 distinct behaviors in three phylactolaemate species and classified them into behaviors involving separate tentacles, groups of tentacles, lophophore arms, the introvert, or multiple zooids. We examined (1) the repertoire of behaviors in each species, and each behavior's (2) absolute frequency, (3) relative frequency and (4) duration in each of the three species, at two flow velocities (0 and 0.2 cm s^?1). Nine feeding behaviors were shared by all three species, but the occurrence of other behaviors in a given species was limited by its morphology. Behaviors involved in particle capture were the most frequent, and were often faster than the reactions involved in particle rejection. By contrast, the absolute frequency of behaviors varied widely among species without clear associations with species form, or function of the behavior. Flow velocity had only minor effects on the feeding behaviors exhibited by a species, or their frequencies or durations. Our results show that phylactolaemates have the same key feeding behaviors of the individual polypides (especially involving separate tentacles) as previously described in gymnolaemate and stenolaemate bryozoans, although their behaviors tend to be carried out more slowly than those of stenolaemates or gymnolaemates. Feeding behaviors involving multiple zooids were nearly absent in the studied phylactolaemates, but are common in gymnolaemates. Freshwater bryozoans appear to be intermediate between stenolaemate and gymnolaemate bryozoans in terms of richness of the repertoire of feeding behaviors.     

Particle retention in suspension-feeding fish after removal of filtration structures

The suspension-feeding cichlids Oreochromis aureus (blue tilapia) and Oreochromis esculentus (ngege tilapia) are able to selectively retain small food particles. The gill rakers and microbranchiospines of these species have been assumed to function as filters. However, surgical removal of these oral structures, which also removed associated mucus, did not significantly affect the total number of 11–200 μm particles ingested by the fish. This result supports the hypothesis that the branchial arch surfaces themselves play an important role in crossflow filtration. Both species selectively retained microspheres greater than 50 μm with gill rakers and microbranchiospines intact as well as removed, demonstrating that neither these structures nor mucus are necessary for size selectivity to occur during biological crossflow filtration. After removal of the gill rakers and microbranchiospines, O. esculentus retained significantly more microspheres 51–70 μm in diameter and fewer 91–130 μm microspheres compared to retention with intact structures, but the particle size selectivity of O. aureus was not affected significantly. These results support conclusions from previous computational fluid dynamics simulations indicating that particle size can have marked effects on particle trajectory and retention inside the fish oropharyngeal cavity during crossflow filtration. The substantial inter-individual variability in particle retention by suspension-feeding fish is an unexplored area of research with the potential to increase our understanding of the factors influencing particle retention during biological filtration.     

Novel synthesis of ZnO by Ice-cube method for photo-inactivation of E. coli

The ZnO particle with varieties of morphology was prepared from ice-cube of zinc ammonium complex at boiling water surface in 1 min induction of thermal shock. The zinc ammonium complex in ice cube was developed using zinc acetate and biologically activated ammonia in 1 hr and kept in the freezer. Temperature gradient behaviour of the water medium during thermal shock was captured by the thermal camera and thermometer. Morphology study revealed a variety of flower-like ZnO particles with variable size from 1.0 to 2.5 μm. Further, ZnO particle morphologies were tuned by adding trisodium citrate and hexamine to obtain uniform spherical (2–3 μm) and flower (3–4 μm) shapes, respectively. XRD patterns revealed that all ZnO samples are of a hexagonal structure. Photocatalytic inactivation of E. coli has been investigated using various particle morphologies of ZnO in an aqueous solution/overcoated glass slide under sunlight. The photo-inactivation of E. coli by ZnO particles in suspension condition was better when compared to a coated glass slide method. AFM study confirmed the destruction of bacterial cell wall membrane by the photocatalytic effect. The particles morphology of photocatalyst is well dependent on antibacterial activity under sunlight.     

Suspension feeding in ciliated protozoa: Feeding rates and their ecological significance

The quantitative uptake of suspended particles has been studied in 14 species of ciliated protozoa in terms of the maximum rate of water cleared at low particle concentrations and of the maximum ingestion rate at high particle concentrations. The results, supported by data from the literature, show that ciliates which feed on larger particles (> 1–5m) compare favorably with metazoan suspension feeders with respect to the ability to concentrate dilute suspensions of particles. Species specialized on smaller food particles (0.2–1m), the size range of most bacteria in natural environments, require a higher concentration of particles. Bacterial population densities which can sustain ciliate growth are found in sediments, waters rich in organic material, and in the early successional stages of decomposing organic material. This is not the case in open waters in general where bacterivorous ciliates cannot play a role as grazers of bacteria.     

Computational fluid dynamics in the oral cavity of ram suspension-feeding fishes

We have modeled steady, three-dimensional flow with a no-slip boundary condition in cylindrical and conical oral cavities possessing vertical or slanted branchial slits. These numerical simulations illustrate the transport of food particles toward the esophagus, as well as the velocity profiles of water exiting the oral cavity via the branchial slits. The maximum and average velocities are highest for flow exiting the most posterior branchial slit. The highest volume flow rates also occur in the most posterior slit for the cylindrical simulations, but occur in the most anterior slit for the conical simulations. Along the midline, there is a pronounced bilaterally symmetrical vortex in the posterodorsal region of the cylindrical and conical oral cavities and a second bilaterally symmetrical vortex in the posteroventral region of the cylinder. Particles entrained in the vortices will recirculate in the posterior oral cavity, increasing the probability of encounter with sticky, mucus-covered surfaces such as the oral roof, gill arches, or gill rakers. The posterodorsal vortex could serve to concentrate particles near the entrances of the epibranchial organs. The ventral vortex could be involved in sequestering dense inorganic particles that sink toward the floor of the oral cavity. All vortices are absent in the conical simulation with vertical branchial slits, indicating that the slanted branchial slits between the gill arches are responsible for the formation of the vortex in the conical oral cavity. Experiments using in vivo flow visualization techniques are needed to determine whether ram suspension feeders, pump suspension feeders, and non-suspension-feeding fishes possess vortices in the posterior oral cavity that contribute to particle transport, food particle encounter with sticky surfaces, and inorganic particle rejection.     

Plasticity of the scleractinian body plan: Functional morphology and trophic specialization ofMycedium elephantotus (Pallas, 1766)

Summary Morphological, histological and behavioral features indicate thatMycedium elephantotus, a zooxanthellate scleractinian species without tentacles, is well adapted for utilizing suspended organic matter for nutrition. The colonies are composed of vertically growing fan-like plates and can reach diameters of more than 1 m in depths below 20 m. The external body surface is coated with a mucus layer (cuticle) which enables the acquisition and accumulation of suspended organic material. The mucus-entangled particles pass to the mouth openings by gravitational transport assisted by water movement. In experiments the corals were able to discriminate between suspended food and mineral particles. Both types of particles were rapidly entangled in fine mucus nets or filaments. Mineral particles were never ingested and instead tumbled down the inclined skeletal plates. In contrast, food particles were actively incorporated when the mucus filaments accidentally touched the stomodaea during the downward gliding. The food-enriched mucus filaments were either transported by ciliary activity into the coelenteron or were sucked into the body cavities by decreasing pressure in the coelenteron caused by contraction of longitudinal, mesenterial muscles. The discriminative reactions to mineral or food particles are probably based on the release of different types of mucus. Nematocysts are infrequent in the oral epidermis, indicating that the capture of living prey plays a subordinate role in nutrition. The mesenterial filaments, in contrast, are densely packed with large nematocysts. Storage products were piled up within the tissues of gastral pockets. The adaptations ofMycedium elephantotus for using suspended food particles may explain the particularly high abundance of this species between ca. 20 and 40 m depth on a steeply inclined fore-reef slope in the Gulf of Aqaba (Red Sea). The evidence indicating the importance of heterotrophic fueling toM. elephantotus is supported by carbonate production rates which are, in contrast to that of many other zooxanthellate scleractinian species, almost constant at depths between 5 and 40 m and which are uneffected by varying light regimes over the year, suggesting that the reduced phototrophic contribution by the zooxanthellae is compensated by mucus suspension feeding.     

A trophic framework for animal origins

Metazoans emerged in a microbial world and play a unique role in the biosphere as the only complex multicellular eukaryotes capable of phagocytosis. While the bodyplan and feeding mode of the last common metazoan ancestor remain unresolved, the earliest multicellular stem‐metazoans likely subsisted on picoplankton (planktonic microbes 0.2–2 μm in diameter) and dissolved organic matter (DOM), similarly to modern sponges. Once multicellular stem‐metazoans emerged, they conceivably modulated both the local availability of picoplankton, which they preferentially removed from the water column for feeding, and detrital particles 2–100 μm in diameter, which they expelled and deposited into the benthos as waste products. By influencing the availability of these heterotrophic food sources, the earliest multicellular stem‐metazoans would have acted as ecosystem engineers, helping create the ecological conditions under which other metazoans, namely detritivores and non‐sponge suspension feeders incapable of subsisting on picoplankton and DOM, could emerge and diversify. This early style of metazoan feeding, specifically the phagocytosis of small eukaryotic prey, could have also encouraged the evolution of larger, even multicellular, eukaryotic forms less prone to metazoan consumption. Therefore, the first multicellular stem‐metazoans, through their feeding, arguably helped bridge the strictly microbial food webs of the Proterozoic Eon (2.5–0.541 billion years ago) to the more macroscopic, metazoan‐sustaining food webs of the Phanerozoic Eon (0.541–0 billion years ago).     

Food availability and gut contents in the ascidian Cnemidocarpa verrucosa at Potter Cove, Antarctica

A high seasonality characterizes Antarctic environments, and generates marked differences in availability and composition of food for benthic filter feeders. During a year-round period at Potter Cove, Antarctica, algal pigment concentration (chlorophyll a) and organic matter were measured in water column and sediment samples. Chemical composition of gut contents of the ascidian Cnemidocarpa verrucosa was also analyzed. Despite the low standing stock, capture and absorption of organic matter were detected year-round, suggesting intake of other particles besides microalgae. The mechanism that provides food to the ascidians and epibenthic communities may be related to the supply of allochtonous particles, bottom resuspension events, and microbial community dynamics. Sea-ice cycles may affect food availability in terms of water-column mixing and benthic resuspension. The scarce primary production and the high amount of sedimented material are not limiting conditions in Potter Cove, which presents a rich ascidian community.