Phagostimulation of Lytechinus variegatus (Lamarck) (Echinodermata: Echinoidea)

Artificial agarose foods containing either starch, glycogen, carragheenan, alginate, casein, gelatin, albumin, or peptones at a concentration of 1% dry weight were not phagostimulatory. Artificial foods containing sugars, especially galactose, at a concentration of 0.005 M were phagostimulatory. Of the common amino acids, only L‐phenylalanine was phagostimulatory at this concentration. The initiation of feeding postures (movement of the spines, tube feet, peristome, and Aristotle's lantern) of Lytechinus variegatus was little affected by the concentration of plant extracts. Many aspects of this behavior were attributable to physical rather than chemical stimulation. Feeding rates of L. variegatus upon artificial agarose foods containing either Thalassia testudinum or Gracilaria verrucosa were directly dependent upon plant tissue concentration. All artificial agarose food‐items were initially moved to the mouth and fed upon, however the time spent feeding was directly dependent upon plant type and tissue concentration. Retention of the artificial food‐items upon the test after feeding was not influenced by plant type or tissue concentration. Differences in consumption of foods by L. variegatus are determined by the degree of persistence in feeding upon encountered items and not be selection before initiating feeding or by retention of food‐items for subsequent feedings.     

Differential inputs from chemosensory appendages mediate feeding responses to glucose in wild-type and glucose-averse German cockroaches, Blattella germanica

Glucose is a universal phagostimulant in many animal species, including the cockroach Blattella germanica. However, some natural populations of B. germanica have been found that are behaviorally deterred from eating glucose. In dose-response studies, glucose was a powerful phagostimulant for wild-type cockroaches, but it strongly deterred feeding in a glucose-averse strain. Both strains, however, exhibited identical dose-response curves to other phagostimulants and deterrents. As a lead to electrophysiological and molecular genetics studies to investigate the mechanisms that underlie glucose-aversion, we used 2 assay paradigms to delineate which chemosensory appendages on the head contribute to the reception of various phagostimulatory and deterrent chemicals. Both simultaneous dual stimulation of the antenna and mouthparts of the insects and 2-choice preference tests in surgically manipulated insects showed that the glucose-averse behavior could be elicited through the gustatory systems of the antennae and mouthparts. The paraglossae alone were sufficient for maximum sensitivity to both phagostimulants and deterrents, including glucose as a deterrent in the glucose-averse strain. In addition to the paraglossae, the labial palps were more important than the maxillary palps in the reception of deterrents (caffeine in both strains and glucose in the glucose-averse strain). The maxillary palps, on the other hand, played a more important role in the reception of phagostimulants (fructose in both strains and glucose in the wild-type strain). Our results suggest that distinct inputs from the chemosensory system mediate opposite feeding responses to glucose in the wild-type and glucose-averse strains.     

Mechanoresponsive neurones in the suboesophageal ganglion of the locust

Abstract. A preparation is described for intracellular recording from the neur-opile of the sub-oesophageal ganglion (SOG) of the locust, while stimulating the labial and maxillary palps with plant material in such a way as to mimic the palpation behaviour which precedes and continues throughout feeding. Twelve neurones responding to simulated palpation were recorded from and stained in the SOG. Axons of three neurones ascended to the brain, six had descending axons and three had all of their processes confined to the suboesophageal ganglion. The major regions of arborization were in the ventrolateral and mediolateral neuropiles of the maxillary and labial neuromeres. All twelve neurones were solely mechanoresponsive. In addition to responding to palpation of one or more of the four palps, five also responded to stimulation of the labrum, one to touching each antenna, and one to mechanical stimulation of each of the six tarsi. In the context of what is known about the role of mechano-stimulation in the control of feeding, and given their particular patterns of input and arborizations, it is suggested that the neurones may be active during food selection and ingestion.     

The role of the maxillary and labial palps in the feeding behaviour of Schistocerca gregaria

The role of the maxillary and labial palps in the initiation and maintenance of feeding has been analysed in female fifth-instar nymphs of Schistocerca gregaria. The ability to perceive single chemicals prior to feeding and to feed to repletion on them is markedly reduced in the absence of the palps. Increasing the sensory input of suboptimal diets by providing an attractive odour results in an increased meal length in insects without palps but not in controls.Insects without their palps reared on a diet of lettuce and bran eat relatively less bran and more lettuce than do the controls, although the total volume of food ingested is the same. The smaller dry weight of food consumed as a result of this greater lettuce intake results in a significantly smaller dry weight gain over the fifth instar.

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Zusammenfassung Bei Fünftlarven von Schistocerca gregaria wurde analysiert, welche Rolle die Maxillar- und Labialpalpen beim Frassbeginn und während des Fressens spielen. Palpenlose Insekten nahmen einzelne Chemikalien vor dem Fressen weniger wahr und frassen auch weniger als Kontrolltiere. Die sensorische Qualität suboptimalen Futters konnte durch einen attraktiven Geruch verbessert werden, was zu verlängerter Fresszeit bei palpenlosen Insekten, nicht aber bei Kontrolltieren, führte.Palpenlose Insekten, die auf einer Diät von Salat und Kleie gehalten wurden, frassen weniger Kleie und mehr Salat als Kontrolltiere, wobei das Volumen der aufgenommenen Nahrung gleich blieb. Durch den erhöhten Salatkonsum wurde die Trockensubstanzmenge der aufgenommenen Nahrung vermindert. Als Folge davon nahmen die Tiere während des fünften Larvenstadiums gesichert weniger an Trockensubstanz zu als Kontrolltiere.

     

Feeding behavior of the cirratulid Cirriformia filigera (Delle Chiaje, 1825) (Annelida: Polychaeta).

Observations of the feeding behavior of Cirriformia filigera (Delle Chiaje, 1825) (Annelida: Polychaeta) from the intertidal zone of S?o Francisco and Engenho D'água beaches (S?o Sebasti?o, State of S?o Paulo) were made in the laboratory. This species, like other cirratulids, is a deposit feeder, feeding mainly on sediment surface with the aid of its grooved and ciliated palps, which are used to capture food particles. The worm lies just beneath the substrate surface in a J-shaped tube. When feeding, it extends up to 4 palps over the sediment surface, capturing food particles which pass down the groove of each palp directly to the mouth. Only fine sand grains are ingested. The worm frequently extends 4 branchial filaments into the overlying water for aeration. When it moves with the prostomium sideways, it collects and transports sand grains that pass backwards along its ventral region until reaching the middle part of its body. Next, the parapodia and palps move the sand grains to the dorsal posterior end of the animal, covering this area with sand. Some sand grains are also ingested as the worm moves.     

Carbon isotope signatures reveal that diet is related to the relative sizes of the gills and palps in bivalves

In marine bivalves, the relative sizes of the gills and palps appear to be a useful functional trait that reflect feeding mode, i.e. suspension feeders have relatively larger gills than palps for pumping, whereas deposit feeders have relatively larger palps than gills for sorting. Also, within a species, the relative sizes of the gills and palps are related to changes in local food conditions. However, there is still no firm evidence showing that differences in the relative gill and palp sizes between species are related to diet selection. Based on the knowledge that carbon and nitrogen isotope signatures of an animals tissues reflect past diet, we compared the relative gill and palp sizes of bivalves from Roebuck Bay, northwestern Australia with their carbon and nitrogen isotope signatures. The carbon isotope signatures distinguished clear differences in diet between bivalves along a gradient from suspension to deposit feeding, and strikingly this pattern was closely followed by the relative sizes of the gills and palps of the bivalves. This study confirms that relative gill and palp sizes in bivalves are a functional trait that can be used to compare resource use between species. Furthermore, these data may suggest that morphospace occupation, as determined by relative gill and palp sizes of bivalves, could reflect a gradient of resource use between species.     

Evolution of cave suspension feeding in Protodrilidae (Annelida)

Protodrilidae belongs in a lineage that until now entirely consisted of deposit‐feeding, highly adapted interstitial annelids. Except for a pair of anterior palps, all protodrilids lack appendages, parapodia and chaetae; and have slender bodies adapted to glide between the sand grains by ciliary motion. The first exception to these characteristics is Megadrilus pelagicus n. sp. inhabiting the water column of the anchialine La Corona cave system in Lanzarote. Its morphology and evolutionary history are here investigated by combining observations from in vivo video recordings and advanced microscopy with phylogenetic analyses. Our studies revealed a unique pelagic, suspension feeding behaviour attained by its long ciliated palps in combination with an autopomorphic dorsal ciliated keel and several longitudinal and transverse ciliary bands. Phylogenetic analyses recovered Megadrilus pelagicus n. sp. nested within Protodrilidae indicating that its unique traits are derived within the family. These traits are traced in the tree topologies in correlation to cave colonization. The evolution of these traits can be functionally explained by the different demands of a pelagic suspension feeding strategy compared to the ancestral deposit‐feeding guild of the family. The origin of this suspension feeding strategy was presumably favoured by the partial isolation of the anchialine ecosystem, connected to the sea only through the highly porous volcanic subterranean bedrock. This crevicular connection limits the amount of predators and turbulence in the cave, but allows continuous water flow into the system carrying organic particles, which is the main source of food when photosynthetic primary production does not occur and sedimentation is limited. These conditions may select for pelagic suspension feeding as the most feasible life‐strategy in anchialine caves, which the dominance of pelagic, suspension feeding crustaceans and annelids in anchialine cave assemblages may also reflect. For species of ancestrally deposit‐feeding lineages entering the cave system, such as the annelid families Protodrilidae and Nerillidae, an adaptive‐shift from interstitial to crevicular habitats seemingly correlates with dramatic morphological changes and speciation. The dramatic changes observed in these primarily interstitial lineages compared to their relatives, point to alternative adaptive evolutionary pathways related to ecological fitness contrary to the previously proposed theories focusing on geological or stochastic processes.     

Morphology of anterior regeneration in two spionid polychaete species: implications for feeding efficiency

Abstract. In marine soft‐sediment habitats, tissue loss by infuanal invertebrates can provide significant energetic input to higher trophic levels, have substantial impacts on individual behavior, growth, and fecundity, and resulting changes in bioturbation rates can secondarily affect community dynamics. The degree to which a community is affected by such sublethal predation depends in part on whether injured individuals can regenerate and on the speed at which they do so. Previously, we demonstrated differences in the rate of anterior segment and palp regeneration by the spionid polychaetes Pygospio elegans and Dipolydora quadrilobata. The current study examined the morphology of the anterior segment and palp regeneration in these species using scanning electron microscopy (SEM) and indirect immunohistochemistry with confocal laser scanning microscopy at 3, 6, 9, and 12 d post‐ablation. Antibodies for acetylated α‐tubulin and the neural tetrapeptide FMRFamide were used to label the regenerating nervous system. SEM revealed that the morphology of anterior tissue regeneration was similar for both species, but the ciliated food groove tended to form sooner on palps in P. elegans than on those of D. quadrilobata. In both species, palp regeneration and ciliated food groove formation were faster when only palps were removed. A shortened ciliated food groove is likely to reduce particle contact and transport efficiency in suspension and deposit feeding. Regenerating palp nerves were initially visible at 3 d following ablation of palps only, but at 6 d following ablation of five anterior segments. Following ablation of anterior segments, the regenerating nervous system was largely complete by 9 d, nuchal organs were innervated by 6 d, and processes of palp sensory cells were visible at 12 d. Contact chemoreception by sensory cells on the palps may be diminished during the early stages of regeneration, but chemoreception of waterborne cues via the nuchal organ should not.     

Taste receptor activity and feeding behaviour reveal mechanisms of white spruce natural resistance to Eastern spruce budworm Choristoneura fumiferana

The pattern of feeding of Eastern spruce budworm Choristoneura fumiferana (Clem.) (Lepidoptera, Tortricidae) is compared on foliage from white spruce Picea glauca (Moench) Voss. (Pinaceae) trees previously determined to be susceptible and resistant to defoliation by budworm. No differences are observed in electrophysiological responses from taste sensilla to aqueous extracts of the two foliage types, nor is there a preference for either extract type in a choice test. Acetone extracts from the two foliage types are both preferred to a control sucrose solution, although neither elicits a preference relative to the other. These results suggest that there is no difference in phagostimulatory power of internal leaf contents of the two foliage types. Longer‐term observation of feeding behaviour in a no‐choice situation shows no difference in meal duration, confirming the lack of difference in phagostimulatory power. However, on average, intermeal intervals are twice as long on the resistant foliage, leading to an overall lower food consumption during the assay. This result suggests an anti‐digestive or toxic effect of the resistant foliage that slows behaviour and limits food intake. Previous research has shown that waxes of the resistant foliage deter initiation of feeding by the spruce budworm and that this foliage contains higher levels of tannins and monoterpenes. The data suggest that the resistant foliage contains a post‐ingestive second line of defence against the spruce budworm.     

The role of the antennae and maxillary palps in mediating food preference by larvae of the tobacco hornworm, Manduca sexta

The role of antennae and maxillary palps in mediating food preferences by Manduca sexta (Johan.) (Lepidoptera: Sphingidae) was examined. Fifth‐instar larvae, having different chemosensory organs remaining after microsurgery, were tested in two‐choice feeding assays using three solanaceous host species Lycopersicon esculentum (Mill.), Datura innoxia (Mill.), and Solanum pseudocapsicum (L.), an acceptable non‐host Raphanus sativus (L.) (Cruciferae), and an unacceptable non‐host Pelargonium hortorum (L.H. Bailey) (Geraniaceae). Larvae had a choice between leaf discs of two plant species or between one species and moist filter paper discs (water). The antennae are fully competent in mediating normal (unimpaired) food preferences for S. pseudocapsicum vs. water and P. hortorum vs. water. Thus, the antennae alone can mediate both acceptance and rejection behaviour. The latter is the first report of such a function in M. sexta. The antennae are partly competent (reduced preference) for S. pseudocapsicum vs. P. hortorum. No antennal competence could be demonstrated using the other plant species tested. The antennae alone are either partly needed (R. sativus vs. water) or not necessary to elicit normal food preferences for the plant species tested. The maxillary palps are fully competent in mediating normal food preference for S. pseudocapsicum vs. water, and D. innoxia vs. water. The palps are partly competent for S. pseudocapsicum vs. P. hortorum and incompetent for the other plant species tested. Thus, the maxillary palps alone could only mediate acceptance behaviour in this study. The need of the maxillary palps in normal food preferences could not be demonstrated. Interestingly, the maxillary palps alone can mediate food preferences for two normally rejected plants, R. sativus and P. hortorum. Similar results are reported for two other non‐host plant species, Vigna unguiculata (Walp.) (Leguminosae) and Vinca rosea (L.) (Apocynaceae). Perhaps, the maxillary palps inform M. sexta mostly about feeding stimulatory chemicals common to most plant species.     

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.     

Do food protein and carbohydrate content influence the pattern of feeding and the tendency to explore of forest tent caterpillars?

This study examines whether the ratio of protein to carbohydrate affects the timing of meals and the propensity to explore of forest tent caterpillars (Malacosoma disstria). The behavior of fourth instar caterpillars was observed on three semi-defined artificial diets varying in protein (p)-carbohydrate (c) ratio. These diets were (a) p14:c28, (b) p28:c14, and (c) p35:c7. The probability of initiating feeding at first contact with the food and the duration of the first feeding event did not vary across diets, suggesting not much difference in phagostimulatory power. There was also no difference in the total time spent eating, at rest and in motion between diets. However, the timing and duration of meals varied significantly; more short meals were observed on the carbohydrate-biased diet. The duration of pauses between meals also increased with food protein content. Furthermore, caterpillars on the carbohydrate-biased diet were more likely to leave the trail leading to the known food source and to discover a second food source, suggesting that protein deprivation promotes exploration. These findings shed insight into the physiological responses to protein and carbohydrate ingestion and demonstrate how post-ingestive effects can favor consumption of foods containing protein without invoking an explicit mechanism of independent nutrient regulation, but simply by influencing the pattern of feeding and the propensity to explore.     

Observations of serotonin and FMRFamide-like immunoreactivity in palp sensory structures and the anterior nervous system of spionid polychaetes

Evidence suggests that ciliated sensory structures on the feeding palps of spionid polychaetes may function as chemoreceptors to modulate deposit-feeding activity. To investigate the probable sensory nature of these ciliated cells, we used immunohistochemistry, epi-fluorescence, and confocal laser scanning microscopy to label and image sensory cells, nerves, and their organization relative to the anterior central nervous system in several spionid polychaete species. Antibodies directed against acetylated alphatubulin were used to label the nervous system and detail the innervation of palp sensory cells in all species. In addition, the distribution of serotonin (5-HT) and FMRFamide-like immunoreactivity was compared in the spionid polychaetes Dipolydora quadrilobata and Pygospio elegans. The distribution of serotonin immunoreactivity was also examined in the palps of Polydora cornuta and Streblospio benedicti. Serotonin immunoreactivity was concentrated in cells underlying the food groove of the palps, in the palp nerves, and in the cerebral ganglion. FMRFamide-like immunoreactivity was associated with the cerebral ganglia, nuchal organs and palp nerves, and also with the perikarya of ciliated sensory cells on the palps.     

Feeding response of the stable fly, Stomoxys calcitrant L., to blood fractions and adenine nucleotides

Abstract Feeding responses of the stable fly ( Stomoxys calcitrans L., Diptera: Muscidae) to blood fractions and saline-based diets were studied in an artificial feeding device. Flies fed equally on whole blood, plasma and erythrocyte fractions while resuspended platelets or the buffy layer did not stimulate feeding. Evidence from filtration studies indicates that plasma contains both less-than-5000D and greater-than-100,000D phagostimulatory components. ATP, ADP and AMP in saline stimulate feeding in this insect in a dose-dependent manner. cAMP also stimulates feeding in the range of concentrations tested. It is argued that the less-than-5000D phagostimulant may be one or more of these adenine nucleotides.     

A note on the watermite Hydrodroma despiciens feeding on chironomid egg masses

SUMMARY. Chironomid eggs are the primary food of aduh and nymphal Hydrodroma despiciens. Observations on feeding behaviour showed that the palps are well adapted for probing the egg mass jelly. Chironomid egg masses also serve as primary sites for male spermatophore deposition.     

Influence of diet on pre-ingestive particle processing in bivalves: II. Residence time in the pallial cavity and handling time on the labial palps

Previous studies have demonstrated that bivalve molluscs respond to changing food conditions through processes such as preferential selection and ingestion of particulate matter. Little is known, however, about the underlying mechanisms accountable for these responses. To further explain feeding processes at the level of the pallial organs, we determined pallial cavity residence times, or the amount of time it took particles to travel from the inhalant aperture to the stomach, in two species of bivalves, Crassostrea virginica and Mytilus edulis, under conditions of differing particle quality, particle concentration, and temperature. From these residence times, particle-handling times on the labial palps were determined. Diets of three different qualities were tested, including Rhodomonas lens cells, particles prepared from ground Spartina sp. detritus, and a 50/50 mixture of both. Bivalves were delivered one of the three diets along with 10-μm fluorescent polystyrene beads (tracer), removed from feeding chambers at intervals from 30 s up to 20 min, and placed in liquid nitrogen to halt particle transport. Digestive systems of bivalves were then dissected and examined for the presence of tracer beads. Particle-residence times in the pallial cavity and handling times on the labial palps of C. virginica were significantly affected by changes in diet type. Particle-handling times on the palps decreased with increasing diet quality and ranged from 2.2 min (100% R. lens) to 22.8 min (100% ground Spartina sp.), accounting for 88% and 99%, respectively, of the total time particles spent in the pallial cavity. In contrast, diet quality had little effect on particle-residence times in the pallial cavity of M. edulis. However, residence times were affected by temperature and diet concentration. Temperature significantly affected residence times at particle concentrations of both 20 and 100 particles μl⁻¹, whereas particle concentration affected residence times at 20 °C, but not at 5 °C. Particle-handling times on the labial palps ranged from less than 1 to 5.5 min, depending on temperature and concentration, accounting for 50% to 82%, respectively, of the total time particles spent in the pallial cavity. We suggest that (1) observed interspecific differences in particle handling on the labial palps may be due to differences in palp morphology and function, and (2) particle sorting and selection on the labial palps is a rate-limiting step of pre-ingestive feeding processes in by bivalves.     

Identification and activity-dependent labeling of peripheral sensory structures on a spionid polychaete

In marine sedimentary habitats, chemoreception is thought to coordinate feeding in many deposit-feeding invertebrates such as polychaetes, snails, and clams. Relatively little is known, however, about the chemosensory structures and mechanism of signal transduction in deposit feeders. Using electron microscopy, confocal laser scanning microscopy (CLSM), and immunohistochemistry, we investigated the structure and function of putative chemosensory cells on the feeding appendages of a deposit-feeding polychaete species, Dipolydora quadrilobata. Tufts of putative sensory cilia were distributed over the prostomium and feeding palps and typically occurred next to pores. Examination of these regions with transmission electron microscopy revealed multiciliated cells with adjacent glandular cells beneath the pores. The sensory cells of prostomium and palps were similar, displaying an abundance of apical mitochondria and relatively short ciliary rootlets. Staining with antiserum against acetylated alpha-tubulin was examined by CLSM, and revealed axonal processes from putative sensory tufts on the palp surface to palp nerves, as well as many free nerve endings. Activity-dependent cell labeling experiments were used to test the sensitivity of putative sensory cells on the palps to an amino acid mixture that elicited feeding in previous behavioral experiments. In static exposures, the number of lateral and abfrontal cells labeled in response to the amino acid mixture was significantly greater than in the controls. Ultrastructural, positional, and now physiological evidence strongly suggests that spionid feeding palps are equipped with sensory cells, at least some of which function as chemoreceptors.     

Aboveground phytochemical responses to belowground herbivory in poplar trees and the consequence for leaf herbivore preference

Belowground (BG) herbivory can influence aboveground (AG) herbivore performance and food preference via changes in plant chemistry. Most evidence for this phenomenon derives from studies in herbaceous plants but studies in woody plants are scarce. Here we investigated whether and how BG herbivory on black poplar (Populus nigra) trees by Melolontha melolontha larvae influences the feeding preference of Lymantria dispar (gypsy moth) caterpillars. In a food choice assay, caterpillars preferred to feed on leaves from trees that had experienced attack by BG herbivores. Therefore, we investigated the effect of BG herbivory on the phytochemical composition of P. nigra trees alone and in combination with AG feeding by L. dispar caterpillars. BG herbivory did not increase systemic AG tree defences like volatile organic compounds, protease inhibitors and salicinoids. Jasmonates and salicylic acid were also not induced by BG herbivory in leaves but abscisic acid concentrations drastically increased together with proline and few other amino acids. Leaf coating experiments with amino acids suggest that proline might be responsible for the caterpillar feeding preference via presumptive phagostimulatory properties. This study shows that BG herbivory in poplar can modify the feeding preference of AG herbivores via phytochemical changes as a consequence of root‐to‐shoot signaling.     

Particle transport in the zebra mussel, Dreissena polymorpha (Pallas)

The capture, transport, and sorting of particles by the gills and labial palps of the freshwater mussel Dreissena polymorpha were examined by endoscopy and video image analysis. More specifically, the morphology of the feeding organs in living zebra mussels was described; the mode and speeds of particle transport on the feeding organs was measured; and the sites of particle selection in the pallial cavity were identified. Particle velocities (outer demibranch lamellae, 90 microm s(-1); inner demibranch lamellae, 129 microm s(-1); marginal food groove of inner demibranchs, 156 microm s(-1); dorsal ciliated tracts, 152 microm s(-1)), as well as the movement of particles on the ctenidia and labial palps of D. polymorpha, are consistent with mucociliary, rather than hydrodynamic, transport. Particles can be sorted on the ctenidia of zebra mussels, resulting in a two-layer transport at the marginal food groove of the inner demibranch. That is: preferred particles are transported inside the marginal groove proper, whereas particles destined for rejection are carried superficially in a string of mucus. Sorting also occurs at the ventral margin of the outer demibranch; desirable particles are retained on the outer demibranch, whereas unacceptable particles are transferred to the inner demibranch and ultimately excluded from ingestion. We suggest that the structure of homorhabdic ctenidia does not preclude particle sorting, and that some ecosystem modifications attributed to zebra mussels may ultimately be due to ctenidial sorting mechanisms.