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1.
The feeding ecology of Axiothella rubrocincta (Johnson) from Tomales Bay, California, is described. This worm inhabits a U-shaped tube of agglutinated sand grains and mucus. Morphological adaptations such as nuchal and anal plaques prevent the tube from becoming clogged. Foreign debris entering the tube is either consumed or incorporated into the tube wall. The proboscis and notosetae are probably used to clean the tube wall. A. rubrocincta combines feeding and burrowing activities to form the funnel and complete the tube: it does not ingest sediment while burrowing. Organic matter deposited into the funnel is buried there by sand slides originating at the rim of the funnel. The concentration of organic matter within the funnel is significantly higher than for non-funnel sediments. A. rubrocincta consumes food from the upper 2 cm of the substratum and is 4.6% efficient as a depositfeeder. It also feeds within the funnel and can ingest large quantities of food. This feeding process is described. A. rubrocincta irrigates its tube at a rate of 5.1 ml sea water/g/h while feeding, and briefly reverses this current to a rate of 0.1 ml/g/h when defaecating. The rhythmic activity patterns are integrated: the mean defaecation and inverse pumping time is 14.4 min at 15 ± 1 C.  相似文献   

2.
Palpation increases the amount of sensory input reaching the central nervous system compared with that obtained from sustained contact but that increase is not essential to allow discrimination. During a meal on favoured food, phagostimulatory input from the palps is not needed to drive feeding. When less favoured food is taken, phagostimulatory input from the palps may enhance feeding. Even with favoured food, the palps are important in registering inhibitory substances.  相似文献   

3.
Ogrinc  N.  Hintelmann  H.  Eckley  C.  Lojen  S. 《Hydrobiologia》2003,496(1-3):207-213
The functional morphology of the feeding palps and prostomium of the spionid polychaetes Streblospio benedicti and S. shrubsolii was studied. Three functional groups of cilia of the feeding palps were found on both species – frontal cilia, latero-frontal cirri and lateral cilia. Frontal cilia line the food groove and transport food particles to the pharynx, and have been reported for all spionid polychaetes except species of the genus Scolelepis. Latero-frontal cirri deflect particles onto the frontal surface and have been observed in several spionid genera including Paraprionospio, Streblospio, Polydora and Dipolydora. Lateral cilia beat in continuous metachronal waves creating lateral vortices that potentially entrain suspended particles, and are known in Paraprionospio and Streblospio. The two species of Streblospio did differ in the distribution of prostomial papillae. These papillae are eversible and thought to function in particle selection as particles on the pharynx come in contact with the papillae. Prostomial papillae were restricted to the peripheral surface of S. benedicti and were widely scattered on all surfaces of the prostomium of S. shrubsolii. A conical tentaculate structure occurs between the branchiae of the first setiger of S. benedicti, but only a low raised elevation is present on S. shrubsolii.  相似文献   

4.
Aspects of feeding, such as food capture and ingestion, as well as mobility of the polychaetes Eurythoe complanata, Marphysa formosa and Diopatra aciculata, from S?o Sebasti?o Channel (S?o Sebasti?o, state of S?o Paulo) were observed in laboratory conditions. Eurythoe complanata, a carnivorous species, fed exclusively on pieces of fish with the aid of strong muscular retractable lips, and detected the presence of food by chemical stimuli. Diopatra aciculata, an omnivorous species, captured and ingested different kinds of food with the aid of its jaws, generating a flow of water through its tube by which it detects the presence of food and oxygenates its gills. Marphysa formosa also used its jaws to bite and lacerate food. These species showed greater or lesser degrees of intolerance to light.  相似文献   

5.
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.  相似文献   

6.
Macoma balthica (L.), an abundant clam, ubiquitous in temperate estuaries across the North Atlantic, is known to practice both alternative basic modes of feeding available to seafloor invertebrates. It either holds its feeding organ, the siphon, at a fixed position just above the sediment surface to filter out food particles suspended in the overlying water or else extends and moves its siphon around to vacuum up deposited food particles on the sediment surface. Previous laboratory experiments have established an understanding of the role of current flow in dictating the choice of whether suspension or deposit feeding will be used by marine invertebrates with the facultative flexibility to choose. Faster flows imply greater fluxes of suspended particles so that the energetic rewards of suspension feeding are enhanced. Slower flows imply reduced renewal rates of suspended foods in the bottom boundary layers and enhanced deposition of food particles on the seafloor so that a switch to deposit feeding is favored. Like early optimal foraging theory, this understanding is based on energetic considerations alone without incorporation of broader implications of how population interactions such as predation and competition influence individual foraging behavior. Feeding behavior of Macoma balthica is influenced in the Neuse River estuary by both hydrodynamics and siphon-cropping by juvenile demersal fishes. Under conditions of identical concentrations of suspended particulates in the water column and organic contents of surface sediments, Macoma exhibited much higher levels of deposit feeding where currents were slower. In addition, exclosure and fish inclosure experiments demonstrated that juvenile demersal fishes influence feeding behavior of Macoma by cropping exposed siphons and inducing reduction in deposit-feeding activity. Effects of croppers were substantial in early to midsummer, when juvenile fish abundances were greatest in trawl samples from this estuarine nursery and before the growing fish exhibited ontogenetic changes in diet away from early concentration on bivalve siphons. Field experiments in which siphon-cropping fish were caged at varying distances off the bottom failed to detect any effective behavioral avoidance by Macoma of cropping in response to proximity of fish. One might have hypothesized that under high risk of cropping, Macoma would switch to suspension feeding and away from deposit feeding, the feeding method entailing more risk of losses to croppers because of greater siphon activity and greater extension of siphons on the sediment surface. Consequently, partial predation by siphon-cropping fishes greatly reduces deposit-feeding activity by Macoma balthica during summer as an apparent direct effect of disfigurement and reduction of siphons, the organ required for efficient deposit feeding. Information on current flows alone would not suffice to predict feeding behavior of this marine invertebrate: the influence of partial predation must also be included.  相似文献   

7.
Rich Mooi 《Zoomorphology》1986,106(2):75-90
Summary There are five major types of non-respiratory podia in the Order Clypeasteroida: accessory, barrel-tipped, food groove, large food groove, and buccal. The anatomy of each type is intimately related to its function in the feeding mechanism of clypeasteroids. Accessory podia are found aborally and orally in some species, only aborally and ambitally in others. Accessory podia are largely sensory and manipulatory, but can be locomotory in the small fibulariids and juvenile sand dollars. Barrel-tipped podia have expanded disk muscles and connective tissue, and are usually found in two sizes, large and small. In species that have them they are usually restricted to the oral surface. These podia collect food and pass it towards the food grooves in the manner of a bucket brigade. Food groove podia are found only in species with food grooves. These podia are small, with reduced tip musculature and expanded secretory tissue for coating food with mucus. They transport food down the food grooves to the mouth. Large food groove podia are simply large versions of ordinary food groove podia. They help move the clumped food into the mouth area towards the buccals, and are found only in the Clypeasteridae and some scutellines. Buccal podia lack tip musculature, but possess tip support fibres and a single type of small secretory cell. They are sensory, and capable of manipulating particles into the mouth. Buccals are present in all families except the Clypeasteridae. Juvenile Echinarachnius less than 3 mm in diameter have only respiratory, accessory and buccal podia. Food groove and barrel-tipped podia start to differentiate from the accessories as the juvenile approaches a diameter of 4 to 5 mm. Clypeasteroid podial diversity increases the efficiency of the food collecting mechanism. The anatomy and distribution of podia on the oral surface of scutellines supports the fact that this surface is the prime food collecting area in all true sand dollars. The podia (not miliary spines) are the major source of mucus used during the feeding process and are the primary feeding appendages.  相似文献   

8.
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.  相似文献   

9.
Mineralogical segregation of sand grains distinguishes the trace fossil Macaronichnus segregatis, which is composed of a felsic burrow infill with a mafic-and mica-rich burrow mantle. This study focuses on determining the mechanism by which M. segregatis trace-makers segregated mineral grains during deposit feeding. A modern opheliid polychaete, Euzonus mucronata, from Pachena Bay, Vancouver Island (Canada), was examined to explain the activities of their ancient counterparts. Microscopic videotaping of deposit feeding allowed for collection of data on ingestion and excretion through visual grain counts of felsic, mafic, and shell components. Normalization of these grain counts to the composition of the host sediment illustrates preferential ingestion of felsic grains over mafic. Shell fragments were generally avoided and visually mantled the burrows, obscuring the paucity of mafic grains in burrow infills. The avoidance of shell fragments is potentially a function of the large grain size, angular shape, surface texture, and/or associated low nutritive value. The preferential ingestion of felsic grains is attributed to en masse feeding in felsic-rich locales identified through sediment probing. This form of mineral segregation likely reflects the specific nature of the sediment and worm population. Accordingly, en mass deposit feeding in selected felsic-rich localities is one possible mechanism used in the construction of Macaronichnus segregatis and M. segregatis-like structures.  相似文献   

10.
An agar sediment test was developed to evaluate the suitability of organic waste streams from the food industry for recovering nutrients by the aquatic worm Lumbriculus variegatus (Lv). The effects of agar gel, sand, and food quantities in the sediment test on worm growth, reproduction, and water quality were studied. Agar gel addition ameliorated growth conditions by reducing food hydrolysis and altering sediment structure. Best results for combined reproduction and growth were obtained with 0.6% agar-gel (20 ml), 10 g. fine sand, 40 g. coarse sand, and 105 mg fish food (Tetramin). With agar gel, ingestion and growth is more the result of addition of food in its original quality. Final tests with secondary potato starch sludge and wheat bran demonstrated that this test is appropriate for the comparison of solid feedstuffs and suspended organic waste streams. This test method is expected to be suitable for organic waste studies using other sediment dwelling invertebrates.  相似文献   

11.
The fiddler crab Uca panamensis (Stimpson, 1959) inhabits rocky shores. We examined its preference for feeding substratum—sand or rock—and its manner of feeding. The crab made its burrow in the sand among rocks but preferred to feed on rocks. The feeding time decreased as the distance between the burrow and the rock increased. We consider this to be a result of exclusive interaction among the crabs because they defended their feeding area on the rocks against others.The crab wetted a small area of rock with water held in the branchial chambers before and during feeding. It pinched up the wetted surface in the minor chelipeds, which have bundles of setae on the posterior tips of the dactyl and pollex, and put the material into its buccal cavity. It never expelled sand pellets while feeding on rock, which indicates that it swallowed the food particles directly, without sorting. The bundles of setae retained water by capillary attraction, which suggests that they capture the suspended fine food particles scraped from the rock. The wetting action may prevent the fine materials from dispersing. We consider that morphological alteration of the minor chelipeds, the application of water from the branchial chambers, and direct swallowing permit the fiddler crab to feed on fine materials attached to rocks.  相似文献   

12.
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.  相似文献   

13.
The influence of the food content and the particle size of the substratum on the distribution and relative abundance of Metapenaeus macleayi (Haswell) has been investigated by periodic trawl sampling for prawns and laboratory studies of their food, feeding, and burrowing behaviour.M. macleayi are opportunistic omnivores; they pick up material from the bottom with their chelipeds and convey it to the mouthparts where edible matter is sorted and ingested. They burrow into the sediment with their pereopods and pleopods, and are usually totally buried beneath the surface. A respiratory water current enters a tube formed by the antennal scales and the antennules, flows over the gills and then out of the carapace; this current is regularly reversed with increased strength, presumably to carry away de-oxygenated water. The results of experiments on substratum preference with adequately nourished juveniles suggest that the particle size of the sediment is more important than the food in the substratum in determining the distribution; the apparent preference of juveniles for a fine sandy substratum both in the laboratory and in the natural habitat may be attributed to the minimum threshold velocity of the sand particles.Adults are most abundant in turbid coastal waters arising from estuarine discharge and here the size of the sediment particles appears to be less important than the food content of the substratum in determining the distribution. The mangrove and reed swamps in the Hunter region play an important rôle in the food requirements of juvenile and adult prawns.  相似文献   

14.
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.  相似文献   

15.
ABSTRACT. Scanning electron microscopic observations of feeding plasmodia show three characteristic features: 1) extension of multilobed pseudopodia protruding from the leading edge of the plasmodium as it advances onto the surface of a food particle, 2) confluence of the lobes to form a sheath-like pseudopodium attached to the surface of the food particle, and 3) protrusion of small nodules with thin lamellar projections from the leading edge of the plasmodium. Sections through freeze-dried preparations of the feeding plasmodium exhibit a highly convoluted under surface in contact with loosened starch grains that appear to be released by extracellular digestion. the cytoplasm, viewed by transmission electron microscopy, contains branched, internally penetrating canals (ca. 2 μm wide) enclosing engulfed starch grains. Starch grains in the deeper part of the canals are more electron dense and appear to be digested. Micropseudopodia (70-80 nm dia.), projecting from the surface of the canals, protrude toward and into the ingested starch grains. Digestive marker enzyme (acid phosphatase) activity was detected cytochemically in food particles penetrated by micropseu-dopodia indicating a digestive role for these structures not reported previously.  相似文献   

16.
The particle size distribution of faecal pellets produced by the tubificid worm Tubifex tubifex in laboratory culture, was measured with a Coulter® Multisizer. The faecal material from worms cultured in a range of sediments was composed of particles with a mean diameter of less than 63 m, and only a few isolated larger particles were found by microscopic analysis. This suggests that this species actively selects the silt-clay fraction, avoiding larger sand particles. A more detailed analysis of faeces revealed that about 75%, by volume, was composed of particles with a mean diameter < 25m, and the mode was < 10m. T. tubifex fed selectively on the organic rich particles of the sediment, and this feeding was independent of particle size. Measurement of the organic content of faeces (measured as % loss on ignition) showed that they had a consistently higher organic content than the sediment, considered as whole sediments or the <63 m sieved fraction. On the basis of these results, we hypothesise that this species exhibits two levels of selectivity in its feeding behaviour. Thus selection is primarily based on particle size, avoiding the ingestion of sand particles and also, on the preferential selection of particles associated with organic material, within the fine (silt-clay) fraction of the sediment.  相似文献   

17.
Scalibregmatidae is a small annelid family of subsurface deposit feeders in sand or mud, which are generally well adapted to infaunal burrowing. The overall morphology of Scalibregmatidae is very similar, with thick bodies, small parapodia, and no prostomial appendages or short horns. The only exception is members of the genera Axiokebuita and Speleobregma that most frequently inhabit crevices or gravel and possess extensive ventral ciliated palps and globular adhesive pygidium. Character reconstruction using maximum likelihood and Bayesian methods show that ciliated palps and adhesive pygidium are synapomorphies of the AxiokebuitaSpeleobregma clade. The most likely transformation series is from horns to ciliated palps, the origin of which correlates with the occurrence of Axiokebuita and Speleobregma in crevices or gravel. The wide spaces among rocks or granules yield high permeability and inertial water flow, preventing deposition of organic matter. Under these flow conditions that differ significantly from those of sand or mud bottom, ciliated palps aid to the collection of suspended particles and an adhesive pygidium provides attachment. With palps being a highly debated character in annelid evolution, it is remarkable that prominent ciliated palps are gained within a lineage of ancestrally nonpalpate annelids, most likely increasing their fitness when colonizing a new environment.  相似文献   

18.
To thoroughly understand the feeding processes of the blue mussel, Mytilus edulis, under the variable environmental conditions it experiences in nature, it is important to examine individually the different components of its feeding system. The ciliated ventral food groove represents one of these components, within which the majority of food particles trapped by the gill are transported to the labial palps and mouth. Any ability of the mussel to adjust food transport rates within this groove could serve as an important feeding regulatory mechanism in response to variations in the environment.

Mucous strand velocities in the ventral groove of the mussel, M. edulis, were determined by video endoscopy over different time periods and during short- and long-term manipulations of ambient particle concentration and temperature. Mucous strand velocity decreased with increasing ambient particle concentration at 14°C, but a similar relationship was not observed at 5°C. The data support the hypothesis that M. edulis possesses compensatory mechanisms to control particle transport at the level of the ventral groove cilia in response to changes in the environment. Furthermore, mucous strand velocity in the ventral groove increased when the ambient temperature of mussels acclimated to 5°C was increased to 15°C during acute and long-term acclimation temperature experiments. This response is consistent with standard physiological responses of ciliary systems to changes in temperature.  相似文献   


19.
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.  相似文献   

20.
1. Nais pardalis Piguet, a small freshwater oligochaete, lives in the surface few cm of sandy sediments in Loch Leven, County of Kinross, Scotland. 2. It forms branching passageways or tubes in the sediment not by any active constructional behaviour, but by the secretion of a material which sticks the sand grains together. 3. The sand grains making up the tubes are exactly the same size as those in the surrounding sediment. 4. Animals select the smallest grains in their environment for ingestion. 5 In the laboratory, rates of ingestion of sand grains measured on different animals vary from 5 to 60 grains/hour and rates of excretion from 3 to 6/hour, but these figures should be viewed with caution. 6. Analysis of the rate of ingestion of sand grains indicates that grains are not ingested randomly in time. Having eaten one grain, an animal is less likely to eat another within the next 10 seconds.  相似文献   

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