Peribuccal organs ofPlacopecten magellanicus andChlamys varia (Mollusca: Bivalvia): Structure,ultrastructure and implications for feeding

In order to better understand the structure of bivalve peribuccal organs and relate this to existing functional paradigms of their role in feeding, the labial palps of two scallop species,Placopecten magellanicus from the Bay of Fundy, Canada (1985 and 1986), andChlamys varia from the Bay of Brest, France (1986), were examined using histological techniques and electron microscopy. The ridged palp surface displays a uniformly dense ciliation with relatively few mucocytes; these are essentially concentrated in the region of the secondary ledge and may, through their secretory activity, determine the fate of particle masses in this area. The mucus secretions of the ridged palp surface are qualitatively different from those of the smooth palp surface. Mucocytes are much more abundant on the smooth palp surface, where it is suggested that their homogeneous secretions attenuate the potentially adverse effects of anteriorly-directed cleansing and swimming currents. Two other cell types are found in the palp epithelia: ciliated cells, which are very numerous on the ridged surface and relatively rare on the smooth surface, and non-ciliated epithelial cells, which are very numerous on the smooth surface and rare on the ridged surface, where they are confined to the palp margin. In addition to the mechanical role of the ciliated cells and mucocytes, the ultrastructural characteristics of the ciliated and non-ciliated epithelial cells indicate a dichotomy of function between the ridged and smooth surfaces. The ridged surface epithelial cells present an ultrastructural specialization in the absorption of dissolved and colloidal matter, suggesting an accessory nutritive role, whereas the smooth surface simple epithelial cells show signs of active molecular synthesis. No specialised sensory cells were observed on the ridged surface; it is therefore not yet possible to conclude whether the labial palps are capable of selection based on individual particle characteristics.     

Particle sorting and formation and elimination of pseudofaeces in the bivalves <Emphasis Type="Italic">Mulinia edulis</Emphasis> (siphonate) and <Emphasis Type="Italic">Mytilus chilensis</Emphasis> (asiphonate)

Mulinia edulis and Mytilus chilensis are suspension-feeding bivalves with homorhabdic gills that live in different sedimentary habitats in the lower and upper intertidal, respectively, in Yaldad Bay, Chile. They are faced with different suspended particle size distributions when feeding, and both eliminate most of the inorganic particles by pseudofaeces production. This study used histology, scanning and transmission electron microscopy, and video endoscopy to compare particle processing on the labial palps and the mechanisms of particle sorting, acceptance, and rejection in the two species. In both species, disaggregation of mucus-bound particles occurs on the plicate surface of the palps. Particles destined for ingestion pass anteriorly from crest to crest and reach the mouth via the palp acceptance tract. Those destined for rejection enter the troughs between the plicae and move ventrally to the palp rejection tract. The palps manipulate the pseudofaeces into a mucous ball, which is transferred to the mantle rejection tract. In Mulinia edulis (an infaunal, siphonate mactrid), the pseudofaeces are stored in a chamber at the base of the inhalant siphon until expelled by intermittent contraction of the siphon wall. In contrast, Mytilus chilensis (an epifaunal, non-siphonate mytilid) releases pseudofaeces continuously when submerged.     

Mode of particle ingestion in five species of suspension-feeding bivalve molluscs

In order to elucidate the mode of particle ingestion and the functional anatomy of the oesophagus in bivalves, a histological study was performed onMytilus edulis (Mytilidae),Crassostrea virginica (Ostreidae),Placopecten magellanicus, Chlamys varia, and juvenilePecten maximus (Pectinidae). Specimens were sampled from various sites in New Brunswick, Canada, and Brittany, France, from 1987 to 1989. The buccal, peribuccal, and oesophageal epithelia of all species contained a dense distribution of actively secreting mucocytes, although these were somewhat less abundant inCrassostrea virginica, which also has the shortest oesophagus. Mucocyte morphology, while constant within a family (Pectinidae), showed clear differences among families. Both acid and neutral mucopolysaccharides were secreted by the epithelial mucocytes of all species. Mucus and mucus-particle masses were observed in the peribuccal and buccal regions, as well as in the oesophageal lumina of all species, even in those specimens which had been maintained without feeding (Placopecten magellanicus) or held out of water for 48 h (C. virginica) prior to dissection and fixation. These results indicate that a basal level of mucus production and transport is continuous on the peribuccal, buccal, and oesophageal ciliated epithelia, regardless of the particle concentration in the external medium. Buccooesophageal glands, generally thought to be absent in the Bivalvia, were observed in one of the species examined (M. edulis). It is concluded that the mode of particle ingestion in these suspension-feeding bivalves is via ciliatransported mucus masses; the presence of buccooesophageal glands inM. edulis suggests a digestive role for the oesophagus in this species.     

Particle sorting in bivalves: in vivo determination of the pallial organs of selection

Benthic particle feeders are exposed to a food supply varying in both quantity and quality. Previous studies have shown that bivalve molluscs deal with such fluctuating particle regimes in a variety of ways, including adjustments in pumping and ingestion rates, and selective rejection of non-nutritive particles as pseudofeces. The actual site of particle selection within the pallial cavity, however, has remained a topic of speculation. During August 1995 and January and August 1996, we exposed the oysters Crassostrea virginica (Gmelin) and C. gigas (Thunberg), and the mussel Mytilus trossulus Gould to a mixture of ground, aged Spartina alterniflora Loisel and similar-sized phytoplankton at three concentrations (10³, 10⁴, 10⁵ particles ml⁻¹). We then examined the ctenidia and labial palps by means of endoscopy and sampled, in vivo, the particulate material from various ciliated tracts, and analyzed the samples with a flow cytometer. We found that in oysters, the ctenidia are responsible for particle sorting, whereas the labial palps play an accessory role in particle selection, or function to control the volume of material to be ingested. In mussels, however, the ctenidia play little role in particle selection and simply transport particulate matter to the palps for further processing. We suggest that selection by the ctenidia of oysters is a function of their architecture (plicate, heterorhabdic). Received: 4 June 1997 / Accepted: 23 January 1998     

A functional interpretation of cilia and mucocyte distributions on the abfrontal surface of bivalve gills

The lack of fundamental data on the abfrontal surface of bivalve gills has prompted a comparative study of cilia and mucocytes on this surface, using scanning electron microscopy and histology on eight species of bivalves, representing seven families and the four major gill types: Mytilus edulis, Modiolus modiolus, Arca zebra, Placopecten magellanicus, Crassostrea virginica, Spisula solidissima, Mercenaria mercenaria, and Mya arenaria. Abfrontal cilia and mucocytes were found in all species studied, with types and densities differing within and between gill types. The three species of homorhabdic filibranchs presented different densities of abfrontal cilia and mucocytes, from very dense in M. edulis to sparse in A. zebra. The heterorhabdic gills had intermediate cilia and mucocyte densities, with highest concentrations of both abfrontal cilia and mucocytes on the principal filaments. The eulamellibranchs showed low ciliary densities together with high mucocyte densities, especially in S. solidissima, where the abfrontal mucocytes were glandular. These results indicate that: (1) the abfrontal surface is a vestigial mucociliary surface; (2) the abfrontal surface cannot participate in water pumping in most species, due to low ciliary densities; and (3) species with high densities of abfrontal mucocytes could utilize abfrontal mucus to reduce drag, especially in the highly fused gills, such as those of the eulamellibranchs. The differing distributions of abfrontal cilia and mucocytes may reflect different selective pressures acting on the gills within the various taxa. Received: 12 February 2000 / Accepted: 10 September 2000     

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

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

The role of mucus in particle processing by suspension-feeding marine bivalves: unifying principles

Contemporary research on bivalve suspension-feeding has revealed a diversity of particle processing mechanisms depending on the anatomy and functioning of the pallial organs involved. On the biochemical level, however, some evidence of homogeneity has emerged concerning the types of mucopolysaccharide associated with particle processing. The present study uses both previous data and original research combining video endoscopy and mucocyte mapping to further explore the relationships between pallial organ topography, functional correlates, direction of current flow, and mucocyte secretion type. Five species representing five different families and all four major gill types are represented: Mytilus edulis, Placopecten magellanicus, Crassostrea virginica, Mya arenaria, and Spisula solidissima. Viscous acid or acid-dominant mucopolysaccharides are used when particle transport occurs on an exposed surface, or on a structure leading directly to such a surface, counter to the prevailing current flow. Associated functions are indiscriminate transport in gill ventral particle grooves and rejection of pseudofeces. Lower-viscosity mixed mucopolysaccharides are used when particle transport is on an enclosed or semi-enclosed surface, leading to other such surfaces, and with the current flow. Associated functions are transport of particles destined for ingestion, and ingestion itself. Low-viscosity neutral mucopolysaccharides are found in regions where reduction of mucus viscosity is important, such as the areas of the labial palps responsible for fluidization of the high-viscosity mucus-particle cord of the gill ventral particle groove prior to particle extraction. There thus appears to be a specialization of mucus type corresponding to functional specialization of the various pallial organs in suspension-feeding marine bivalves. Received: 10 October 1996 / Accepted: 7 May 1997     

In situ rates of body growth and palp regeneration of a spionid polychaete following simulated sublethal predation

Sublethal predation can be a significant trophic pathway in sediment communities, and the regeneration of damaged tissue affects the activity, growth, and reproduction of prey. A field experiment was conducted 12–18 July 2010 in the Tijuana Estuary, California (+32.56617°, −117.13152°), to measure in situ rates of body growth and palp regeneration of the polychaete Polydora cornuta after simulating sublethal predation by removing 2, 1, or 0 palps from labeled individuals. After 3 days in the field, individuals that had 2 palps removed grew significantly slower than worms that had 0 palps removed; 1-palp worms had intermediate growth. After 6 days in the field, rates of body growth were faster than those of worms recovered after 3 days, with similar trends among the three palp treatments. During the 3-day transplantation, removed palps regenerated to half the initial length of unmanipulated palps. After 6 days, palp lengths did not differ significantly among treatment groups, indicating complete regeneration. The regeneration rate of removed palps was significantly faster than the growth rate of undamaged palps, and palp regeneration rate did not differ significantly between worms that had 1 or 2 palps removed.     

Functional morphology and feeding behavior of Scolelepis squamata (Polychaeta: Spionidae)

Functional morphology and feeding behavior of Scolelepis squamata (Müller) were studied. Gut contents consisted of unconsolidated sedimentary particles, fecal pellets of other species, and a wide variety of embryos, larvae, and juveniles. Unlike other spionid polychaetes the palps of S. squamata lack a median, ciliated groove. Particles captured by the palps were brought to the pharynx by a complete contraction of the palp. In the presence of a current, S. squamata helically coiled their palps, and fed almost exclusively on suspended and resuspended particles. In contrast to most spionid polychaetes the palps of S. squamata are not deciduous. Various aspects of the morphology and feeding behavior of S. squamata are discussed in terms of their functional and ecological significance.     

New observations of the gills of Placopecten magellanicus (Mollusca: Bivalvia), and implications for nutrition

The internal anatomy and microanatomy of the gill of Placopecten magellanicus Gmelin collected in May and November 1985 from Chamcook Bay, New Brunswick, Canada, was studied using thin-section light microscopy and transmission electron microscopy. Most of the spurs show no evidence of organic union, and hence do not participate in vascular exchange. However, the dorsal bend shows both ciliary and organic interfilamentar union. The internal structure and the hemocytes of the dorsal respiratory expansion are presented. The epithelium consists of three distinct cell types, bounded by apical microvilli. All regions of the gill contain an epithelial basal membrane, which is greatly convoluted in the interconnecting vessels of the dorsal respiratory expansion. The significance of these observations is discussed in relation to possible roles in respiration, transmembrane transport and nutrition. The apical surface of all ciliated cells is covered with an acellular matrix composed of clear spherical vesicles, which may serve a mechanical function for which mucus would be unsuited. The significance of the abundance of mucocytes on the abfrontal surface of the principal filaments is discussed in terms of the escape response of pectinid bivalves.     

Anterior regeneration in the spionid polychaetes<Emphasis Type="Italic"> Dipolydora quadrilobata</Emphasis> and<Emphasis Type="Italic"> Pygospio elegans</Emphasis>

Like many other benthic infaunal invertebrates, spionid polychaetes often lose portions of their bodies to predators, which affects their activities. Feeding palp loss and tail loss have been studied in several spionids, but the capacity for anterior tissue regeneration has not been compared in different species. The present study examines anterior tissue regeneration in two species, Dipolydora quadrilobata (Jacobi 1883) and Pygospio elegans Claparède 1863, in two laboratory experiments. Tissue removal treatments included removal of palps only, removal of anterior tissue through the first setiger, anterior tissue through the fifth setiger, all anterior tissue through half of the gill-bearing setigers, and all anterior tissue through the last gill-bearing setiger. Regeneration was monitored by capturing images of the worms and digitizing the area of regenerated anterior tissue or counting the number of segments that grew over time. Worms of both species regenerated anterior tissue regardless of the amount removed. Morphogenesis during regeneration followed a similar pattern in these two species regardless of the amount of anterior tissue lost, progressing from wound healing to formation of a recognizable prostomium and peristomium (“head”) by 6 days post-ablation. Palp and setal growth, addition of segments, and formation of nuchal organs and the ciliated food groove followed so that worms appeared to have re-grown “normal,” but smaller, “heads” and palps by 9–12 days following ablation. Over the course of 16 days, worms that lost more segments regenerated less tissue relative to their initial intact size and did so more slowly. There was no significant palp growth during the first 3 days following ablation. Rate of segment addition was directly related to the degree of tissue loss in D. quadrilobata. P. elegans added segments at similar rates whether 50 or 70% of the original segments was removed.     

Gill function and particle transport in Placopecten magellanicus (Mollusca: Bivalvia) as revealed using video endoscopy

The technique of endoscopic video observation was used to study feeding processes of Placopecten magellanicus (Gmelin), collected from Bull Arm, Newfoundland in August 1991 and 1992, under near-natural feeding conditions. The fate of captured particles depended on the extent of ingestive or handling capacity saturation. Under low (1 to 10 particles l^-1) to medium (10 to 20 particles l^-1) particle concentrations, most particles were incorporated in continuous anteriorly directed slurries in the dorsal ciliated tracts of the gill arch and dorsal bends. As particle concentration or exposure time to the lower particle concentrations increased, four endogenous mechanisms of ingestion volume control were increasingly observed: (1) rejection of dense mucus-particle masses from the principal filament troughs onto the ventrally beating cilia and associated currents of the ordinary filament plicae, counter to and below the incoming pallial current maintained by the principal filament cilia; (2) intermittent stopping of the anteriorward flow in the dorsal ciliated tracts; (3) reduction or cessation of input from the principal filaments to the dorsal ciliated tracts; (4) detachment of the dorsal bends from the mantle to establish a shunt from the infrabranchial to the suprabranchial cavity. Chemical and histochemical tests of purified fluid withdrawn from the dorsal ciliated tracts indicate that mucus is present at all particle concentrations. Mucus therefore participates both in normal feeding and in ingestion volume control on the bivalve gill, although different mechanisms, and types of mucus, effect transport of material in the dorsal (feeding) and ventral (cleaning) ciliated tracts.     

Effects of simulated sublethal predation on the growth and regeneration rates of a spionid polychaete in laboratory flumes

Most spionid polychaetes switch from deposit feeding to suspension feeding as current speed and the flux of suspended food increase. Growth rates of juvenile Polydora cornuta are strongly affected by flow and can be as rapid as 60% day⁻¹ in moderate currents. Feeding palps that extend above the sediment–water interface during suspension feeding are especially vulnerable to sublethal predation, but individuals with damaged posteriors are also common. We performed a series of laboratory flume experiments to test the effects of sublethal tissue damage on the growth and regeneration rates of P. cornuta juveniles. Replicated experiments were conducted at three flow speeds in counter-rotating annular flumes containing field-collected sediment and a nonliving algal slurry as deposited and suspended food. In the first set of experiments, we removed 2, 1, or 0 of worms’ two feeding palps and measured the relative growth rates of worm bodies and palps after 3 days in the flumes. Worms that lost both palps grew significantly slower than the other two groups, but the growth rate of worms that had one undamaged palp was not significantly different from worms that had two undamaged palps. Faster flow speeds significantly increased rates of body growth, and there was a significant interaction between flow and the effect of palp loss. During the 3-day experiments, damaged palps fully regenerated and often grew larger than they were prior to being removed. Damaged palps also grew significantly faster than undamaged palps. The second set of experiments tested the effects of removing a worm’s posterior region (~18% of body volume). The growth rates of these damaged and undamaged worms did not differ significantly. By the end of a 3-day flume experiment, damaged worms had grown 6× larger than they were prior to the posterior damage. The rapid regeneration of damaged palps and posterior tissue in moderate flows that allow suspension feeding suggests that sublethal predation on spionids might be more frequent than previously estimated and will have little impact on the growth of juvenile recruits.     

New observations of the gills of Placopecten magellanicus (Mollusca: Bivalvia), and implications for nutrition

The organization, general anatomy, and surface microanatomy of all regions of the gills of a representive bivalve mollusc, Placopecten magellanicus Gmelin, were studied using stereo-microscopic, histological, and scanning electron microscopic techniques. Individuals were collected in May and November 1985 from Chamcook Bay, New Brunswick, Canada. In addition to correcting earlier accounts of this structure, a number of new observations are reported. The orientation of the ciliated spurs appears to be responsible for the sinusoidal arrangement of the gill filaments. Micrographs showing the structure of the dorsal respiratory expansion are presented. The entire abfrontal surface of the principal filament, including the dorsal respiratory expansion, is densely ciliated and mucosecretory. These characteristics may aid in the establishment of a respiratory current and in the prevention of gill damage during escape responses. All nonciliated regions of the gill filaments are covered with microvilli, thus greatly increasing the surface area of the gill. The feeding mechanism is discussed in relation to the dorsal and ventral ciliated tracts. Symbiotic ciliate protozoans are constantly dislodged from the gill filaments and transported via the ventral mucus string to the buccal region. The nutritional implications of these observations are discussed.     

Particle-size selection by Pseudopolydora paucibranchiata (Polychaeta: Spionidae) in suspension feeding and in deposit feeding: influences of ontogeny and flow speed

Pseudopolydora paucibranchiata Okuda suspension feeds and deposit feeds at the sediment-water interface, where it is exposed to a variety of particles differing in physical characteristics and nutritional value. In flume experiments (conducted in August 1994 and May 1995) with two sizes of either suspended or deposited beads, I measured particle-size selection separately in each feeding mode. The same influences of palp width and of ambient flow speed were observed in each mode. At velocities 0.74 cm s^-1 there were no relationships between palp width and the proportion of gut contents composed of large beads. At velocities 1.8 cm s^-1 worms with narrower palps ingested relatively fewer large beads (and more small beads) than did worms with wider palps. Palp width and body length were linearly related, and results were similar when analyzed with body length as the independent variable. As flow speed increased, selectivity changed in a worm-size-specific manner: worms with a palp width -1. Assuming that in the field (1) particle size is the principle criterion for selection, and (2) the amount of digestible food component in deposited and suspended particles, respectively, is related to particle surface area and volume, I hypothesize that changes in selectivity as velocity rises can cause juveniles to experience a decreasing profitability of suspension feeding and a simultaneously increasing profitability of deposit feeding. Juveniles could maintain a diet of high food value despite flow variations by adjusting the proportion of time they spend suspension feeding relative to deposit feeding.     

背角无齿蚌不同组织的基因组DNA甲基化分析

用甲基化敏感扩增多态性(MSAP)对背角无齿蚌(Anodonta woodiana)腮、唇瓣、闭壳肌、外套膜和斧足五个组织基因组DNACCGG区域的甲基化水平进行了分析.结果表明,背角无齿蚌腮基因组DNA甲基化比例为47.9%,唇瓣甲基化比例为35.5%,闭壳肌甲基化比例为50%,外套膜甲基化比例为46.3%,斧足甲基化比例为56%;基因组中CCGG区域存在甲基化现象.不同组合甲基化比例不同说明该区域甲基化可能参与到基因的调控中.通过比较不同地点的采集的背角无齿蚌,发现污染严重地区(太湖三山岛水域)采集到的样品和非污染地区(南泉养殖水域)采集的蚌样甲基化区域略有变化.其中的相关性有待进一步的研究.     

Isolation,characterization and glucose metabolism of glycogen cells (=vesicular connective-tissue cells) from the labial palps of the marine mussel Mytilus edulis

A method has been developed to isolate glycogen cells (=vesicular connective tissue cells) from the labial palps of the marine mussel Mytilus edulis L. These cells have a modal density of 1.14 g ml^-1 and this property has been exploited to separate them from pronase-dissociated cell dispersions by density-gradient centrifugation. The isolated cells were mainly spherical, with a small peripheral nucleus. The quasitotality of the cell was filled with -like glycogen particles with cellular organelles being limited to the cell periphery. The major recipients of [U-¹⁴C] glucose-derived carbon were glycogen and amino acids. Only small quantities of ¹⁴C radiolabel were recovered from lipids, protein and CO₂. Glucose incorporation into glycogen increased linearly over 6 h and showed saturation kinetics with respect to exogenous glucose concentration.     

Effects of the toxic dinoflagellate <Emphasis Type="Italic">Gymnodinium catenatum</Emphasis> on uptake and fate of paralytic shellfish poisons in the Pacific giant lions-paw scallop <Emphasis Type="Italic">Nodipecten subnodosus</Emphasis>

Juvenile Pacific giant lions-paw scallops Nodipecten subnodosus were fed the toxic dinoflagellate Gymnodinium catenatum, a producer of paralytic shellfish poison (PSP), supplied with Isochrysis galbana (a nontoxic microalgae). Short-term (<24 h) experiments were performed to determine clearance and ingestion rates of G. catenatum. Kinetics of PSP was examined in longer-term experiments (>2 days). At high food concentrations, juvenile scallops showed production of pseudofeces, partial shell valve closure, and reduction in feeding. According to HPLC analysis, the only toxins present in the dinoflagellate G. catenatum and in the scallops were the gonyautoxins (GTXs), except in the labial palps and digestive gland, where trace amounts of saxitoxin (STX) were present in scallops. These tissues could play an important role in toxin biotransformation. The ranking of toxin concentration in tissues was: digestive gland > labial palps > intestine > gills > mantle > adductor muscle, where the total contribution of viscera was more than 80% of the total toxin body burden. Juvenile scallops exhibited no apparent detrimental physiological responses during the long-term feeding experiment. The dinoflagellate may contribute nutrients to the scallop, in addition to the microalgae I. galbana. The dinoflagellate may enhance cell uptake and byssus production. Once PSP accumulated during the first 12 days, it was slowly eliminated. The limited capacity for accumulating toxins in the adductor muscle favors domestic marketing of scallops.     

An electron-microscope investigation of the cells lining the outer surface of the mantle in some marine molluscs

The ultrastructural details of the cells lining the outer surface of the mantle in Mytilus edulis (L), Cardium edule (L), and Nucula sulcata Bronn. are described. Along the length of the outer mantle fold and the general outer surface of the mantle, the epithelial cells differ, with a progressive reduction in the protein synthetic apparatus and an increase in glycogen, while large numbers of mitochondria are evident in the apical and basal regions.     

Experimental analysis of byssus thread production by Mytilus edulis and Modiolus modiolus in sediments

Laboratory experiments have been conducted on byssus thread production by two species of mussel, Mytilus edulis L. and Modiolus modiolus (L.), representing the epibyssate and endobyssate species of the Mytilidae, respectively. Mussels were placed in seven particle sizes of sediment ranging from 50 m to 16 mm for 12 d. The number of byssus threads per mussel, length of threads, number of threads per particle and size of pads were then recorded. Modiolus modiolus (endobyssate) produced more threads than Mytilus edulis (epibyssate). M. edulis produced most threads in the size range 2 to 16 mm and Modiolus modiolus in the size range 500 to 1000 m. M. modiolus produced longer threads than Mytilus edulis. Both species produced longer threads in particle sizes finer than 2 mm. M. edulis produced smaller pads than Modiolus modiolus. Both species produced larger pads in sediments coarser than 1 000 m. Results were also analysed in terms of attachment units (a thread attached to one or more particles, or a particle to which more than one thread is attached). The thread:particle ratio of the attachment units ranged from ca. 1:10 in the finest sediments to ca. 10:1 in the coarsest sediments. Mytilus edulis attached more threads to single particles (72% of attachment units) than Modiolus modiolus (37% of attachment units). M. modiolus had a wider spread of ratios in the finer particle sizes than Mytilus edulis. The significance of our results are discussed in relation to the ecology and palaeoecology of epibyssate and endobysste species.