Functional responses associated with pallial organ variations in the Pacific oyster Crassostrea gigas (Thunberg, 1793)

The plasticity and function of the pallial organs were studied in the Pacific oyster Crassostrea gigas from three sites of Bourgneuf Bay (French Atlantic coast, 46-47°N, 1-2°W) characterized by different turbidity conditions. Labial palp area was closely and positively related to the turbidity gradient. No clear pattern was established between the gill area and the gradient of suspended particulate matter (SPM). The functional responses induced by these morphological variations were investigated in the laboratory by means of ecophysiological experiments and endoscopic observations. Oysters with different pallial organ areas were supplied with mixed suspensions of heat-killed Tetraselmis suecica and living Skeletonema costatum added to different concentrations of kaolinite to simulate low (SPM = 8.5 ± 0.4 mg l^− 1) and high (SPM = 48.3 ± 1.4 mg l^− 1) turbidity conditions. At each SPM concentration, heat-killed T. suecica were preferentially rejected in pseudofaeces compared to S. costatum, indicating a preingestive particle selection. At low seston load, clearance rate (CR) was closely and positively related to gill area and particle selection occurred only on the gills, between the ventral grooves and dorsal tracts. At higher seston load, palps exhibited a particle-sorting capacity dependent on gill area. Indeed, with small gills, an increase in selection efficiency (SE) and CR was positively related to palp area. On the other hand, large gills processed the particles without an effect of palps but with a decrease in CR. The functional responses associated with pallial organ variations clearly showed that the preingestive particle processing in oysters is an integrated mechanism dependant on the gill and labial palp areas.     

Filter feeding,deviations from bilateral symmetry,developmental noise,and heterochrony of hemichordate and cephalochordate gills

We measured gill slit fluctuating asymmetry (FA), a measure of developmental noise, in adults of three invertebrate deuterostomes with different feeding modes: the cephalochordate Branchiostoma floridae (an obligate filter feeder), the enteropneusts Protoglossus graveolens (a facultative filter feeder/deposit feeder) and Saccoglossus bromophenolosus (a deposit feeder). FA was substantially and significantly low in B. floridae and P. graveolens and high in S. bromophenolosus. Our results suggest that the gills of species that have experienced a relaxation of the filter feeding trait exhibit elevated FA. We found that the timing of development of the secondary collagenous gill bars, compared to the primary gill bars, was highly variable in P. graveolens but not the other two species, demonstrating an independence of gill FA from gill bar heterochrony. We also discovered the occasional ectopic expression of a second set of paired gills posterior to the first set of gills in the enteropneusts and that these were more common in S. bromophenolosus. Moreover, our finding that gill slits in enteropneusts exhibit bilateral symmetry suggests that the left‐sidedness of larval cephalochordate gills, and the directional asymmetry of Cambrian stylophoran echinoderm fossil gills, evolved independently from a bilaterally symmetrical ancestor.     

INCREASED NUMBERS OF ANNULATE LAMELLAE IN MYOCARDIUM OF CHICK EMBRYOS INCUBATED AT ABNORMAL TEMPERATURES

Annulate lamellae have been observed in the myocardium of 18-day-old chick embryos maintained at the normal temperature of 100°F and at 90°F during the last week of incubation. An increased number of annulate lamellae was observed in heart muscle of embryos incubated at 90°F. This is probably caused by a persistent production of these organelles, since annulate lamellae are present in greater frequency than in 11-day-old embryos incubated at 100°F. In the hypertrophic hearts of 18-day-old embryos incubated at 90°F, the annulate lamellae were associated with a net increase of protein content and an elevated concentration of myocardial glycogen. It is suggested that the increased number of annulate lamellae is a sequela of reduced environmental temperature during incubation.     

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.     

Gross structure of the respiratory organs and dimensions of the gill in the mud–skipper, Periophthalmodon schlosseri (Bleeker)

In Periopkrlialnwdon scldosseri the respiratory organs consist of the gills, the suprabranchial and opercular chambers. The gills are more suited for aerial than aquatic respiration as is shown by the presence of the vascular papillae, blood sinusesand dilated blood vessels in their lamellae. The gill lamellae possess a surface coat of sulphated mucopolysaccharides that prevents water loss during exposure to the air. The filaments of the outer hemibranchs in the first gill arch are reduced to nearly one quarter of those of its posterior hemibranch. The gill area in relation to body weight shows a high slope value ( b =0·93).     

Early development and microhabitat of the monogenean Horricauda rhinobatidis, with observations on the related Troglocephalus rhinobatidis, from Rhinobatos batillum from Queensland,Australia

The small monocotylid monogenean Horricauda rhinobatidis is abundant on the gills of its host Rhinobatos batillum, whereas the larger, related monogenean Troglocephalus rhinobatidis is comparatively uncommon. Young specimens of Horricauda live between the host's secondary gill lamellae. Post-oncomiracidia have 14 marginal booklets but as the larvae develop these are supplemented first by a pair of hamuli and then by muscular ventral loculi followed by six forwardly-directed, dorsal spines. By impaling secondary gill lamellae these spines may serve to prevent parasites from being dislodged by gill ventilating currents. Before reaching sexual maturity the parasites leave the secondary gill lamellae and establish themselves in the septal canals. It is uncommon to find more than one adult specimen of Horricauda in each septal canal. The significance of this in relation to sperm exchange is discussed. Like Troglocephalus, Horricauda has eight head sacs, and the suggestion is made that these sacs may play a part in feeding.     

Ciliary ultrastructure of protobranchs (Mollusca, Bivalvia)

Abstract. The external epithelial cilia and other surface structures of the nuculoid protobranchs Nuculana pernula and Nucula nitidosa were studied. The gill lamellae and labial palps are partly covered with very long cilia. These have a modified slender distal portion, an ordinary metazoan-type basal body, a basal foot. and a single, long cross-striated rootlet. In cilia on the gills of N. nitidosa , the basal foot is thick and attaches to the next basal body directly behind. Unciliated surface areas on the gills, labial palps, and foot are covered with a dense brushborder of microvilli. We observed no specific homologies between the cilia of the protobranchs studied and the epidermal cilia of the enigmatic Xenoturbella bocki , hence the recent hypothesis of a close connection of the latter to the protobranch bivalves is questioned.     

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.     

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.     

Larval development and metamorphosis in the marine pulmonate Amphibola crenata (Mollusca: Pulmonata)

The development of the free-swimming veliger of Amphibola is followed from hatching to settlement, and the larval structures compared with those of post-metamorphic juveniles and adult snails. Observations of living specimens and light-microscope sections were combined with scanning electron microscopy to build up a composite picture of veliger structure.
Four stages in the development of veligers are recognized, each being characterized by the appearance of organ systems such as the mantle cavity, larval heart, adult heart and kidney, and larval pallial gland. At or after metamorphosis, the larval systems (heart, kidney and pallial gland) disappear, and the developing adult organs move to the positions characteristic of adult snails.
Organogenesis in Amphibola veligers is compared with that of prosobranch and opisthobranch larvae, and with that of pulmonate larvae with direct development. The closest similarity is seen to be with opisthobranch veligers.     

Structural organization of the gills in pipefish (Teleostei,Syngnathidae)

Summary The morphology and structural features of the gills of the two Western Baltic pipefish Nerophis ophidion and Syngnathus rostellatus were investigated using scanning electron microscopy. The general anatomy of the gills complies with the general pattern in fish. Several adaptations though, show the highly specialized nature of pipefish gills. The filaments are extremely short, few in number and carry only a few lamellae due to the limited space in the branchial cavity. The lamellae have a widely projecting form yet still have a small area in comparison to other fish. Gill irrigation is performed by a specialized pumping mechanism which forces respiratory water through the small but densely packed gill sieve. Although both species live in the same habitat and belong to the same family, differences in gill morphology were found and are related to different lifestyles. S. rostellatus is the more active species and therefore has more filaments per gill arch, more lamellae per filament, wider projecting lamellae and a more extreme utilisation of available space in the gill cavity through a very densely packed gill sieve. N. ophidion has a stationary mode of life and therefore has a less extreme gill anatomy.     

Impact of seston characteristics on qualitative particle selection sites and efficiencies in the pseudolamellibranch bivalve Crassostrea gigas

To date, knowledge of the qualitative particle selection sites and conditions in the widely-distributed bivalve Crassostrea gigas is incomplete, having focussed either on heterogeneous particles, or on particles intentionally too large to enter the gill principal filament tracts. We used endoscope-directed sampling and the intact diatom-empty, cleaned frustule approach to unambiguously establish qualitative selection sites and the influence of seston quality (varying proportions of intact diatoms and empty, cleaned frustules) and quantity (particle loads) on the degree of qualitative selection. Normally-feeding oysters were presented test mixtures of the naturally-occurring Actinoptychus senarius (small enough to enter the gill principal filaments), and the potential selection sites (gill: dorsal and ventral collecting tracts; labial palps: anteriorly-deposited pseudofaeces), were sampled for comparison with the proportions and concentrations of the ambient medium. Qualitative selection was demonstrated at both the gills and labial palps. Gill selection efficiency was shown to be directly proportional to seston quality and quantity, using a technique independant of pseudofeces mucus content. The oyster gill is thus able to increase ingested food quality when environmental food quality is low and / or when seston concentrations are high, which is typical of oyster habitats. Palp selection efficiency was directly proportional to seston quality, but at the highest concentration tested, no qualitative selection was observed on the labial palps, probably due to overload on these smaller organs. The partial functional redundancy of these key processing organs in heterorhabdic species such as oysters and scallops may enhance their success in high-turbidity habitats.     

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.     

Interspecific variations in the surface ultrastructure of the gills of freshwater mullets

SEM studies were made on the gills of freshwater mullets,Rhinomugil corsula andSicamugil cascasia, to correlate surface ultrastructure of various gill units with their probable functions. Two types of lamellated gill rakers of the former fish are suited for plankton feeding and the short, stumpy and transversely beaded gill rakers of the latter reflect the varied food and feeding habit of the fish.R. corsula has numerous mucous glands on the epithelium covering the gill arch and gill filaments,S. cascasia has fewer. In accordance with the differences in the density and distribution of the mucous glands, the microridged epithelial cells also show variations in their architectural plan. In both species the epithelium of the secondary lamellae is smooth, probably an adaptation for better gaseous exchange.     

A morphological study on posterior gills of the mangrove crab Ucides cordatus

Morphological and histological studies on posterior gills of the mangrove crab Ucides cordatus showed that the 5th gill (of 7) has a larger surface area and a greater number of lamellae compared to the 6th gill. Regular separation of gill lamellae, important when the gill is in air, is maintained by enlargements of the marginal canals. Conical, spine-like structures along the efferent vessel of both 5th and 6th gills were also observed. In addition, pillar cells, a discontinuous lamellar septum and a hypobranchial artery were observed. The presence of valve-like structures near the efferent vessel was also indicated. These structures, together with the pillar cells, may have a role in directing the hemolymph flow towards certain gills during particular physiological states. Localization of osmoregulatory epithelia in the lamellae of both gills was inferred from dimethylaminostyrylethylpyridiniumiodine staining. Apparently gills 5 and 6 have osmoregulatory epithelial cell patches of similar area, corresponding to 43% and 38% of the total lamellae area, respectively. However, their localization is quite different. Gill number 5 osmoregulatory patches seem to be restricted to the afferent region of the lamella whereas in gill number 6, they are more dispersed over the entire lamella. These differences may be related to the particular functional characteristics of these gills.     

Embryonic encapsulation and maternal incubation: Requirements for survival of the early stages of the estuarine gastropod Crepipatella dilatata

During their reproductive period, females of Crepipatella dilatata deposit their embryos in capsules that they then brood in the pallial cavity until juveniles emerge several weeks later, after passing through a transient veliger “larval” stage. Artificially excapsulated veligers of this species experimentally exposed to a wide range of salinities (5, 10, 15, 20, 25, and 30 psu) for six hours showed reduced activity at salinities of 15 and 20 psu, whereas encapsulated veligers exposed to those same salinities showed no reduction of activity. Artificially excapsulated veligers showed high mortality at salinities of 5 and 10 psu; encapsulated embryonic stages also showed high mortalities at 5 psu and serious sublethal effects at 10 psu in tests excluding maternal protection, showing that encapsulation alone does not provide complete protection from low salinity stress. Natural tidal cycles in the Quempillén River estuary also reduced embryonic survival at salinities of ≤ 10 psu when the capsules were exposed without maternal protection. In contrast, encapsulated embryos protected by their mothers survived well regardless of the salinity to which they were exposed, under both natural and laboratory-simulated estuarine tidal cycles. C. dilatata are able to develop in the estuary only because of maternal protection, since salinity levels in this environment sometimes decline to as low as 7 psu. Successful embryonic development in this estuary reflects the capacity of C. dilatata adults to detect dangerously low salinity levels and then seal themselves off from the environment for up to 50 hrs (O. Chaparro pers. obs.) when the salinity drops below 22.5 psu, allowing salinity to remain above this level within the pallial cavity despite continued salinity declines in the surrounding seawater.     

Gross morphology and surface ultrastructure of the gills of Odontesthes argentinensis (Actinopterygii, Atherinopsidae) from a Southwestern Atlantic coastal lagoon

Odontesthes argentinensis was collected from Mar Chiquita Coastal Lagoon, the Southernmost coastal Atlantic Lagoon of Argentina. The morphology of the gills was analyzed by scanning electron microscopy. The morphology of the superficial structures of the gill filaments and pharyngeal region of the gill arch was discussed and related to their functional aspects. The gills arches are structurally similar to those of other teleosts and bring out the osmoregulatory capacity of this species. The epithelium that covers the surface of the filaments and the pharyngeal region of the gill arch is formed by polygonal pavement cells with conspicuous microridges. These folds in the membrane are not denoted in the epithelium of the respiratory lamellae. Apical crypts of chloride cells are present on the afferent and interlamellar filament surfaces, but are absent elsewhere on the gill arch. The highest density of mucous cells is observed into the gill filament and the pharyngeal region which indicates the existence of a protective strategy of the respiratory lamellae and the pharynx. The epithelium of the gill arches and the rakers is studded with spines. There are taste buds along the whole pharyngeal region that may be associated with their participation in tasting at this zone.     

The mucous coat on gill lamellae of rainbow trout (Oncorhynchus mykiss)

The existence of a layer of mucus covering the gill lamellae of healthy rainbow trout (Oncorhynchus mykiss) was investigated. Using cryo-scanning electron microscopy, a smooth, undulating, thin layer was observed which completely covered gill filaments and lamellae, thereby obscuring epithelial microridges. After processing cryopreserved gill arches in glutaraldehyde for conventional scanning electron microscopy, the layer was no longer present and epithelial microridges were clearly visible. The identity of this layer was investigated using cryopreserved gills which were treated in one of two ways. First, gills were incubated with a rabbit antiserum to gill mucus, with normal rabbit serum, or with phosphate-buffered saline. Following fixation in glutaraldehyde and processing, only the gill tissue incubated with the mucus-specific antiserum was still covered with the smooth layer. The layer was also retained on the gills of fish anesthetized in a solution containing mucusspecific antiserum and then processes in glutaraldehyde for conventional scanning electron microscopy. The tenacious nature of the mucous layer was demonstrated by its stability following exposure to formalin and a cationic detergent. Second, the presence of this layer was confirmed on gill tissue which was cryopreserved, followed by freeze-substitution and vapor fixation, and then examined by transmission electron microscopy.     

Myoanatomy and serotonergic nervous system of the ctenostome Hislopia malayensis: evolutionary trends in bodyplan patterning of ectoprocta

Background

Near the end of the nineteenth century the hypothesis was presented for the homology of book lungs in arachnids and book gills in the horseshoe crab. Early studies with the light microscope showed that book gill lamellae are formed by outgrowth and possibly some invagination (infolding) of hypodermis (epithelium) from the posterior surface of opisthosomal limb buds. Scorpion book lungs are formed near the bilateral sites of earlier limb buds. Hypodermal invaginations in the ventral opisthosoma result in spiracles and sac-like cavities (atria). In early histological sections of embryo book lungs, widening of the atrial entrance of some lamellae (air channels, air sacs, saccules) was interpreted as an indication of invagination as hypothesized for book gill lamellae. The hypodermal infolding was thought to produce the many rows of lamellar precursor cells anterior to the atrium. The ultrastructure of scorpion book lung development is compared herein with earlier investigations of book gill formation.

Results

In scorpion embryos, there is ingression (inward migration) of atrial hypodermal cells rather than invagination or infolding of the atrial hypodermal layer. The ingressing cells proliferate and align in rows anterior to the atrium. Their apical-basal polarity results in primordial air channels among double rows of cells. The cuticular walls of the air channels are produced by secretion from the apical surfaces of the aligned cells. Since the precursor cells are in rows, their secreted product is also in rows (i.e., primordial air channels, saccules). For each double row of cells, their opposed basal surfaces are gradually separated by a hemolymph channel of increasing width.

Conclusions

The results from this and earlier studies show there are differences and similarities in the formation of book lung and book gill lamellae. The homology hypothesis for these respiratory organs is thus supported or not supported depending on which developmental features are emphasized. For both organs, when the epithelial cells are in position, their apical-basal polarity results in alternate page-like channels of hemolymph and air or water with outward directed hemolymph saccules for book gills and inward directed air saccules for book lungs.