The functional morphology of Mactrinula reevesii (Bivalvia: Mactroidea) in Hong Kong: adaptations for a deposit-feeding lifestyle

The shallow subtidal mactrid bivalve Mactrinula reevesii is a deposit-feeder in the southern and south-eastern oceanic waters of Hong Kong. Buried obliquely, large quantities of fine sediment are taken into the mantle cavity and sorted on enormous labial palps. The small ctenidia probably have little value in collecting material, amounts taken in being too large. The mid gut is long and complexly folded inside the visceral mass. It is also capable of distension, although superficial visceral muscles maintain internal tonus. The rectum is narrow and creates compact faecal pellets.
Most interest is in the ventral mantle margin which is, posterior to the pedal gape and the base of the inhalant siphon, united by a sheet of cuticle. There is no fourth pallial aperture. There are, however, two pairs of flaps extending along the posterior third of the internal ventral mantle surface. These arch over left and right mantle rejection tracts which transfer unwanted material to the base of the inhalant siphon for final expulsion. The mantle flaps prevent such material from being returned to the anterior end of the mantle cavity, for palp reprocessing, when new material arrives. They, thus, maximize sorting efficiency by separating unsorted from sorted and rejected material.
Other mactrids have similar mantle flaps which they use in different ways, including the channelling of unwanted material to a fourth pallial aperture for expulsion, as in Lutraria lutraria. The Mactridae have thus evolved a unique method of increasing the efficiency of pseudofaecal waste rejection which has thereby facilitated the deposit-feeding lifestyle by the diverse representatives of this family.     

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.     

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.     

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.     

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.     

The biology and functional morphology of Arca noae (Bivalvia: Arcidae) from the Adriatic Sea, Croatia, with a discussion on the evolution of the bivalve mantle margin

In the Croatian Adriatic, Arca noae occurs from the low intertidal to a depth of 60 m; it can live for > 15 years and is either solitary or forms byssally attached clumps with Modiolus barbatus. The shell is anteriorly foreshortened and posteriorly elongate. The major inhalant flow is from the posterior although a remnant anterior stream is retained. There are no anterior but huge posterior byssal retractor muscles and both anterior and posterior pedal retractors. The ctenidia are of Type B(1a) and the ctenidial–labial palp junction is Category 3. The ctenidia collect, filter and undertake the primary sorting of potential food in the inhalant water. The labial palps are small with simple re‐sorting tracks on the ridges of their inner surfaces. The ciliary currents of the mantle cavity appear largely concerned with the rejection of particulate material. The mantle margin comprises an outer and an (either) inner or middle fold. The outer fold is divided into outer and inner components that secrete the shell and are photo‐sensory, respectively. The latter bears a large number of pallial eyes, especially posteriorly. The inner/middle mantle fold of A. noae, possibly representative of simpler, more primitive conditions, may have differentiated into distinct folds in other recent representatives of the Bivalvia.     

THE FUNCTIONAL MORPHOLOGY OF THE DIGESTIVE SYSTEM OF LYONSIA HYALINA CONRAD, 1831 (BIVALVIA: ANOMALODESMATA: PANDOROIDEA)

The anatomy and histology of the digestive tract of the suspensionfeeding bivalve Lyonsia hyalina were examined using microdissectionand conventional light microscopy. Lyonsia hyalina has a typeIV stomach which contains a major typhlosole that does not penetratethe left pouch, as in other members of the Pandoroidea. Theventral and posterior sorting areas of the stomach are sitesof vigorous ciliary activity. The gastric shield is locatedon the left and posterior stomach walls, underlain by tall basophiliccells with microvilli that project into the gastric shield.The style sac and midgut are combined, and contain the morphologicalcell types A-D seen in other bivalves. Many ciliated cells ofthe digestive tract appear to have high densities of apicalmitochondria. The ducts within the digestive diverticula arelined by epithelia containing a conspicuous brush border. Bothcrypt cells and digestive cells exist in the digestive tubules.The presence of numerous fragmentation spherules throughoutthe digestive diverticula indicates that intracellular digestionoccurs there. (Received 21 April 1992; accepted 1 October 1992)     

PHOSPHATIZED SOFT-TISSUE IN TRIASSIC BIVALVES

Abstract: Exceptionally preserved specimens of the genera Myophoria, Neoschizodus and Trigonodus (Bivalvia, Palaeoheterodonta, Trigonioida) from carbonate sediments of the Muschelkalk (Anisian, Ladinian) contain phosphatized soft‐tissues. This is the oldest record of soft‐tissue preservation in bivalves and the first from the German Muschelkalk. The phosphatized remains are here interpreted as relics of the originally chitinous gill supports, the gill axis, the labial palps (?), the adductor muscles, the pedal retractor muscles, the mantle margin including the radial mantle musculature, and the ‘siphons’. According to microprobe analysis, the mineral replacing the soft‐tissue is mainly apatite and, more rarely, francolite. Additionally, quartz filled voids within the gill supports, and in one sample it occurs in minute crystals in the phosphatized remains of the adductor muscle of Neoschizodus. Myophoria, Neoschizodus and Trigonodus were soft‐bottom dwellers and five of the specimens were discovered in life position. This is indicated by geopetal structures in three specimens.     

Sexual dimorphism of adult labial palps of the peach fruit moth Carposina sasakii Matsumura (Lepidoptera: Carposinidae) with notes on their sensilla

The labial palps and their sensilla of the peach fruit moth Carposina sasakii Matsumura, a serious pest of pome fruits in eastern Asia, were investigated using light microscopy and scanning and transmission electron microscopy. The labial palps are three‐segmented and exhibit distinct sexual dimorphism in length: much longer in the female than in the male. Four types of sensilla were found on the labial palps: campaniform, squamiform, chaetic and flattened sensilla. The campaniform sensilla are present on the first segment alone. The squamiform sensilla are located on all the three segments. The chaetic sensilla are mainly present on the third segment. The flattened sensilla are grooved with wall pores and situated in a labial palp‐pit organ, which is located at the apex of the distal segment of the labial palp. The sexual dimorphism of the labial palp and the labial palp‐pit organ was briefly discussed.     

On the habits and nature of association of the copepod Pseudomyicola spinosus with the rock oyster Crassostrea glomerata in New Zealand

Both adults and early stages of Pseudomyicola spinosus (Cyclopoidea, Myicolidae) occur on the labial palps as well as in the gut of the rock oyster Crassostrea glomerata. There is evidence to suggest movements of the copepod to and from the gut. Clear evidence of damage to the epithelial walls of the gut is seen in several regions, particularly in confined portions of the stomach near the openings of the ducts of the digestive diverticula. In addition to mechanical injuries, changes from a columnar to a low squamous epithelium have been observed in one instance; an increase of leucocytes has also been observed in another specimen. The evidence suggests that the myicolid copepod is a parasite in the gut of the oyster.     

Ultrastructure of sensilla on the maxillary and labial palps of three species (Coleoptera: Staphylinidae: Staphylininae)

Most species of Staphylinidae are predators in an agroecosystem. They acquire prey information from their environment through receptors. In this study, the sensilla on maxillary and labial palps of Philonthus kailiensis, Philonthus lewisius and Quedius robustus were examined with scanning electron microscopy to identify and analyse the external morphology and distribution of the sensilla to enhance our knowledge of the sensilla of Staphylinidae and provide a rationale of taxonomical studies on the two genus. Results showed that the sensilla are classified into six types: Böhm bristles, sensilla chaetica, sensilla furcate, sensilla coeloconica, sensilla placodea and sensilla basiconica. No sexual dimorphism exists among the three species. The relationships and functions of sensilla on maxillary and labial palps were also speculated. There may be a certain correlation between the sensilla on maxillary and labial palps of the staphylinid and its habitat.     

Sensilla of the antennae and the labial and maxillary palps of Blattella germanica (L.) (Dictyoptera: Blattellidae): Their classification and distribution

The various sensilla on the antennae and on the labial and maxillary palps of Blattella germanica (L.) were studied. Thick-walled chemoreceptors with fluted shafts and articulated bases are located on the antennae and on the labial and maxillary palps. Thin-walled chemoreceptors, without fluted shafts or articulated bases, are restricted to the flagellar segments of the antennae and to the distal segments of the palps. Antennae of adult males have more thin-walled chemoreceptors than do those of females. Hair-plate sensilla are found at the scape-head and scape-pedicel joints, and at the joints of segments on the palps. Campaniform sensilla are concentrated as a ring around the distal margin of the pedicel, and are also scattered singly on the scape, pedicel, and flagellar segments of the antennae, and on the first segment of the maxillary palps. Occasionally, a few sensilla coeloconica and cold receptor sensilla are found on the antennal flagellum.     

Spirochetes in gastropods from Lake Baikal and North American freshwaters: new multi-family, multi-habitat host records

We describe the first records of spirochetes in the gut of fourteen species of continental gastropods from a range of habitats and representing six families (Amnicolidae, Baicaliidae, Bithyniidae, Pyrgulidae, Lithoglyphidae and Benedictiidae). The bacteria were mainly found in the crystalline style sac, as has been reported in marine bivalves. The surveyed habitats include water bodies in North America and Eurasia, including deep water hydrothermal vent and gas hydrate zones in Lake Baikal. Spirochetes were present both in mature and young snails, but were not detected in embryos before hatching, indicating lateral transfer. The surveyed gastropods range in trophic strategy, including phyto-, detrito- and bacteriophagous grazers and filter feeders. Our results indicate that spirochetes are commensal in the surveyed gastropods with potential limited benefit and no detriment to the host animal. We suggest that the specialized internal habitat of the crystalline style sac in molluscs is likely to reveal unrecognized spirochete diversity that will shed new light on gastropod trophic ecology and spirochete diversity.     

Anatomy of the feeding apparatus of the nurse shark,Ginglymostoma cirratum

The anatomy of the feeding apparatus of the nurse shark, Ginglymostoma cirratum, was investigated by gross dissection and computer axial tomography. The labial cartilages, jaws, jaw suspension, muscles, and ligaments of the head are described. Palatoquadrate cartilages articulate with the chondrocranium caudally by short, laterally projecting hyomandibulae and rostrally by ethmoorbital articulations. Short orbital processes of the palatoquadrates are joined to the ethmoid region of the chondrocranium by short, thin ethmopalatine ligaments. In addition, various ligaments, muscles, and the integument contribute to the suspension of the jaws. When the mouth is closed and the palatoquadrate retracted, the palatine process of the palatoquadrate is braced against the ventral surface of the nasal capsule and the ascending process of the palatoquadrate is in contact with the rostrodorsal end of the suborbital shelf. When the mandible is depressed and the palatoquadrate protrudes slightly rostroventrally, the palatoquadrate moves away from the chondrocranium. A dual articulation of the quadratomandibular joint restricts lateral movement between the mandible and the palatoquadrate. The vertically oriented preorbitalis muscle spans the gape and is hypothesized to contribute to the generation of powerful crushing forces for its hard prey. The attachment of the preorbitalis to the prominent labial cartilages is also hypothesized to assist in the retraction of the labial cartilages during jaw closure. Separate levator palatoquadrati and spiracularis muscles, which are longitudinally oriented and attach the chondrocranium to the palatoquadrate, are hypothesized to assist in the retraction of the palatoquadrate during the recovery phase of feeding kinematics. Morphological specializations for suction feeding that contribute to large subambient suction pressures include hypertrophied coracohyoideus and coracobranchiales muscles to depress the hyoid and branchial arches, a small oral aperture with well‐developed labial cartilages that occlude the gape laterally, and small teeth. J. Morphol. 241:33–60, 1999. © 1999 Wiley‐Liss, Inc.     

小菜蛾成虫下唇须感器的超微结构及其神经元中枢投射

【目的】明确小菜蛾Plutella xylostella成虫下唇须感器的形态结构及感器神经元的投射。【方法】利用光学显微镜观察和扫描电子显微镜观察下唇须结构和感器类型,利用神经回填技术和激光共聚焦显微镜观察下唇须感器神经元在脑部的投射。【结果】小菜蛾成虫下唇须共3节,其上存在Böhm氏鬃毛、钟形感器、鳞形感器、锥形感器、微毛形感器5种不同类型的感器和一个陷窝器结构。Böhm氏鬃毛短小尖细,钟形感器形如顶部凹陷的圆帽,两种感器均分布于下唇须第1节,且大小上均无雌雄二型差异;鳞形感器形同柳叶,锥形感器粗而直,均散生于下唇须的第2和3节,两种感器在大小上均存在雌雄二型差异,其中雌性的鳞形感器显著大于雄性的,根据其雌雄二型差异现象推测雌蛾的鳞形感器可能与感受寄主植物挥发物有关;下唇须第3节中上部具有一个圆形陷窝器结构,雄虫的陷窝器内径为5.68±0.33μm,雌虫的为6.03±0.23μm,雌雄间无显著性差异;凹坑内长有表面光滑的微毛形感器。小菜蛾下唇须感器神经元主要投射于脑部咽下神经节、每个触角叶的下唇须陷窝器神经纤维球和腹神经索3条通路。【结论】阐明了小菜蛾下唇须感器的类型、分布和形态特征及其感器神经元在脑部的投射形态,为深入了解小菜蛾下唇须感器的生理和功能奠定了基础。     

Functional validation of the carbon dioxide receptor in labial palps of Helicoverpa armigera moths

Adult moths possess an organ in their labial palps, the labial-palp pit organ, which is specialized for sensing carbon dioxide (CO₂). They use CO₂ as a cue to detect healthy plants and find food or lay eggs on them. The molecular bases of the CO₂ receptor in Drosophila melanogaster and Aedes aegypti have been reported, but the molecular mechanisms of the CO₂ receptor in Lepidoptera remains elusive. In this study, we first re-examined three putative Helicoverpa armigera CO₂ gustatory receptor genes (HarmGr1, HarmGr2, and HarmGr3), and then analyzed expression patterns of them. RT-PCR results verified they were predominantly expressed in the labial palps of H. armigera. Thus, we used in situ hybridization to localize the expression of three genes in the labial palps. We found that all three genes were co-expressed in the same cells of the labial palps. Next, we employed the Xenopus laevis oocyte expression system and the two-electrode voltage-clamp recording to study the function of the three genes. Results showed that only oocytes co-expressing HarmGr1 and HarmGr3 or co-expressing HarmGr1, HarmGr2 and HarmGr3 gave robust responses to NaHCO₃. Finally, we confirmed that the sensory cells in labial palps of both females and males show dose dependent responses to CO₂ stimuli by using single sensillum recording. Our work uncovers that HarmGr1 and HarmGr3 are indispensable and sufficient for CO₂ sensing in labial palps of H. armigera.     

七星瓢虫唇须感器分布和统计特征的雌雄差异

应用扫描电镜观察七星瓢虫Coccinella septempunctata的唇须和附节,以便确认在感应农药残留时起主要作用的化学感受器。研究证明分布在下颚须上的感器类型包括两种锥形感器（兼有机械感受器和化学感受器的功能）和一种钟形感器。同时本文首次报道了对这些感器精确的形态测量,明确了其性二型性。对下颚须的测量表明其左右末节的长度差异与七星瓢虫的性别之间有明显的相关性。文中还记述了下唇须上一种新的钟形感器,其在每个下唇须上的数量大约为12～17个。经测量这些机械感受器的平均直径为2.4 μm。早期研究中所记述的锥形感器被证实是下唇须上仅有的化学感应器,并且首次证明这些锥形感器的数量与性别相关,雄性平均为18个,雌性平均为16个。附节上也存在性别差异： 第2跗节跗垫的宽度在雄性中是从后向前逐渐增大,而在雌性中刚好相反。本文还就这些差异的意义以及新发现感器的功能进行了讨论。     

Ultra morphology of sensilla on the head and mouth parts of the earwig Anisolabis martima (Bonelli) (Dermaptera: Carcinophoridae)

The earwig Anisolabis maritima is a cosmopolitan insect with a worldwide distribution, omnivorous in feeding, eating the eggs of different insects. Recently recorded in Saudi Arabia as a beneficial predator on eggs and newly hatched larvae of the red palm weevil Rhyncophrous ferrugineus, scanning electron microscopy (SEM) observations revealed the presence of different kinds of receptor sensillae occurring on the head and mouth parts of A. maritime. The sensillae are the same, but differ in numbers on both sexes. The head surface bears three subtypes of trichoid sensillae, one type of ceoloconica. Labrum, labium, labial palps and maxillary palps, possess different kinds of sensillae. Three subtypes of trichoid have been identified on the labrum. The labium contained one type of tricoid and basiconic sensillae. There are great numbers and numerous sensillae cheatica differing in length distributed on the maxillary and the labial palps on all segments between other stutter spines. The last terminal segment of maxillary or labial palps bears the greatest number of the sensillae cheatica. Adult A. maritima may be introduced in the future as a beneficial predator of eggs of R. ferrugineus as a tool for integrated pest management.     

Beak gape dynamics during song in the zebra finch

Bird song is a complex communication behavior that requires the coordination of several motor systems. Sound is produced in the syrinx and then modified by the upper vocal tract, but the specific nature and dynamics of this modification are not well understood. To determine the contribution of beak movements to sound modification, we studied the beak gape patterns in zebra finches (Taeniopygia guttata). Subsyringeal air sac pressure and song were recorded together with changes in beak gape, which were monitored with a magneto-sensitive transducer. Beak gape was positively correlated with fundamental frequency, peak frequency, and subsyringeal air sac pressure in all but one bird. For harmonic stacks, peak frequency increased with increasing beak gape, and the relationship between fundamental frequency and beak gape was no longer significant. Experimentally holding the beak open or closed had acoustic consequences consistent with the model in which beak movements change upper vocal tract length and, thus, the filter properties. Beak gape was positively correlated with sound amplitude in all but two birds. The relationship between beak aperture and amplitude may, however, be indirect because air sac pressure is correlated with amplitude and beak gape. The beak is opened quickly and to its widest aperture immediately prior to the onset of sound and at rapid transitions in sound, suggesting that beak movements may affect vibratory behavior of the labia.     

Expression of odorant-binding and chemosensory proteins and spatial map of chemosensilla on labial palps of Locusta migratoria (Orthoptera: Acrididae)

The sensilla on labial palps in Locusta migratoria were observed and mapped using light microscopy, scanning and transmission electron microscopy. A dome region on the tip of the fourth segment (distal segment) of labial palps is mainly covered with sensilla chaetica (about 98%), and few sensilla basiconica (2%). The total number of both types of sensilla is significantly higher in females than in males. Sensilla chaetica can be further subdivided into three groups containing 6, 7 or 10 neurons. Immunocytochemical localization of odorant-binding protein (OBP) and chemosensory proteins (CSPs) was performed on ultrathin sections of sensilla on labial palps. The antiserum against odorant-binding protein from Locusta migratoria (LmigOBP) only labelled sensilla basiconica, with gold granules only found in the sensillum lymph. Chemosensory protein instead was specifically present in the outer sensillum lymph of all three subgroups of sensilla chaetica with antiserum against CSP-I from Schistocerca gregaria (SgreCSP-I). In contrast these three subgroups were never labelled with antiserum against CSP-II from Locusta migratoria (LmigCSP-II). In addition, a few sensilla chaetica could not be stained with any of the antisera used.