淡水贝类贝壳多层构造形成研究

对几种淡水贝（包括蚌、螺）进行形态及组织学观察，并通过实验方法重现贝壳三种物质，即：角质、棱柱质、珍珠质的生成过程，结果表明：外套膜外表皮细胞是由相同类型细胞组成，这些相同细胞在不同的作用条件下形成贝壳多层构造。     

Mechanics of sculpture formation in Magadiceramus? rangatira rangatira (Inoceramidae,Bivalvia) from the Upper Cretaceous of New Zealand

The surface sculpture of the inoceramid bivalve Magadiceramus? rangatira rangatira consists of commarginal ribs and curious, transverse wrinkles. The wrinkles typically are at a high angle or orthogonal to the shell margin (‘antimarginal’) and thus differ from purely radial structures. They show features of distribution and morphology that reveal them to be products of margin‐parallel compression of the shell‐secreting mantle and its adjacent, flexible, uncalcified periostracum. The interaction of wrinkles with commarginal ribs indicates that the ribs also formed as folds of the mantle margin. During growth, commarginal folding caused withdrawal of the entire mantle margin towards the umbo, with a consequent reduction in perimeter length. Measurement of specimens indicates that fabrication of the commarginal ribs resulted in the magnitude of commarginal shortening that is required for the formation of transverse wrinkles. We infer that early in ontogeny, at the first development of these sculptures, the wrinkles resulted entirely from mantle contraction and resultant commarginal shortening. With subsequent growth, total wrinkling included a component of ‘pre‐wrinkling’ inherited from the preceding growth stage; the contribution of pre‐wrinkling to total wrinkling increased with shell size. The proposed mechanical model is two‐phase. First, the transversely corrugated (pre‐wrinkled) mantle and periostracum advanced and secreted a slightly concave growth increment. Secondly, the mantle subsequently contracted to create a commarginal rib and increase the number and amplitude of transverse wrinkles. This model is consistent with a homogeneous mantle lacking any differentiated and specialized rib‐constructing segments.     

Intermediary metabolism and shell growth in the brachiopod Terebratalia transversa

Juvenile Terebratalia transversa (Brachiopoda) metabolize carbohydrates in the anterior-most marginal mantle at a rate of 0.46 μM glucose/g/hr (in vitro incubation of mantle in C¹⁴-glucose in a carrying medium of 10^-3 M non-radioactive glucose). The rate declines to 0.18μM glucose/g/hr in full-grown specimens. Carbohydrate metabolism in the marginal (anterior-most) mantle averages approximately 3.7 times greater than metabolism in (a portion of the ‘posterior’) mantle situated between the coelomic canals and the marginal mantle. This ratio remains constant in specimens of all sizes (i.e. an ontogenetic trend in the ratio is absent at p≤ 0.05). Organic acids are not detectable within the mantle (HPLC techniques) even after simulated anoxia (N₂ bubbling during mantle incubation). Glucose metabolism in vitro declines in both the marginal and ‘posterior’ mantles during anoxia and the metabolic ratio between marginal/‘posterior’ mantles becomes 1/1. We found no difference (at p≤ 0.05) in mean metabolic activity or in sue-related metabolic trends among populations from depths ranging between mean sea level and 70 m. However, the activity within the ‘posterior’ mantle was more variable in specimens from 70 m than in those from shallower habitats (10 m - mean sea level). The size of the specimens analyzed was most variable in the groups obtained from the shallowest habitats and least variable at 70 m depth. Our results may help define the energetics of fossil as well as living brachiopod shell growth. Brachiopod shell growth is known to be very slow relative to that of bivalves and our results indicate that this is a result of the animals' slow metabolism. The inflation of the valves in T. transversa is, in part, a function of the high ratio of intermediary metabolism in the marginal vs‘posterior’ mantle (i.e. parallels the relative growth rates at the shell margin vs‘posterior’ areas). We found that the bivalve, Chlamys hastata, which is commonly associated with T. transversa, has a lower ratio of metabolic activities in the ventral/dorsal mantle areas than the brachiopod has in the anterior/posterior. The difference produces a flatter shell in the bivalve in accord with allometric principles. The higher metabolic rate in the marginal vs‘posterior’ brachiopod mantle and its more pronounced decline with anaerobiosis is reflected in the greater definition of growth increments in the outer shell layer. Our results do not support recent generalizations that correlate shell thickness of a wide variety of invertebrates inversely with metabolic rate. Growth rate as determined from width of shell growth increments is a better index of metabolic rate. Although the genetic basis of glucose metabolism is unknown, the observed metabolic variability is consistent with suggestions that populations of marine organisms living in stable offshore environments are genetically more variable but morphologically more uniform than populations from shallow water. Furthermore, our results support suggestions that bivalved molluscs and brachiopods are very different metabolically, but the data are neutral with respect to theories of competitive exclusion of the two taxa throughout geologic history.     

The development of growth lines on articulate brachiopods

Three types of growth lines are recognised on articulate brachiopod shells: (1) very fine diurnal growth lines formed by calcite increments at the shell margin, (2) seasonal growth lines, formed by inward reflection (doubling back) of the mantle edge, seen as concentric steps on the shell surface and marked by re-orientation of growth vectors evidenced by secondary shell fibres, (3) disturbance lines, formed by abrupt regression of the mantle edge, also seen as concentric steps on the shell surface, but indicated by a dislocation in the shell fabric. Lamellose and spinose ornaments of the sort seen in Tegulorhynchia are essentially genetically controlled. Periodic outgrowths from the outer mantle lobe secrete frills of primary shell that project from the shell surface and form short hollow spines where they cross the radial ornament. In longitudinal section spine formation is seen to involve gradual increase in the rate of secretion of primary shell followed by retraction, and often collapse, of the mantle outgrowth, accompanied by regression. Reflection of the mantle edge usually follows spine formation.     

Bivalves with 'concrete overcoats': Granicorium and Samarangia

Two veneroidean bivalves Granicorium indutum from Australia and Samarangia quadrangularis from the tropical Indo-Pacific region, cement a thick, hard layer of sand over most of their shells. In Granicorium this layer forms low commarginal ribs while in Samarangia it forms more prominent radial features. Sand grains are cemented to the shell and to each other with growths of a crystalline aragonitic cement similar in morphology to inorganic marine cements. Both species secrete mucus layers at the growing shell margin which initially hold the sediment grains together and form a substrate for the nucleation and growth of calcium carbonate crystals. The ribs of Samarangia are formed by the accretion of successive sheets of spherulitic growths. In G. indutum , the middle and outermost of two inner mantle folds are large, glandular and capable of considerable extension beyond the shell margin. Mucus secreted by the folds contains abundant bacteria and small calcium carbonate crystals. It is proposed that initial nucleation of the calcium carbonate cement takes place within this biofilm possibly mediated by the bacteria. The function of the sand layers is unknown but predation resistance and protection of the shells from endobionts are the most likely possibilities.     

Origins,bottlenecks, and present‐day diversity: Patterns of morphospace occupation in marine bivalves

It has long been known that species should not be distributed randomly in morphospace (a multidimensional trait space), even under simple models of evolution. However, recent studies suggest that position in morphospace can affect aspects of evolution such as the durations of clades and the species richness of their constituent taxa. Here we investigate the dynamics of morphospace occupancy in living and fossil marine bivalves using shell size and aspect ratio, two functionally important traits. Multiple lines of evidence indicate that the center of a family's morphospace today represents a location where taxonomic diversity is maximized, apparently owing to lower extinction rates. Within individual bivalve families, species with narrow geographic ranges are distributed throughout the morphospace but widespread species, which are generally expected to be extinction resistant, tend to be concentrated near the center. The morphospace centers of most species‐rich families today (defined as the median value for all species in the family) tend to be close to the positions of the family founders, further suggesting an association between position in morphospace and net diversification rates. However, trajectories of individual subclades (genera) are inconsistent with the center of morphospace being an evolutionary attractor.     

A possible mechanism for the formation of annual growth lines in bivalve shells

We report a unique shell margin that differed from the usual shell structure of Pinctada fucata. We observed empty organic envelopes in the prismatic layer and the formation of the nacreous layer in the shell margin. All the characteristics of the growing margin indicated that the shell was growing rapidly. To explain this anomaly, we propose the concept of “jumping development”. During jumping development, the center of growth in the bivalve shell jumps forward over a short time interval when the position of the mantle changes. Jumping development explains the unusual structure of the anomalous shell and the development of annual growth lines in typical shells. Annual growth lines are the result of a discontinuity in the shell microstructure induced by jumping development.     

A clockwork mollusc: Ultradian rhythms in bivalve activity revealed by digital photography

Time-lapse digital images can be acquired and archived using web-cameras, allowing non-invasive analysis of behavior patterns of bivalve molluscs at ultradian (sub-daily) time-scales over long intervals. These records can be analyzed directly by a human operator or through properly calibrated image analysis software. Preliminary results using species of marine and freshwater bivalves identify several ultradian biological rhythms of similar duration. Wavelet analysis indicates strong periodicity in mantle and siphon activity in the 3 to 7 min range, with longer duration shell contraction periods at 60-90 min. The recurrence of these rhythms among marine and freshwater bivalve species maintained under constant (but differing) conditions suggests the influence of common intrinsic drivers (chemico-physical mechanisms or biological clocks). Sub-daily growth increments preserved in the shells of rapidly growing bivalve species are potentially related to these biological rhythms, with implications for shell growth, biomineralization, and the temporal resolution of paleoclimate proxy data.     

Characteristics of shell microstructure and growth analysis of the Antarctic bivalve <Emphasis Type="Italic">Laternula elliptica</Emphasis> from Lützow-Holm Bay,Antarctica

Growth performance of the Antarctic bivalve Laternula elliptica was examined both by shell microstructural observation and by applying a fluorescent substance, tetracycline, as a shell growth marker. The shell was composed of two calcareous layers: the thick outer layer was homogeneous or granular in structure and the thin inner layer was nacreous. The architecture of Antarctic L. elliptica was different from that of temperate L. marilina, and the ratio of thickness between the outer and inner layers appeared to be different. The growth rate of the nacreous layer was analyzed to be very low. High correlations were found between the major axis of chondrophore and both shell length and shell dry weight, respectively. It is suggested that the chondrophore is an appropriate growth indicator, and combining the information of growth increments with the fluorescent method may be useful in estimating the bivalve growth performance in the Antarctic sea.     

Correlation of stable oxygen isotope temperature record with light attenuation profiles in reef-dwellingTridacna shells

Molluscs are known to record environmental changes in their carbonate shells in detail. This paper reports the findings of a high-resolution analysis of stable oxygen isotopic compositions and light transmission properties of a shell of the reef-dwelling Pacific giant clamTridacna gigas. Our findings reveal that the annual growth rates and the longevity ofTridacna specimens can be readily determined by measuring the annual light attenuation pattern within the shell. Annual seasonal changes in water temperature are reflected with high resolution in the stable oxygen isotope ratios and in the light attenuation values of the aragonite shell. The inner shell ofT. gigas deposited below the pallial line revealing undisturbed shell accretion with high growth rates shows the maximum seasonal oxygen isotope range and the highest resolution in light attenuation changes. We suggest that this is the best part of the shell to reconstruct former seasonal surface water temperatures in tropical environments. Scanning electron microscopy (SEM) studies suggest that the annual growth patterns observed in transmitted light are generated by a complex pattern of daily growth increments with varying sizes of skeletal crystallites and varying amounts of organic carbon.     

Structure and formation of the adventitious tube of the Japanese watering-pot shell Stirpulina ramosa (Bivalvia, Anomalodesmata, Clavagellidae) and a comparison with that of the Penicillidae

Abstract. Stirpulina ramosa is the only extant endobenthic representative of the Clavagellidae and is restricted to the waters of Japan. A single intact adventitious tube of this species has been obtained and its structure is described. The right valve is 16 mm long and located within the adventitious tube. It has an opisthodetic ligament located on resilifers. There are anterior and posterior adductor muscle scars, a thick pallial line, and pallial and pedal gape (right valve only) sinuses. The left shell valve is but 9 mm long and is united into the fabric of the adventitious tube via the intermediary of a shelly saddle. Internally, only the anterior adductor muscle scar and a small element of the pallial line scar are identifiable on the left valve. The posterior adductor and the rest of the pallial line scar (including a pallial sinus) are, remarkably, located on the adventitious tube beyond the shell valve margin. The adventitious tube of S. ramosa is formed in a manner wholly dissimilar from that of Brechites vaginiferus (Penicillidae). In B. vaginiferus, the tube is secreted as a single entity from the general outer mantle surface, including the siphons, covering the body. As a consequence, both shell valves are incorporated into the structure of the tube and the watering pot is bilaterally symmetrical. In S. ramosa, the tube and watering pot are secreted from the mantle margin and surface surrounding and extending from the left shell valve, so that only the left valve is incorporated into its structure. A dorsally derived mantle element is progressively extended over to the right side of the body, meeting a ventrally derived counterpart that passes beneath it, forming a pleat in the calcareous structure of the right side of the tube that they secrete. This pleat extends into the complex of watering‐pot tubules and forms the pedal gape. The watering pot is thus Ω shaped. The ventrally derived mantle element forms a sinusoidal crest on the right‐hand base of the watering pot, creating a pedal gape sinus scar on the right valve. The Clavagellidae radiated widely in the Mesozoic, leaving behind a rich fossil record for Stirpulina. Only S. ramosa, however, has survived until the present. In contrast, the Cenozoic Penicillidae has a poor fossil record, but there is a rich variety of extant endobenthic watering‐pot shells. It has been argued hitherto that the two families represent a remarkable example of convergent evolution. In view of the success of the Penicillidae and thus the endobenthic, tube‐dwelling lifestyle, however, it is hard to understand why Stirpulina has largely died out—even S. ramosa being known by but one or two specimens. A study of the anatomy of S. ramosa might one day answer this question.     

Age and growth rate determinations of an intertidal bivalve, Phacosoma japonicum, using internal shell increments

A venerid bivalve Phacosoma japonicum (Reeve) occurring commonly in the Japanese coastal area preserves periodic growth lines in the shell cross-section. Long-term shell growth patterns of this species have been traced for many individuals on the intertidal flat of the Seto Inland Sea, west Japan. Sclerochronological analysis of these individuals and specimens collected monthly shows that several growth cessation marks within their shells are formed during the winter of each year prior to spawning. Hence the marks were used for age and growth rate determinations. As large individuals showed little shell growth for more than two years after the formation of 7 or 8 annual increments, this species probably has a lifespan of more than ten years. Shell growth patterns of this species based on annual increments can be accurately approximated by a von Bertalanffy curve. The number of microgrowth increments formed during a year tends to decrease with age, although it varies markedly among specimens of the same age. Furthermore, even in summer during rapid shell growth, the microgrowth increments do not represent daily and/or sub-daily tidal rhythms in many specimens. The results of this study and those by several authors strongly suggest that the annual increments are the key for age and growth rate determinations of both living and fossil bivalve species.     

The relationship of the mantle and shell of the Polyplacophora in comparison with that of other Mollusca

The structure and growth of the polyplacophoran shell, characteristically consisting of eight plates surrounded by a girdle, is examined in the light of current views on the relationships of mantle and shell in the Bivalvia. The periostracum and outer and inner calcareous layers of the shell of the latter group are homologous with the cuticle, tegmentum and articulamentum respectively of the shell of the Polyplacophora. The margin of the mantle consists of a large marginal fold, which secretes the cuticular girdle, and a small accessory fold bearing mucous cells. These are functionally comparable with all three folds of the mantle margin found in other molluscs, although anatomically the marginal fold of the chitons probably represents only the inner surface of the outer fold of the mantle margin.
The cuticle not only forms the girdle, which bears calcified spines or spicules, but also extends between the shell plates. The principal part of the cuticle consists largely of mucopolysaccharide material but there is also a thin discrete inner region which is similar chemically to the periostracum of other molluscs. The cuticle, possibly without spines, probably covered the entire dorsal surface of a primitive placophoran and beneath this, plates developed. As these grew the cuticle became worn away except marginally and between the plates. It is suggested that a covering of mucus over the visceropallium may have been the forerunner of the molluscan shell and the possible evolutionary relationships of the shell throughout the Mollusca are discussed.     

The relationship of the mantle and shell of the Polyplacophora in comparison with that of other Mollusca

The structure and growth of the polyplacophoran shell, characteristically consisting of eight plates surrounded by a girdle, is examined in the light of current views on the relationships of mantle and shell in the Bivalvia. The periostracum and outer and inner calcareous layers of the shell of the latter group are homologous with the cuticle, tegmentum and articulamentum respectively of the shell of the Polyplacophora. The margin of the mantle consists of a large marginal fold, which secretes the cuticular girdle, and a small accessory fold bearing mucous cells. These are functionally comparable with all three folds of the mantle margin found in other molluscs, although anatomically the marginal fold of the chitons probably represents only the inner surface of the outer fold of the mantle margin.
The cuticle not only forms the girdle, which bears calcified spines or spicules, but also extends between the shell plates. The principal part of the cuticle consists largely of mucopolysaccharide material but there is also a thin discrete inner region which is similar chemically to the periostracum of other molluscs. The cuticle, possibly without spines, probably covered the entire dorsal surface of a primitive placophoran and beneath this, plates developed. As these grew the cuticle became worn away except marginally and between the plates. It is suggested that a covering of mucus over the visceropallium may have been the forerunner of the molluscan shell and the possibleevolutionary relationships of the shell throughout the Mollusca are discussed.     

Rib fabrication in Ostreoidea and Plicatuloidea (Bivalvia,Pteriomorphia) and its evolutionary significance

Ribs of Ostreoidea and Plicatuloidea are defined as antimarginal, that is, perpendicular to the margin throughout growth. Morphogenetically, these ribs are unique, since, unlike radial ribs, they are secreted by a homogeneous mantle margin. Based also on the reconstructed shell secretion cycle in Bivalvia, we propose that ribs of Ostreoidea and Plicatuloidea are formed by a mantle margin which, upon extension from the shell margin, stretches and folds by taking the preformed ribs as templates. In extending perpendicular to the margin (as in all Bivalvia growing isometrically), such a mantle extends the rib pattern antimarginally. Ribs of this kind are purely mechanical structures, as their arrangement depends on the mechanical properties of the mantle and on the environmental conditions. This explains the high irregularity of such ribbing patterns. The presence of antimarginal ribs in both the Ostreoidea and Plicatuloidea sheds light on their origin. The first known oyster, Actinostreon cristadifformis, probably derived from an antimarginally ribbed Prospondylidae gen. indet. in the Late Permian or Early Triassic. Antimarginally ribbed Triassic species formerly included in Placunopsis originated both the Dimyidae Atreta in the Late Triassic and Enantiostreon in the Mid Triassic, which was transitional to Plicatulidae. Therefore, Dimyidae and Plicatulidae are closely connected and grouped under Plicatuloidea, to which Ostreoidea is phylogenetically unrelated.     

Direct innervation of the mantle edge gland by neurosecretory axons in Helisoma duryi (Mollusca: Pulmonata)

Summary The mantle edge gland of Helisoma duryi is innervated by neurosecretory axons from the pallial nerves. Synaptoid contacts occur between axons and gland cells, and there is ultrastructural evidence for the release of neurosecretory material. The mantle edge gland contributes to the deposition of periostracum during shell formation, and direct neurosecretory innervation may control shell growth and regeneration.Supported by a National Research Council of Canada Grant (A-4673) and Negotiated Grant D-61     

Finite element analysis of a double membrane tube (DMS-tube) and its implication for gastropod shell morphology

Molluscan shells, including those of Gastropoda, are formed by accretionary growth at the mantle edge. The mantle is a thin membrane of skirt-like shape, which extends minutely beyond the aperture, and its edge adds a shell increment to the aperture margin so that each increment copies a configuration of the mantle edge at that time. Thus, regulation of shell morphogeny is almost equivalent to the factors which control the mantle form at the moment of shell growth. Form of the mantle skirt is considered to be kept in a state of balance between the force of its internal stress and forces acting on it such as fluid pressure or muscle contraction. The expansion behavior of the mantle skirt has been numerically analyzed by using an elastic model (DMS-tube), which represents the fundamental structure of the mantle tissue as a double membrane structure with internal springs (DMS). Four characteristic expansion patterns of the DMS-tube have been detected: (1) general outward expansion; (2) developing a ridge-like fold on an initial longitudinal protrusion of the tube edge; (3) drastic shift of the expanded state from a uniformly curved to an elliptical shape in outline, owing to the existence of a fixed boundary condition on the tube wall; and (4) constricted protrusion on the open region of the shell wall surrounding the DMS-tube. These results have the potential for answering the following questions relating to the morphogenesis of gastropod shells. How does the mantle skirt usually make contact with the inner surface of the shell wall so as to ensure continuous accretion of shell materials to the aperture margin? What is the cause of spiral ridges? Why do open coiling or minimally overlapping shells have generally circular apertures, while shells with apertures overlapped by whorls have non-uniformly curved apertural lips? What is the cause of long closed spines and why do they always appear on spiral ridges?     

SPIRAL GROWTH: THE 'MUSEUM OF ALL SHELLS' REVISITED

Raup's model of shell growth, now in standard use, is operationalonly for strictly conispiral shells. The pertinence of evolutionaryinterpretations of the distribution of existing shells in themorphospace defined by Raup's parameters is questioned. A simple,more general model accounts for the non-isometric growth ofmany shells. Some aspects of the distribution of existing shellsin the morphospace derived from the new model are discussed. (Received 18 November 1996; accepted 11 March 1997)     

Astronomically controlled deep-sea life in the Late Cretaceous reconstructed from ultra-high-resolution inoceramid shell archives

The periodicity of the mutual position of celestial bodies in the Earth-Moon-Sun system is crucial to the functioning of life on Earth. Biological rhythms affect most of the processes inside organisms, and some can be recorded in skeletal remains, allowing one to reconstruct the cycles that occur in nature deep in time. In the present study, we have used ultra-high-resolution elemental ratio scans of Mg/Ca, Sr/Ca and Mn/Ca from the fossil, ca. 70 Ma old inoceramid bivalve Inoceramus (Platyceramus) salisburgensis from deep aphotic water and identified a clear regularity of repetition of the geochemical signal every of ~0.006 mm. We estimate that the shell accretion rate is on average ~0.4 cm of shell thickness per lunar year. Visible light–dark lamination, interpreted as a seasonal signal corresponding to the semilunar-related cycle, gives a rough shell age estimate and growth rate for this large bivalve species supported by a dual feeding strategy. We recognize a biological clock that follows either a semilunar (model A) or a tidal (model B) cycle. This cycle of tidal dominance seems to fit better considering the biological behaviour of I. (P.) salisburgensis, including the estimated age and growth rate of the studied specimens. We interpret that the major control in such deep-sea environment, well below the photic zone and storm wave base, was due to barotropic tidal forces, thus changing the water pressure.     

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.