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.     

Electron microscopic studies of growth margins of articulate brachiopods

Summary The brachiopod shell is secreted by the mantle epithelium lining the internal surfaces of its two valves. Growth lines, seen on their external surfaces, have been interpreted in terms of mantle regression and transgression from the valve margins. This scanning electron microscope study of the shell microstructure in recent brachiopods confirms these views and shows the skeletal evidence upon which such interpretations can be made. Electron micrographs reveal that from a growth line a plane dips posteriorly into the shell substance along which normal skeletal secretion was interrupted. Commonly a mosaic of secondary fibres, similar to that seen on the inside surface of the valve, is preserved upon this regression plane, most of the inside surface of which is covered by primary shell, usually extending posteriorly well into the secondary shell layer. The regression plane marks the area from which the mantle withdrew and the area over which shell secretion was interrupted. During mantle transgression primary shell was deposited over much of this surface, before the redevelopment of secondary fibres, so that the old internal surface of the valve was preserved as a false mosaic within the shell. In this way it is possible to recognise the extent of mantle regression and to note the position of the primary — secondary shell secreting junction of the mantle at the time when shell secretion was resumed.     

Shell microstructure in mesozoic articulate brachiopods

The shell microstructure was studied based on extensive material of Triassic, Jurassic, and Cretaceous brachiopods from the orders Rhynchonellida, Terebratulida, and Thecideida. Evolutionary changes in relationships between the shell layers of thecideids were revealed; the narrowing of fibers in historical development of rhynchonellids was established; the importance of the prismatic layer for taxonomy was recognized; and differences in width and shape of fibers within the superfamilies of rhynchonellids and terebratulids were determined.     

Body volumes and internal space constraints in articulate brachiopods

Brachiopods were once dominant in all the oceans of the world. but their distributions are non more restricted. There are few species which are found in shallow warm habitats and these are predominantly small. They have exceptionally low metabolic rates and exhibit low energy lifestyles. The majority of living articulate brachiopods are punctate (possessing mantle extensions. or caeca. which traverse the shell). Evidence produced hei-e suggests that the evolution of these phenomena may have been strongly affected by architectural constraints placed on articulate brachiopods by the use of the lophophore for feeding and respiration. They are essentially space limited because of the large volume needed for this organ. In some punctate brachiopods over 75% of their total body volume may be occupied by the lophophore and mantle cavity. This figure is only 60% in an impunctate (no caeca) species and may be only 20% in bivalve molluses. The implications are that caeca evolved to reduce pressure on space requirements, that maximum sizes may be set by the scaling patterns of space allocation and metabolic efficiency is a consequence of space constraints. Current distribution patterns may be strongly affected by the low metabolism and low energy lifestyles. The relative success of small brachiopods in warm shallow seas may have been facilitated by the scaling patterns of space allocations which show small specimens to have similar mantle cavity volumes to bivalve molluscs.     

Predatory asteroids and the decline of the articulate brachiopods

Various causes, such as increased predation pressure, the lack of planktotrophic larvae, a 'resetting' of diversity, increased competition from benthic molluscs and the decline of the Palaeozoic fauna, have been suggested to explain the failure of the brachiopods to reradiate following the Permo-Triassic mass extinction. Increased predation pressure has hitherto appeared improbable, because typical predators of brachiopods, such as teleostean fish, brachyuran crabs and predatory gastropods, did not undergo major radiation until the late Mesozoic and early Cenozoic. However, new evidence strongly suggests that one important group of predators of shelly benthic organisms, the asteroids, underwent a major radiation at the beginning of the Mesozoic. Although asteroids appeared in the early Ordovician, they remained a minor element of the marine benthos during the Palaeozoic acme of the brachiopods. However, these early asteroids lacked four important requirements for active predation on a bivalved epifauna: muscular arms (evolved in the early Carboniferous); suckered tube feet, a flexible mouth frame and an eversible stomach (all evolved in the early Triassic). Thus radiation of the Subclass Neoasteroidea coincided with both their improved feeding capability and the decline of the articulates. The asteroids were the only group of predators of brachiopods that underwent a major adaptive radiation in the earliest Mesozoic. The asteroids may therefore have contributed to inhibiting a Mesozoic reradiation of the brachiopods. Epifaunal species lacking a muscular pedicle may have been particularly vulnerable. Unlike bivalve molluscs, modern brachiopods show only a limited range of adaptations to discourage asteroid predation. □ Asteroidea, Brachiopoda, evolution, predation, functional morphology.     

The origin and geometry of radial ribbing patterns in articulate brachiopods

Ackerly, S. C. 1992 07 15: The origin and geometry of radial ribbing patterns in articulate brachiopods.
Geometric models for simple. radial ribbing in articulate brachiopods include (1) ribs radiating isometrically from the shell umbo. (2) divergence of thc ribs from some 'point' within the shell, and (3) reorientation of the ribs at right angles to the shell margin. Analyses of the Orthida, the ancestral taxon of articulate brachiopods, indicate that rib geometries are isometric in Early Cambrian taxa (model 1). but that by the Early Ordovician rib orientations are generally perpendicular to the shell margin (model 3). A combination of functional and morphogenetic Factors explains the ribbing geometries observed in orthide brachiopods.     

The relationship between major lamellae and epithelial regressions in some articulate brachiopods

Brunton, C. Howard C. & Alvarez, Fernando 1989 07 15: The relationship between major lamellae and epithelial regressions in some articulate brachiopods. Lethaia , Vol. 22, pp. 247–250. Oslo. ISSN 0024–1164.
Hiller (1988, Lethaia , Vol. 212) proposed three relationships between the secretory epithelium of articulate brachiopods and the shell surface ornamentations of growth lines, lamellae and spines. None of his models satisfy the growth of strongly lamellose athyrid shells and we propose a fourth involving strong regressions effecting both primary and secondary shell layers. In Recent Tegulorhynchia we suggest a growth function for the 'frayed' shell of Hiller occurring immediately in front of the spines.     

Large extinctions of articulate brachiopods in the paleozoic and their ecological and evolutionary consequences

The largest Paleozoic extinctions of articulate brachiopods occurred at the Frasnian—Famennian boundary in the Late Devonian and at the Permian—Triassic boundary. Both extinctions affected taxa of all levels, including orders, but differed in scale, course, and ecological and evolutionary consequences. The Frasnian—Famennian extinction event was selective and evolutionary activity after the crisis varied in different orders. However, in the Early Carboniferous, the brachiopod diversity was mostly restored in comparison with the Devonian maximum. In particular groups, preadaptation played a role in changes in diversity and reconstruction of communities. The brachiopod composition at this boundary changed sharply. The extinction event at the end of Permian was global and accompanied by changes in the biota. Later, in the Meso-Cenozoic, the brachiopod diversity was not restored, and bivalves acquired primary importance in various bottom communities of different sea zones where Paleozoic brachiopods previously dominated. Extinction of brachiopods at this boundary was long and gradual. The symptoms of the ecological crisis in the development of Permian brachiopods are recognized beginning from the Roadian Age, which was probably the onset of this crisis.     

Taphonomic and paleoecologic implications of flow-induced forces on concavo-convex articulate brachiopods: an experimental approach

The Orientation of benthic marine organisms may be disturbed by flow-induced forces (i.e. drag and lift) caused by wave and current activity. Drag and lift are partly a function of organism size and shape. Consequently, morphology may affect stability (defined as resistance to reorientation, flipping, or entrainment) both during the life of an organism and after its death. An understanding of drag-and-lift effects is therefore essential to the interpretation of paleoecology and biostratinomic processes. An experimental method for quantifying the relative effects of flow-induced forces is described. These forces are measured during flume experiments using transducers and plaster replicas of fossils. As an illustration of the method's potential for taphonomic research, results from experiments investigating the effects of concavo-convex morphologies of articulate brachiopods are presented. Concave-up and convex-up orientations are commonly used to infer paleohydraulic conditions. Two geniculate brachiopods (Rafinesquina alternata and Leptaena richrnondensis) and three flattened forms (a second morphotype of Rafinesquina altemata, Strophodonta demissa , and Tropidoleptus carinatus) were tested in convex-up and concave-up postures and in three azimuthal orientations (hingeline oriented upstream, hingeline downstream, and hingeline parallel to flow). Concave-up orientations consistently exhibit higher drag than convex-up orientations, and this supports the common observation that valved fossils are typically found convex up in paleoenvironments dominated by traction transport. The presence of geniculation significantly increases drag. Lift is relatively insignificant for all models in most orientations. □ Taphonomy, paleoecology, brachiopods, flow-induced forces, transport.     

Microborings in Recent brachiopods and the functions of caeca

Microborings in the primary shell layer of Recent brachiopods are clearly seen to avoid endopunctamicroscopic canals pervading the shell fabric and housing papillose extcnlions of the mantle (the caeca). This avoidance confirms the suggestion that the caecal contents inhibit boring organisms (Owen & Williams 1969; Proc. R. Soc. Loud. B, 172 ), and as such the caecum can be considered as an important instrument in protecting the brachiopod shell. A comparison of the relative fecundity of co-habitating impunctate and cndopunctate New Zealand brachiopods provides indirect evidence that the caecum may indeed also function in a nutrient storage capacity. Brachiopods, microborings, primary shell layer, endopuncta, defence, storage.     

The protegulum and juvenile shell of a Recent articulate brachiopod: patterns of growth and chemical composition

Patterns of shell formation and the chemical composition of the shell deposited during early post-larval life were investigated in laboratory-reared cultures of the Recent articulate brachiopod Terebraralia transversa (Sowerby). A non-hinged protegulum averaging 148 pm in length is secreted by the mantle within a day after larval metamorphosis. The inner surface of the protegulum exhibits finely granular, non-fibrous material. A rudimentary periostracum constitutes the outer layer of the primordial shell. and concentrically arranged growth lines are lacking. By four days post-metamorphosis, a brephic type of juvenile shell develops from periodic additions of shell material to the anterior and lateral edges of the protegulum. Imbricated secondary fibers occur throughout the inner layer of the newly formed juvenile shell, and a rudimentary hinge apparatus is present posteriorly. The external surface of the shell exhibits concentric growth lines anterior to the caudally situated protegulum, and unbranched punctae begin to form in the subperiostracal region of the shell. At 23 days post-metamorphosis, the shell weighs an average of 1.7 μg and measures 318 μm in length. Electron microprobe analyses reveal that the protegulum is calcified. Minor amounts of sulfur, magnesium, iron, chlorine, aluminum, and silicon are also present in protegula and juvenile shells. Based on electron diffraction data, the mineral phase of juvenile shells consists of calcite, and protegula also appear to contain calcite.     

Comparative larval myogenesis and adult myoanatomy of the rhynchonelliform (articulate) brachiopods Argyrotheca cordata, A. cistellula, and Terebratalia transversa

Background

Primary agametic-asexual reproduction mechanisms such as budding and fission are present in all non-bilaterian and many bilaterian animal taxa and are likely to be metazoan ground pattern characters. Cnidarians display highly organized and regulated budding processes. In contrast, budding in poriferans was thought to be less specific and related to the general ability of this group to reorganize their tissues. Here we test the hypothesis of morphological pattern formation during sponge budding.

Results

We investigated the budding process in Tethya wilhelma (Demospongiae) by applying 3D morphometrics to high resolution synchrotron radiation-based x-ray microtomography (SR-μCT) image data. We followed the morphogenesis of characteristic body structures and identified distinct morphological states which indeed reveal characteristic spatiotemporal morphological patterns in sponge bud development. We discovered the distribution of skeletal elements, canal system and sponge tissue to be based on a sequential series of distinct morphological states. Based on morphometric data we defined four typical bud stages. Once they have reached the final stage buds are released as fully functional juvenile sponges which are morphologically and functionally equivalent to adult specimens.

Conclusion

Our results demonstrate that budding in demosponges is considerably more highly organized and regulated than previously assumed. Morphological pattern formation in asexual reproduction with underlying genetic regulation seems to have evolved early in metazoans and was likely part of the developmental program of the last common ancestor of all Metazoa (LCAM).     

Commissural asymmetry in brachiopods

Consistent asymmetric folding of the commissure is a characteristic feature of a small but significant number of brachiopod species. The feature may be obligate or facultative and is almost entirely confined to rhynchonellids, most of which are Mesozoic. The detailed nature of the asymmetry is very variable, but does not extend to internal hard parts such as crura. Taken as a whole, asymmetric brachiopod species show no preference for any particular environment or geographic region, and in no circumstances seem to have been markedly more or less successful than symmetric species. We are thus led to suggest that asymmetry was a genetically based condition which cropped up periodically in brachiopod evolution, and which possibly was selected neither particularly for nor against.     

'Flapping valves' in brachiopods

M. J. S. Rudwick and others postulate 'rhythmic-flow' feeding for the Permian richtho-feniacean bi-achiopods, whereas R. E. Grant claims that they fed by normal ciliary action. Suspension-feeding has two components, current generation and food capture; normal brachiopod lophophores do both, but this is neither universal nor compulsory among animals. Opening and closing the richthofeniid shell generated a 'tidal-flow' current precisely analogous to respiratory currents in mammals; this is neither inefficient nor 'self-defeating', as Grant claims. Grant's analysis fails because he chose the wrong mechanical analogy (a pump). Morphological features of richthofeniids are better explained on a tidal-flow hypothesis than on a ciliary-flow model, and all the data adduced by Grant in rejecting the former is compatible with it or favorable to it. It explains morphological features that are bizarre mysteries on the ciliaiy-current model, and is therefore superior even though it implies that these Permian brachiopods were radically innovative.     

Recent advances in the genetic investigation of osteoarthritis

Osteoarthritis (OA) demonstrates considerable clinical heterogeneity, generating heated debate over whether OA is a single disease or a complex mix of disparate diseases and concerning which tissues are principally involved in disease initiation and progression. Epidemiological studies have demonstrated a major genetic component to OA risk. However, these studies have also revealed differences in risk between males and females and for disease at different skeletal sites. This observation has resulted in the concept of genes for specific sites rather than a generalised OA phenotype. Recent breakthroughs have shed considerable light on the nature of OA genetic susceptibility. Many candidate genes have been confirmed, such as the interleukin-1 gene cluster and the oestrogen alpha-receptor gene ESR1. Genome-wide linkage scans have revealed several regions harbouring novel loci, some of which are beginning to yield their genes.     

Recent advances in the investigation of Antarctic macroalgae

 The paper summarizes our present knowledge of Antarctic macroalgae with regard to the following topics: explorative history of Antarctic macroalgae, the environment of macroalgae in the Antarctic, life histories and physiological anatomy, seasonal development, light demands and depth zonation, temperature demands and geographical distribution and the effect of desiccation, salinity and temperature on supra- and eulittoral algae. A baseline is visible, but much more work is necessary, especially to assess the susceptibility of macroalgae and coastal ecosystems in Antarctica to global climatic changes. Received: 29 June 1995/Accepted: 3 October 1995