The ontogeny of shell secretion in Terebratalia transversa (Brachiopoda, Articulata). II. Formation of the protegulum and juvenile shell

The fine structure of the shell and underlying mantle in young juveniles of the articulate brachiopod Terebratalia transversa has been examined by electron microscopy. The first shell produced by the mantle consists of a nonhinged protegulum that lacks concentric growth lines. The protegulum is secreted within a day after larval metamorphosis and typically measures 140-150 micron long. A thin organic periostracum constitutes the outer layer of the protegulum, and finely granular shell material occurs beneath the periostracum. Protegula resist digestion in sodium hypochlorite and are refractory to sectioning, suggesting that the subperiostracal portion of the primordial shell is mineralized. The juvenile shell at 4 days postmetamorphosis possesses incomplete sockets and rudimentary teeth that consist of nonfibrous material. The secondary layer occuring in the inner part of the juvenile shell contains imbricated fibers, whereas the outer portion of the shell comprises a bipartite periostracum and an underlying primary layer of nonfibrous shell. Deposition of the periostracum takes place within a slot that is situated between the so-called lobate and vesicular cells of the outer mantle lobe. Vesicular cells deposit the basal layer of the periostracum, while lobate cells contribute materials to the overlying periostracal superstructure. Cells with numerous tonofibrils and hemidesmosomes differentiate in the outer mantle epithelium at sites of muscle attachments, and unbranched punctae that surround mantle caeca develop throughout the subperiostracal portion of the shell. Three weeks after metamorphosis, the juvenile shell averages about 320 micron in length and is similar in ultrastructure to the shells secreted by adult articulates.     

Development of the pedicle in the articulate brachiopod Terebratalia transversa (Brachiopoda,Terebratulida)

Summary The development of the pedicle in the articulate brachiopod Terebratalia transversa has been examined by electron microscopy. The posterior half of the free-swimming larva comprises a non-ciliated pedicle lobe that contains the primordium of the juvenile pedicle at its distal end. During settlement at five to six days post-fertilization, the pedicle lobe secretes a sticky sheet that attaches the larva to the substratum. As metamorphosis proceeds, the epithelium in the posterior half of the pedicle lobe produces a thin overlying cuticle, and the pedicle primordium develops into a stalk-like anchoring organ. The juvenile pedicle protrudes through the gape that occurs between the posterior margins of the shell valves. A cup-like canopy, called the pedicle capsule, lines the posterior end of the shell and surrounds the newly formed pedicle. The core of the juvenile pedicle is filled with a solid mass of connective tissue. Numerous tonofibrils occur in the pedicle epithelium, and the overlying cuticle consists of amorphous material covered by a thin granular fringe. By one year post-metamorphosis, a body cavity develops anterior to the pedicle. Two pairs of adjustor muscles extend from the posterior end of the shell and traverse the cavity to insert in the pedicle. The connective tissue core of the pedicle in sub-adult specimens lacks muscle cells but contains numerous fibroblasts and collagen fibers. Three regions are recognizable in the connective tissue compartment of the adult pedicle: a subepithelial layer of non-fibrous connective tissue, a central fibrous zone, and a proximal mass of tissue that resembles cartilage.List of abbreviations as adhesive sheet - bc body cavity - bv brachial valve of shell - cf collagen fibrils - ct connective tissue - cu cuticle - di diductor muscle - ec epithelial cell - f fibroblast - fz fibrous zone - g gut - gc granular cell - gd gastric diverticulum - ht hinge tooth - ia interarea of pedicle valve - icl inner cuticular layer - lo lophophore - lu lumen of gut - m mesenchyme - ma mantle - ml mantle lobe - ocl outer cuticular layer - p periostracum - pc pedicle capsule - pce pedicle capsule epithelium - pcl pedicle collar of shell - pcn pedicle connectives - pd pedicle - pe pedicle epithelium - pl pedicle lobe - pv pedicle valve of shell - pzc proximal zone of cartilage-like tissue - s substratum - sel subepithelial layer - t tendon - tf tonofibril - vam ventral adjustor muscle     

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.     

Systematic position and composition of the superfamily Notanoplioidea Gill, 1969 (Brachiopoda, Articulata)

The systematic position of several genera of articulate brachiopods grouped around Notanoplia Gill is controversial. Notanoplia was established in 1950 within the order Chonetida; this genus and several externally similar genera, which were established later, were referred by different authors to the orders Orthida, Strophomenida, Chonetida, Atrypida, Spiriferida, or considered as incertae sedis. The morphological analysis of the shell of Notanoplia and the related genera Boucotia and Costanoplia supports the placement of these genera in a separate superfamily, Notanoplioidea, within the order Strophomenida. Other genera similar in the shell outline and shape to Notanoplia and conditionally considered as related to it probably belong to other strophomenid superfamilies or other orders.     

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).     

The ontogeny of pulmonary surfactant secretion in the embryonic green sea turtle (Chelonia mydas).

Pulmonary surfactant, consisting predominantly of phosphatidylcholine (PC), is secreted from Type II cells into the lungs of all air-breathing vertebrates, where it functions to reduce surface tension. In mammals, glucocorticoids and thyroid hormones contribute to the maturation of the surfactant system. It is possible that phylogeny, lung structure, and the environment may influence the development of the surfactant system. Here, we investigate the ontogeny of PC secretion from cocultured Type II cells and fibroblasts in the sea turtle, Chelonia mydas, following 58, 62, and 73 d of incubation and after hatching. The influence of glucocorticoids and thyroid hormones on PC secretion was also examined. Basal PC secretion was lowest at day 58 (3%) and reached a maximal secretion rate of 10% posthatch. Dexamethasone (Dex) alone stimulated PC secretion only at day 58. Triiodothyronine (T(3)) stimulated PC secretion in cells isolated from days 58 and 73 embryos and from hatchling turtles. A combination of Dex and T(3) stimulated PC secretion at all time points.     

The ontogeny of the Late Permian terebratulids of the family Beecheriidae Smirnova (Brachiopoda)

Based on the study of the ontogenetic changes of cardinalia and brachidium in the genera Beecheria Hall et Clarke, 1893 and Calycelasma Smirnova, 2007 (family Beecheriidae), a complex development of the brachidium, which corresponds to the angustothyridid type, and the formation of crural plates, which functioned as the outer hinge plates at the early developmental stages, are revealed.     

The ontogeny of the Late Permian terebratulids (Brachiopoda) Gruntelasma Smirnova, Grigorjevaelasma Smirnova, and Campbellelasma Smirnova and their taxonomic position

A specific type of ontogenetic change combining elements of dielasmoid and angustothyridid stages of brachidium development is established for the Late Permian genera Gruntelasma Smirnova, 2004, Grigorjevaelasma Smirnova, 2004, and Campbellelasma Smirnova, 2004 from the Russian Platform. On this basis, these genera are assigned to the superfamily Compositelasmatoidea Smirnova, family Compositelasmatidae.     

The ontogeny of swimming behavior in the scyphozoan, Aurelia aurita. I. Electrophysiological analysis.

1. Electrical correlates of behavioral activity were observed in the lip and tentacles of the polyp, but none were detected during column contraction. The tentacles are the most electrically active tissue, and the potentials are conducted along the length of the tentacle, but conduction to other parts of the animal were not observed. 2. Although the tentacles of the polyp and the rhopalia of the medusa are probably homologous, the development of pacemaker activity during strobilation is not a smooth transition from tentacle contraction potentials (TCPs) to marginal ganglion potentials (MGPs). This result indicates that each pacemaker activity develops de novo. 3. Two types of behavior were observed in the polyp: local responses, and coordinated activity which involved integrated responses in several body parts. The coordinated responses indicate that neurological coordination can take place in the polyp. Furthermore, feeding and spasm in the ephyra are similar to feeding and the protective response in the polyp. This similarity suggests that both coordinated responses in the polyp are coordinated by interneural facilitation in the diffuse nerve net (DNN) as in the ephyra. 4. Swimming in the ephyra is a medusoid behavior but feeding and spasm are coordinated by the DNN and are polypoid responses. Therefore, the ephyra is a mixture of polypoid and medusoid behaviors. As the ephyra matures into an adult medusa both polypoid responses are lost, but the DNN remains to modulate pacemaker output and control marginal tentacle contractions. As development proceeds from polyp, to ephyra, to medusa, each subsequent stage acquires some new behavior while retaining some aspect from the previous stage.     

I. I. Shmal'gauzen's concept of the evolution of ontogeny

The concept of evolution of ontogenesis by I. I. Shmal'gauzen is presented as a result of reviewing some of his theoretical works. This concept appears to be the most consistent development of classical darwinism on the basis of a basically new (as compared with synthetic theory of evolution) evolutionary-synthetic approach. This latter has been based on the idea of the organism integrity in onto- and phylogenesis and the involvement of the organismic level (ontogenesis), together with the population and biocoenotic ones, in the evolutionary process as a whole.     

Radioimmunoassay of cytidine 3',5' monophosphate (cCMP). I. Development of the assay.

A method is presented for the production of reagents for a radioimmunoassay for cCMP. cCMP was succinylated at the 2′0 position with [1,4 ¹⁴C] succinic anhydride, and the monosuccinyl cCMP coupled to Keyhole limpet hemocyanin and injected into rabbits. Antibodies to cCMP were produced that showed minimal crossreactivity with other cyclic nucleotides. Monosuccinyl cCMP was coupled to tyrosine methyl ester, then labeled with ¹²⁵I, and used as the radiolabeled ligand in the immunoassay of cCMP. By use of this assay, the concentration of cCMP in various tissues of rat and guinea pig have been determined.     

The interphotoreceptor space. I. Postnatal ontogeny in mice and rats

The postnatal ontogeny of the retinal interphotoreceptor space of mice and rats was studied by electron microscopy to elucidate the cytological developments in the surrounding cells relevant to the accumulation of extracellular glycosaminoglycans and glycoproteins constituting the interphotoreceptor matrix. This extracellular material at birth is principally the cell coat on all the immature cells that border the space at that time, but later additional weblike strands are seen in the space. The cells delimiting the space in the adult are the pigment epithelium (PE), the photoreceptors, and the glial cells of Müller. The Golgi complex of the PE is large at birth but involutes by day 15. Melanogenesis is the principal activity in this cell at birth but as the melanin granules mature, lysosomes gradually accumulate. Growth of the apical microvilli of the PE is most pronounced between day 5 and 15, which is also the time of rapid expansion of the interphotoreceptor space. The Golgi complex of the photoreceptor enlarges during this interval also, and remains voluminous thereafter. Müller's cells insert only slender apical processes lacking in secretory vesicles, into the interphotoreceptor space. All the adult cells have a cell coat on the surfaces facing the interphotoreceptor space. Secretory vesicles were not identified in any of the cells impinging on the space.     

Evo-devo and the I.I. Schmalhausen concept of the evolution of ontogeny

Evo-devo is considered a special field of knowledge emphasizing the role of developmental processes and mechanisms in evolution and integrating the data of many disciplines studying various structural levels of biosphere organization. A mechanical approach to estimation of events is regarded as a specific feature of the evo-devo concept. In our opinion, the I.I. Schmalhausen concept of the evolution of ontogeny, opposing the reductionist approach of many contemporary evo-devo adherents, can be regarded as the fundamental basis of evo-devo.     

A granulosis ofEupsilia transversa Hufn. (Lepidoptera,Noctuidae) in West Norway

Résumé La NoctuelleEupsilia transversa Hufn. était abondante dans l'Ouest de la Norvège en 1962, la chenille causant de graves dégats aux arbres fruitiers. En 1963, une infestation de granulose apparut sur les chenilles et en deux années réduisit la population à un niveau très bas. Le syndrome de la larve malade est décrit. Les résultats de quelques essais d'infection à petite échelle permettent de considérer ce virus comme capable de réduire effectivement la population deE. transversa dans les conditions de laboratoire. La granulose semble être léthale à tous les stades du ravageur sauf le dernier.      

Embryonic development of the shell in biomphalaria glabrata (Say).

During embryogenesis of the fresh water snail Biomphalaria glabrata (Say) (Pulmonata, Basommatophora) shell formation has been studied by light and electron microscopical techniques. The shell field invagination (SFI), the secretion of the first shell layers, the development of the shell-forming mantle edge gland and spindle formation have been investigated. During embryonic development at 28 degrees C environmental temperature, the shell field invaginates after 35 h. After 40 h the SFI is closed apically by cellular protrusions and scale-like precursors of the periostracum. The first electron translucent layer of the periostracum stems from electron dense vesicles of the cells which lie at the opening of the SFI. A second electron dense layer appears some hours afterwards. When the shell appears birefringent in the polarizing microscope (45 h of development) calcium can be detected in it using energy dispersive x-ray analysis. As calcification occurs the intercrystalline matrix appears under the periostracum and the SFI begins to open. In embryos of 60 h the mantle cavity appears at the left caudal side. When the mantle edge groove develops (65 h of development) lamellate units are added to the outer layer of the periostracum, but no distinct lamellar layer is formed in B. glabrata. In addition to the lamellar cell and the periostracum cell, a secretory cell can be observed in the developing groove. After 65 h of development, spindle formation starts and the shell begins to coil in a left hand spiral. After 5 days of development the embryos are ready to leave the egg capsules.     

The periesophageal celom of the articulate brachiopod Hemithyris psittacea (Rhynchonelliformea,Brachiopoda)

The celomic system of the articulate brachiopod Hemithyris psittacea is composed of the perivisceral cavity, the canal system of the lophophore, and the periesophageal celom. We study the microscopic anatomy and ultrastructure of the periesophageal celom using scanning and transmission electron microscopy. The periesophageal celom surrounds the esophagus, is isolated from the perivisceral cavity, and is divided by septa. The lining of the periesophageal celom includes two types of cells, epithelial cells and myoepithelial cells, both are monociliary. Some epithelial cells have long processes extending along the basal lamina, suggesting that these cells might function as podocytes. The myoepithelial cells have basal myofilaments and may be overlapped by the apical processes of the adjacent epithelial cells. The periesophageal celom forms protrusions that penetrate the extracellular matrix (ECM) of the body wall above the mouth and the ECM that surrounds the esophagus. The canals of the esophageal ECM form a complicated system. The celomic lining of the external circumferential canals consists of the epithelial cells and the podocyte‐like cells. The deepest canals lack a lumen; they are filled with the muscle cells surrounded by basal lamina. These branched canals might perform dual functions. First, they increase the surface area and might therefore facilitate ultrafiltration through the podocyte‐like cells. Second, the deepest canals form the thickened muscle wall of the esophagus and could be necessary for antiperistalsis of the gut. J. Morphol., 2011. © 2010 Wiley‐Liss, Inc.