The sphaeridia of sea urchins: ultrastructure and supposed function (Echinodermata,Echinoida)

Summary Sphaeridia are minute skeletal appendages of the echinoid test which are considered to be sense organs, organs of equilibrium, according to their shape. The sphaeridium forms a functional unit with the tubercle to which it adheres. The tubercle is encircled by a basiepithelial nerve ring of the epidermis. A circle of regularly arranged myocytes stretches from the tubercle to the sphaeridium. The muscles are distant from each other. The myofibrillar processes enter the pore space of the sphaeridial skeleton to which they are anchored by tendons; tendons are absent in the tubercle region. The cell bodies of the myocytes lie opposite to the nerve ring outside the skeleton. In this region the muscle cells and the nerve ring are in contact with each other, their basal laminae fuse. Tensions of the various myocytes are dependent on the position of the top-heavy sphaeridium. The nerve ring contains neurones which are provided with a cilium which lies close to the contact region with the myocytes. This arrangement leads to the assumption that the nerve cells in question have a proprioceptor function. Unique filter cells are present in the pore spaces of the sphaeridium and the tubercle. They possibly detoxicate the extracellular fluid that surrounds the myocytes. Phagocytes loaded with spacious phagosomes are crowded in the adjacent pore spaces. They are possibly extruded via the epidermis. Filter cells and phagocytes have obviously to do with the metabolism within the sphaeridium-tubercle-system.     

Functional anatomy of the valves in the ambulacral system of sea urchins (Echinodermata,Echinoida)

Summary The water vascular system of sea urchins is examined with special reference to the valves positioned between the radial vessel and the ampullae of the tube feet. The lips of the valve protrude into the ampulla. Thus the valve functions mainly like a check valve that allows the unidirectional flow of fluid towards the ampulla. Each ampulla-tube foot compartment acts as a semi-autonomous hydraulic system. The lumina of the ampulla and the tube foot are lined with myoepithelia except for the interconnecting channels that pierce the ambulacral plate. The contraction of the ampulla results in an increasing hydraulic pressure that protrudes the tube foot, provided that the valve is closed. The retraction of the tube foot results in a backflow of fluid independent of the condition of the valve. The lips of the valve are folds of the hydrocoel epithelium. The pore slit lies in the midline. The perradial faces of the lips are covered with the squamous epithelium of the lateral water vessel. The ampullar faces are specialized parts of the ampulla myoepithelium. Turgescent cells which form incompressible cushions take the place of the support cells. The valve myocytes run parallel to the pore slit and form processes that run along the base of the ampulla and the perradial channel up to the podial retractor muscle. The findings lead to the hypothesis of multiple control of the ampulla-tube foot system: (1) The mutual activity of the ampulla and the tube foot is indirectly controlled by the lateral and podial nerves which release transmitter substances that diffuse through the connective tissue up to the muscle layers. (2) A muscle-to-muscle conduction causes the simultaneous contraction of the ampulla or the podial retractor muscles. (3) The valve muscles are directly controlled by the processes of the valve myocytes which make contact with the podial retractor. In extreme conditions a backflow of hydrocoel fluid towards the radial water vessel occurs.     

Ultrastructure and organization of the epineural canal and the nerve cord in sea urchins (Echinodermata,Echinoida)

Summary The radial nerve cord ofMespilia globulus has been examined as an example of echinoid nerve cords. In the radius of echinoids only the ectoneural component of the nerve cord is present which is a derivative of the ectoderm. The nerve cord runs in the interior of the body and is accompanied by the epineural canal. In echinoids, the neuroepithelium makes up the upper and side walls of the epineural canal. Each lateral branch of the nerve cord forms a sort of neural tube. It encloses a branch of the epineural canal which represents an open connection with the sea water. Thus, the epineural canal exhibits numerous openings which probably allow sea water to flow back and forth. This organization is unique in echinoderms. — The neuroepithelium exhibits the organization of an epidermis with well-developed nervous elements. Glial cells are not present. The support cells are the true epithelial cells. Their monociliated cell bodies border the lumen and, by means of cytoplasmic stems that contain a bundle of filaments, they reach up to the basal lamina. The nerve cells and their trunk of nerve fibres fill the spaces between the support cells. — Three types of nerve cells can be distinguished according to their polarity: (1) Primary sensory cells that project a cilium into the epineural canal, the axon hillock region is at the opposite pole. (2) Subluminal cells whose cilium originates in the axon hillock region. (3) Neurones that lie within the trunk of nerve fibres. They are highly stretched in the direction of the nerve cord and are also provided with a cilium. Types 2 and 3 may be homologized with the basal nerve cells of the epidermis. They are possibly multipolar. — The lateral nerve cords make contact with the ampulla and pass the ambulacral plate parallel to the channel that connects the ampulla and the tube foot. The activity of the tube foot-ampulla system is possibly controlled by means of transmitter substances that diffuse through the connective tissue layer between the nerve cord and the myoepithelia of the ampulla and the tube foot respectively.     

Pourtalesiid sea urchins (Echinodermata: Echinoidea) of the northern Mid-Atlantic Ridge

Five species of deep-sea pourtalesiid echinoids were collected by the RV G.O. Sars MAR-ECO expedition to the northern part of the Mid-Atlantic Ridge: Solenocystis imitans new genus, new species, Echinosigra phiale (Thomson, 1872), Echinosigra (Echinogutta) fabrefacta Mironov, 1974, and two unidentified species of Pourtalesia. Three types of plastron plating are distinguished within the family Pourtalesiidae; in Solenocystis new genus the plating is similar to that in Spatagocystis, a genus known only from the deep-sea Antarctic. Each of three groups with distinctive plastron plating shows a continuous morphological gradation from a less elongated test without subanal rostrum to an extraordinarily elongated test with large rostrum and wide subanal fasciole; the ratio width/length of test varies from 0.86 to 0.12. Some morphological characters of the pourtalesiids are regarded as adaptations to burrowing and appear to be correlated with test elongation. The family is represented in the northern Atlantic only by 'advanced' genera (Echinosigra, Pourtalesia and Solenocystis) with very elongated tests.     

Four lines of spermatid development and dimorphic spermatozoa in the sea urchin Anthocidaris crassispina (Echinodermata, Echinoida)

 The process of sperm development in the sea urchin Anthocidaris crassispina was studied by light and electron microscopy. Similar to other echinoids studied, a single flagellum, striated rootlet and nuage-like materials were present in spermatogonia of A. crassispina. Spermatocytes near the diplotene stage showed intracellular localization of the axoneme which appeared to be a retracted flagellum prior to cell division. Fibrous filaments were associated with a proximal centriole in spermatocytes and spermatids and might be involved in movement of the proximal centriole. An acrosomal vesicle was developed and a residual body was formed in spermatids. The special development patterns in A. crassispina attributed to the presence of two patterns of tail development and two patterns of mitochondrial development during spermiogenesis. These four lines of spermiogenesis resulted in the formation of four morphological types of sperm cell, i.e. sperms with: (1) a symmetrical midpiece and posterior tail, (2) an asymmetrical midpiece and posterior tail, (3) a symmetrical midpiece and bent tail and (4) an asymmetrical midpiece and bent tail. Sperm cells with bent tails (type 3+4) were probably still at the late spermatid stage because results of scanning electron microscopy demonstrated gradual detachment and eventual straightening of the bent tail, and their percentage occurrence in the sperm population decreased significantly (P<0.05) towards the spawning season of A. crassispina. Spermatozoa with a symmetrical midpiece were dominant (averaging 70% occurrence in the sperm population) over those with an asymmetrical midpiece. The dimorphic spermatozoa in A. crassispina (types 1, 2) are both considered to be euspermatozoa as their morphology is typical for Echinoida. Accepted: 4 May 1998     

Ontogeny and origin of the brooding system in Antarctic urechinid sea urchins (Echinodermata,Holasteroida)

Summary Echinoids usually broadcast gametes, and do not generally engage in a high degree of parental care. However, when they do, juveniles are typically maintained among the spines, or in shallow, external depressions in the test itself. The brooding Antarctic holasteroids Urechinus mortenseni and Plexechinus nordenskjoldi are bizarre exceptions: females develop an elaborate brooding system in which a small number of direct developing young are protected. Ontogeny of post-natal brooding urechinids is marked by profound divergence in the growth trajectories of male and female apical systems. In females, this leads to dramatic departures from the patterns found in all other echinoids. Otherwise, coronal skeleton allometry of males and females is almost identical. Juveniles in brood pouches grow larger than the diameter of the apical aperture through which they must pass to reach the external environment. The apical plates, from which the brooding system is suspended, hinge downward to enlarge the aperture, allowing the young to emerge from the female. A possible origin for the brooding system suggests derivation by centripetal plate addition from the ocular plates in the coronal skeleton. We develop a contrasting model for the origin of the brooding system that relies on a proposed homology between genital and periproctal elements of the apical system of echinoids and the more highly developed dorsal skeleton of other echinoderm classes.     

Ultrastructural investigation of matrix-mediated biomineralization in echinoids (Echinodermata,Echinoida)

Summary The formation of echinoderm endoskeletons is studied using echinoid teeth as an example. Echinoid teeth grow rapidly. They consist of many calcareous elements each produced by syncytial odontoblasts. The calcification process in echinoderms needs (1) syncytial sclerocytes or odontoblasts, (2) a spacious vacuolar cavity within this syncytium, (3) an organic matrix coat in the cavity. As long as calcite is deposited, the matrix does not touch the interior face of the syncytium. The cooperation between syncytium, interior cavity and matrix coat during the mineralization process is discussed. The proposed hypothesis applies to the formation of larval skeletons, echinoderm ossicles and echinoid teeth.When calcite deposition ceases the syncytium largely splits up into filiform processes, and the skeleton is partly exposed to the extracellular space. However, the syncytium is able to reform a continuous sheath and an equivalent of the cavity and may renew calcite deposition.The tooth odontoblasts come from an apical cluster of proliferative cells, each possessing a cilium. The cilium is lost when the cell becomes a true odontoblast. This suggests that cilia are primitive features of echinoderm cells. The second step in calcification involves the odontoblasts giving rise to calcareous discs which unite the hitherto single tooth elements. During this process the odontoblasts immure themselves. The structures necessary for calcification are maintained until the end of the process.The mineralizing matrix is EDTA-soluble. The applied method preserves the matrix coating the calcite but more is probably incorporated into the mineral phase and dissolved with the calcite.Abbreviations A adhesive point (LNC) - B adaxial bag - bb basal body (ci) - CA calcareous deposits - cb cytoplasmic bladder (cp) - ce centriole - ci cilium - cp cable-like cell process - cv condensing vacuole - dp distal processes (sh) - E epithelium of the tooth - ex extracellular space - f extracellular fibrils - ga gasket (sh) - ic interior cavity - L lamellae (LNC) - LNC lamellae needle complex - m mitochondrium - mc matrix coat - MF main fold (P) - MI mitosis - mt microtubules - N nucleus - O odontoblast - P primary plate - Ph phagocyte - PR proliferative cell - pr prism - rb reserve body - RER rough endopl. reticulum - rl rootlet (ci) - RY relatively youngest plate - s satellite (bb, ce) - sh synplasmic sheath (O) - SP secondary plate - sv smooth-walled vesicle - TF transversal fold (P) - U umbo (P) - v Golgi vesicle - Y youngest tooth element     

The molecular evolution of sperm bindin in six species of sea urchins (Echinoida: Strongylocentrotidae)

The acrosomal protein bindin attaches sperm to eggs during sea urchin fertilization. Complementary to ongoing functional biochemical studies, I take a comparative approach to explore the molecular evolution of bindin in a group of closely related free-spawning echinoid species. Two alleles of the mature bindin gene were sequenced for each of six species in the sea urchin family Strongylocentrotidae. The nucleotide sequences diverged by at least 1% per Myr at both silent and replacement sites. Two short sections flanking the conserved block show an excess of nonsynonymous substitutions. Each is homologous to a region that had been identified as a target of selection in other sea urchin comparisons. A large proportion of the bindin-coding sequence consists of a highly variable repeat region. Bindin sequences, even including the large intron, could not resolve the branching order among five of the species.      

Origin and evolutionary plasticity of the gastric caecum in sea urchins (Echinodermata: Echinoidea)

Background  

The digestive tract of many metazoan invertebrates is characterized by the presence of caeca or diverticula that serve secretory and/or absorptive functions. With the development of various feeding habits, distinctive digestive organs may be present in certain taxa. This also holds true for sea urchins (Echinodermata: Echinoidea), in which a highly specialized gastric caecum can be found in members of a derived subgroup, the Irregularia (cake urchins, sea biscuits, sand dollars, heart urchins, and related forms). As such a specialized caecum has not been reported from "regular" sea urchin taxa, the aim of this study was to elucidate its evolutionary origin.     

Evolution of a novel muscle design in sea urchins (Echinodermata: Echinoidea)

The sea urchin (Echinodermata: Echinoidea) masticatory apparatus, or Aristotle's lantern, is a complex structure composed of numerous hard and soft components. The lantern is powered by various paired and unpaired muscle groups. We describe how one set of these muscles, the lantern protractor muscles, has evolved a specialized morphology. This morphology is characterized by the formation of adaxially-facing lobes perpendicular to the main orientation of the muscle, giving the protractor a frilled aspect in horizontal section. Histological and ultrastructural analyses show that the microstructure of frilled muscles is largely identical to that of conventional, flat muscles. Measurements of muscle dimensions in equally-sized specimens demonstrate that the frilled muscle design, in comparison to that of the flat muscle type, considerably increases muscle volume as well as the muscle's surface directed towards the interradial cavity, a compartment of the peripharyngeal coelom. Scanning electron microscopical observations reveal that the insertions of frilled and flat protractor muscles result in characteristic muscle scars on the stereom, reflecting the shapes of individual muscles. Our comparative study of 49 derived "regular" echinoid species using magnetic resonance imaging (MRI) shows that frilled protractor muscles are found only in taxa belonging to the families Toxopneustidae, Echinometridae, and Strongylocentrotidae. The onset of lobe formation during ontogenesis varies between species of these three families. Because frilled protractor muscles are best observed in situ, the application of a non-invasive imaging technique was crucial for the unequivocal identification of this morphological character on a large scale. Although it is currently possible only to speculate on the functional advantages which the frilled muscle morphology might confer, our study forms the anatomical and evolutionary framework for future analyses of this unusual muscle design among sea urchins.     

Macrostructure and evolution of the digestive system in Echinoida (Echinodermata)

The structure of the digestive system in Echinoida has long been puzzling since comparative studies have suggested that a derived structure, the siphon, has apparently evolved twice independently. New observations on the digestive system in five species of Cidaroida, four species of the Diadematoida and three species of Echinothurioida are presented. The results show that the four diadematoid species have a siphon and the three species of Echinothurioida have a siphonal groove, contrary to previous assertions. These observations make the macrostructure of the echinoid digestive system fully consistent with more recent phylogenetic hypotheses based on molecular and general morphological data, and support the idea that a siphon has evolved only once, in the stem lineage of the Acroechinoidea.     

Molecular genetic markers of intra- and interspecific divergence within starfish and sea urchins (Echinodermata)

A fragment of the mitochondrial COI gene from isolates of several echinoderm species was sequenced. The isolates were from three species of starfish from the Asteriidae family (Asterias amurensis and Aphelasterias japonica collected in the Sea of Japan and Asterias rubens collected in the White Sea) and from the sea urchin Echinocardium cordatum (family Loveniidae) collected in the Sea of Japan. Additionally, regions including internal transcribed spacers and 5.8S rRNA (ITS1–5.8S rDNA–ITS2) were sequenced for the three studied starfish species. Phylogenetic analysis of the obtained COI sequences together with earlier determined homologous COI sequences from Ast. forbesii, Ast. rubens, and Echinocardium laevigaster from the North Atlantic and E. cordatum from the Yellow and North Seas (GenBank) placed them into strictly conspecific clusters with high bootstrap support (99% in all cases). Only two exceptions–Ast. rubens DQ077915 sequence placed with the Ast. forbesii cluster and Aph. japonica DQ992560 sequence placed with the Ast. amurensis cluster–were likely results of species misidentification. The intraspecific polymorphism for the COI gene within the Asteriidae family varied within a range of 0.2-0.9% as estimated from the genetic distances. The corresponding intrageneric and intergeneric values were 10.4-12.1 and 21.8-29.8%, respectively. The interspecific divergence for the COI gene in the sea urchin of Echinocardium genus (family Loveniidae) was significantly higher (17.1-17.7%) than in the starfish, while intergeneric divergence (14.6-25.7%) was similar to that in asteroids. The interspecific genetic distances for the nuclear transcribed sequences (ITS1–5.8S rDNA–ITS2) within the Asteriidae family were lower (3.1-4.5%), and the intergeneric distances were significantly higher (32.8-35.0%), compared to the corresponding distances for the COI gene. These results suggest that the investigated molecular-genetic markers could be used for segregation and identification of echinoderm species.     

Males and females gonad fatty acids of the sea urchins Paracentrotus lividus and Arbacia lixula (Echinodermata)

The aim of this study was to analyze male and female gonad fatty acids of two sea urchin species, Paracentrotus lividus and Arbacia lixula, from the south coast of Spain. Additionally, we investigated possible differences between two locations. The ovaries of both species showed higher percentages of 14:0, 16:0, 16:1n-7, 18:2n-6, 18:3n-3 and 18:4n-3 than testes and lower levels of 18:0, 22:1n-9, 20:4n-6 and 22:5n-3. In P. lividus but not in A. lixula, the level of 20:5n-3 was higher in testes than in ovaries. These differences between sexes probably indicate different requirements of males and females during gametogenesis although the presence of a large number of gametes in the mature gonad may also have influences on fatty acid composition. Significant differences in gonad fatty acid profiles where also found when individuals of P. lividus collected at a location of the Mediterranean region were compared with specimens collected at the Atlantic coast. The most remarkable changes were the lower levels of 14:0, 18:1n-7, 20:1n-9, 20:4n-6 and 22:4n-6 and the higher values of 20:1n-11, 20:5n-3 and 22:6n-3 found in males and females of the Mediterranean specimens compared to those of the Atlantic coast. These differences probably reflect the differences in potential food sources at each location.     

Fine structure and presumed functions of the pedicellariae of Echinocardium cordatum (Echinodermata,Echinoida)

Summary Tridactylous, trifoliate, and globiferous pedicellariae occur on the body surface of Echinocardium cordatum. Tridactyles have three forms: the typical, the rostrate, and the large forms. Both typical and rostrate tridactyles and trifoliates occur all around the echinoid body (trifoliates are, however, 4 times more numerous than tridactyles). Large tridactylous and globiferous pedicellariae are restricted to the peribuccal area.As a general rule tridactyles and trifoliates are similar in morphology. The distal part of the valves forms an open blade and bears lateral teeth and/or denticles (single or in combs). The stalk consists of a rigid proximal part supported by an axial rod and a flexible distal part which includes an axial fluid-filled cavity. The cavity is surrounded by muscle fibers and acts as an hydroskeleton, allowing the undulating-coiling movements of the flexible part of the stalk. Trifoliates are always active while tridactyles react only to direct or indirect mechanical stimulation.The valves of the globiferous pedicellariae have a tubular distal part whose upper opening is surrounded by teeth. There is no differentiated venom gland but a cluster of epithelial glandular cells located at the level of the valve upper opening. A small ciliary pad occurs just below the glandular cluster. Globiferous stalks are not flexible, being supported for their full length by an axial rod. Globiferous pedicellariae appear to be sensitive only to chemical stimulation.The presumed functions of E. cordatum pedicellariae are (1) cleaning of the body surface and ciliary structures (trifoliates), (2) protection against sedimenting particles (tridactyles), and (3) defense of the peribuccal area against potential small predators (globiferous pedicellariae).     

Functional morphology of coronal and peristomeal podia in Sphaerechinus granularis (Echinodermata,Echinoida)

Summary Coronal podia of Sphaerechinus granularis are anchoring (adhering) appendages involved in either locomotion or capture of drift materials. Adhesion is not due to the presumed sucker action of the disc but relies entirely on secretions of the disc epidermis. Peristomeal podia function in wrapping together food particles or food fragments in an adhesive material thus facilitating their capture by the Aristotle's lantern. In both types of podia, the disc epidermis is made up of four cell types: non-ciliated secretory cells (NCS cells) that contain graules whose content is at least partly mucopolysaccharidic in nature, ciliated secretory cells (CS cells) containing granules of unknown nature, ciliated non-secretory cells (CNS cells) and support cells. The cilia of CS cells are subeuticular whereas those of CNS cells, although also short and rigid, traverse the cuticle and protrude in the outer medium. All these cells are presumably involved in an adhesive/de-adhesive process functioning as a duogland adhesive system. Adhesive secretion would be produced by NCS cells and de-adhesive secretion by CS cells. These secretions would be controlled through stimulations by the two types of ciliated cells (receptor cells) which presumably interact with the secretory cells by way of the nerve plexus. This model of adhesion/de-adhesion fits well with the activities of both coronal and peristomeal podia. The secretion of NCS cells would make up a bridge of adhesive material between a podium and the substratum (coronal podia) or would coat and gather food particles (peristomeal podia), respectively. The de-adhesive material enclosed in the granules of CS cells would allow the podia (either coronal or peristomeal) to easily become detached from the substratum and to always remain clear of any particles.Research Assistant, National Fund for Scientific Research (Belgium)     

Molecular heterotopy in the expression of <Emphasis Type="Italic">Brachyury</Emphasis> orthologs in order Clypeasteroida (irregular sea urchins) and order Echinoida (regular sea urchins)

The expression patterns of Brachyury (Bra) orthologs in the development of four species of sand dollars (order: Clypeasteroida), including a direct-developing species, and of a sea urchin species (order: Echinoida) were investigated during the period from blastula to the pluteus stage, with special attention paid to the relationship between the expression pattern and the mode of gastrulation. The sand dollar species shared two expression domains of the Bra orthologs with the Echinoida species, in the vegetal ring (the first domain) and the oral ectoderm (the second domain). The following heterotopic changes in the expression of the Bra genes were found among the sand dollar species and between the sand dollars and the Echinoida species. (1) The vegetal ring expressing Bra in the sand dollars was much wider and was located at a higher position along the AV axis, compared with that in the Echinoida species. The characteristic Bra expression in the vegetal ring of the sand dollar embryos was thought to be involved in the mode of gastrulation, in which involution continues from the beginning of invagination until the end of gastrulation. (2) Two of the three indirect-developing sand dollar species that were examined exhibited a third domain, in which Bra was expressed on the oral side of the archenteron. (3) In the direct-developing sand dollar embryos, Bra was expressed with an oral-aboral asymmetry in the vegetal ring and with a left-right asymmetry in the oral ectoderm. In the Echinoida species, Bra was expressed in the vestibule at the six-armed pluteus stage.Edited by N. Satoh     

Comparative morphology of the axial complex and interdependence of internal organ systems in sea urchins (Echinodermata: Echinoidea)

Background

The axial complex of echinoderms (Echinodermata) is composed of various primary and secondary body cavities that interact with each other. In sea urchins (Echinoidea), structural differences of the axial complex in "regular" and irregular species have been observed, but the reasons underlying these differences are not fully understood. In addition, a better knowledge of axial complex diversity could not only be useful for phylogenetic inferences, but improve also an understanding of the function of this enigmatic structure.

Results

We therefore analyzed numerous species of almost all sea urchin orders by magnetic resonance imaging, dissection, histology, and transmission electron microscopy and compared the results with findings from published studies spanning almost two centuries. These combined analyses demonstrate that the axial complex is present in all sea urchin orders and has remained structurally conserved for a long time, at least in the "regular" species. Within the Irregularia, a considerable morphological variation of the axial complex can be observed with gradual changes in topography, size, and internal architecture. These modifications are related to the growing size of the gastric caecum as well as to the rearrangement of the morphology of the digestive tract as a whole.

Conclusion

The structurally most divergent axial complex can be observed in the highly derived Atelostomata in which the reorganization of the digestive tract is most pronounced. Our findings demonstrate a structural interdependence of various internal organs, including digestive tract, mesenteries, and the axial complex.     

Domes,arches and urchins: The skeletal architecture of echinoids (Echinodermata)

Summary A combination of simple membrane theory and statical analysis has been used to determine how stresses are carried in echinoid skeletons. Sutures oriented circumferentially are subject principally to compression. Those forming radial zig-zags are subject to compression near the apex and tension near the ambitus. Radial and circumferential sutures in Eucidaris are equally bound with collagen fibers but in Diadema, Tripneustes, Psammechinus, Arbacia and other regular echinoids, most radial sutures are more heavily bound, and thus stronger in tension. Psammechinus, Tripneustes and several other echinoids have radial sutures thickened by ribs which increase the area of interlocking trabeculae. Ribs also increase flexural stiffness and carry a greater proportion of the stress. Further, ribs effectively draw stress from weaker areas pierced by podial pores, and increase the total load which can be sustained.Allometry indicates that regular echinoids become relatively higher at the apex as size increases, thus reducing ambital stresses. Some spatangoids with very high domes (eg Agassizia) maintain isometry, but others (eg Meoma) become flatter with size. Both holectypoids (Echinoneus) and cassiduloids (Apatopygus) maintain a constant height to diameter relationship. Flattening, and consequently ambital tensile stress, is greatest in the clypeasteroids. In this group the formation of internal buttresses which preferentially carry stress, reaches maximum development. A notable exception, however, is the high domed Clypeaster rosaceus.In this analysis it was assumed that local buckling or bending does not occur. The test of some echinoids (e.g. Diadematoida) have relatively wide sutures swathed in collagen, which allows local deformation. Others (e.g. Arbacia) have rigid sutures with reduced collagen. In Psammechinus and other members of the Order Echinoida, in addition to rib formation, inner and outer surface trabeculae are thickened so that the individual plates are stiffened. Some spatangoids (Meoma, Paleopneustes) have extensive sutural collagen, but the cassiduloid Apatopygus has collagen confined to junctions of sutures, and elsewhere the joints are strengthened and stiffened by fusion of trabeculae. Fusion of surface trabeculae is almost complete in the holectypoid, Echinoneus, and the sutures are obscured.     

Functional morphology of regular echinoid tests (Echinodermata,Echinoida): a finite element study

Functional morphology of the calcareous test ofEchinus esculentus was investigated by parametric finite element analysis, an engineering technique developed for numerical analysis of the behaviour of complex structures responding to external forces. Finite element models of the test were generated by methods of Computer Aided Geometric Design (CAGD) to calculate the mechanical responses to different types of loading. The load cases included vertical, concentrated load at the apex, vertical, distributed load on the upper third of the test, internal pressure and tensile forces as introduced into the test by tube feet activity. The objectives were the shape of the test, the distribution of material and the alternating zones of porous and non-porous plates within the test.—Echinoid tests resist external loading without showing any specific points of failure. The thickened margins of the periproct and peristome apertures account for load-bearing capacity as well as the thickned meridional structures which carry a greater portion of stress than the thinner parts of the test. Distribution of material is not a response to concentrated loads on the apex nor to self-weight. Taken strictly, echinoid tests are not thin (or membrane) shells. Under loading, bending moments occur which influence the stress state in the entire test. The pneu hypothesis could not be confirmed. Adaptation of the test shape or of the distribution of material as a response to internal pressure does not exist. Tests of regular echinoids are especially well adapted to the mechanical activity of the ambulacral tube feet, i.e. the shape of the test, its flattening towards the substrate, the outward bulge of the ambulacra and the differential distribution of material within the test.     

The ultrastructure of spines in sea urchins of the family Strongylocentrotidae

The ultrastructure of primary spines (microscopic relief of the surface of radial wedges on the spines and cross-sections of the spines) was studied by scanning electron microscopy in seven sea urchin species of the family Strongylocentrotidae. The spines were taken from the ambitus area of equally sizes individuals with a test diameter of 50 ± 5 mm. According to the number of wedges on their spines, the studied species can be divided into three groups: Strongylocentrotus intermedius, S. pallidus (18–25 wedges), S. droebachiensis, S. polyacanthus, Allocentrotus fragilis (24–32), and Mesocentrotus franciscantus, S. nudus (45–70). The species visibly differ in the microrelief of the wedges, which can be longitudinally streaked, with protuberances, with cross-dentate or non-dentate wedges, or with cross-series of denticles; in some species, the relief is absent. In S. intermedius, spines with smooth surface of the wedges, longitudinally streaked, with sparse protuberances, and with numerous cross-series of denticles only distally, only proximally, or over the entire length of the spine have been found. Wedge surface is convex or flattened in cross-sections; wedge shape in cross-section is rectangular (S. droebachiensis, S. intermedius, S. polyacanthus), triangular (S. pallidus), trapezoid (S. fragilis), or ansiform (M. franciscanus, M. nudus). Species of the genus Mesocentrotus are readily distinguished from the other species by the stereome of their spines: wider than the height of the wedges and more homogeneous, without regular concentric circles. Data on the ultrastructure of primary spines confirm the generic status of Mesocentrotus Tatarenko et Poltaraus, 1993 and do not support the recognition of Allocentrotus Mortensen, 1942 as a distinct genus.