The juxtaligamental cells of Ophiocomina nigra (Abildgaard) (Echinodermata: Ophiuroidea) and their possible role in mechano-effector function of collagenous tissue

Summary The intervertebral ligament of the brittlestar Ophiocomina nigra contains numerous cellular processes which belong to perikarya located on the outer surfaces of the ligament. These are described as the juxtaligamental cells and have been studied by light and electron microscopy. The cells are mainly concentrated in four pairs of ganglion-like nodes associated with the intervertebral ligament and in similar nodes adjacent to every other major connective tissue component of the arm. Although their histochemistry and ultrastructure indicate a neurosecretory function, they are anomalous in containing unusually large electron-dense granules probably associated with calcium. The ganglion-like nodes are innervated by hyponeural nerves, though synaptic contacts with the juxtaligamental cells have yet to be demonstrated.The function of the cells is discussed and it is suggested that they may be involved in the rapid loss of tensile strength which the intervertebral ligament sustains during arm autotomy. They may achieve this by controlling the availability of Ca²⁺ ions to the extracellular compartment of the ligament.A version of this paper was read at the U.K.-Eire Echinoderms Colloquium, Bedford College, London, in July 1978This work was conducted mainly at University Marine Biological Station, Millport, during tenure of a N.E.R.C. research studentship. I am grateful to Professor N. Millott for his keen supervision, to Professor D.R. Newth for permission to use the electron microscope in the Department of Zoology, University of Glasgow, where Maureen Gardner provided expert assistance, and to Professor R.M. Kenedi for Facilities in the University of Strathclyde. I have benefited from discussion with J.L.S. Cobb, V.W. Pentreath, and especially A.M. Raymond, University of St. Andrews, who allowed me to mention his unpublished observations.     

Ultrastructure of the tube foot sensory-secretory complex in Ophiocomina nigra (Echinodermata,Ophiuridea)

Summary The ultrastructure of the epidermal layer of both the oral and arm podia of the brittle star Ophiocomina nigra is described. Despite external differences, little variation occurs in their internal structure. The podial epidermis, which is overlain by a three-layered cuticle, consists of five cell types: support, mucous, sensory, adhesive secretory and monociliated neurosecretory-like cells. Areas of specialisation are superimposed on this basic plan. These comprise four cells forming cohesive units, made up of two adhesive secretory, one sensory and one monociliated neurosecretory-like cells. The two adhesive secretory cells may be identical or vary in the structure of their secretory packets. The sensory cells are of the normal type bearing a short cilium with a 9+2 microtubular arrangement. The monociliated neurosecretory-like cells contain many small dense vesicles and a short sub-cuticular cilium of irregular microtubular structure. Together, they appear to form a sensory-secretory complex which functions in adhesion both for feeding and locomotion. A system in which the secretion of the monociliated neurosecretory-like cell may control adhesive secretion is proposed.     

Morphological and mechanical aspects of fission in Ophiocomella ophiactoides (Echinodermata,Ophiuroida)

Summary During fission the ophiuroid Ophiocomella ophiactoides splits across the disc into two halves each of which regenerates to form a complete individual. This paper describes the gross anatomy of the fission plane and the histology, ultrastructure, and mechanical properties of key structures transected during fission.Rupture of the disc integrument appears not to be facilitated by a pre-determined plane of weakness. Comparison of naturally split and artificially split animals showed that at fission a mechanism operates which restricts breakage to the interradial plane of two jaws. The interradial plane is subtended mainly by collagenous ligaments and by muscles linked to the skeletal components by basal lamina-like tendinous fibres. No fission-related adaptations could be identified in the oesophagus, circumoral nerve ring, or circumoral water vascular canal.On the basis of creep tests on isolated preparations of the disc integrument and jaw-frame and the responses of these preparations to ionic manipulation, it is suggested that their behaviour is dominated by collagenous elements that can undergo actively controlled changes in their mechanical properties. A hypothesis is presented which proposes a role for such a mechanism in the initiation and facilitation of fission.Contribution 318 of the Discovery Bay Marine Laboratory, University of the West Indies     

Arm autotomy in brittlestars (Echinodermata: Ophiuroidea)

Although ophiuroid arm shedding has long been accepted as an example of autotomy, there has been little investigation of the phenomenon to substantiate this. This paper describes the outwardly visible aspects of autotomy and the function of the internal components of the arm during detachment. Observations are focussed on Ophiocomina nigra, some comparisons being made with eight other species.
Ophiuroid autotomy is characterized by its occurrence close to the point of stimulation, its rapidity, and by the pattern of intervertebral muscle separation at the insertions which is constant for a given species. Evidence is presented showing the important role played by the intervertebral ligament. Both this and the muscle insertions are collagenous, and it is suggested that they and the other intersegmental connective tissues facilitate autotomy by undergoing a drastic, nervously mediated loss in tensile strength which enables the animal to part from its arm with a minimum of effort. Comparable properties have been ascribed to other echinoderm connective tissues, and their role in asteroid and holothurian autotomy has been acknowledged, but such a mechanism has not previously been suggested for ophiuroid arm autotomy.     

Morphology and histology of the disc autotomy plane in Ophiophragmus filograneus (Echinodermata,Ophiurida)

Summary Normal and autotomized specimens of Ophiophragmus filograneus were studied by gross dissection, light microscopy, transmission electron microscopy, and scanning electron microscopy to determine how autotomy occurs at the major region of disc attachment. Each of the 10 genital bars is attached to an arm by a broad ligament consisting of a thick outer layer of collagenous connective tissue and an inner layer of neurosecretory cells innervated by a lateral ectoneural branch of the radial nerve. Neurosecretory cell processes extend into the collagenous layer. During autotomy the collagenous fibers separate, weakening the ligament near its insertion along the genital bar. Collagen fiber separation is probably caused by neurosecretions. There is no evidence for ossicle dissolution. The results indicate that the histological organization of the genital bar ligaments is identical to that of mutable collagenous tissues in other echinoderms and that the autotomy process is similar to that described for other echinoderms.     

Comparative morphology of echinoderm calcified tissues: Histology and ultrastructure of ophiuroid scales (Echinodermata,Ophiuroida)

Summary The calcified body wall of an ophiuroid was investigated by a new method and compared with that of other echinoderms. The previous opinion that the epidermis of ophiuroid arm shields consists of a reduced syncytium continuous with the underlying dermis is incorrect. The epidermis is distinctly separated from the dermis by a basal layer and consists of (1) supporting cells which bear the cuticle, (2) ciliated cells (hitherto unknown and probably sensory), (3) gland cells, and (4) nerve cells with the basal nerve plexus. The overall structure of the epidermis is a three-dimensional tube system (marked by the basal lamina) which penetrates the dermal tissue of the scale's pore space and continues with nerve cords situated below the scale. This arrangement is unique in echinoderms.The dermal sclerocytes largely conform with those of the echinoid Eucidaris. The mineral skeleton is produced intracellularly or intrasyncytially. Moreover, dermal sclerocytes were found to release extracellular microfibrils which have nothing to do with calcite deposition. The attachment of the cuticle to the dermis is achieved by means of epidermal coupling areas. Collagen fibers fasten the scale to the underlying connective tissue sheath. The supposed fibrocytes within this sheath resemble sclerocytes. Each collagen bundle is provided with a strand of nerve fibers which, in contrast to the basal nerve plexus, are naked. They are said to infuence the mechanical properties of the connective tissue.Structures associated with cilia occur in cell types which normally lack a cilium. This finding suggests that most echinoderm cells are potentially monociliate.Abbreviations A apical shield - asp secretory products - B bacteria - bb basal body - bl basal lamina - C ciliated cell - ca coupling area - ci cilium, - cf collagen fibrils - cs cell surface - CTS connective tissue sheath - cu _i inner cuticular layer - cu _m middle cuticular layer - dp distal processes (Sc) - EC epineural canal - G Golgi complex - gv granular vesicle - H haemal vessel - hb homogeneous body - hl horizontal lamina (Su) - j cell junction - L lateral shield - le boundary layer (Sc) - lo distal lobe (Su) - M intervertebral muscle or its attachment - m mitochondrium - mf microfibrils - mu mucus - mv microvilli - mvb multivesicular body - N nerve cell - n nucleus - nf neurofibrils - ng neurogranules - nn naked neurofibrils - O oral shield - P tube foot - Pc phagocyte - pg pigment granules - rl rootlet - RN radial nerve - RV radial vessel - Sc sclerocyte - sh cytoplasmic sheath (Sc) - sj septate junction - Su supporting cell - sv secretory vesicle - T calcite trabeculum - V vertebral ossicle - v vesicle (Su)     

Juxtaligamental system of the disc and oral frame of the ophiuroid Amphipholis kochii (Echinodermata: Ophiuroidea) and its role in autotomy

Abstract. The ophiuroid Amphipholis kochii is able to detach its central disc from the underlying oral frame in response to external stimuli. In this article we supply new observations on the microanatomy and ultrastructure of the autotomy plane, and of the juxtaligamental system which is believed to bring about connective tissue changes that underpin the detachment process. We correct previous confusion over the innervation of juxtaligamental nodes involved in disc autotomy, provide evidence that juxtaligamental cells are a population of specialized nerve cells, and present observations on changes in the ultrastructure of juxtaligamental cells during autotomy, which support the view that they are responsible for connective tissue disruption.     

Fine structure of syzygial articulations before and after arm autotomy in florometra serratissima (Echinodermata: Crinoidea)

Summary About 1 s after appropriate stimulation, arms of Florometra serratissima break at articulations called syzygies that are specialized for autotomy. The fine structure of unreacted and of newly broken syzygies is described. The unreacted syzygy includes (1) ligament fibers consisting of collagen fibrils interconnected by interfibrillar strands and (2) axons filled with presumed neurosecretory granules. The newly broken syzygy includes (1) ruptured ligament fibers consisting of swollen collagen fibrils associated with interfibrillar globules and (2) axons containing few presumed neurosecretory granules, some of which are fixed in the act of exocytosis; moreover, the calcareous skeleton adjacent to the broken syzygy is partly eroded. The observations before and after breaking suggest that the autotomy mechanism may comprise the following sequence of events: rapid neural transmission from stimulation site to syzygy triggers a massive exocytosis of granules from presumed neurosecretory axons; the released neurosecretions (which could include chelating agents, strong acids, proteolytic enzymes or enzyme activators) etch the skeleton and lower the tensile strength of the ligament fibers by weakening the collagen fibrils and/or the interfibrillar material; breakage of the ligament fibers, the major connective tissue of the articulation, is quickly followed by rupture of all the other tissues at the syzygy.     

The nonfeeding auricularia of Holothuria mexicana (Echinodermata,Holothuroidea)

Among echinoderms, nonfeeding larvae usually are simplified in body shape, have uniform ciliation, and have lost the larval gut. A few species have nonfeeding larvae that express some remnant features of feeding larvae like ciliated bands and larval skeleton or larval arms, but typically their larval mouth never opens and their gut does not function. Still other species have retained the feeding larval form, a functional gut, and can feed, but they do not require food to metamorphose. The present note describes the development of a tropical holothurian, Holothuria mexicana, which hatches as a gastrula that is already generating coelomic structures. A translucent auricularia forms with a mouth that opens but becomes reduced soon thereafter. In form and ciliation this auricularia resembles a feeding larva, but it does not respond to food. A doliolaria forms on day 4 and the pentactula on day 6 post‐fertilization. Further study of this larva and that of its closely related congener, Holothuria floridana, is warranted.     

The ultrastructure of the morula cells of Eupentacta quinquesemita (Echinodermata: Holothuroidea) and their role in the maintenance of the extracellular matrix

The ultrastructure of the morula cells of Eupentacta quinquesemita and the distribution of these cells in the dermal connective tissue are described. Morula cells are abundant in the dermis and appear to function in the maintenance of the extracellular matrix (ECM) as a source of ground substance material. The synthetic activity of these cells is described in detail. Morula cells are filled with large secretory vesicles containing three electrondense materials which are derived from rough endoplasmic reticulum and Golgi activity. The synthetic product of these cells contains glycosaminoglycans and is secreted into the ECM by degranulation. The ultrastructural and histochemical similarity of the degranulation product to the ECM ground substance suggests that they are comprised of the same material. Morula cells appear to function primarily in connective tissues where ground substance predominates. The cells often contain secretory vesicles at various stages of formation, all of which eventually mature and degranulate. The synthetic pathway of the morula cells appears to result ultimately in the complete disruption and death of the cells. The function of morula cells in the holothuroid ECM is discussed, and the synthetic activity of the cells is compared with that of other secretory cells.     

Biodiversity dynamics and their driving factors during the Cretaceous diversification of Spatangoida (Echinoidea, Echinodermata)

Variations in recorded diversity over time present a scrambled signal that is modulated by a large number of variables: the potential of particular life forms to generate evolutionary innovations, external constraints induced by the environment in its broad sense, the heterogeneity of the fossil record and the analytical artefacts due to sampling bias. A key question is how to characterise and quantify the separate input of any given factor in the overall diversity signal. This paper explores the structure of diversity data for spatangoid heart urchins and the sensitivity of recorded diversity to different factors of analytical bias (length of geological periods, proportion of palaeogeographical realms explored, accessible area of outcrops and historical determinism). Unexpectedly, recorded diversity of spatangoids is not proportional to the duration of stages. Bias implied by time scale is negligible compared to bias of sampling or historical determinism. Diversity at any given time is dependent on its recent history (autocorrelation). For spatangoids, a high correlation between diversity at time t_i and t_i−1 suggests that recorded diversity has an evolutionary significance. A nearly constant rate of diversification is hypothesised for the Cretaceous. A relative poor fossil record during the Turonian and the Coniacian interrupts the main trend of diversification. The number of species counted for a single time interval depends on the number of palaeogeographical realms considered. In conjunction with ecological and phylogenetic data, this relation suggests an evolutionary signal in which western Tethys acted as a centre of origination. Diversity at a single location is constrained ecologically and diversification is controlled by migration into new realms. Recorded diversity and available area of outcrop seem to be correlated, but alternative interpretations can be drawn, including large-scale bias in the fossil record or operation of similar causes (e.g., effect of sea-level fluctuation). Comparing recorded diversity with separate factors independently leads to conflicting results. A multivariate approach suggests that the main trend in recorded diversity might be partially related to evolutionary signal or biases connected with the heterogeneity of the fossil record. Results from other approaches (phylogeny, morphological disparity) are consistent with and emphasise the evolutionary significance of the recorded diversity of spatangoids.     

The echinoderms (Echinodermata) from El Salvador

A list of echinoderms from El Salvador (tropical eastern Pacific) is presented. The results were obtained from field surveys (between the years 2000 and 2004), the scarce literature sources, and Internet information. A total of 37 species and six genera are reported. The most abundant echinoderms in rocky shores were: Phataria unifascialis, Echinometra vanbrunti, Holothuria kefersteini, as well as Astropecten armatus in soft bottoms.     

The compass depressors ofParacentrotus lividus (Echinodermata,Echinoida): ultrastructural and mechanical aspects of their variable tensility and contractility

Summary The compass depressors are bands of soft tissue which connect the compass ossicles of the echinoid lantern to the inner edge of the test. They are essentially ligaments with on one side a thin layer of muscle cells. The ligamentous component consists mainly of a parallel array of collagen fibrils with interspersed 12 nm microfibrils. The most notable cellular constituents are granule-containing cell bodies and their processes which resemble the juxtaligamental cells that have been found in all echinoderm mutable collagenous tissues and which may control the tensility of these tissues. The muscle cells occupy about 8% of the total cross-sectional area of the compass depressor and are located in a richly innervated pseudostratified myoepithelium. When subjected to constant low loads in creep tests the compass depressor stretches to a fixed length beyond which there is no further extension. The length at this creep limit coincides with the maximum length to which the compass depressor is stretched by natural movements of the intact lantern. Stress-strain tests show that treatment with 1 mM acetylcholine or 100 mM K⁺ ions can increase reversibly the stiffness of the compass depressor to an extent that cannot be due to contraction of the myoepithelium, suggesting that the mechanical properties of the ligament are under physiological control. Tension-length data on the myoepithelium suggest that it generates a maximum active tension when the compass depressor is stretched to the creep limit. The implications of these results for the function of the compass depressors are discussed.     

Regeneration of the epithelial lining of the stomach after autotomy of a disk in the brittle star Amphipholis kochii (Lütken) (Echinodermata: Ophiuroidea)

The regeneration of the epithelial lining of the stomach of the brittle star Amphipholis kochii after autotomy of the aboral part of the disk was studied. It was shown that a part of the stomach epithelium remained after autotomy. Its cells participated in regeneration of the epithelium during restoration of the lost digestive system. There was a partial dedifferentiation of the epithelium cells of stomach, their migration and proliferation; cells of the stomach retained some specialized cytoplasmic structures, secretory vacuoles, and pinocytic vesicles. Migration of ectodermal cells of the esophagus in the damaged area was also recorded.     

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.     

Polykristalliner Calcit bei Seeigeln (Echinodermata,Echinoidea)

Zusammenfassung Es wurde erstmals für Echinodermen primär polykristalliner Calcit nachgewiesen, und zwar im Cortex der Primärstacheln der Cidaridae, dem sekundären Zahnskelet von Clypeaster und in den akzessorischen Kalkstrukturen, die im Kauabschnitt die Furche der Diadematiden-Zähne ausfüllen. Es gibt bei anderen Seeigelfamilien keine Bildungen, die dem Cidariden-Cortex oder den akzessorischen Kalkstrukturen der Diadematiden homolog sind. Das polykristalline sekundäre Zahnskelet von Clypeaster ist dagegen dem monokristallinen sekundären Zahnskelet anderer Seeigel homolog.Der Mg-Gehalt des Calcits liegt in den feinkristallinen Zonen (mit Ausnahme des Cortex) im allgemeinen höher; die höchsten Werte finden sich in den Steinteilen der Zähne, gleichgültig ob das sekundäre Zahnskelet mono- oder polykristallin ist.Polykristalline Teile sind im allgemeinen härter als monokristalline Teile. Die Steinteile der Seeigelzähne sind die härtesten Skeletteile von Echinodermen überhaupt; ihre VickersHärte übertrifft weit diejenige von solidem Calcit. Im Steinteil ist das feinkristalline Gefüge von Calcit eng mit organischer Matrix verbunden, und es wird vermutet, daß darauf die besonders hohe Härte der Steinteile beruht.

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Polycristalline calcite in sea urchins

Summary For the first time primary polycrystalline calcite in Echinoderms is shown in the cortex of primary spines of Cidaridae, in the secondary tooth skeleton of Clypeaster and in the accessory calcareous structures filling the crevice fold in the chewing areas of Diadematoidae teeth. Other Echinoid families lack formations homologous to the cortex of Cidaridae and accessory calcareous structures of Diadematoidae. On the other hand the polycrystalline secondary tooth skeleton of Clypeaster is homologous to the monocrystalline one of the other sea urchins.With the exception of cortex the Mg-content in calcite—analyzed by microprobe and X-ray powder method—is generally greater in macrocrystalline parts. The highest Mg-contents are found in the stone parts of teeth irrespective of whether the secondary tooth skeleton is monocrystalline or polycrystalline.Polycrystalline parts are usually harder than monocrystalline ones. The stone parts of Echinoid teeth are the hardest skeleton parts of Echinoderms on the whole; their hardness is much greater than that of solid calcite. It is supposed that the strong interlacing of the microcrystalline calcite and organic matter causes the enormous hardness of the stone part.

Mit Unterstützung durch die Deutsche Forschungsgemeinschaft.     

Potential for paired vestibules in plutei (Echinodermata, Echinoidea)

Abstract. Echinoids are generally thought to develop only a single ectodermal invagination (V1), which expands into a vestibule and contributes to the formation of the juvenile rudiment on the left side of the larval body. However, in 2 echinoid species, Strongylocentrotus purpuratus and Dendraster excentricus, a second invagination (V2) was observed to form on the right side of the larval body exactly opposite V1. Formation of the invaginations was followed and characterized using light microscopy and antibodies directed against the S2amide echinoderm neuropeptide as a morphological tag. V2 consistently formed later than V1, and appeared capable of contributing to the formation of a second rudiment when it received the appropriate mesodermal influence from contact with a hydrocoel. Otherwise, in both species, V2 formed a skeletal element that has not been previously described in this location. Because of the location, shape, and timing of its formation, V2, like V1, appears to have the innate ability of the larval ectoderm to contribute to juvenile rudiment formation.