A comparative study of the life style of two Jurassic irregular echinoids

The irregular echinoids Plesiechinus ornatus (Buckman) (Pygasteroida) and Galeropygus agariciformis (Forbes) (Cassiduloida) occur together in beds of the murchisonoe Zone, Bajocian, outcropping in the Cheltenham region of Gloucestershire. These species were largely restricted to different lithofacies within the carbonate shelf environment. Both adopted a hidden mode of life but achieved this by different techniques. Plesiechinus had fairly short spines and strongly muscular podia over the whole corona and was able to cover itself with coarse substrate particles. The oral tubercles are bilaterally symmetrical and are radially arranged. The oral spines are thought to have pulled sediment out from beneath the test, excavating a small depression for it. Galeropygus bore a dense covering of very small spines and its tube feet were differentiated into aboral respiratory podia and oral suckered podia. It had a preferred anterior direction of locomotion and is thought to have buried itself completely by excavating and ploughing into the substrate as it moved forward. Plesiechinus fed using only its lantern and postulated peristomial tube feet, whereas Galeropygus was a continugus sediment swallower and used its phyllode tube feet and peristomal lip spines in transferring particles towards the mouth.     

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.     

Experimental morphology of coronar growth in regular echinoids

Summary The coronar growth of a cidaroid and an aulodont echinoid are investigated by means of tetracycline labelling. The results are compared with earlier investigations on a stirodont and on a camarodont echinoid in order to evaluate the general features of coronar growth. In all echinoids new coronar plates are added at the apical end of the corona throughout the life cycle. The plates are shifted towards the peristome and they grow peripherically.In cidaroids ambulacral (A-) plates are detached from the firm corona. They are transformed into scales covering the peristomial field. The interambulacral (IA-) plates, however, are partially reabsorbed at the peristomial margin. In this manner the oldest solitary interambulacral plates are lost. The subsequent plates are arranged in pairs. The cidaroids thus show interradial growth even at their peristomial margin. This is unique to echinoids.In non-cidaroids there is a perignathic girdle made up of paired ambulacral auricles with interambulacral ridges in between. In some species the ridge is a solitary element. Therefore interradial growth cannot occur in the peristomial margin. In other species the ridge consists of several elements, but it also grows as a whole. Slight resorption of calcite occurs in places at the peristomial margin. In other places, however, calcite is added onto the peristomial edge. In non-cidaroids, therefore, the widening of the peristome is achieved solely by means of lateral growth in the plates bordering the peristome. The shift of the coronar plates from apicad to orad in noncidaroids is a relative shift only.In all echinoids the coronar plates are arranged in meridional columns. All plates grow up to the peristome. Their growth rates are relatively uniform towards the adambulacral sutures (which run between A- and IA-columns). Their growth rates towards the perradius and the interradius respectively are high in younger plates which are positioned above the ambitus, and decrease rapidly in plates located below the ambitus. Near the peristome the interradiad and perradiad growth rates are always considerably lower than adradiad growth rates. Perradial and interradial growth serve to adjust the plates in size and shape to their respective position in the corona.     

Localization of S1- and S2-like immunoreactivity in the nervous system of the brittle star Amphipholis squamata (Delle Chiaje 1828).

The recent isolation and characterization of the SALMFanide neuropeptides S1 GFNSALMFamide; and S2 (SGPYSFNSGLTFamide) from the sea stars. Asterias rubens and Asterias forbesi have initiated numerous studies on their morphological localization and distribution within the phylum Echinodermata. It has been shown by immunocytochemistry and radioimmunoassay that these peptides are widely distributed in the nervous system of some asteroids, echinoids and ophiuroids. A physiological approach has also shown that S1 and S2 potentiate the luminescence of the small ophiuroid Amphipholis squamata. In the present study. S1- and S2-like immunoreactivity have been localized in A. squamata by immunocytochemistry on both wholemount preparation and histological sections. The results reveal a widespread neuronal distribution of S1-like immunoreactivity in the circumoral ring, radial nerve cord, and tube feet. S1-like immunoreactivity was found to be associated with axons and cell bodies in both the ectoneural and hyponeural components of the nervous. S2-like immunoreactivity was detected only in the ectoneural plenus of the circumoral ring and radial nerve cord.     

Is the adhesive material secreted by sea urchin tube feet species‐specific?

Sea urchin adoral tube feet are highly specialized organs that have evolved to provide efficient attachment to the substratum. They consist of a disk and a stem that together form a functional unit. Tube foot disk tenacity (adhesive force per unit area) and stem mechanical properties (e.g., stiffness) vary between species but are apparently not correlated with sea urchin taxa or habitats. Moreover, ultrastructural studies of sea urchin disk epidermis pointed out differences in the internal organization of the adhesive secretory granules among species. This prompted us to look for interspecific variability in the composition of echinoid adhesive secretions, which could explain the observed variability in adhesive granule ultrastructure and disk tenacity. Antisera raised against the footprint material of Sphaerechinus granularis (S. granularis) were first used to locate the origin of adhesive footprint constituents in tube feet by taking advantage of the polyclonal character of the generated antibodies. Immunohistochemical assays showed that the antibodies specifically labeled the adhesive secretory cells of the disk epidermis in the tube feet of S. granularis. The antibodies were then used on tube foot histological sections from seven other sea urchin species to shed some light on the variability of their adhesive substances by looking for antibody cross‐reactivity. Surprisingly, no labeling was observed in any of the species tested. These results indicate that unlike the adhesive secretions of asteroids, those of echinoids do not share common epitopes on their constituents and thus would be “species‐specific.” In sea urchins, variations in the composition of adhesive secretions could therefore explain interspecific differences in disk tenacity and in adhesive granule ultrastructure. J. Morphol., 2011. © 2011 Wiley Periodicals, Inc.     

Evolutionary convergence in<Emphasis Type="Italic"> Otx</Emphasis> expression in the pentameral adult rudiment in direct-developing sea urchins

Convergence is a significant evolutionary phenomenon. Arrival at similar morphologies from different starting points indicates a strong role for natural selection in shaping morphological phenotypes. There is no evidence yet of convergence in the developmental mechanisms that underlie the evolution of convergent developmental phenotypes. Here we report the expression domains in sea urchins of two important developmental regulatory genes ( Orthodenticle and Runt), and show evidence of molecular convergence in the evolution of direct-developing sea urchins. Indirect development is ancestral in sea urchins. Evolutionary loss of the feeding pluteus stage and precocious formation of the radially symmetric juvenile has evolved independently in numerous sea urchin lineages, thus direct development is an evolutionary convergence. Indirect-developing species do not express Otx during the formation of their five primordial tube feet, the ancestral condition. However, each direct-developing urchin examined does express Otx in the tube feet. Otx expression in the radial arms of direct-developing sea urchins is thus convergent, and may indicate a specific need for Otx use in direct development, a constraint that would make direct development less able to evolve than if there were multiple molecular means for it to evolve. In contrast, Runt is expressed in tube feet in both direct- and indirect-developing species. Because echinoderms are closely related to chordates and postdate the protostome/deuterostome divergence, they must have evolved from bilaterally symmetrical ancestors. Arthropods and chordates use Otx in patterning their anterior axis, and Runt has multiple roles including embryonic patterning in arthropods, and blood and bone cell differentiation in vertebrates. Runt has apparently been co-opted in echinoderms for patterning of pentamery, and Otx in pentameral patterning among direct-developing echinoids. The surprisingly dynamic nature of Otx evolution reinvigorates debate on the role of natural selection vs shared ancestry in the evolution of novel features.     

Phylogeny and classification of the Asteroidea (Echinodermata)

Post-Palaeozoic asteroids share a large number of derived characters of the ambulacral column and the mouth frame, and constitute the crown group of the monophyletic group Asteroidea. This crown group is here called the Neoasteroidea (new subclass). The stem species of the crown group lived in the Permian or early Triassic and so the evolution of the asteroids parallels that of the echinoids. Character distribution within the Neoasteroidea, especially morphology of the skeleton, digestive system, larvae and tube feet, allows subdivision into four orders (Paxillosida, Notomyotida, Valvatida, Forcipulatida). The latter three orders possess the synapomorphy of suckered tube feet and are united as the Surculifera (new superorder); the Paxillosida are their primitive sister group. Palaeozoic asteroids represent the stem group of the class, and may be divided into plesions according to the order of appearance of synapomorphies with the crown group. Classification of Palaeozoic asteroids requires much further study. The appearance of new characters within the crown group asteroids, such as suckered tube feet, implies that these were absent in the stem group. The range of life-habits possible in Palaeozoic asteroids can thus be partly deduced from evidence provided by living asteroids. Palaeozoic asteroids are deduced to have lacked suckered tube feet and were presumably unable to evert the stomach; hence they were precluded from life on hard substrates and extraoral feeding on epifaunal organisms. It is suggested that they lived on soft substrates by deposit feeding, scavenging and predation on small benthos.     

Tube foot preservation in the Devonian crinoid Codiacrinus from the Lower Devonian Hunsrück Slate,Germany

Fossilized tube feet are described on Codiacrinus schultzei Follmann from the Lower Devonian Hunsrück Slate of Germany. This is the first definitive proof of tube feet on any fossil crinoid. Three lightly pyritized, flattened tube feet are preserved in a single interray of this cladid crinoid. The tube feet were at least 7 mm long. Their preservation is very similar to the tube feet reported previously from a Hunsrück ophiuroid, except that the Codiacrinus tube feet have small papillae, similar to living crinoids.     

Membrane Dynamics of the Contractile Vacuole Complex of Paramecium1

ABSTRACT. Membrane dynamics of the contractile vacuole complex of Paramecium were investigated using conventional electron microscopy of cells so that the vacuoles were serial-sectioned longitudinally and transversely. During systole, vacuolar membrane collapses first into flattened cisternae which undergo further modification into a mass of interconnected small membrane tubules. These tubules retain their connections with the radiating microtubular ribbons; consequently they are found only in the poleward hemisphere. Permanent connections between ampullae and the collapsed vacuole membrane could not be verified nor was a sphincter-like mechanism for closing such a junction observed. Membranes of the ampullae and the collecting canals also collapse to varying extents into arrays of tubules that remain bound to microtubular ribbons during diastole. Thus vacuole, ampullae, and collecting canal membranes all assume tubular forms when internal volume is at a minimum. Having failed to observe a microfilamentous encasement of the vacuole, we suggest that an alternative mechanism for the “contractile” function should be sought. One such is based on fluid volume increase and fluid flow within transiently interconnected tubular membrane systems that cycle between a tubular and a planar membrane form as internal volume is periodically increased and reduced. The driving force for this mechanism might best be sought in the molecular structure of the membranes of the contractile vacuole complex.     

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.     

NGIWYamide-induced contraction of tube feet and distribution of NGIWYamide-like immunoreactivity in nerves of the starfish Asterina pectinifera

NGIWYamide, a neuropeptide recently isolated from sea cucumbers, was tested on tube feet of the starfish Asterina pectinifera. NGIWYamide (10(-6)-10(-4) M) caused contraction of isolated tube feet. NGIWYamide-like immunoreactivity (NGIWYa-LI) was investigated with an antiserum against NGIWYamide. NGIWYa-LI was found in the radial nerve cord (RNC), the marginal nerve, and the tube feet. Both ectoneural and hyponeural parts of the RNC showed NGIWYa-LI. Immunoreactive cell bodies were found in both parts of RNC. Extensive labeling in the basal region of the ectoneural part suggests that a substantial proportion of axons in this part contains NGIWYamide or a similar substance. In tube feet, NGIWYa-LI was found in the sub-epithelial nerve plexus and in the basal nerve ring. Double labeling along with 1E11, a neuron-specific monoclonal antibody developed from A. pectinifera, indicated that the structures with NGIWYa-LI are neurons. These results suggest that NGIWYamide or an NGIWYamide-like peptide exists in starfish and functions as a neurotransmitter or a neuromodulator.     

Scolicia and its producer in shallow-marine deposits of the Miocene Chenque Formation (Patagonia,Argentina): functional morphology and implications for understanding burrowing behavior

Abstract

Scolicia is one of the most conspicuous trace fossils in lower shoreface deposits of the Miocene Chenque Formation of Patagonia, Argentina. This ichnotaxon consists of horizontal, sinuous or meandering trace fossils with a laminated backfill and two parallel strings located at the base. Abundant body fossils attributed to Brisaster iheringi occur in close association to these trace fossils. The echinoids are very well preserved, and most of the specimens have some areas with their spines attached in life position. In particular, preservation of the subanal tufts of spines, associated to isopores (which are interpreted as being associated to funnel building tube feet), supports the interpretation that these organisms are the producers of Scolicia isp. A shaft connecting the burrow with the sediment-water interface was not observed in vertical sections of the excavations, although on bedding plane surfaces, some small circular tube outlets are recognized. However, the great diversity and abundance of ichnofossils co-occurring in these deposits may preclude adequate identification of vertical structures. These deposits provide an excellent opportunity to integrate trace fossil data, body fossil information and observations from modern analogues in order to perform a morpho-functional analysis of Scolicia.     

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.     

Completeness of a fossil record: the Pleistocene echinoids of the Antilles

Diversity of Pleistocene marine invertebrates with single component, robust, easily identifiable skeletons shows a good correspondence to that of the Recent when compared between similar environments in the same region. Such comparisons are less easily made for multi-element skeletons. In the Antilles there are 24 extant species of echinoids in 0–30  m water depth. Considering all available fossils - complete tests, disarticulated spines, and test fragments and ossicles - the Pleistocene echinoids of the Antillean region have a high similarity with the extant shallow water fauna (c. 63% specific and 72% generic similarity). Many of the extant, shallow water, regular echinoid taxa are known from the Pleistocene, although irregular echinoids are less well represented. These results are comparable with those determined for coeval benthic invertebrates from this and other geographic regions, and indicate a high degree of similarity from a survey which is so far limited to only a few of the islands. Not all Pleistocene units considered herein were deposited in shallow water, but they include autochthonous specimens of taxa whose depth range extends beyond shallow water at the present day or presumed allochthonous specimens derived by downslope transport from shallower water, or both.     

Xylem-Phloem Exchange Via the Rays: The Undervalued Route of Transport

The radial solute exchange between xylem and phloem is an importantdeterminant of the nutrient distribution and C/N economy ofseed plants. Nevertheless, our knowledge of the mechanism ofxylem-phloem exchange is very limited. This paper aims to integrateanatomical and physiological data about xylem-phloem exchangeand focuses on the mechanisms of radial transport. Electrophysiologicalmapping and intracellular injection of fluorescent dyes demonstratethat stem tissue is organized in symplastic subunits specializedin either longitudinal or lateral transport. Radial orientation,anatomical and physiological organization, strong metabolicactivities, relatively negative membrane potentials, and highcapacities of uptake make the rays the most likely routes forsymplastic xylem-to-phloem transport. Parallel apoplastic transportmay occur through radial canacules between the ray cells. Thesymplastic xylem-to-phloem transport includes a number of steps:(a) passage across the vessel/parenchyma interface, (b) transferin the ray, and (c) delivery from the ray into the sieve tube/companioncell complex. Large contact pits in the vessel walls give accessto the ray cells, by which the solutes are accumulated throughsubstrate/proton co-transport. Dense pitting on the tangentialwalls and other ultrastructural features suggest preferentialsymplastic transport in the radial direction. Auxiliary endo/exocytosisis under dispute. The transport step from the ray to the sievetube/companion cell complex remains to be elucidated. In phloem-to-xylemtransport, the unloading from the phloem along the stem is essentiallyapoplastic, but under specific conditions it may be symplastic.Further transfer to the xylem probably occurs via the symplasticroute through the ray. During radial transport, important metabolicinterconversions take place into either transport or storagecompounds. In many trees, the metabolism of the ray cells undergoescyclic, annual fluctuations. Complex season-bound deposition/mobilizationprocesses in the ray cells corresponding with seasonal changesin membrane permeability and cellular organization, reflecta master-system designed to co-ordinate the C/N economy throughoutthe year. Key words: Xylem-phloem exchange, radial transport     

Localisation of gamma aminobutyric acid (GABA)-like immunoreactivity in the echinoderm Asterias rubens

Gamma amino butyric acid (GABA) is believed to be the principal inhibitory neurotransmitter in the mammalian central nervous system, a function that has been extended to a number of invertebrate systems. We have used a specific antiserum raised against GABA to demonstrate GABA-like immunoreactivity in the radial nerve cord (RNC), tube feet and the digestive system of the asteroid Asterias rubens. In the RNC, immunoreactivity was restricted to ectoneural fibres and cell bodies while in the tube feet fibres were revealed in the basal nerve ring and longitudinal nerve. In the gut, extensive labelling was apparent in the basi-epithelial plexus as well as in mucosal perikarya.     

ON THE TUBE ULTRASTRUCTURE AND ORIGIN OF CALCIFICATION IN SABELLIDS (ANNELIDA, POLYCHAETA)

Abstract:  Tube ultrastructure of Jurassic and Cretaceous Glomerula is very similar to that of Recent Calcisabella , supporting the synonymy of these genera and the early Mesozoic origin of calcification in sabellids. Tube structure of serpulids differs from that of Glomerula ; calcareous tubes probably evolved convergently within Sabellida. The tube wall in Recent Glomerula piloseta is composed of subparallel lamellae of aragonitic, irregular spherulitic prisms in the inner layer, and spherulites in the outer layer. Calcified lamellae are separated by organic films of different thickness. The structure of the internal tube layer in Glomerula piloseta , and the structure of entire wall in fossil Glomerula , are similar to the tube structure of Dodecaceria (Polychaeta, Cirratulidae). The irregular spherulitic prisms of Glomerula are similar to those found in the external layer of Hydroides dianthus and the internal layer of Spiraserpula caribensis .     

Comparative Morphology of Tube Feet Among the Asteroidea: Phylogenetic Implications

Tube-foot morphology has been included among a variety of taxonomiccriteria for the Asteroidea over the past twenty-five years.Other than a few families belonging to the order Paxillosida,which are thought to have pointed, non-suckered tube feet thatare used for digging and burial in soft sediments, the presumptionhas been that asteroids have flat-tipped, suckered tube feet.This has become an accepted model despite the fact that thecomparative morphology of asteroid tube feet has not been considered.In the present study we examine tube-foot morphology of 45 speciesof Asteroidea representing 19 families. Our analysis confirmsthat members of the Luidiidae and Astropectinidae (order Paxillosida)lack suckers on the tips of their pointed tube feet. We demonstratethat there is considerable variation in tube-foot morphologyamong members of the Asteroidea including an entirely new typeof flat-tipped, non-suckered tube foot in species belongingto the order Valvatida. The external morphology of tube feetin species belonging to the order Velatida could not be distinguishedfrom "typical" flat-tipped, suckered tube feet; nonetheless,histological sections revealed a distinctive internal morphology.Finally, we report the first observations of the tube-foot morphologyof representatives of deep-sea asteroids belonging to the ordersNotomyotida and Brisingida, a group that also lacks the typicalflat-tipped, suckered tube-foot morphology. The results of ourstudy demonstrate that the current tube-foot morphology modelneeds to be reconsidered, as there is considerably greater variationthan was previously believed to be the case. Moreover, we concludethat while tube-foot morphologies show consistent similaritieswithin orders, tube-foot morphology is less appropriate as ataxonomic character below this level.