Functional morphology of synostosial articulations in the crinoid column

The rarest articulation found in the crinoid column is the synostosis, in which adjacent articular facets are essentially planar. 'Synostoses' in the columns of post-Palaeozoic isocrinids are more correctly termed cryptosymplexies. comprising symplectial articulations which have become infilled by secondary calcite. Cryptosymplexial joints are totally inflexible and are the preferred sites of stem autotomy. Synostoses sensu stricto appear to have been limited to early Palaeozoic taxa, but this form of articulation was soon succeeded by the symplexy. Synostoses were probably commonest in meric columns which evolved from hohlwurzels and in which the main flexibility was along meric sutures, rather than between columnals. With the evolution of the holomeric columnal. a more flexible articulation between columnals was a functional necessity in order that the stem did not develop into a stiffened rod. The solution was the evolution of the symplexy. *** Crinoids, columnals. synostosis, functional morphology evolution.     

Early stalked stages in ontogeny of the living isocrinid sea lily Metacrinus rotundus

The early stalked stages of an isocrinid sea lily, Metacrinus rotundus, were examined up to the early pentacrinoid stage. Larvae induced to settle on bivalve shells and cultured in the laboratory developed into late cystideans. Three‐dimensional (3D) images reconstructed from very early to middle cystideans indicated that 15 radial podia composed of five triplets form synchronously from the crescent‐shaped hydrocoel. The orientation of the hydrocoel indicated that the settled postlarvae lean posteriorly. In very early cystideans, the orals, radials, basals and infrabasals, with five plates each in the crown, about five columnals in the stalk, and five terminal stem plates in the attachment disc, had already formed. In mid‐cystideans, an anal plate appeared in the crown. Late cystideans cultured in the field developed into pentacrinoids about 5 months after settlement. These pentacrinoids shared many crown structures with adult sea lilies. On the other hand, many features of the stalk differed from those in adult isocrinids, while sharing many characteristics with the stalk of feather star pentacrinoids, including disc‐like proximal columnals, high and slender median columnals, synarthrial articulations developmentally derived from the symplexial articulations, limited formation of cirri only in the proximal columnal(s), and an attachment disc. On the basis of these findings, phylogenetic relationships among extant crinoid orders are discussed.     

Cretaceous and Paleogene Bourgueticrinina (Echinodermata,Crinoidea) of the USSR

The theca and the columnals of twenty-two lateCretaceous species of the genus Bourgueticrinus (including two new species), of one species of the genus Democrinus, of four species of the genus Conocrinus (including one new species) and of one species of the genus Paleobathycrinus known from USSR are figured and shortly described. Biometric reconstructions of stems for B. aequalis, B. bellus, B. danicus, B. fischeri and C. tauricus nov. sp. are given. A new index is used to suggest the relative flexibility of stem. Decrease in dimension and flexibility of the arms in several species take place simultaneously with an increase of stem flexibility. This fact seems to be closely related to environmental changes.     

Columnal ontogeny in articulate crinoids and its implications for their phylogeny

Simms, Michael J. 1969 01 15: Columnal ontogeny in articulate crinoids and its implications for their phylogeny. Lethaia , Vol. 22, pp. 61–68. Oslo. ISSN 0024–1164.
Detailed doamentation of columnal ontogeny in several Lower Jurassic isocrinids reveals a number of features which are of phylogenetic significance. Columnal proportions are determined by two largely independent growth vectors. Columnal diameter is primarily a function of cup size at the time of formation, though it may subsequently undergo considerable increase through lateral accretion. Columnal height is largely independent of cup size except for a gradual increase with the growth of the whole animal. Columnal diameter increases at a much greater rate than columnal height and so columnals are relatively lower in large individuals. Heterochronous variation in vertical growth rate or the extent of lateral accretion in columnals may significantly alter their shape. The smallest isocrinid columnals found (0.35 mm diameter) have synarthrial articula. The transformation to a typical pentaradiate symplexy, via a triradiate stage, occurs at a diameter of about 0.75 mm. Synarthrial articula are unknown in millericrinid columnals, which instead have symplectial articula at small diameters (0.45 mm). The presence of synarthrial articula in the stems of juvenile isocrinids, and their absence in millericrinids. supports previous hypotheses concerning derivation of bourgueticrinids from an isocrinid, rather than a millericrinid, ancestor.     

A scanning electron microscope technique for restoring deformed fossils

Deformed fossils which originally had a regular geometric outline, such as crinoid columnals, can be restored using a scanning electron microscope technique called tilt correction. This magnifies the specimen in one direction only. It is unsuitable for specimens where deformation has occurred unevenly and can also distort small structures while restoring the whole specimen. D Crinoid columnals, scanning electron microscope, tilt correction.     

Functional morphology of synarthrial articulations in the crinoid stem

Synarthrial fulcral ridges are found in crinoid columnals from the mid-Ordovician to the present and in all four subclasses. Similar articulations did not become common in the cirri until the Mesozoic. Synarthrial stem articulations fall into two broad groups. Type I articulations have a fulcral ridge in the centre of the articular facet. In elliptical ossicles this ridge corresponds to either the long (IA) or short (IB) axis of the facet. Although both are functionally similar, Type IA ossicles are more common. Type II articulations have an excentric fulcral ridge, parallel to either the long (IIA) or short (IIB) axis of the articular facet. Type IIA articulations are found in crinoid stems capable of coiling. Type II articulations are particularly common in the cirri of articulates and are well adapted for attachment to hard and soft substrates. Columnals with Type I articulations often have divergent fulcra, giving the stem flexibility in all directions, but this feature is not seen in cirri or in coiled stems, where it would impair normal functions. Only the cirri of isocrinids and comatulids are muscular, so the movement of columns with fulcra must be passive.     

Discovery of two rare species of stalked crinoids from Okinawa Trough, southwestern Japan, and their systematic and biogeographic implications

A recent submersible dive on the southern slope of Aguni Knoll, central Okinawa Trough has led to a discovery of two rare species of stalked crinoids, Proisocrinus ruberrimus and Naumachocrinus hawaiiensis, both of uncertain taxonomic affinity. The specimens of Proisocrinus ruberrimus, here termed "Moulin rouge" as a pseudonym due to its conspicuous red color, were collected from depths of approximately 1,800 m and are characterized by a long, slender stalk. The proximal columnals are pentalobate in cross section, with isocrinid-like articular facets; several bear rudimentary cirri. The following columnals lose the cirri and rapidly become almost circular; their facets bear radiating crenulations similar to those of Millericrinida. The characters observed in the proximal stalk indicate that the 'Moulin Rouge' (P. ruberrimus) belongs to the order Isocrinida rather than to the order Millericrinida. Naumachocrinus hawaiiensis was collected at a depth of 1,440 m. The long, cylindrical crown and number of arms indicate that Naumachocrinus should be classified in the Bathycrinidae.     

Phenotypic variability, deformations and post-traumatic reactions of the stem ofEncrinus cf.liliiformis Lamarck (Crinoidea) from the Middle Triassic of Qingyan, south-western China

Well-preserved stem remains ofEncrinus cf.liliiformis Lamarck 1801 from the Upper Anisian and lowest Ladinian (Middle Triassic) of Qingyan, Guizhou Province, south-western China, are described. The characteristic morphological features of the columnals, especially the sculptural characteristics of the articular facets, vary in the different parts of the stem. Numerous fine, short, peripheral crenulae and perilumina with a tendency to pentalobate shapes in the proximal stem region change to few long, coarse crenulae and circular perilumina in the distal stem part. The holdfasts exhibit various differentiations: the basic discoid type occurs on flat surfaces, encrusting holdfasts are characteristic of irregular substrates; juvenile holdfasts often show an irregular outline, adult ones have a more even, circular shape. Some special and unusual structures allow palaeoecological interpretations: Holdfasts and stems joined by callous growth probably result from the lack of suitable hard substrates for attachment on the soft seafloor; the larvae often had to settle very closely to each other on a relatively small substrate. After breakage of stem,Encrinus was able to continue living despite the traumatic loss of its basal fixation. Broken ends underwent skeletal regeneration. Some specimens strongly suggest that juvenile individuals sometimes even were able to re-attach secondarily, e.g. by coiling around stems of other crinoids and cementing to these by callous outgrowths.     

Stalked crinoids from a Jurassic tidal deposit in western North America

This is the first systematic and paleoecological study of a crinoidal limestone (encrinite) from the Jurassic System of North America. The encrinite is part of a shallow-water tidal facies of the Middle Jurassic Carmel Formation located at Mount Carmel Junction (southwestern Utah, U.S.A.) and may represent one of the youngest shallow-water encrinites in the geological record. In the past, the crinoid at this locality was referred to as Pentacrinus asteriscus, a name used to describe almost all of the crinoid columnals found throughout the Jurassic of the U.S. western interior. However, systematic work indicates that the crinoid is Isocrinus nicoleti and is the first non-endemic crinoid to be reported from North American Jurassic strata. Although articulated pinnules and arms have been found, I. nicoleti occurs predominantly as well-preserved, partially articulated columnals. The crinoids occur within a tidal complex consisting of ooid shoal, tidal channel, and lagoonal facies. The unique environmental and ecological conditions which existed in the southernmost end of the Jurassic North America seaway may have allowed for the development of this crinoid colony and subsequent deposition of the encrinite.     

The skeleton of the stalked stages of the comatulid crinoid Antedon bifida (Echinodermata)

Summary The cystidean calyx of Antedon bifida (Pennant, 1777) consists of two superposed series of five inter-radially located plates called basals (lower plates) and orals (upper plates). A single radianal and five radial plates develop in early pentacrinoids. They are radially located between the orals and the basals and alternate with them. During pentacrinoid life, radials grow continuously while orals progressively regress. Basals fuse to the proximal forming the so-called rosette, and the radianal grows till the anal cone is formed and then regresses. The first primibrachials arise in 25 day pentacrinoids. Arms of detached juveniles usually have 13 secundibrachials, with IIBr13 each having a developing pinnule. No infrabasal develops between the series of basals and the upper columnal.During pentacrinoid life, the stalk progressively changes from homeomorphic to xenomorphic while most columnal articulations change from symmorphies to synarthries. Synarthrial fulcral ridges can rotate about 60° from one columnal articulation to another, allowing the stalk to be flexible in every direction. Formation of new columnals is continuous until the proximal appears (i.e. in 65 day pentacrinoids). The number of columnals in the fully-formed pentacrinoid stalk varies from 17 to 24 depending on the individual. Developing cirri are seen in 70 day pentacrinoids. Detached juveniles have two well-developed series of five cirri, which usually consist of ten cirral ossicles.     

Alpine and Arctic tundra shrub populations show similar ontogenetic growth trends but differing absolute growth rates and lifespan

The majority of terrestrial primary production is performed by plants, the ontogenetic growth trends of which significantly influence biomass and carbon dynamics. Here, we present a study of ontogenetic trends in primary (apical) and secondary (stem thickening) growth of plants in Arctic (Svalbard, Norway) and alpine (Krkonoše, Czechia) populations of the black crowberry (Empetrum nigrum), the dominant plant species of certain tundra communities. The environmental conditions in alpine areas are more favourable for plant growth than those in the High Arctic, where temperatures are lower, there is less precipitation and soils are shallower, among other differences. These differences were clearly reflected in significant distinctions in absolute growth rates and shrub age between the populations under study. However, we found almost no divergence in ontogenetic growth trends between the populations (based on ring width measurements made from the base to the top of plants, known as serial sectioning). In both populations, primary and secondary stem base growth decrease over the course of ontogeny whereas secondary stem top growth and basal area increment increase. No significant differences in the slope of the trends were found in either primary or secondary stem base growth. Trends of the growth ratio between basal area increment and primary growth revealed neither absolute nor relative differences between the populations. Ontogenetic trends in the shrubs analysed were surprisingly stable despite the prominently dissimilar environmental conditions. Empetrum plants have adapted to the different environments by altering their absolute growth rate only. This adaptation has probably also resulted in the different longevity of plants constituting the study populations, confirming the theory that slower-growing plants live longer. Primary growth and secondary stem base growth seem to be more basic characteristics of plant growth compared to basal area increment and secondary growth at the apex because the latter two seem to be dependent on the absolute growth rate.     

Pelmatozoan arms from the Middle Cambrian of Australia: bridging the gap between brachioles and brachials?

The early Middle Cambrian Monastery Creek Phosphorite (Beetle Creek Formation, Queensland, Australia) contains an assemblage of disarticulated echinoderm ossicles that are exquisitely preserved. Amongst this material we recognize pelmatozoan brachials, radials, basals and holomeric columnals. Although we cannot reconstruct the complete animal with precision, these elements represent the oldest known pelmatozoan with crinoid-like appendages. Key elements include isotomously to heterotomously branched uniserial appendage plates with a tripartite adoral food groove, a longitudinal central canal interpreted as housing entoneural nerve, and differentiated articulation facets. There are also epispire-bearing radials bearing one to four arm insertion-facets, each one pierced by a central neural canal. These canals run internal towards the oral area beneath the external food groove. Co-occuring material includes single truncated cone-shaped basals and holomeric columnals, both with a similar articulation pattern, and irregular, epispire-bearing thecal plates. This mosaic of crinozoan (uniserial isotomous to heterotomous arms with neural canal), blastozoan (epispire-bearing thecal plates, appendage leading to oral thecal food groove without direct connection with body cavity) and apomorphic characters (circumoral instead of basal entoneural plexus) is unexpected and demonstrates that crinoid-like pelmatozoans with uniserial, branched arms appeared significantly earlier than the Tremadocian, when the first articulated crinoid skeletons are found. It also raises questions about the polyphyletic appearance of feeding appendages among pelmatozoan echinoderms.     

Cyrtocrinids (Cyrtocrinida,Crinoidea) and other associated crinoids from the Jurassic (Kimmeridgian–Tithonian)–Cretaceous (Berriasian–Barremian) of the Carpathian Foredeep basement (western Ukraine)

Upper Jurassic (Kimmeridgian–Tithonian) and Cretaceous (Berriasian–Barremian) strata of the Ukrainian part of the Carpathian Foredeep basement are rich, at least locally, in crinoid remains. Crinoids belonging to cyrtocrinids (Cyrtocrinida) are represented by whole cups, isolated remains of disarticulated cups, brachial plates and columnals. They are assigned to the following taxa: Cyrtocrinida indet., Eugeniacrinites cf. cariophilites (von Schlotheim), Lonchocrinus sp., Phyllocrinus stellaris Zaręczny, Ascidicrinus pentagonus (Jaekel), Gammarocrinites sp., Psalidocrinus armatus (Zittel), Psalidocrinus sp., and Hemibrachiocrinidae gen. indet. Cyrtocrinids are associated with other stalked (isocrinids, Isocrinida and millericrinids, Millericrinida) and stemless (saccocomids, Roveacrinida) crinoids. Columnals, pluricolumnals, brachial plates, and cirrals of isocrinids are assigned to Balanocrinus sp., Isocrinina fam. et subfam. indet., and columnals of millericrinids to Millericrinida indet. Free-living roveacrinids are assigned to Saccocoma sp. and Crassicoma sp. Knowledge on Jurassic and Cretaceous crinoids formerly described from Ukraine is discussed. Although majority of crinoids described herein seems to be allochthonous, autochthonous forms were also found with certainty in some intervals. These include some cyrtocrinids, which dominate in shallow-water environments of the Ukrainian Carpathian Foredeep basement. Isocrinids are also common in this shallow marine environment, whereas sessile saccocomids are assigned to low-energy, mud-supported bottom, open marine, outer-platform/upper slope, and relatively deep environments.     

Contrôle de la sécrétion stéréomique dans les stolons d'un Crinoïde nouveau du Dévonien inferieur d'Algérie

The discovery of numerous distal stem fragments of Eurax marhoumensis n. sp. brings new data on the formation of the stolons and the control of stereomic secretion. The presence of primary fixations in the dististele demonstrates the composite nature of stolonifcrous scgmcnts. The creeping zone is formed by gradual incorporation of the lower columnals of the mesistele into the stolon. Stereom growth rings observed on weathered facets or transverse sections are markers of the morphofunctional evolution of the columnals during different growth stages. The fixation organs are true stereomic outgrowths without axial canal or Segmentation; they grow from a single columnal and remain in connexion with it. The morphology of stolons results partly from the topography of the substrate and partly from control by stereomic secretion which influences the columnal thickness. The introduction of wedge-shaped columnals in curves of the stem and on the development of fixation devices. Epizoan attachment (bryozoans, corals, crinoids) induces different stereomic reaction depending on the growth stage of the crinoid column. The stereom of the host may proliferate as a sheet, abortive ossicles or shapeless stercomic masses which tend to limit the growth of the epizoans and incorporate them in the column stercom. □Crinoid, column, morphology, stereomic secretion, Devonian, Algeria. Le découverte de nombreux fragments pédonculaires distaux d'Eurax marhoumensis n. sp. apporte des données nouvelles sur le processus de la formation des stolons et sur le contrôle de la sécrétion stéréomique. L'existence de fixations primaires dans la dististéle démontre la nature composite des segments stoloniaux. Les zones rampantes se forment par incorporation progressive des columnales de la partie inférieure du pédoncule dans les stolons. Les cernes de croissance du stéréome observées sur les facettes altérées ou sur les sections constituent des témoins de leur évolution morphofonctionnelle au cours des différents stades de la croissance. Les organes de fixation correspondent à de véritables expansions stéréomiques dépourvues de canal axial et de segmentation; ils bourgeonnent àpartir d'une seule columnale avec laquelle ils rcstent en conncxion. La morphologie des stolons résultc en partie de la topographie du substratum et du contrôle de la sécrétion stéréomique qui se traduit par I'ajustement de l'epaisseur des columnales, l'introduction d'articles cunéiformes dans les courbures et le développement des organes de fixation. L'implantation d'épizoaires (Bryozoaires, Coraux, Crinoïdes) provoque des réactions stéréomiques différentes suivant le stade de croissance du pédoncule du Crinoïde. Le stéréome de I'hôte peut proliférer sous formc d'une lame, d'articles abortifs ou d'amas difformes qui tendent a limiter le développement de I'épizoaire et à l'incorporer dans le pédoncule. □Crinoïde, pédoncule, morphologie. Sécrétion stéréomique, Dévonien, Algérie.     

Devonian (Eifelian) disparid crinoids from the Zeravashan range,southern Tien Shan,Uzbekistan

No Devonian disparid crinoids have been described from Uzbekistan, although parahexacrinid thecae and assorted columnals have been described in several papers in the past 50 years. Discovery of a cup of Pisocrinus and a thecae of Haplocrinites are the first of these genera known from Uzbekistan. The stratigraphic range of Haplocrinites is revised to be late Early Devonian (Emsian) to Early Carboniferous (Tournaisian) because Silurian species assigned to the genus are based on loose ossicles that morphologically do not belong to the genus. The paleogeographic ranges of Pisocrinus and Haplocrinites are extended into Uzbekistan. Haplocrinites uzbekistanensis n. sp. is described.     

叉孢苏铁茎的解剖学研究

对叉孢苏铁茎的解剖结构进行了仔细的研究,结果表明:叉孢苏铁茎具有发达的皮层,皮层及髓中具有黏液道,有外韧型和周韧型的维管束分布在皮层中,但主要以外韧型为主,在同一切面上能看到横、纵、斜3种不同走向的维管束,所有皮层维官束韧皮部组成成份比较单一。维管组织区由多个生长环组成,每个生长环都具有木质部、维管形成层和韧皮部组成,各生长环韧皮部所占比例均大于木质部,在具4生长环的下部茎的维管组织中,环与环之间间隔2～10层不等的薄壁细胞。从纵向变化上看,从上至下,维管组织区的厚度及生长环数逐渐递增,在同一组织中,含晶及含单宁物质的异细胞呈从无到有,从少到多的分布趋势。     

New discoveries of Cambrian pelmatozoan echinoderm ossicles from North China

A variety of echinoderm ossicles have been discovered in the middle to upper Cambrian rocks of the North China Craton. Material was collected from two localities: Shuangqiao (SQ) section (Paibian) in Hebei Province and Linyi (LY) (Guzhangian?–Paibian?) in Shandong Province. Six types of ossicles can be recognized, including rounded thecal plates, and columnals with a narrow lumen and ridged articulating facets. The combination of characters suggests an unknown stalked echinoderm, probably an eocrinoid, with relatively advanced columnal morphology.     

Epi- and endobiontic organisms on Late Jurassic crinoid columns from the Negev Desert, Israel: Implications for co-evolution

Columns of the articulate crinoids Millericrinus and Apiocrinites from the Upper Jurassic (Upper Callovian) Zohar and Matmor formations of the Negev Desert of Israel display abundant encrusting organisms of about ten species, as well as diverse trace fossils produced by endobionts. Pluricolumnals were colonized by epi- and endobiontic organisms both during life and post-mortem. Skeletonized encrusting organisms include abundant ostreid bivalves (which evidently colonized both live and dead crinoid columnals), two types of serpulid worms, encrusting foraminifera, three species of bryozoans, and small encrusting sclerosponges. Several types of borings are present: Trypanites (possibly produced by sipunculids), Gastrochaenolites (crypts of boring lithophagid bivalves), elliptical barnacle? borings, and channel-like annelid? borings. In addition, approximately 16% of the pluricolumnals display circular parabolic embedment pits assignable to the ichnogenus Tremichnus. They are associated with substantial deformation of the containing columnals and were probably the work of host-specific ectoparasitic organisms. Discovery of Tremichnus on Jurassic crinoids extends the range of this trace by almost 100 million years, providing evidence for one of the longest-ranging host-parasite interactions documented thus far (over 200 million years). The relationship of epibionts to the Jurassic crinoids thus ranged from simple utilization of dead hard substrate to probable opportunistic commensalism in forms that colonized the live upright stems, as in some oysters, through host-specific parasitism in the case of Tremichnus.     

Middle Cambrian cystoid (sensu lato) stem columnals from Bornholm, Denmark

A rich material of echinoderm fragments from two Middle Cambrian stratigraphical levels on Bornholm are preserved due to phosphatization of the original calcitic stereom. Preservation of echinoderms in this way - not previously recorded from the Middle Cambrian - permits detailed analysis of the three-dimensional stereom structure. Identifiable are fragments of stylophorans and eocrinoids. Stem columnals, most likely from eocrinoids, show a wide and advanced morphological variation indicating articulation similar to that of crinoids. The material from the Exsulans Limestone/Kalby marl ( Ptychagnostus gibbus Zone) represents stem-bearing cystoids older than Akadocrinus from Bohemia. The Andrarum Limestone ( Sole-nopleura brachymetopa Zone) contains echinoderm fragments from a higher stratigraphical level, a level correlatable with that from which the oldest North American stem–bearing cystoid, Eustypocystis , has been recorded.     

Endogenous fluorescence signatures in living pluripotent stem cells change with loss of potency

The therapeutic potential of stem cells is limited by the non-uniformity of their phenotypic state. Thus it would be advantageous to noninvasively monitor stem cell status. Driven by this challenge, we employed multidimensional multiphoton microscopy to quantify changes in endogenous fluorescence occurring with pluripotent stem cell differentiation. We found that global and cellular-scale fluorescence lifetime of human embryonic stem cells (hESC) and murine embryonic stem cells (mESC) consistently decreased with differentiation. Less consistent were trends in endogenous fluorescence intensity with differentiation, suggesting intensity is more readily impacted by nuances of species and scale of analysis. What emerges is a practical and accessible approach to evaluate, and ultimately enrich, living stem cell populations based on changes in metabolism that could be exploited for both research and clinical applications.