The improbability of a muscular crinoid column

Donovan, Stephen K. 1989 07 15: The improbability of a muscular crirroid column. Lethaia , Vol. 22, pp. 307–315. Oslo. ISSN 0024–1164.
It has sometimes been assumed that the stems of ancient crinoids were muscular. However, the only contractile fibres found in the stems of extant crinoids are in the cirri of isocrinids and comatulids. On uniformitarian grounds, it is improbable that any ancient crinoid had tissues in the stem that would have aided active orientation, apart from the non-muscular catch connective tissue. This conclusion is supported by a detailed functional analysis of the various forms of column and cirrus articulation. The axial canal has been seen as the probable site of any column musculature, but this is functionally impossible with symplexial and synostosial articulations, both of which are poorly adapted for muscular flexure. Columns that articulate synarthrially are much more flexible, but modern bourgueticrinids lack any stem musculature, despite having such stalks. The only crinoid columns that were functionally adapted to utilize contractile fibres were those of certain coiled-stemmed taxa, where the axial canal was not in the plane of the synarthrial fulcra. However, it is unlikely that such stems, even if muscular, possessed any advantage over a non-muscular column. The only pelmatozoans with a probable muscular column were not crinoids, but the glyptocystitid rhombiferans. * Crinoids, stem, columnal, cirrus, catch connective tissue, contractile fibres, functional morphology .     

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.     

Chromatography of human prothrombin from Nitschmann fraction III on Q Sepharose Fast Flow using axial and radial flow column

An axial column (Hitrap Q 5 ml, 2.5 2 1.6 cm) and a radial flow column (3.5 2 5 cm) packed with Q Sepharose Fast Flow media had been evaluated for the separation of human prothrombin. Nitschmann fraction III dissolved in buffered saline (0.10 M sodium chloride buffered with 0.06 M Tris/HCl to pH 7.5) was the starting material. Effects of sample flow rate of the two columns were screened. Under radial flow conditions using the radial column, sample flow rate up to 15 ml/min (i.e. 18 bed volumes/h) was achieved and the operating pressure was below 0.2 MPa eventhough the elution velocity was 30 ml/min. Breakthrough capacity was determined by analyzing the total protein and prothrombin activity of the target protein-containing fraction under subsaturating conditions and both columns had almost the same breakthrough capacity per ml media, indicating that the sample loading was independent of radial column geometry. It was concluded that the radial column is an attractive alternate to traditional axial packed bed column, exhibiting very good potential for use in the separation of human prothrombin.     

Evolutionary history of regeneration in crinoids (Echinodermata)

The fossil record indicates that crinoids have exhibited remarkable regenerative abilities since their origin in the Ordovician, abilities that they likely inherited from stem-group echinoderms. Regeneration in extant and fossil crinoids is recognized by abrupt differences in the size of abutting plates, aberrant branching patterns, and discontinuities in carbon isotopes. While recovery is common, not all lost body parts can be regenerated; filling plates and overgrowths are evidence of non-regenerative healing. Considering them as a whole, Paleozoic crinoids exhibit the same range of regenerative and non-regenerative healing as Recent crinoids. For example, Paleozoic and extant crinoids show evidence of crown regeneration and stalk regrowth, which can occur only if the entoneural nerve center (chambered organ) remains intact. One group of Paleozoic crinoids, the camerates, may be an exception in that they probably could not regenerate their complex calyx-plating arrangements, including arm facets, but their calyxes could be healed with reparative plates. With that exception, and despite evidence for increases in predation pressure, there is no compelling evidence that crinoids have changed though time in their ability to recover from wounds. Finally, although crinoid appendages may be lost as a consequence of severe abiotic stress and through ontogenetic development, spatiotemporal changes in the intensity and frequency of biotic interactions, especially direct attacks, are the most likely explanation for observed patterns of regeneration and autotomy in crinoids.     

Survival of crinoid stems following decapitation: evidence from the Ordovician and palaeobiological implications

It was recently discovered that the stems of extant crinoids may survive after detachment of the crown, presumably feeding by the absorption of nutrients through the ectoderm. Previously, only one analogous, albeit morphologically dissimilar, pattern of crownless survival has been recognized from the fossil record. Certain Upper Ordovician (Cincinnatian) crinoid pluricolumnals from Kentucky, Ohio and Indiana, derived from the disparids Cincinnaticrinus spp., have rounded terminations reminiscent of some modern bourgueticrinid overgrowths. Such specimens have hitherto been interpreted as distal terminations of mature individuals that have become detached from their attachment structures and taken to an eleutherozoic existence. However, it is considered more probable that they represent overgrowths of the column following predatory decapitation. If this new interpretation is correct, then post-decapitation survival of crinoid stems is now recognized for most of the history of the crinoids, 'lethal' predation on crinoid crowns occurred at least as early as the Late Ordovician and ancient crinoid populations can no longer be determined merely by counting crowns.     

Disarticulation patterns in Ordovician crinoids: Implications for the evolutionary history of connective tissue in the Crinoidea

Taphonomic information is examined to evaluate the early history of connective tissues in the Crinoidea. The pattern of stalk segmentation of Middle and Late Ordovician crinoids is consistent with the two-ligament (intercolumnal and through-going ligaments) pattern present in living isocrinid crinoids and interpreted for fossil isocrinids, holocrinids, and Lower Mississippian crinoids. A single rhombiferan was also examined; its taphonomic pattern is also indicative of this style of tissue organization. Furthermore, the taphonomy of all Middle and Late Ordovician crinoids may reflect that they lacked discretely organized muscles between arm brachials, which is consistent with the hypothesis that muscles evolved as a connective tissue between plates only once within the Crinoidea, during the Early Devonian. These data indicate that the two-ligament organization of the stalk is a primitive feature among the Crinoidea and perhaps even among stalked echinoderms. Therefore, the autotomy function of this column-tissue organization among living crinoids is an exaptation. On the other hand, discretely organized muscles as connective tissue in crinoid arms is a derived trait that first appeared during the middle Paleozoic; this adaptation proved very successful for the advanced cladid crinoids.     

SOFT-TISSUE PRESERVATION OF THE HIND GUT IN A NEW GENUS OF CLADID CRINOID FROM THE MISSISSIPPIAN (VISEAN, ASBIAN) AT ST ANDREWS, SCOTLAND

Abstract:  Soft-tissue preservation of the hind gut, or anal sac, in the tegmen of Tubulusocrinus (gen. nov.) doliolus (Wright) from the Mississippian (Visean, Asbian) Pittenweem Formation at St Andrews, Scotland, is the first of its kind known in crinoids; it sheds important new light on the nature of the cladid tegmen. Many cladid crinoids had a calcite-plated anal sac that may have functioned like a chimney to prevent fouling of the ambulacra. The tubular, uncalcified anal sac of Tubulusocrinus may have functioned like a hose by pointing down current between the arms to avoid fouling of the ambulacra. Its smaller size may also have required less energy to produce than a heavy-plated sac. The hind gut is apparently preserved by haematite replacement, which was probably altered from original authigenic pyrite that formed soon after catastrophic burial.     

Train crash crinoids revisited

‘Train crash crinoids’ represent an unusual mode of preservation of crinoid columns, superficially resembling to the carriages of a crashed train. They were exhumed from the White Peak (Mississippian limestones) of the Peak District in the Treak Cliff area of Castleton, Derbyshire, north‐central England, and were presented by Broadhurst & Simpson as part of a varied suite of observations supporting the recognition of an ancient apron reef (= fore‐reef slope). A re‐examination of these specimens shows that they were not live crinoids at or near the reef crest, but were recently dead specimens, recumbent on the sediment surface. Something triggered their movement into deeper water, and some travelled downslope parallel to the direction of movement and broke into a ‘train crash pattern’ when the lower end impacted on an immovable object.     

Complete anaerobic mineralization of pentachlorophenol (PCP) under continuous flow conditions by sequential combination of PCP‐dechlorinating and phenol‐degrading consortia

Complete mineralization of 50 µM of pentachlorophenol (PCP) was achieved anaerobically under continuous flow conditions using two columns connected in series with a hydraulic retention time of 14.2 days, showing the highest reported mineralization rate yet of 3.5 µM day^?1. The first column, when injected with a reductive PCP dechlorinating consortium, dechlorinated PCP to mainly phenol and traces of 3‐chlorophenol (3‐CP) using lactate supplied continuously as an electron donor. The second column, with an anaerobic phenol degrading consortium, decomposed phenol and 3‐CP under iron‐reducing conditions with substantial fermentative degradation of organic compounds. When 20 mM of lactate was introduced into the first column, the phenol degradation activity of the second column was lost in a short period of time, because the amorphous Fe(III) oxide (FeOOH) that had been packed in the column before use was depleted by lactate metabolites, such as acetate and propionate, flowing into the second column from the first column. The complete mineralization of PCP was maintained for a long period by reducing the lactate concentration to 4 mM, effectively extending the longevity of second‐column activity with no depletion of FeOOH for more than 200 pore volumes (corresponding to 3,000 days). The carbon balance showed that 50 µM PCP and 4 mM lactate in the influent had transformed to CO₂ (81%) and CH₄ (3%) and had contributed to biomass growth (8%). A comparison of the microbial consortia introduced into the columns and those flowing out from the columns suggested that the introduced population did not flow out during the experiments, although the microbial composition of the phenol column was considered to be affected by the inflow of microbes from the PCP dechlorination column. These results suggest that a sequential combination of reductive dechlorinating and anaerobic oxidizing consortia is useful for anaerobic remediation of chlorinated aromatic compounds in a microbial permeable reactive barrier. Biotechnol. Bioeng. 2010;107: 775–785. © 2010 Wiley Periodicals, Inc.     

Efficacy of a pentaiodide resin disinfectant on Cryptosporidium parvum (Apicomplexa: Cryptosporidiidae) oocysts in vitro

The resin-I5 column developed at Kansas State University was tested for efficacy against oocysts of Cryptosporidium parvum (Apicomplexa: Cryptosporidiidae). Cesium chloride gradient-purified oocysts were passed through 1.0-cm-diameter columns with lengths of 2.5, 5.0, and 10.0 cm at 23 C. Following column passage, oocyst viability was determined both in vitro by excystation and in vivo by the ability to establish infections in suckling mice. Oocysts were found to be retained by the pentaiodide resin in a linear fashion, probably by electrostatic interactions. Linear regression analysis revealed 100% of the oocysts should be removed in such a manner using a column length of greater than or equal to 25.7 cm. When compared to untreated control oocysts, less than 12% of the oocysts that passed through the columns appeared to be affected by the resin, as assessed by excystation. Inoculation of suckling mice with these column-treated oocysts supported the excystation data and revealed the coccidian to be viable. These results indicate that oocysts of C. parvum are retained on the pentaiodide column in a 1-hit manner and that, although killing of parasites may occur within the column, the greatest effect that the column may have on the parasite is as an electrostatic retention device.     

Reusability of immunoaffinity columns for determination of fumonisins in maize

Eighteen maize samples were assayed for fumonisin B1 (FB1) and B2 content by immunoaffinity column coupled with high performance liquid chromatography (HPLC). The FumoniTest columns were used once for the isolation of fumonisins (single-use column method). In the second part of the assay the columns were regenerated. After elution with methanol, PBS solution was left on the column for one day at room temperature to regenerate the columns (regenerated column method). The efficiency of columns regenerated twice was tested by determining FB, recovery and the reproducibility of the determinations. The recovery rate of FB1 proved to be 82% by the single-use column method (RSD: 5.7%) and 82.6% (RSD: 5.6 %) by the regenerated column method; 500-8,000 ng FB1 loaded onto the columns did not affect column performances. Nearly identical values were obtained when the FB1 content of fumonisin-containing maize samples was determined by both methods. The results indicate that the FumoniTest columns can be regenerated by the method applied at least twice without decrease in column performance. The fumonisin affinity, capacity and specificity of the regenerated columns were not changed. Thus, columns regenerated in this way can be used for determining the fumonisin content of maize samples at least three times.     

Infestation of Middle Devonian (Givetian) camerate crinoids by platyceratid gastropods and its implications for the nature of their biotic interaction

One of the classic examples of biotic interactions preserved in the fossil record is that between crinoids and infesting platyceratid gastropods. This relationship, spanning an interval from the Middle Ordovician to the end of the Permian, is recognized by the firm attachment and positioning of platyceratids over the anal vent of their hosts. Several hypotheses have been proposed to explain this interaction; the most widely accepted is that the gastropods were coprophagous commensals, feeding on crinoid excrement without any significant detriment to their hosts. The purpose of this investigation was to test this hypothesis. Two species of Middle Devonian camerate (Monobathrida, Compsocrinina) crinoids, Gennaeocrinus variabilis Kesling & Smith 1962 and Corocrinus calypso (Hall 1862), were used in this investigation. The data consisted of 426 individuals of G. variabilis collected near Rockport, Michigan, 30 of which were infested, and 188 individuals of C. calypso collected near Arkona, Ontario, Canada, of which 25 were infested. Length and volume were measured for each crinoid to determine whether a significant difference existed in the size of infested versus uninfested individuals. The results indicated that for both species of crinoids individuals infested by snails were significantly smaller than uninfested individuals (p < 0.05). We explored a variety of scenarios to explain this pattern and conclude that they falsify the null hypothesis that the crinoid-gastropod relationship was strictly commensal. The smaller size of the infested crinoids is interpreted as a consequence of nutrient-stealing by the parasitic gastropods, a strategy that finds analogs in modern seas. Moreover, the absence of platyceratids on the largest crinoids suggests that large size may have inferred immunity from lasting infestation.     

Fractionation of lymphocytes involved in the generation of cell-mediated cytotoxicity over insolubilized conjugated histamine columns.

Spleen cells from normal BALB/c mice were cultured in vitro with irradiated C57BL/6 stimulating cells. Five days later the T cell-mediated cytotoxic activity of the effector cells was assessed with a 51Cr-release assay that used H-2bEL-4 tumor cells as targets. Before the BALB/c responding lymphocytes were sensitized they were fractionated by passing the spleen cells over insolubilized histamine rabbit serum albumin Sepharose columns (H-RSA-S) or over rabbit serum albumin Sepharose (RSA-S) control columns. Fractionation of cells over the H-RSA-S columns depleted or significantly reduced the cytotoxic potential of the unretained cells. All cytotoxic potential was recovered when the cells that adhered to the H-RSA-S were eluted from the columns. In contrast, no effect on responsiveness was detected after the cells had been fractionated over the control column. The loss of response potential by the cells that did not adhere to H-RSA-S could not be accounted for by removal of macrophages nor by the concentration of cells with suppressor activity in the effluent. These cell fractionation studies raise the possiblity but do not prove that cytotoxic precursor cells may express amine receptors that could be responsible for their retention by insolubilized histamine columns.     

Late Ordovician extinction of North American and British crinoids

After reaching a diversity peak in the Caradocian, North American Ordovician crinoids underwent a gradual decline to a nadir in the early Ashgillian (Maysvillian). This interval, recording extinction of the Cleiocrinidae, Merocrinidae, Ottawacrinidae, Hybocystitidae, and several lineages of camerate crinoids, was apparently caused by major environmental shifts in seas of eastern North America resulting from a westward-prograding wedge of terrigenous clastics derived from the Taconic Highlands, possibly coupled with a marine transgression in the Maysvillian that allowed colder water slope biofacies to invade the craton. Crinoids suffered a major episode of extinction in the late Ashgillian (late Richmondian/Rawtheyan). This event, preceding the end of the Ordovician by at least one stage or 2 to 4 million years, resulted in extinction of 12 families of crinoids including the Xenocrinidae, Tanaocrinidae, Reteocrinidae, Archaeocrinidae, Anthracocrinidae, Cincinnaticrinidae, Iocrinidae, Anomalocrinidae, Carabocrinidae, Cupulocrinidae, Porocrinidae, and Hybocrinidae. Glacio-eustatic lowering of sea level may have triggered this crisis by partially draining the North American craton, resulting in changes in oceanic circulation, salinity, and temperature. Latest Ordovician (Hirnantian) carbonates of the North American mid-continent region contain pelmatozoan assemblages from which Silurian crinoids radiated. These taxa were largely unaffected by a minor extinction event at the Ordovician/Silurian boundary.     

A new holocrinid (Articulata) from the Paris Biota (Bear Lake County,Idaho, USA) highlights the high diversity of Early Triassic crinoids

After the end-Permian crisis and the extinction of their four Paleozoic subclasses, crinoids rapidly recovered. This group is classically believed to have radiated from a small surviving clade and to have diversified during the Middle and Upper Triassic from two lineages. Nevertheless, recent findings suggested that several lineages of crinoids had already diversified during the Early Triassic, and that their diversity has been overlooked. Here we describe a new form of holocrinid, Holocrinus nov. sp., from the earliest Spathian (Early Triassic) of southeastern Idaho (USA). So far, the exceptional completeness of sampled specimens, with skeletal elements of arms and stem in connection, is unique for the Early Triassic. They show that derived morphological features had already evolved ∼1.3 million years after the Permian–Triassic boundary, supporting the scenario of a rapid Early Triassic diversification of crinoids.     

The root of the problem: palaeoecology of distinctive crinoid attachment structures from the Silurian (Wenlock) of Gotland

The holdfast (attachment structure) is the most understudied aspect of the palaeoecology of the endoskeleton of fossil crinoids. A new collection of well-preserved holdfasts from a recently reopened quarry at Hunninge, Gotland, in Homerian (upper Wenlock) strata includes several morphologies. The most common are terminal dendritic radicular holdfasts (TDRHs) that may be derived from the cladid Ennallocrinus d'Orbigny. These have a consistent morphology of five, equally spaced, long radices that spread across the sea floor. These crinoids were gregarious, and TDRHs in a group commonly show the same radice orientations. The radices have a large axial canal compared with those of modern crinoids; each included, at least, nervous tissues. Taken together, these features suggest that, apart from attachment, these distinctive TDRHs may have served a sensory function. Other holdfasts in this assemblage also show monospecific aggregations, perhaps suggesting biochemical attraction such as that shown by certain other sessile invertebrates such as barnacles.     

Application of microbore HPLC in combination with tandem MS for the quantification of rosuvastatin in human plasma

The potential of microbore high-performance liquid chromatography (HPLC) in combination with tandem mass spectrometry (MS/MS) for the sensitive detection of rosuvastatin (Crestor) in human plasma was investigated. Three microbore HPLC columns with internal diameters (i.d.) of 0.5, 1.0 and 2.0 mm were evaluated for column efficiency and mass sensitivity, and compared to a conventional 4.6 mm i.d. column. The 2.0 and 1.0 mm i.d. columns performed very well while the 0.5 mm i.d. column was slightly less efficient, this is probably due to a lower packing density. Good results with respect to gains in mass sensitivity compared to the conventional analytical column were achieved with the 2.0 and 1.0 mm columns. Thus, the 2.0 mm i.d. column had an improved signal-to-noise (S/N) ratio of 16 whilst the 1.0 mm i.d. column had an improved S/N ratio of greater than 70. Experiments with the 1.0 mm i.d. HPLC column were performed to determine the robustness of the microbore method for human plasma extracts after sample preparation using solid-phase extraction (SPE). A number of problems were encountered with extracts including high backgrounds, the blocking of the column and a rapid deterioration in column performance. The blocking of the column by particulates was solved by off-line filtration of the sample extracts. Peak tailing of the analytes and high background, both of which were due to endogenous interferences in the extracts, were eliminated using gradient elution. Using these approaches over 500 injections of plasma extracts were achieved without significant deterioration in assay performance. Quantities of rosuvastatin of 0.3 pg on-column could be detected and cross-validation experiments demonstrated that the conventional and the microbore HPLC-MS/MS methods provided similar information on the concentration of rosuvastatin but with greatly reduced sample consumption using the microbore method.     

Different bioreactor configurations for the decolourisation of the azo dye reactive black 5 by Geotrichum sp. CCMI 1019

Decolourisation of the azo dye Reactive Black 5 by Geotrichum sp. CCMI 1019 was studied using stirred tank reactors (STR) and two types of bubble columns (porous plate (PP) bubble column and aeration tube (AT) bubble column). For the bubble columns, the k_La increased with the gas fractional hold-up (ε_G) and the aeration rate. A linear relationship between ε_G and superficial gas velocity was obtained for all reactors. At same aeration rates, the PP bubble columns showed higher k_La and hold-up values than the AT bubble column. In the STRs, large and dense aggregates were formed which adhered to surfaces whereas bubble columns gave smaller and less compact pellets.

Manganese peroxidase and laccase were detected in the extracellular media in all reactors. However, laccase was only detected after the onset of decolourisation, suggesting that additional enzymes may be involved. Mn peroxidase activity was detected (about 46 U/ml) in both the STRs and AT bubble columns but higher values (110 U/ml) were obtained with the PP bubble columns.

Out of the three reactor systems studied, the AT bubble columns gave the most favourable results for Reactive Black 5 decolourisation. Rapid and complete colour removal was obtained throughout the visible spectrum. Bubble columns are simple in design as well as operation and may be useful for the bioremediation of textile wastewater.     

花卉柱式无土栽培

于玻璃温室内的160 m2面积上,以干湿交替型盆钵组装起高200 cm, 直径15 cm的立柱77根,然后组成立柱“树林”并种花,25个科的共53种草花用同一营养系统管理,生长都良好,立柱“树林”象是花的“森林”,显示了良好的生态关系。在此基础上挑选不同种盆花组成不同情趣的家庭阳台花柱。阳台花柱具若干优点：新型盆钵具良好的水、气、肥协调关系,适合多种花卉生长;柱高任选,便手提携带;柱体能环绕中轴旋转使植物受光均匀;花柱底部是具中岛结构式底盆,起蓄水和稳定作用,还便于多根立柱串联扩大栽培量;有人工和自动浇灌两种系统,省工省时且干净卫生。     

Effect of column dimensions and flow rates on size-exclusion refolding of beta-lactamase

We have investigated the effect of changing the column diameter and length on the size exclusion chromatography (SEC) refolding of beta-lactamase from Escherichia coli-derived inclusion bodies (IBs). Inclusion bodies were recovered and solubilised in 6 M GdnHCl and 5 mM DTT. Up to 16 mg of denatured, solubilised beta-lactamase was loaded onto size exclusion columns packed with Sephacryl S-300 media (fractionation range: 10(4)-1.5 x 10(6) Da). beta-Lactamase was refolded by eluting the loaded sample with 1 M urea in 0.05 M phosphate buffer, pH 7 at 23 degrees C. The following columns were studied: 26 x 400, 16 x 400 and 26 x 200 mm, with a range of mobile phase flow rates from 0.33 to 4.00 ml/min. beta-Lactamase was successfully refolded in all three columns and at all flow rates studied. The beta-lactamase activity peak coincided with the major protein peak. Reducing the column diameter had little effect on refolding performance. The enzyme activity recovered was relatively independent of the mobile phase linear velocity. Reducing the column length gave a poorer resolution of the protein peaks, but the enzyme activity peaks were well resolved. Calculation of the partition coefficients for beta-lactamase activity showed that the 26 x 400 column gave the greatest refolding performance.