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Crinoidea И Их Система. Blastozoa

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Crinoidea И Их Система. Blastozoa Crinozoa Echinodermata П/т Homalozoa (Camb.-Dev.) • П/т Echinozoa – Ctenocystoidea – Camptostromatoidea (Camb.) – Homostelea – Helicoplacoidea (Camb.) – Stylophora – Edrioasteroidea (Camb.-Carb.) – Homoiostelea – Edrioblastoidea (Ord.) – Cyclocystoidea (Ord.-Dev.) – П/т Blastozoa – Ophiocystoidea (Ord.-Carb.) – Eocrinoidea (Camb.-Sil.) – Echinoidea (Ord.-pr.) – Blastoidea (Camb.-Perm.) – Holothuroidea (Ord.-pr.) – Parablastoidea (Ord.) – Cystoidea (Ord.-Dev.) • Rombifera • П/т Asterozoa (Stelleroidea) • Diploporita • П/т Crinozoa (Pelmatozoa) – Somasteroidea (Ord.) – Paracrinoidea (Ord.-Sil.) – Asteroidea (Ord.-pr.) – Crinoidea (Camb.-pr.) – Ophiuroidea (Ord.-pr.) CRINOID TERMINOLOGY Monocyclic Crinoid Cup -первые чл. Рук - радиалия - базалия Dicyclic Crinoid Cup - первые чл. рук - радиалия - базалия - инфрабазалия Морфология чашечки коматулид Morphology of the calyx in comatulids. A, B: Species lacking a calyx sensu strictu, all ossicles of body and arms lying in the same plane. A: Aboral aspect of Comanthus suavia. Bar=0.3 mm. B: Aboral aspect of the macerated disc of Comanthus mirabilis formed of centrodorsale (ce), radialia (ra), and first brachials (primibrachials, pb). Note the absence of cirrus sockets. Bar=1 mm. C, D: Promachocrinus kerguelensis, a species with a large conical centrodorsale bearing numerous cirri. With their outer face, 10 wedge-shaped radialia (in D) make the arms rise in an inclined angle. With the inner surface they from a bow housing the viscera. Bars in C and D = 2mm. E, F: Tropiometra afra. E: X-ray photograph showing the disc- like centrodorsale (ce) seen from the side, perforated by thin canals (arrowheads) through which extensions of the chambered organ and accompanying nerves reach the cirri at the disc’s periphery. Bar = 3 mm. F: Insight into the partly macerated calyx seen obliquely from an oral point of view; radialia (ra), pimibrachialia (pb), and secundibrachialia (sb) contribute to the formation of the calyx wall. Bar=3 mm (from Heinzeller, Welsch, 1994). Морфология стебелька и цирр Morphology of stalk and cirri. A: Metacrinus interruptus. Bar=1 cm. B: X-ray photograph of the growth zone of the stalk (Cenocrinus asteria). The nodals (bright discs) are formed in short distances at the stalk-calyx border and become more distant from each other by intussusceptionally formed new columnals. Bar=2 mm. C: X-ray photograph (Metacrinus interuptus) of a maximally bowed stalk showing the central canal and, at the left side, the crenulated surface of the columnals. Bar=2 mm. D: X-ray photograph of a nodal (Endoxocrinus parrae). Arrowheads point to cirrus canals. Bar=1 mm. E: Cirri of Colobometra perspinosa. Bar=5 mm. F, G: Cirral articulation (Tropiometra afra); F, distal face; and G, proximal face of the cirrals together forming one joint. Bar=0.5 mm. H: X-ray photograph of a cirrus rising from the stalk of Endoxocrinus parrae showing the cirral articulations from the side. Bar=1 mm (from Heinzeller, Welsch, 1994). Строение рук Строение рук A: X-ray photograph of an arm of Clarkgomanthus littoralis in oral-aboral projection. Note the wedge-like shape of the brachials. Bar=1 mm. C: X-ray photographs. Position of syzygies (stars) in an arm of Comanthus suavia. Bar=2.5 mm. D: External site of a syzygy (arrowhead) in Himerometra robustipinnia, recognizable by the dotted line. Bar=1 mm. E: REM picture of a syzygial face of Tropiometra afra showing the typical crests. Bar=300 mkm. E (6): REM. Brachial ossicle of Tropiometra afra, showing a muscular-ligamentary articulation face with oblique fulcrum and, forming on its shoulder, a synarthrial joint (arrow) with the first pinnular ossicle. Bar=0.5 mm. al, aboral ligament; f, fulcrum; mf, mucsular facet; ol, oral ligament (from Heinzeller, Welsch, 1994). Тегмен, рот и анус, пищесобирающие борозды Ambularcra, mouth and anus. A: Oral aspect of an endocyclic specis (Antedon mediterranea). Ray A points to the left side; the mouth is situated in the disc’s center. Bar=2 mm. B: Food grooves on arm and pinnules of Capillaster multiradiatus. Bar=1 mm. C: Oral aspect of an exocyclis specis (Comatella nigra), ray A to the left. The anal tube occupies the disc’s center. Bar=2 mm. D: REM picture of tentacles lining a food groove (Lamprometera palmata). Bar=50 mkm (from Heinzeller, Welsch, 1994). Амбулакральные ножки иглокожих: a - схема продольного среза через присасывательную ножку Echinus. b - ловчая ножка криноидов (Antedon). Одна из папилл увеличена. c - копательная ножка морских звезд (Phanerozona). d - присасывательная ножка морских звезд (Spinulosa). e - ножка офиур. Показано ампулообразное расширение амбулакрального канала. f-i - амбулакральные ножки морских ежей. Вид со стороны присасывательного диска. Показана эволюция скелетных элементов ножек спатангоидов: f - Echinus, g - Brissopsis, h - Schizaster, I - Echinocardium. j - дыхательная ножка спатангоидов. k - роющая ножка спатангоидов. Показан скребок и слизепродуцирующие папиллы. l - питающая ножка спатангоидов. m - присасывательная ножка голотурий. (Nichols, 1969) Anatomy of Crinoidea (Nichols, 1969) Пищеварительная система Top: reconstruction of the gut as revealed from serial sections; left: endocyclic type (Antedon bifida); right: exocyclic type (Capillaster multradiatus). Bottom: Spiral lines representing gut axis running from mouth (m) to anus (a). Stars mark the points where the outer spiral turns being. A to E represent the direction of the rays (from Heinzeller, Welsch, 1994). Schematic drawing of the crinoid body plan as represented by a vertical section through ray E (left) and interray BC (right), oral side at the top, not to scale. A: Coelomic spaces clear; all other structures including the lumen of the gut stippled. ab, aboral canal; as, axial sinus; ci, cirral coelomic canals; co, chambered organ; cos, septum of the chambered organ; ge, genital coelom; ma, madreporic canals (possible axocoelomic remnant); oec, oral extension of the chamber; or, oral canal; ra, radial canal; ri, ring canal; se, subesophageal coelom; si, subintestinal coelom; ste, subtegminal coelom; sto, stone canal. Arrows indicate direction of coelomic fluid flow. The area marked by the star is subdivided by numerous somatocoelomic clefts (not drawn). B: Hemal spaces (black). al, lacuna in the axial organ; cl, cirral lacuna; gl, genital lacuna; pep, periesophageal plexus; rl, redial lacuna; sel, subesophagela lacuna; sil, subintestinal lacuna; spo, spongy organ; stp, subtegminal plexus. C: Principal skeletal (densely stippled) and neural (black) elements. Ossicles: ba, basale; ce, centrodorsale; ci, cirrale; pbr, primibrachiale; ra, radiale. Nervous system: anm, aboral nerve mass; bn, brachial nerve; ep, esophageal plexus; eri, ectoneural ring; hyr, hyponeural ring (no nerves drawn); ip, intestinal plexus; nr, nerve ring. Representing the neural plexuses, which are to be found in all coelomic epithelia, cp indicates the coelomic plexus in the periesophageal and peri-intestinal coelomic epithelia, gp that in the genital coelomic epithelium (from Heinzeller, Welsch, 1994). Саккулы Sacculi. A-C: Light microscopy. A: Antedon bifida. The sacculus contains numerous transformed cells with granular inclusions, stained according to Tramezzani et al., 1964. Bar=20 mkm. B: Lamprometra palmata. Bar=25 mkm. C: Antedon bifida. A sacculus empties its granule–filled cells (star) into the water. Bar=25 mkm. D: Low power electron micrograph, Antedon bifida. ep, flattened epithelium in the saccular periphery; g, granular inclusions in transformed cells; l, saccular lumen. Bar=2 mkm (from Heinzeller, Welsch, 1994). Location of gonads Location of gonads in adult crinoids. Each diagram shows one arm with its pinnules and an adjacent portion of the central mass. Gonads are solid black, and non-gametogenic parts of the reproductive system (genital strands) are dotted lines (the more distal parts of the latter, not shown here, continue in the arm axis and extend into the distal pinnules). The upper 5 diarams apply to stalkless crinoids: a) most species, b) some species, c) Thaumatocrinus investigatoris only, d) the genus Notocrinus, and e) some, but not all, species in the genera Comanthina and Comanthus. The lower 4 diagrams apply to stalked crinoids: f) the genera Bathycrinus, Rhizocrinus, and Hyocrinus, g) Metacrinus angulatus, g) Holopus rangii and Cyathidium foresti, and i) Endoxocrinus parrae. Abbreviations: CM, central mass; OP, oral pinnules; GP, genital pinnules; DP, distal pinnulis. (Holland, 2002) Development of Crinoidea (Nichols, 1969) The later development of Oxycomanthus japonicus Diagrams a-e are dorsal views of larvae with anterior ends pointed toward the top of the page; diagrams f-j are side views. Mesenchyme cells are indicated by stippling. Abbreviations: AP, adhesive pit; EH, enterohydrocoel; ES, enteric sac; HP, hydropore; L1, axocoel; L2, hydroceol; L3, left somatocoel; R3, Right somatocoel; RD, rectal diverticulum; SC, primary stone canal; VI, vestibular invagination; VS, vestibular sac. a) Relatively early uniformly ciliated larva; the archenteron is separating into an anterior enterohydrocoel and a posterior precursor of the somatocoels. b) Mid uniformly ciliated larva; the left somatocoel is separating from the right somatocoel. c) Late uniformly ciliated larva; the former enterohydrocoel is dividing into the axocoel (L1), hydrocoel (L2), and enteric sac. d) Early doliolaria larva with 5 ciliated bands encircling the body and an apical tuft; the axocoel has established communication with the exterior via the hydropore,
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