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Psolus fabricii Skeletal structures In all echinoderms the epidermis ...

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<strong>Skeletal</strong> <strong>structures</strong><br />

<strong>In</strong> <strong>all</strong> <strong>echinoderms</strong> <strong>the</strong> <strong>epidermis</strong> covers <strong>the</strong> body and contains ossicles<br />

<strong>In</strong> sea urchins and sand dollars, ossicles are attached to each o<strong>the</strong>r and form a rigid test.<br />

Muscles are weakly developed.<br />

<strong>In</strong> sea cucumbers, ossicles are scattered in <strong>the</strong> fleshy <strong>epidermis</strong>. Layered muscles are<br />

developed.<br />

<strong>In</strong> sea stars and ophiurods,<br />

skeletal plates articulate and<br />

<strong>all</strong>ow arm motion.<br />

<strong>Psolus</strong> <strong>fabricii</strong><br />

B-Spines of <strong>the</strong> sand dollar<br />

Echinarachnius parma<br />

C-<strong>Skeletal</strong> ossicles from central discs<br />

of 4 ophiurid spp.<br />

D-Ossicles from holothurian <strong>Psolus</strong><br />

chintinoides


<strong>Skeletal</strong> <strong>structures</strong><br />

<strong>Skeletal</strong> plates remain single<br />

or fuse. They give rise to<br />

spines, tubercles, and<br />

pincer-like pedicellariae<br />

(starfish, urchins)<br />

Pedicellariae respond<br />

external stimuli.<br />

<strong>In</strong>dependent from nervous<br />

system. Function: defense,<br />

cleaning, predation<br />

Stylasterias forreri<br />

E-Kinds of echinoid pedicellariae surrounding a spine<br />

F,G-Pedicellariae used for prey capture in starfish Stylasterias forreri<br />

H-Generalized pedicellariae<br />

I-2 types of muscle systems in pedicellariae<br />

J-A movable spine


Support and Locomotion<br />

General body shape maintained by skeletal elements except in sea cucumbers (hydrostatic skeleton)<br />

Crinoids can be sessile (sea lilies) or crawl and swim. They move with oral side away from<br />

substrate and use cirri for temporary attachment or balance<br />

Ophiurids use flexible articulated arms for crawling and clinging. Tube feet no suckers and ampullae<br />

Asteroids use podia. No metachronal waves. Some excrete mucus for defense<br />

Luidia ciliaris<br />

Urchins use both podia and spines.<br />

Holothurians crawl by podia or by action of body w<strong>all</strong> muscles<br />

Crinoid Antenon<br />

A,B- Resting, moving<br />

C-Starfish with tube feet in<br />

motion<br />

D-Changes in position of<br />

individual podia<br />

Amphiura


Feeding And Digestion<br />

Digestion is both extracellular and<br />

intracellular<br />

ASTEROIDS<br />

1. Ecology: Mostly carnivores,<br />

many feed on bivalves, barnacles,<br />

o<strong>the</strong>r starfish.<br />

Tube feet are used to open <strong>the</strong><br />

valves<br />

2. Anatomy<br />

Short gut extends from oral mouth<br />

to aboral anus. Mouth in <strong>the</strong><br />

center of <strong>the</strong> peristomial<br />

membrane.<br />

Pyloric ceca (digestive glands) of <strong>the</strong> pyloric<br />

stomach. The gut lined with ciliated epi<strong>the</strong>lium<br />

3. Digestion: Cardiac stomach is everted to engulf prey<br />

outside <strong>the</strong> mouth. It can be slipped through <strong>the</strong> gape<br />

between clam valves. Digestion begins extern<strong>all</strong>y<br />

Extracellular digestion occurs in <strong>the</strong> stomach. Absorption, phagocytosis, and intracellular digestion<br />

occur in <strong>the</strong> pyloric ceca<br />

4. DOM<br />

30% or food needs met by DOM absorption


1. Ecology: Most are carnivores, deposit feeders, or suspension feeders<br />

Some capture (lasso) sm<strong>all</strong> invertebrates with <strong>the</strong>ir arms<br />

Many use tube feet to move suspended or deposited particles (food bolus) to <strong>the</strong> mouth<br />

2. Anatomy: Gut is a blind<br />

pouch with mouth on oral<br />

surface, with no rectum<br />

or anus<br />

Digestion and absorption<br />

occur in <strong>the</strong> stomach<br />

Tentacular scale near<br />

basis of each podium.<br />

Trap plankton<br />

Amphiura brachiata<br />

OPHIUROIDS


1. Ecology<br />

Regular urchins are grazers<br />

Irregular urchins are deposit<br />

feeders<br />

Aristotle's lantern used to<br />

scrape algae, etc from rocks<br />

It’s a complex masticatory<br />

apparatus, just inside <strong>the</strong> mouth<br />

with 5 protractible teeth<br />

Made of 5 triangular plates<br />

(pyramids), moved by<br />

comminator muscles<br />

Tube feet are used to move<br />

organic particles to <strong>the</strong> mouth<br />

2. Anatomy<br />

Mouth at center of oral surface<br />

A long intestine extends to anus<br />

Siphon bypasses <strong>the</strong> stomach<br />

ECHINOIDS


HOLOTHUROIDEA<br />

1. Ecology<br />

Most are suspension or deposit feeders<br />

Direct deposit feeders ingest sediment<br />

Sediment is pushed into mouth by tentacles<br />

Suspension feeders and selective deposit feeders capture individual<br />

organic particles with adhesive papillae on tentacles<br />

Tentacles are stuffed into <strong>the</strong> mouth one by one.<br />

Evisceration: Partial or complete expulsion of digestive tract, gonads<br />

Chemical or physical stress, defense mechanism?<br />

Discharge of Cuverian tubules: defensive action, sea cucumber<br />

points <strong>the</strong> anus at predator, and discharges <strong>the</strong> tubules by rupturing<br />

<strong>the</strong> hindgut. All tissues can be regenerated.<br />

2. Anatomy<br />

Body is elongate on oral-aboral<br />

axis<br />

Mouth at oral end, anus at aboral<br />

Mouth surrounded by 10-30<br />

dendritic tentacles<br />

Tentacles are modified tube feet<br />

Respiratory trees-internal<br />

<strong>structures</strong> where water is<br />

pumped via anus for gas<br />

exchange


Cuvierian tubules released<br />

Neothyonidium magnum<br />

C-Parastichopus feeding-deposit feeder<br />

D-Cucumaria showing feeding tentacles<br />

E-Mucus-secreting papillae on tentacles of Aslia lefevrei<br />

F-Cucumaria minuta<br />

G-Psolidium suspension feeder


1. Ecology: Suspensions feeders similar to ancestral <strong>echinoderms</strong><br />

2. Anatomy: Feed with oral surface up, unlike most<br />

Echinoderms. Arms surround <strong>the</strong> mouth<br />

Arms and ambulacra are branched<br />

Mouth at center of oral surface, anus is also oral,<br />

but displaced to periphery, gut is U-shaped.<br />

3. Feeding<br />

Food particles are<br />

captured by podia<br />

Particles are transferred<br />

to <strong>the</strong> mouth by ciliary<br />

currents<br />

The original function<br />

of tube feet was for<br />

feeding, not locomotion<br />

CRINOIDS<br />

A-Central disc and base of one arm<br />

B-An arm with open ambulacral (food) groove<br />

C-Oral surface of Antedon. Note alphabetical<br />

position of ambulacral radii


Asexual and sexual reproduction (by far <strong>the</strong> most common)<br />

A. ASEXUAL<br />

Most <strong>echinoderms</strong> are capable of regeneration of lost parts. Cut-off arms soon die,<br />

however Linckia can regenerate from single arms<br />

Brittle stars and crinoids cast off whole arms<br />

(autotomy) when disturbed<br />

Reproduction<br />

<strong>In</strong> asexual reproduction, disk divides (fissiparity)<br />

Best developed in brittle stars and starfish<br />

Arm with 20% of disk can produce<br />

a new star<br />

B. SEXUAL<br />

Gonochoric, few are hermaphrodites<br />

External fertilization. Spawning usu<strong>all</strong>y nocturnal.<br />

Both sexes release gametes into <strong>the</strong> sea<br />

A few species brood; female retains eggs and embryos<br />

Some brittle stars brood in <strong>the</strong> genital bursae<br />

Linckia guildingi


Sexual Reproduction<br />

Gonads are housed in genital sinuses. Sea Cucumbers with 1 gonad.<br />

Crinoids with no distinct gonads. Gametes arise from <strong>the</strong> peritoneum of genital canals.<br />

No gonoducts. Gametes are released by rupturing <strong>the</strong> pinnule w<strong>all</strong>s.<br />

Ophiurids with 1 to many gonads attached next to bursal slits. Gametes are released into<br />

bursae and released though slits<br />

Starfish, urchins with<br />

multiple gonads<br />

Gonoducts lead to<br />

interambulacral<br />

gonopores<br />

Regular sea urchins<br />

with 5 gonads (located<br />

on 5 genital plates<br />

Brooding in <strong>all</strong> groups<br />

in polar and boreal<br />

species.


Development<br />

A. GENERAL<br />

Classical deuterostome development<br />

Radial, regulative (indeterminate) cleavage<br />

Coeloblastula<br />

Gastrulation by invagination<br />

Enterocoelous coelom and mesoderm formation<br />

Tricoelomic<br />

Blastopore becomes <strong>the</strong> anus<br />

B. LARVAE<br />

Dipleurula is <strong>the</strong> first larva (a gastrula)<br />

Becomes a specialized larva for each class<br />

Larvae are plesiomorphic<strong>all</strong>y bilater<strong>all</strong>y<br />

symmetrical, have a recurved gut and transparent<br />

ectoderm, and feed by upstream particle capture<br />

using <strong>the</strong> ciliated band (planktotrophic).<br />

Metamorphosis is typic<strong>all</strong>y radical and occurs<br />

during settlement onto <strong>the</strong> benthos.<br />

Bilateral larvaradial adult<br />

A-Vitellaria (crinoid)<br />

B,C-Bipinnaria and later brachiolaria of starfish<br />

D-Ophiopluteus of an ophiurid<br />

E-Auricularia of sea cucumber


Phylogeny<br />

Current hypo<strong>the</strong>sis: lineage leading to <strong>the</strong> Crinoidea branched most bas<strong>all</strong>y,<br />

Echinoidea and Holothuroidea are sister taxa.<br />

Current debate: Are Ophiuroidea and Asteroidea sister taxa? The two best-supported<br />

hypo<strong>the</strong>ses are:<br />

"Asterozoan" hypo<strong>the</strong>sis proposed by Ba<strong>the</strong>r (1900), and supported by Mooi and David<br />

(1997). Mitochondrial gene order supports this hypo<strong>the</strong>sis (Smith et al. 1993), but crinoid<br />

info is needed to clarify <strong>the</strong> polarity of <strong>the</strong> character transformations.<br />

“Cryptosyringid” hypo<strong>the</strong>sis proposed by MacBride (1906). Smith 1988 advocated <strong>the</strong><br />

same topology.<br />

Both hypo<strong>the</strong>ses are supported equ<strong>all</strong>y, by 18S and 26S rRNA analyses (summarized in<br />

Littlewood et al. 1997).

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