Tian S , Egertová M , Elphick MR .

	Figure 1. Anatomy of the starfish Asterias rubens. (A) Aboral view of a specimen of A. rubens; the white line indicates the approximate position of the section shown in (C). (B) Oral view of a specimen of A. rubens; the white line indicates the approximate position of the section shown in (D). (C) Trichrome-stained transverse section of an arm from a specimen of A. rubens. The body wall comprises ossicles (which appear white here) that are surrounded and interlinked by collagenous tissue (stained blue). The body wall has several types of appendages, which include spines and papulae. The body wall is lined internally by a coelomic epithelium, which is underlain along its midline aborally by a longitudinally orientated muscle known as the apical muscle. Linked to the aboral body wall by mesenteries are digestive organs known as pyloric caeca. The oral side of the body wall comprises ambulacral and adambulacral ossicles and two rows of tube feet, which are linked to bulb-shaped ampullae located internal to the body wall. The V-shaped radial nerve cord runs between the two rows of paired tube feet; lateral to the outer rows of tube feet are the marginal nerves. (D) Trichrome-stained transverse section of the central disk region, which is largely filled by the highly folded cardiac stomach. The cardiac stomach is linked orally to a short esophagus, which in this preparation is everted through the oral opening (mouth) that is surrounded by a peristomial membrane. Lateral to the peristomial membrane is the cirumoral nerve ring. Aboral to the cardiac stomach is the smaller pyloric stomach, which is linked via pyloric ducts to the pyloric caeca located in the arms [see (C)]. Aboral to the pyloric stomach is a short rectum (not seen here), which has associated rectal caeca. Abbreviations: ado, adambulacral ossicle; am, apical muscle; ao, ambulacral ossicle; ce, coelomic epithelium; co, carinal ossicle; conr, circumoral nerve ring; cs, cardiac stomach; es, esophagus; lsm, longitudinal supra-ambulacral muscle; me, mesentery; mn, marginal nerve; mo, marginal ossicle; pa, papula; pc, pyloric caeca; pd, pyloric duct; pm, peristomial membrane; ps, pyloric stomach; rc, rectal caeca; ro, reticular ossicle; sp, spine; tf, tube foot; tsm, transverse supra-ambulacral muscles. Scale bars: (A,B) = 1 cm; (C,D) = 210 µm.
	Figure 2. Localization of Asterias rubens gonadotropin-releasing hormone precursor (ArGnRHP) mRNA in the nervous system of A. rubens using in situ hybridization. (A,B) Transverse section of a radial nerve cord incubated with antisense probes, showing stained cells (arrowheads) in both the hyponeural and ectoneural regions. The inset of (A) shows absence of staining in a transverse section of a radial nerve cord incubated with sense probes, demonstrating the specificity of staining observed with antisense probes. A higher magnification image of the boxed area is shown in (B). (C) Longitudinal parasagittal section of a radial nerve cord showing stained cells (arrowheads) in both the hyponeural and ectoneural regions. A higher magnification image of the boxed area is shown in (D). (E) Transverse section of the central disk region showing stained cells (arrowheads) in both the hyponeural and the ectoneural regions of the circumoral nerve ring; note, however, that during tissue processing the hyponeural region has been displaced from its natural position adjacent to the ectoneural region. (F) Stained cells (arrowheads) in the marginal nerve, which is located lateral to the outer row of tube feet on each side of the arms. Abbreviations: ec, ectoneural region; hy, hyponeural region; mn, marginal nerve; tf, tube foot. Scale bars: [(A), inset, (C,E)] = 50 µm; (B) = 10 µm; (D) = 20 µm; (F) = 30 µm.
	Figure 3. Localization of Asterias rubens gonadotropin-releasing hormone precursor (ArGnRHP) mRNA in the tube feet, terminal tentacle and digestive system of A. rubens using in situ hybridization. (A) Longitudinal section of a tube foot showing stained cells (arrowheads) in the subepithelial layer of the tube foot stem. (B) Stained cells (arrowheads) located in the subepithelial layer above the tube foot sucker and near the basal nerve ring. (C) Stained cell (arrowhead) located in the optic cushion, which is located at the base of the terminal tentacle. (D) Stained cells (arrowheads) located in a lateral lappet and the body wall epithelium surrounding the terminal tentacle. (E) Stained cell (white arrowhead) in the mucosal layer of the cardiac stomach. (F) Stained cell (arrowhead) located close to the basiepithelial nerve plexus of the cardiac stomach. (G) Stained cell (arrowhead) located close to the basiepithelial nerve plexus of the pyloric stomach. (H) Stained cells (arrowheads) located in the mesenteries associated with the pyloric caeca. Abbreviations: bnr, basal nerve ring; ct, collagenous tissue; ep, epidermis; ll, lateral lappet; lu, lumen; m, muscle; me, mesentery; mu, mucosa; pc, pyloric cecum; vml, visceral muscle layer. scale bars: (A,B,D,H) = 30 µm; (C,E,F,G) = 20 µm.
	Figure 4. Localization of Asterias rubens corazonin precursor (ArCRZP) mRNA in the nervous system of A. rubens using in situ hybridization. (A,B) Transverse section of a radial nerve cord incubated with antisense probes, showing stained cells (arrowheads) in the ectoneural region but not in the hyponeural region. The inset of (A) shows absence of staining in a transverse section of a radial nerve cord incubated with sense probes, demonstrating the specificity of staining observed with antisense probes. A higher magnification image of the boxed area is shown in (B). (C) Longitudinal parasagittal section of a radial nerve cord showing stained cells (arrowheads) in the ectoneural region but not in the hyponeural region. (D) Transverse section of the central disk region showing stained cells (arrowheads) in the ectoneural region of the circumoral nerve ring but not in the hyponeural region, which has been displaced from its natural position adjacent to the ectoneural region during tissue processing. (E) High magnification image of the stained cells (arrowheads) in the ectoneural region of the circumoral nerve ring. Abbreviations: ct, collagenous tissue; ec, ectoneural region; hy, hyponeural region; pm, peristomial membrane; rhs, radial hemal strand. Scale bars: [(A) inset (C)] = 60 µm; (B,E) = 20 µM; (D) = 120 µm.
	Figure 5. Localization of Asterias rubens corazonin precursor (ArCRZP) mRNA in the marginal nerve, tube foot and body wall-associated structures of A. rubens using in situ hybridization. (A) Stained cells in the marginal nerve. (B) Stained cells (arrowheads) located within or beneath the external epithelium layer at the junction between adjacent tube feet. (C) Stained cells (arrowheads) located near to the tube foot sucker. (D) Stained cells (arrowheads) located in the external epidermis on the oral side of the body wall. (E) Transverse section of the apical muscle showing stained cells (arrowheads) in the coelomic epithelium and close to the circular muscle layer. The boxed region of (E) is shown at higher magnification in the inset. Abbreviations: ae, adhesive epidermis of tube foot sucker; bw, body wall; ct, collagenous tissue; lu, lumen; mn, marginal nerve; tf, tube foot. Scale bars: (A,E,D) = 50 µm; (B) = 100 µm; (C) = 60 µm; [(D,E) insets] = 20 µm.
	Figure 6. Localization of Asterias rubens corazonin precursor (ArCRZP) mRNA in the digestive system of A. rubens using in situ hybridization. (A) Transverse section through the central disk region showing stained cells (arrowheads) in the cardiac stomach (dashed line) and pyloric stomach (solid line). Stained cells are more abundant in the pyloric stomach and in the aboral (uppermost) region of the cardiac stomach than in the oral (lowermost) region of the cardiac stomach. (B) High magnification image showing stained cells in the cardiac stomach located both in the mucosal layer (white arrowhead) and close to the basiepithelial nerve plexus (black arrowheads). (C) High magnification image showing stained cells in the pyloric stomach located both in the mucosal layer (white arrowhead) and close to the basiepithelial nerve plexus (black arrowheads). (D) Transverse section of a pyloric duct showing stained cells in the oral (lowermost) region. The boxed area is shown at higher magnification in (E). (E) High magnification image of a pyloric duct showing stained cells in the mucosal layer (white arrowheads). (F) Stained cells (arrowhead) located close to the basiepithelial nerve plexus of a pyloric duct. Abbreviations: cs, cardiac stomach; ct, collagenous tissue; lu, lumen; me, mesentery; mu, mucosa; ps, pyloric stomach; vml, visceral muscle layer. Scale bars: (A) = 100 µm; (B,C,E,F) = 20 µm; (D) = 60 µm.
	Figure 7. Localization of Asterias rubens gonadotropin-releasing hormone (ArGnRH) immunoreactivity in the nervous system of A. rubens. (A) Longitudinal section of a radial nerve cord showing immunostaining in both the hyponeural and ectoneural regions. Clusters of stained cells (arrowheads) can be seen in the hyponeural region. Immunostained fibers (asterisk) can be seen in neuropile of the ectoneural region. (B) Horizontal section through the radial nerve cord and circumoral nerve ring of a juvenile specimen. Immunostained cell bodies (arrow heads) can be seen in the epithelial layer of the ectoneural region. The neuropile of the radial nerve cord and circumoral nerve ring contains a meshwork of immunostained fibers but the density of the stained fibers varies, with a higher density in some areas (white asterisk) and a lower density in other areas (black asterisk). (C) Horizontal section of a radial nerve cord [aboral to the section shown in (B)] showing the segmental structure of the radial nerve cord. Immunostained fibers can be seen in the ectoneural region (asterisk) and immunostained cells (arrowheads) can be seen in the hyponeural region. The boxed region is shown at higher magnification in the inset. (D) Longitudinal section of the marginal nerve showing immunostained cells (arrowhead) and processes (asterisk). Abbreviations: bw, body wall; conr, circumoral nerve ring; ec, ectoneural region; hy, hyponeural region; mn, marginal nerve; rnc, radial nerve cord; rhs, radial hemal strand; tf, tube foot. Scale bars: (A–D) = 60 µm; [(C) inset] = 20 µm.
	Figure 8. Localization of Asterias rubens gonadotropin-releasing hormone (ArGnRH) immunoreactivity in the radial nerve cords and marginal nerves of A. rubens. (A) Transverse section of a radial nerve cord showing stained cells (arrowheads) in both the hyponeural and ectoneural regions. The intensity of immunostaining in the neuropile of the ectoneural region varies, with the region at the apex of the V-shaped nerve cord intensely stained (asterisk) and the lateral regions less intensely stained. The inset shows absence of immunostaining in a radial nerve cord section incubated with affinity-purified ArGnRH-antibodies preabsorbed with the antigen peptide (ArGnRH-ag), demonstrating the specificity of immunostaining observed with the ArGnRH antibody. (B) High magnification image of a radial nerve cord showing immunostained cell bodies in the hyponeural and ectoneural regions. Note that in the neuropile of the ectoneural region there is a band of immunostaining (white asterisk), which is located opposite immunostained hyponeural cell bodies. The boxed area is shown at higher magnification in (C). (C) Immunostained processes (white arrow) derived from monopolar cell bodies (arrowheads) can be seen here in the hyponeural region. Stained axonal profiles (black arrows) can be seen at the margin of the intensely stained region of the ectoneural neuropile (asterisk). The layer of collagenous tissue that separates the hyponeural and ectoneural regions is unstained. (D) High magnification image of the ectoneural region of a radial nerve cord showing immunostained bipolar-shaped cells in the subcuticular epithelium (arrowhead). Immunostained processes extending into the ectoneural neuropile and toward the cuticular layer (white arrows) can also be seen here. (E) High magnification image showing immunostaining in the lateral region of a radial nerve cord, with staining in the ectoneural neuropile (asterisk) and in the processes of hyponeural neurons that project from the radial nerve cord around the lateral wall of the radial perihemal canal (white arrows). (F) Immunostaining in the marginal nerve. The boxed area is shown at higher magnification in the inset, where a stained bipolar-shaped cell can be seen in the epithelial layer (arrowhead) and stained processes can be seen in the underlying neuropile. Abbreviations: amus, ambulacral muscle; ct, collagenous tissue; cut, cuticular layer; ec, ectoneural region; hy, hyponeural region; rhs, radial hemal strand; tf, tube foot. Scale bars: (A,F) = 50 µm; [(A) inset] = 120 µm; [(B–F) inset] = 20 µm.
	Figure 9. Localization of Asterias rubens gonadotropin-releasing hormone (ArGnRH) immunoreactivity in the circumoral nerve ring of A. rubens. (A) Immunostaining in a transverse section of a circumoral nerve ring, with immunostained processes projecting from the circumoral nerve ring into the peristomial membrane and the adjacent body wall (white arrows). A cluster of immunostained cells can be seen in the ectoneural epithelium of the medial region of the nerve ring (arrowheads) and an intensely stained region of ectoneural neuropile (asterisk) can be seen at the apex of the nerve ring. (B) High magnification image of a circumoral nerve ring showing immunostained cells in the hyponeural and ectoneural regions (arrowheads) with stained processes (white arrows). Immunostained processes of ectoneural cells project into the underlying neuropile (white asterisk). (C) High magnification image of the ectoneural region showing immunostained bipolar-shaped cells in the subcuticular epithelium (arrowheads) with immunostained processes (white arrows) extending toward the cuticular layer and the intensely stained neuropile (white asterisk). (D) High magnification image of the hyponeural region showing immunostained monopolar-shaped cells (arrowhead) with immunostained processes (white arrows) adjacent to the unstained collagenous tissue layer. Abbreviations: bw, body wall; ct, collagenous tissue; cut, cuticular layer; ec, ectoneural region; hr, hemal ring; hy, hyponeural region; pm, peristomial membrane. Scale bars: (A) = 50 µm; (B) = 20 µm; (C,D) = 10 µm.
	Figure 10. Localization of Asterias rubens gonadotropin-releasing hormone (ArGnRH) immunoreactivity in the tube feet and terminal tentacle of A. rubens. (A) Longitudinal section of a tube foot showing immunostaining in the subepithelial nerve plexus (white arrows) and basal nerve ring (black arrows). (B) Transverse section of a tube foot showing immunostained processes in the subepithelial nerve plexus (white arrow) and in the basal nerve ring (black arrow). (C) Immunostained cells (arrowheads) and processes (white arrows) at the junction between a tube foot and the radial nerve cord. (D) Immunostaining at the junction between adjacent tube feet. (E) Immunostained cells (arrowheads) and processes in the optic cushion, which is located at the base of the terminal tentacle. Immunostained processes (white arrows) can also be seen here in the basiepithelial plexus of the body wall epithelium surrounding the terminal tentacle. Abbreviations: bnr, basal nerve ring; ct, collagenous tissue; ec, ectoneural region; lu, lumem; m, muscle; oc, optic cushion; rnc, radial nerve cord; tf, tube foot; tt, terminal tentacle. Scale bars: (A) = 50 µm; (B,D,E) = 60 µm; (C) = 30 µm.
	Figure 11. Localization of Asterias rubens gonadotropin-releasing hormone (ArGnRH) immunoreactivity in the digestive system of A. rubens. (A) Horizontal section through the central disk region near to the mouth, showing immunostaining in the peristomial membrane and esophagus. (B) Horizontal section through the central disk region showing immunostaining in the pentaradially symmetrical (asterisks) oral region of the cardiac stomach. (C) Horizontal section through the central disk showing immunostaining in the folded pouches of the cardiac stomach. (D) High magnification image of the cardiac stomach showing immunostained bipolar-shaped cells (white arrowheads) in the mucosal layer. (E) High magnification image of the cardiac stomach showing immunostaining in the basiepithelial nerve plexus (white arrow) and the visceral nerve plexus (black arrow) that is closely associated with the visceral muscle layer. (F) High magnification image showing immunostaining in a thickening of the basiepithelial nerve plexus that underlies a cardiac stomach intrinsic retractor muscle. (G) Horizontal section through the central disk showing immunostaining (white arrows) in the pyloric stomach and the pyloric ducts. (H) High magnification image showing immunostaining in a pyloric duct (white arrow) and in the pyloric stomach (black arrow). (I) High magnification image showing immunostaining in the basiepithelial nerve plexus of the mucosal layer and in a few processes associated with the visceral muscle layer. (J) High magnification image showing immunostaining in the basiepithelial nerve plexus (white arrow) and visceral nerve plexus (black arrow) on the oral side of a pyloric duct. (K) High magnification image showing immunostaining (white arrows) in a mesentery. Abbreviations: ct, collagenous tissue; es, esophagus; irm, instrinsic retractor muscle; lu, lumen; mu, mucosa, pc, pyloric caeca; pd, pyloric duct; pm, peristomial membrane; ps, pyloric stomach; vml, visceral muscle layer. Scale bars: (A,B) = 60 µm; (C,G) = 120 µm; (D,E,I,J) = 20 µm; (F,H) = 50 µm; (K) = 30 µm.
	Figure 12. Localization of Asterias rubens gonadotropin-releasing hormone (ArGnRH) immunoreactivity in body wall-associated structures in A. rubens. (A) Immunostaining in a transverse section of the apical muscle. (B) Immunostained processes (white arrows) can be seen here in the basiepithelial plexus underlying the coelomic epithelium of the body wall and in the apical muscle. (C) Immunostained processes (white arrows) in the basiepithelial nerve plexus beneath the external epithelium of the body wall. The inset shows a higher magnification image of the boxed region. Abbreviations: am, apical muscle; bw, body wall; ce, coelomic epithelium; ct, collagenous tissue; sp, spine. Scale bars: (A,B) = 80 µm; (C) = 60 µm; [(C) inset] = 20 µm.
	Figure 13. Comparison of the pharmacological effects of Asterias rubens gonadotropin-releasing hormone (ArGnRH) and A. rubens corazonin (ArCRZ) on cardiac stomach, apical muscle, and tube foot preparations from the starfish A. rubens. (A) Representative recordings from a single cardiac stomach preparation showing the contraction-inducing effects of ArGnRH (red line) and ArCRZ (purple line) at 10−6 M. Peptide application is labeled with an upward pointing arrowhead, washing of the preparation is labeled with a downward pointing arrowhead. (B) Graphs showing the dose-dependent effects of ArGnRH (red) and ArCRZ (purple) on cardiac stomach, expressed as mean percentages (±SEM) of the contraction induced by 10−7 M NGFFYamide. Data were collected from at least five independent experiments. The effect of ArGnRH was significantly larger than the effect of ArCRZ when tested at 10−7 and 10−6 M. (C) Representative recordings from a single apical muscle preparation showing the contraction-inducing effects of ArGnRH (red line) and ArCRZ (purple line) at 10−6 M. Peptide application is labeled with an upward pointing arrowhead, washing of the preparation is labeled with a downward pointing arrowhead. (D) Graphs showing the dose-dependent effects of ArGnRH (red) and ArCRZ (purple) on apical muscle preparations, expressed as mean percentages (±SEM) of the contraction induced by 10−6 M acetylcholine. Data were collected from at least five independent experiments. The effect of ArCRZ was significantly larger than the effect of ArGnRH when tested at 10−9, 10−8, 10−7, and 10−6 M. (E) Representative recordings from a single tube foot preparation showing the contraction-inducing effects of ArGnRH (red line) and ArCRZ (purple line) at 10−6 M. Peptide application is labeled with an upward pointing arrowhead, washing of the preparation is labeled with a downward pointing arrowhead. (F) Comparison of the effects of ArGnRH and ArCRZ on tube foot preparations, expressed as mean percentages (±SEM) of the contraction induced by 10−6 M acetylcholine. The data were collected from at least three independent experiments. Student’s t-tests were performed to determine whether there were significant differences in the effects of ArGnRH and ArCRZ when tested at the same concentration (***p < 0.001, **p < 0.01; *p < 0.05).

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