{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,9,27]],"date-time":"2026-09-27T16:57:01Z","timestamp":1790528221915,"version":"4.1.0"},"reference-count":66,"publisher":"The Company of Biologists","issue":"7","content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2010,4,1]]},"abstract":"<jats:title>SUMMARY<\/jats:title><jats:p>Sea star tube feet consist of an enlarged and flattened distal extremity (the disc), which makes contact with the substratum, and a proximal contractile cylinder (the stem), which acts as a tether. In this study, the different forces brought into play during tube foot functioning were investigated in two related species. The tube feet of Asterias rubens and Marthasterias glacialis attach to glass with a similar mean tenacity (0.24 and 0.43 MPa, respectively), corresponding to an estimated maximal attachment force of 0.15 and 0.35 N. The contraction force of their retractor muscle averages 0.017 N. The variation of the retractor muscle contraction with its extension ratio follows a typical bell-shaped length\u2013tension curve in which a maximal contraction of approximately 0.04 N is obtained for an extension ratio of approximately 2.3 in both sea star species. The tensile strength of the tube foot stem was investigated considering the two tissues that could assume a load-bearing function, i.e. the retractor muscle and the connective tissue. The latter is a mutable collagenous tissue presenting a fivefold difference in tensile strength between its soft and stiff state. In our experiments, stiffening was induced by disrupting cell membranes or by modifying the ionic composition of the bathing solution. Finally, the force needed to break the tube foot retractor muscle was found to account for 18\u201325% of the tube foot total breaking force, showing that, although the connective tissue is the tissue layer that supports most of the load exerted on the stem, the contribution of the retractor muscle cannot be neglected in sea stars. All these forces appear well-balanced for proper functioning of the tube feet during the activities of the sea star. They are discussed in the context of two essential activities: the opening of bivalve shells and the maintenance of position in exposed habitats.<\/jats:p>","DOI":"10.1242\/jeb.037903","type":"journal-article","created":{"date-parts":[[2010,3,12]],"date-time":"2010-03-12T21:59:06Z","timestamp":1268431146000},"page":"1162-1174","source":"Crossref","is-referenced-by-count":34,"title":["Evaluation of the different forces brought into play during tube foot activities in sea stars"],"prefix":"10.1242","volume":"213","author":[{"given":"Elise","family":"Hennebert","sequence":"first","affiliation":[{"name":"Universit\u00e9 de Mons \u2013 UMONS, Acad\u00e9mie Universitaire Wallonie-Bruxelles, Laboratoire de Biologie Marine, Mons, Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Delphine","family":"Haesaerts","sequence":"additional","affiliation":[{"name":"Universit\u00e9 Libre de Bruxelles, Acad\u00e9mie Universitaire Wallonie-Bruxelles, Laboratoire de Biologie Marine, Bruxelles, Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Philippe","family":"Dubois","sequence":"additional","affiliation":[{"name":"Universit\u00e9 Libre de Bruxelles, Acad\u00e9mie Universitaire Wallonie-Bruxelles, Laboratoire de Biologie Marine, Bruxelles, Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Patrick","family":"Flammang","sequence":"additional","affiliation":[{"name":"Universit\u00e9 de Mons \u2013 UMONS, Acad\u00e9mie Universitaire Wallonie-Bruxelles, Laboratoire de Biologie Marine, Mons, Belgium"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","reference":[{"key":"2021042506080621000_R1","volume-title":"Molecular Biology of the Cell","author":"Alberts","year":"2002","edition":"4th edn"},{"key":"2021042506080621000_R2","first-page":"230","article-title":"Cellular physiology of skeletal, cardiac and smooth muscle","volume-title":"Medical physiology","author":"Apkon","year":"2003"},{"key":"2021042506080621000_R3","first-page":"99","article-title":"Influence of salinity on the ability of starfishes Asterias rubens L. to attach to substrate","volume":"22","author":"Berger","year":"1996","journal-title":"Biologiya Morya"},{"key":"2021042506080621000_R4","doi-asserted-by":"crossref","first-page":"15","DOI":"10.1016\/0300-9629(85)90261-0","article-title":"Some properties of the action potentials conducted in the spines of the sea urchin Diadema antillarum","volume":"81A","author":"Berrios","year":"1985","journal-title":"Comp. Biochem. Physiol."},{"key":"2021042506080621000_R5","doi-asserted-by":"crossref","first-page":"69","DOI":"10.1242\/jeb.117.1.69","article-title":"The mechanical properties of the autotomy tissues of the holothurian Eupentacta quinquesemita and the effects of certain physico-chemical agents","volume":"117","author":"Byrne","year":"1985","journal-title":"J. Exp. Biol."},{"key":"2021042506080621000_R6","first-page":"135","article-title":"Mechanical design of mussel byssus: load cycle and strain rate dependence","volume":"18","author":"Carrington","year":"2004","journal-title":"Am. Malacol. Bull."},{"key":"2021042506080621000_R7","doi-asserted-by":"crossref","first-page":"180","DOI":"10.1002\/lno.1957.2.3.0180","article-title":"The feeding behaviour of the sea star Evasterias troscheli","volume":"2","author":"Christensen","year":"1957","journal-title":"Limnol. Oceanogr."},{"key":"2021042506080621000_R8","doi-asserted-by":"crossref","first-page":"483","DOI":"10.1007\/978-1-4613-1955-9_17","article-title":"Neurobiology of the Echinodermata","volume-title":"Nervous Systems in Invertebrates","author":"Cobb","year":"1987"},{"key":"2021042506080621000_R9","doi-asserted-by":"crossref","first-page":"955","DOI":"10.4319\/lo.1998.43.5.0955","article-title":"The menace of momentum: dynamic forces on flexible organisms","volume":"43","author":"Denny","year":"1998","journal-title":"Limnol. Oceanogr."},{"key":"2021042506080621000_R10","doi-asserted-by":"crossref","first-page":"353","DOI":"10.1002\/jmor.1051490305","article-title":"Aspects of skeletal mechanics of the starfish Asterias forbesii","volume":"149","author":"Eylers","year":"1976","journal-title":"J. Morphol."},{"key":"2021042506080621000_R11","doi-asserted-by":"crossref","first-page":"764","DOI":"10.2307\/1931318","article-title":"On the methods used by the starfish Pisaster ochraceus in opening three types of bivalve molluscs","volume":"36","author":"Feder","year":"1955","journal-title":"Ecology"},{"key":"2021042506080621000_R12","first-page":"1","article-title":"Adhesion in echinoderms","volume-title":"Echinoderm Studies","author":"Flammang","year":"1996"},{"key":"2021042506080621000_R13","doi-asserted-by":"crossref","first-page":"1251","DOI":"10.1017\/S0025315400038807","article-title":"Measurement of the adhesion of the podia in the asteroid Asterias rubens (Echinodermata)","volume":"77","author":"Flammang","year":"1997","journal-title":"J. Mar. Biol. Assoc. UK"},{"key":"2021042506080621000_R14","doi-asserted-by":"crossref","first-page":"35","DOI":"10.2307\/1542163","article-title":"The role of podial secretions in adhesion in two species of sea stars (Echinodermata)","volume":"187","author":"Flammang","year":"1994","journal-title":"Biol. Bull."},{"key":"2021042506080621000_R15","doi-asserted-by":"crossref","first-page":"2383","DOI":"10.1242\/jeb.201.16.2383","article-title":"A study of the temporary adhesion of the podia in the sea star Asterias rubens (Echinodermata, Asteroidea) through their footprints","volume":"201","author":"Flammang","year":"1998","journal-title":"J. Exp. Biol."},{"key":"2021042506080621000_R16","doi-asserted-by":"crossref","first-page":"195","DOI":"10.1007\/BF00221081","article-title":"Ultrastructure of sea urchin tube feet. Evidence for connective tissue involvement in motor control","volume":"177","author":"Florey","year":"1977","journal-title":"Cell Tissue Res."},{"key":"2021042506080621000_R17","volume-title":"Techniques Histologiques","author":"Gabe","year":"1968"},{"key":"2021042506080621000_R18","doi-asserted-by":"crossref","first-page":"121","DOI":"10.1098\/rstb.2001.1022","article-title":"Elastic proteins: biological roles and mechanical properties","volume":"357","author":"Gosline","year":"2002","journal-title":"Phil. Trans. R. Soc. Lond."},{"key":"2021042506080621000_R19","doi-asserted-by":"crossref","first-page":"71","DOI":"10.1242\/jeb.125.1.71","article-title":"Effects of ionic environment on viscosity of catch connective tissue in holothurian body wall","volume":"125","author":"Hayashi","year":"1986","journal-title":"J. Exp. Biol."},{"key":"2021042506080621000_R20","doi-asserted-by":"crossref","first-page":"108","DOI":"10.1016\/j.jsb.2008.06.007","article-title":"Micro- and nanostructure of the adhesive material secreted by the tube feet of the sea star Asterias rubens","volume":"164","author":"Hennebert","year":"2008","journal-title":"J. Struct. Biol."},{"key":"2021042506080621000_R21","first-page":"81","article-title":"Comparative physiology of echinoderm muscle","volume-title":"Echinoderm Studies","author":"Hill","year":"1993"},{"key":"2021042506080621000_R22","doi-asserted-by":"crossref","first-page":"351","DOI":"10.1201\/9780203970881.ch57","article-title":"Active state in echinoderm muscle","volume-title":"Echinoderms: M\u00fcnchen","author":"Hill","year":"2004"},{"key":"2021042506080621000_R23","volume-title":"The Invertebrates, Echinodermata","author":"Hyman","year":"1955"},{"key":"2021042506080621000_R24","first-page":"117","article-title":"Food and feeding mechanisms: Asteroidea","volume-title":"Echinoderm nutrition","author":"Jangoux","year":"1982"},{"key":"2021042506080621000_R25","doi-asserted-by":"crossref","first-page":"575","DOI":"10.1242\/jeb.30.4.575","article-title":"The forces exerted by the tube feet of the starfish during locomotion","volume":"30","author":"Kerkut","year":"1953","journal-title":"J. Exp. Biol."},{"key":"2021042506080621000_R26","doi-asserted-by":"crossref","first-page":"114","DOI":"10.2307\/1539188","article-title":"How sea stars open bivalves","volume":"111","author":"Lavoie","year":"1956","journal-title":"Biol. Bull."},{"key":"2021042506080621000_R27","volume-title":"A Functional Biology of Echinoderms","author":"Lawrence","year":"1987"},{"key":"2021042506080621000_R28","doi-asserted-by":"crossref","first-page":"88","DOI":"10.2307\/1542806","article-title":"Walking versus breathing: mechanical differentiation of sea urchin podia corresponds to functional specialization","volume":"198","author":"Leddy","year":"2000","journal-title":"Biol. Bull."},{"key":"2021042506080621000_R29","doi-asserted-by":"crossref","first-page":"1280","DOI":"10.1126\/science.4071053","article-title":"Myofibrils bear most of the resting tension in frog skeletal muscle","volume":"230","author":"Magid","year":"1985","journal-title":"Science"},{"key":"2021042506080621000_R30","doi-asserted-by":"crossref","first-page":"197","DOI":"10.2307\/1542085","article-title":"The functional morphology of starfish tube feet: the role of a crossed-fiber helical array in movement","volume":"188","author":"McCurley","year":"1995","journal-title":"Biol. Bull."},{"key":"2021042506080621000_R31","first-page":"41","article-title":"The stiffness change of the holothurian dermis caused by chemical and electrical stimulation","volume":"70C","author":"Motokawa","year":"1981","journal-title":"Comp. Biochem. Physiol."},{"key":"2021042506080621000_R32","doi-asserted-by":"crossref","first-page":"255","DOI":"10.1111\/j.1469-185X.1984.tb00409.x","article-title":"Connective tissue catch in echinoderms","volume":"59","author":"Motokawa","year":"1984","journal-title":"Biol. Rev."},{"key":"2021042506080621000_R33","first-page":"219","article-title":"Functional morphology of the water vascular system","volume-title":"Physiology of Echinodermata","author":"Nichols","year":"1966"},{"key":"2021042506080621000_R34","doi-asserted-by":"crossref","first-page":"207","DOI":"10.1016\/0022-0981(94)00158-A","article-title":"Attack behaviour and predatory success of Asterias rubens L. related to differences in size and morphology of the prey mussel Mytilus edulis L","volume":"186","author":"Norberg","year":"1995","journal-title":"J. Exp. Mar. Biol. 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Ser."},{"key":"2021042506080621000_R41","doi-asserted-by":"crossref","first-page":"2555","DOI":"10.1242\/jeb.01683","article-title":"Adhesion of echinoderm tube feet to rough surfaces","volume":"208","author":"Santos","year":"2005","journal-title":"J. Exp. Biol."},{"key":"2021042506080621000_R42","doi-asserted-by":"crossref","first-page":"2277","DOI":"10.1242\/jeb.01641","article-title":"The tube feet of sea urchins and sea stars contain functionally different mutable collagenous tissues","volume":"208","author":"Santos","year":"2005","journal-title":"J. Exp. Biol."},{"key":"2021042506080621000_R43","doi-asserted-by":"crossref","first-page":"259","DOI":"10.1002\/jmor.10187","article-title":"Comparative histological and immunohistochemical study of sea star tube feet (Echinodermata, Asteroidea)","volume":"263","author":"Santos","year":"2005","journal-title":"J. Morphol."},{"key":"2021042506080621000_R44","doi-asserted-by":"crossref","first-page":"9","DOI":"10.1007\/978-1-4020-6695-5_2","article-title":"The echinoderm tube foot and its role in temporary underwater adhesion","volume-title":"Functional Surfaces in Biology","author":"Santos","year":"2009"},{"key":"2021042506080621000_R45","doi-asserted-by":"crossref","first-page":"133","DOI":"10.1093\/oso\/9780199632237.003.0007","article-title":"Soft composites","volume-title":"Biomechanics. Materials. A Practical Approach","author":"Shadwick","year":"1992"},{"key":"2021042506080621000_R46","doi-asserted-by":"crossref","first-page":"91","DOI":"10.3354\/meps261099","article-title":"Influence of chronic, low-level hydrodynamic forces on subtidal community structure","volume":"261","author":"Siddon","year":"2003","journal-title":"Mar. Ecol. Prog. Ser."},{"key":"2021042506080621000_R47","doi-asserted-by":"crossref","first-page":"167","DOI":"10.1007\/978-3-540-31049-5_9","article-title":"The biochemistry and mechanics of gastropod adhesive gels","volume-title":"Biological Adhesives","author":"Smith","year":"2006"},{"key":"2021042506080621000_R48","first-page":"1","article-title":"The activities of tube feet of Asterias rubens L. I. The mechanics of movement and of posture","volume":"88","author":"Smith","year":"1947","journal-title":"Q. J. Microsc. Sci."},{"key":"2021042506080621000_R49","first-page":"887","article-title":"The effects of calcium chelation and cell perforation on the mechanical properties of sea-urchin ligaments","volume-title":"Echinoderms through time","author":"Szulgit","year":"1994"},{"key":"2021042506080621000_R50","doi-asserted-by":"crossref","first-page":"1539","DOI":"10.1242\/jeb.203.10.1539","article-title":"Dynamic mechanical characterization of a mutable collagenous tissue: response of sea cucumber dermis to cell lysis and dermal extracts","volume":"203","author":"Szulgit","year":"2000","journal-title":"J. Exp. Biol."},{"key":"2021042506080621000_R51","first-page":"121","article-title":"The catch apparatus of the sea-urchin spine II. Response to stimuli","volume":"11","author":"Takahashi","year":"1967","journal-title":"J. Fac. Sci. Univ. Tokyo IV"},{"key":"2021042506080621000_R52","doi-asserted-by":"crossref","first-page":"29","DOI":"10.2307\/3593098","article-title":"Mechanical properties of the isolated catch apparatus of the sea urchin spine joint: muscle fibers do not contribute to passive stiffness changes","volume":"208","author":"Takemae","year":"2005","journal-title":"Biol. Bull."},{"key":"2021042506080621000_R53","doi-asserted-by":"crossref","first-page":"45","DOI":"10.1086\/BBLv216n1p45","article-title":"Low oxygen consumption and high body content of catch connective tissue contribute to low metabolic rate of sea cucumbers","volume":"216","author":"Takemae","year":"2009","journal-title":"Biol. Bull."},{"key":"2021042506080621000_R54","doi-asserted-by":"crossref","first-page":"1817","DOI":"10.1242\/jeb.199.8.1817","article-title":"Morphology and biomechanics of the microfibrillar network of sea cucumber dermis","volume":"199","author":"Thurmond","year":"1996","journal-title":"J. Exp. Biol."},{"key":"2021042506080621000_R55","doi-asserted-by":"crossref","first-page":"1951","DOI":"10.1242\/jeb.198.9.1951","article-title":"Evidence that calcium-dependent cellular processes are involved in the stiffening response of holothurian dermis and that dermal cells contain an organic stiffening factor","volume":"198","author":"Trotter","year":"1995","journal-title":"J. Exp. Biol."},{"key":"2021042506080621000_R56","doi-asserted-by":"crossref","first-page":"357","DOI":"10.1042\/bst0280357","article-title":"Towards a fibrous composite with dynamically controlled stiffness. Lessons from echinoderms","volume":"28","author":"Trotter","year":"2000","journal-title":"Biochem. Soc. Trans."},{"key":"2021042506080621000_R57","doi-asserted-by":"crossref","first-page":"89","DOI":"10.1007\/BF00390534","article-title":"Prey capture by Pisaster brevispinus (Asteroidea: Echinodermata) on soft substrate","volume":"48","author":"Van Veldhuizen","year":"1978","journal-title":"Mar. Biol."},{"key":"2021042506080621000_R58","volume-title":"Comparative Biomechanics \u2013 Life's Physical World","author":"Vogel","year":"2003"},{"key":"2021042506080621000_R59","doi-asserted-by":"crossref","first-page":"1","DOI":"10.1080\/10236248409387032","article-title":"Variable tensility in echinoderm collagenous tissues: a review","volume":"11","author":"Wilkie","year":"1984","journal-title":"Mar. Behav. Physiol."},{"key":"2021042506080621000_R60","first-page":"61","article-title":"Mutable collagenous tissues: extracellular matrix as mechano-effector","volume-title":"Echinoderm Studies","author":"Wilkie","year":"1996"},{"key":"2021042506080621000_R61","doi-asserted-by":"crossref","first-page":"159","DOI":"10.1242\/jeb.205.2.159","article-title":"Is muscle involved in the mechanical adaptability of echinoderm mutable collagenous tissue?","volume":"205","author":"Wilkie","year":"2002","journal-title":"J. Exp. 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