{"status":"ok","message-type":"work","message-version":"1.0.0","message":{"indexed":{"date-parts":[[2026,8,24]],"date-time":"2026-08-24T23:33:51Z","timestamp":1787614431907,"version":"build-2736575974"},"reference-count":44,"publisher":"The Company of Biologists","issue":"12","content-domain":{"domain":[],"crossmark-restriction":false},"short-container-title":[],"published-print":{"date-parts":[[2005,6,15]]},"abstract":"<jats:title>SUMMARY<\/jats:title><jats:p>The dynamics of steady swimming were examined in the shortfin mako(Isurus oxyrinchus), a member of the cartilaginous fish family Lamnidae, a family known for their morphological adaptations for high-performance locomotion and their similarity in hydromechanical design to tunas. Patterns of red muscle (RM) strain (i.e. relative length change) and activation were quantified at two axial positions (\u223c0.4 and 0.6L,where L is total body length), using sonomicrometry and electromyography (EMG), and correlated with simultaneous measurements of dorsal midline kinematics during steady swimming (\u223c0.5\u20131 Ls\u20131). RM strain varied longitudinally with strain amplitudes ranging from 5.5\u00b11.1% (s.e.m.) in the anterior to 8.7\u00b10.9% in the posterior. We found no significant longitudinal variation in patterns of RM activation, with mean onset of activation occurring at 83\u201384\u00b0 (90\u00b0 is peak length) and offset at 200\u2013210\u00b0 at both body positions. Likewise, duty cycles were similar:35.5\u00b11.0% in the anterior and 32.2\u00b11.6% in the posterior. Comparison of the timing of waves of dorsal midline curvature and predicted strain relative to measured RM strain revealed a phase shift between RM shortening and local body bending. Furthermore, when the body is bent passively, RM shortens synchronously with the surrounding white muscle (WM)and skin, as expected. During active swimming, peaks in RM strain were delayed relative to peaks in WM strain by a mean of \u223c10% of the tailbeat cycle,with one individual as high as \u223c17% in the anterior and nearly 50% in the posterior. The longitudinal consistency in the EMG\/strain phase relationship in the mako is similar to that in the leopard shark, suggesting a consistent trend among sharks using different locomotor modes. However, unlike in the leopard shark, RM shortening in the mako is physically uncoupled from deformation of the surrounding body during steady swimming, a characteristic shared between the mako and tunas.<\/jats:p>","DOI":"10.1242\/jeb.01618","type":"journal-article","created":{"date-parts":[[2005,6,6]],"date-time":"2005-06-06T18:08:14Z","timestamp":1118081294000},"page":"2377-2387","source":"Crossref","is-referenced-by-count":34,"title":["Patterns of red muscle strain\/activation and body kinematics during steady swimming in a lamnid shark, the shortfin mako (<i>Isurus oxyrinchus<\/i>)"],"prefix":"10.1242","volume":"208","author":[{"given":"Jeanine M.","family":"Donley","sequence":"first","affiliation":[{"name":"Marine Biology Research Division, Scripps Institution of Oceanography,University of California, San Diego, La Jolla, CA 92093-0202, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Robert E.","family":"Shadwick","sequence":"additional","affiliation":[{"name":"Marine Biology Research Division, Scripps Institution of Oceanography,University of California, San Diego, La Jolla, CA 92093-0202, USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Chugey A.","family":"Sepulveda","sequence":"additional","affiliation":[{"name":"Pfleger Institute of Environmental Research, Oceanside, CA 92054,USA"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Peter","family":"Konstantinidis","sequence":"additional","affiliation":[{"name":"Department of Zoology, University of Tu\u0308bingen, Auf der Morgenstelle 28, 72076 Tu\u0308bingen, Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]},{"given":"Sven","family":"Gemballa","sequence":"additional","affiliation":[{"name":"Department of Zoology, University of Tu\u0308bingen, Auf der Morgenstelle 28, 72076 Tu\u0308bingen, Germany"}],"role":[{"vocabulary":"crossref","role":"author"}]}],"member":"237","reference":[{"key":"2021042510251505300_REF1","doi-asserted-by":"crossref","unstructured":"Altringham, J. D. and Block, B. A. (1997). Why do tuna maintain elevated slow muscle temperatures? Power output of muscle isolated from endothermic and ectothermic fish. J. Exp. Biol.200,2617-2627.","DOI":"10.1242\/jeb.200.20.2617"},{"key":"2021042510251505300_REF2","doi-asserted-by":"crossref","unstructured":"Altringham, J. D. and Ellerby, D. J. (1999). Fish swimming: Patterns in muscle function. J. Exp. Biol.202,3397-3403.","DOI":"10.1242\/jeb.202.23.3397"},{"key":"2021042510251505300_REF3","doi-asserted-by":"crossref","unstructured":"Altringham, J. D. and Johnston, I. A. (1990). Modelling muscle power output in a swimming fish. J. Exp. Biol.148,395-402.","DOI":"10.1242\/jeb.148.1.395"},{"key":"2021042510251505300_REF4","doi-asserted-by":"crossref","unstructured":"Altringham, J. D., Wardle, C. S. and Smith, C. I.(1993). Myotomal muscle function at different locations in the body of a swimming fish. J. Exp. Biol.182,191-206.","DOI":"10.1242\/jeb.182.1.191"},{"key":"2021042510251505300_REF5","unstructured":"Bernal, D., Sepulveda, C. A., Mathieu-Costello, O. and Graham,J. B. (2003). Comparative studies of high performance swimming in sharks I. Red muscle morphometrics, vascularization and ultrastructure. J. Exp. Biol.206,2831-2843."},{"key":"2021042510251505300_REF6","doi-asserted-by":"crossref","unstructured":"Biewener, A. A. and Gillis, G. B. (1999). Dynamics of muscle function during locomotion: accommodating variable conditions J. Exp. Biol.202,3387-3396.","DOI":"10.1242\/jeb.202.23.3387"},{"key":"2021042510251505300_REF7","unstructured":"Compagno, L. J. V. (1998). The living marine resources of the western central Pacific. In FAO Identification Guide for Fishery Purposes (ed. K. E. Carpenter and V. H. Niem),pp. 1274-1278. Rome: FAO."},{"key":"2021042510251505300_REF8","doi-asserted-by":"crossref","unstructured":"Coughlin, D. J. (2000). Power production during steady swimming in largemouth bass and rainbow trout. J. Exp. Biol.203,617-629.","DOI":"10.1242\/jeb.203.3.617"},{"key":"2021042510251505300_REF9","doi-asserted-by":"crossref","unstructured":"Coughlin, D. J. (2002). Aerobic muscle function during steady swimming in fish. Fish Fish.3, 63-78.","DOI":"10.1046\/j.1467-2979.2002.00069.x"},{"key":"2021042510251505300_REF10","doi-asserted-by":"crossref","unstructured":"Coughlin, D. J. and Rome, L. C. (1996). The roles of pink and red muscle in powering steady swimming in scup, Stenotomus chrysops.Am. Zool.36,666-677.","DOI":"10.1093\/icb\/36.6.666"},{"key":"2021042510251505300_REF11","doi-asserted-by":"crossref","unstructured":"Coughlin, D. J., Valdes, L. and Rome, L. C.(1996). Muscle length changes during swimming in scup:Sonomicrometry verifies the anatomical high-speed cine technique. J. Exp. Biol.199,459-463.","DOI":"10.1242\/jeb.199.2.459"},{"key":"2021042510251505300_REF12","doi-asserted-by":"crossref","unstructured":"D'Aout, K., Curtin, N. A., Williams, T. L. and Aerts, P.(2001). Mechanical properties of red and white swimming muscles as a function of the position along the body of the eel Anguilla anguilla.J. Exp. Biol.204,2221-2230.","DOI":"10.1242\/jeb.204.13.2221"},{"key":"2021042510251505300_REF13","doi-asserted-by":"crossref","unstructured":"Donley, J. M. and Shadwick, R. E. (2003). Steady swimming muscle dynamics in the leopard shark Triakis semifasciata.J. Exp. Biol.206,1117-1126.","DOI":"10.1242\/jeb.00206"},{"key":"2021042510251505300_REF14","doi-asserted-by":"crossref","unstructured":"Donley, J. M., Sepulveda, C. A., Konstantinidis, P., Gemballa,S. and Shadwick, R. E. (2004). Convergent evolution in mechanical design of lamnid sharks and tunas. Nature429, 61-65.","DOI":"10.1038\/nature02435"},{"key":"2021042510251505300_REF15","doi-asserted-by":"crossref","unstructured":"Ellerby, D. J. and Altringham, J. D. (2001). Spatial variation in fast muscle function of the rainbow trout Oncorhynchus mykiss during fast-starts and sprinting. J. Exp. Biol.204,2239-2250.","DOI":"10.1242\/jeb.204.13.2239"},{"key":"2021042510251505300_REF16","doi-asserted-by":"crossref","unstructured":"Ellerby, D. J., Altringham, J. D., Williams, T. and Block, B. A. (2000). Slow muscle function of pacific bonito (Sarda chiliensis) during steady swimming. J. Exp. Biol.203,2001-2013.","DOI":"10.1242\/jeb.203.13.2001"},{"key":"2021042510251505300_REF17","doi-asserted-by":"crossref","unstructured":"Gemballa, S. and Ro\u0308der, K. (2004). From head to tail: The myoseptal system in basal actinopterygians. J. Morphol.259,155-171.","DOI":"10.1002\/jmor.10175"},{"key":"2021042510251505300_REF18","doi-asserted-by":"crossref","unstructured":"Gemballa, S. and Treiber, K. (2003). Cruising specialists and accelerators \u2013 Are different types of fish locomotion driven by differently structured myosepta? Zoology106,203-222.","DOI":"10.1078\/0944-2006-00116"},{"key":"2021042510251505300_REF19","doi-asserted-by":"crossref","unstructured":"Gemballa S., Ebmeyer L., Hagen K., Hoja K., Treiber K., Vogel F. and Weitbrecht, G. W. (2003). Evolutionary transformations of myoseptal tendons in gnathostomes. Proc. R. Soc. London B Biol. Sci.270,1229-1235.","DOI":"10.1098\/rspb.2003.2345"},{"key":"2021042510251505300_REF20","doi-asserted-by":"crossref","unstructured":"Gillis, G. (1998). Neuromuscular control of anguilliform locomotion: patterns of red and white muscle activity during swimming in the American eel Anguilla rostrata.J. Exp. Biol.201,3245-3256.","DOI":"10.1242\/jeb.201.23.3245"},{"key":"2021042510251505300_REF21","doi-asserted-by":"crossref","unstructured":"Graham, J. B., Dewar, H., Lai, N. C., Lowell, W. R. and Arce, S. M. (1990). Aspects of shark swimming performance determined using a large water tunnel. J. Exp. Biol.151,175-192.","DOI":"10.1242\/jeb.151.1.175"},{"key":"2021042510251505300_REF22","doi-asserted-by":"crossref","unstructured":"Grillner, S. and Kashin, S. (1976). On the generation and performance of swimming in fish. In Neural control of locomotion (ed. R. M. Herman, S. Grillner, P. S. G. Stein and D. G. Stuart), pp. 181-201. New York: Plenum Press.","DOI":"10.1007\/978-1-4757-0964-3_8"},{"key":"2021042510251505300_REF23","doi-asserted-by":"crossref","unstructured":"Hammond, L., Altringham, J. D. and Wardle, C. S.(1998). Myotomal slow muscle function of rainbow trout Oncorhynchus mykiss during steady swimming. J. Exp. Biol.201,1659-1671.","DOI":"10.1242\/jeb.201.10.1659"},{"key":"2021042510251505300_REF24","unstructured":"He, P., Wardle, C. S. and Arimoto, T. (1990). Electrophysiology of the red muscle of mackerel, Scomber scombrus and its relation to swimming at low speeds. In The 2nd Asian Fisheries Forum (ed. R. Hirano and I. Hanyu), pp.469-472. Manila: Asian Fisheries Society."},{"key":"2021042510251505300_REF25","doi-asserted-by":"crossref","unstructured":"Hess, F. and Videler, J. J. (1984). Fast continuous swimming of saithe (Pollachius virens): A dynamic analysis of bending movements and muscle power. J. Exp. Biol.109,229-251.","DOI":"10.1242\/jeb.109.1.229"},{"key":"2021042510251505300_REF26","doi-asserted-by":"crossref","unstructured":"Jayne, B. C. and Lauder, G. V. (1993). Red and white muscle activity and kinematics of the escape response of the bluegill sunfish during swimming. J. Comp. Physiol. A.173,495-508.","DOI":"10.1007\/BF00193522"},{"key":"2021042510251505300_REF27","doi-asserted-by":"crossref","unstructured":"Jayne, B. C. and Lauder, G. V. (1995). Red muscle motor patterns during steady swimming in largemouth bass: Effects of speed and correlations with axial kinematics. J. Exp. Biol.198,1575-1587.","DOI":"10.1242\/jeb.198.7.1575"},{"key":"2021042510251505300_REF28","doi-asserted-by":"crossref","unstructured":"Johnson, T. P. and Johnston, I. A. (1991). Power output of fish muscle fibres performing oscillatory work: effects of acute and seasonal temperature change. J. Exp. Biol.157,409-423.","DOI":"10.1242\/jeb.157.1.409"},{"key":"2021042510251505300_REF29","doi-asserted-by":"crossref","unstructured":"Katz, S. L. (2002). Design of heterothermic muscle in fish. J. Exp. Biol.205,2251-2266.","DOI":"10.1242\/jeb.205.15.2251"},{"key":"2021042510251505300_REF30","doi-asserted-by":"crossref","unstructured":"Katz, S. L. and Shadwick, R. E. (1998). Curvature of swimming fish midlines as an index of muscle strain suggests swimming muscle produces net positive work. J. Theor. Biol.193,243-256.","DOI":"10.1006\/jtbi.1998.0696"},{"key":"2021042510251505300_REF31","doi-asserted-by":"crossref","unstructured":"Katz, S. L., Shadwick, R. E. and Rapoport, H. S.(1999). Muscle strain histories in swimming milkfish in steady and sprinting gaits. J. Exp. Biol.202,529-541.","DOI":"10.1242\/jeb.202.5.529"},{"key":"2021042510251505300_REF32","doi-asserted-by":"crossref","unstructured":"Katz, S. L., Syme, D. A. and Shadwick, R. E.(2001). Enhanced power in yellowfin tuna. Nature410,770-771.","DOI":"10.1038\/35071170"},{"key":"2021042510251505300_REF33","doi-asserted-by":"crossref","unstructured":"Knower, T., Shadwick, R. E., Katz, S. L., Graham, J. B. and Wardle, C. S. (1999). Red muscle activation patterns in yellowfin and skipjack tunas during steady swimming. J. Exp. Biol.202,2127-2138.","DOI":"10.1242\/jeb.202.16.2127"},{"key":"2021042510251505300_REF34","doi-asserted-by":"crossref","unstructured":"Rome, L. C. and Swank, D. (1992). The influence of temperature on power output of scup red muscle during cyclic length changes. J. Exp. Biol.171,261-282.","DOI":"10.1242\/jeb.171.1.261"},{"key":"2021042510251505300_REF35","doi-asserted-by":"crossref","unstructured":"Rome, L. C., Funke, R. P. and Alexander, R. M.(1990). The influence of temperature on muscle velocity and sustained performance in swimming carp. J. Exp. Biol.154,163-178.","DOI":"10.1242\/jeb.154.1.163"},{"key":"2021042510251505300_REF36","unstructured":"Rome, L. C., Sosnicki, A. and Choi, I. H.(1992). The influence of temperature on muscle function in the fast swimming scup. 2. The mechanics of red muscle. J. Exp. Biol.163,281-295."},{"key":"2021042510251505300_REF37","doi-asserted-by":"crossref","unstructured":"Rome, L. C., Swank, D. and Corda, D. (1993). How fish power swimming. Science261,340-343.","DOI":"10.1126\/science.8332898"},{"key":"2021042510251505300_REF38","doi-asserted-by":"crossref","unstructured":"Shadwick, R. E., Steffensen, J. F., Katz, S. L. and Knower,T. (1998). Muscle dynamics in fish during steady swimming. Amer. Zool.38,755-770.","DOI":"10.1093\/icb\/38.4.755"},{"key":"2021042510251505300_REF39","doi-asserted-by":"crossref","unstructured":"Shadwick, R. E., Katz, S. L., Korsmeyer, K. E., Knower, T. and Covell, J. W. (1999). Muscle dynamics in skipjack tuna:timing of red muscle shortening in relation to activation and body curvature during steady swimming. J. Exp. Biol.202,2139-2150.","DOI":"10.1242\/jeb.202.16.2139"},{"key":"2021042510251505300_REF40","doi-asserted-by":"crossref","unstructured":"Syme, D. A. and Shadwick, R. E. (2002). Effects of longitudinal body positions and swimming speed in mechanical power of deep red muscle from skipjack tuna (Katsuwonus pelamis). J. Exp. Biol.205,189-200.","DOI":"10.1242\/jeb.205.2.189"},{"key":"2021042510251505300_REF41","doi-asserted-by":"crossref","unstructured":"van Leeuwen, J. L., Lankheet, M. J. M., Akster, H. A. and Osse,J. W. M. (1990). Function of red axial muscles of carp(cyprinus-carpio): recruitment and normalized power output during swimming in different modes. J. Zool.220,123-145.","DOI":"10.1111\/j.1469-7998.1990.tb04298.x"},{"key":"2021042510251505300_REF42","doi-asserted-by":"crossref","unstructured":"Wardle, C. S. and Videler, J. J. (1993). The timing of the electromyogram in the lateral myotomes of mackerel and saithe at different swimming speeds. J. Fish Biol.42,347-359.","DOI":"10.1111\/j.1095-8649.1993.tb00338.x"},{"key":"2021042510251505300_REF43","doi-asserted-by":"crossref","unstructured":"Wardle, C. S., Videler, J. J. and Altringham, J. D.(1995). Tuning in to fish swimming waves: Body form, swimming mode and muscle function. J. Exp. Biol.198,1629-1636.","DOI":"10.1242\/jeb.198.8.1629"},{"key":"2021042510251505300_REF44","doi-asserted-by":"crossref","unstructured":"Williams, T. L., Grillner, S., Smoljaninov, V. V., Wallen, P.,Kashin, S. and Rossignol, S. (1989). Locomotion in lamprey and trout: the relative timing of activation and movement. J. Exp. Biol.143,559-566.","DOI":"10.1242\/jeb.143.1.559"}],"container-title":["Journal of Experimental Biology"],"original-title":[],"language":"en","link":[{"URL":"http:\/\/journals.biologists.com\/jeb\/article-pdf\/208\/12\/2377\/1545154\/2377.pdf","content-type":"application\/pdf","content-version":"vor","intended-application":"syndication"},{"URL":"http:\/\/journals.biologists.com\/jeb\/article-pdf\/208\/12\/2377\/1545154\/2377.pdf","content-type":"unspecified","content-version":"vor","intended-application":"similarity-checking"}],"deposited":{"date-parts":[[2021,7,11]],"date-time":"2021-07-11T13:12:02Z","timestamp":1626009122000},"score":1,"resource":{"primary":{"URL":"https:\/\/journals.biologists.com\/jeb\/article\/208\/12\/2377\/15490\/Patterns-of-red-muscle-strain-activation-and-body"}},"subtitle":[],"short-title":[],"issued":{"date-parts":[[2005,6,15]]},"references-count":44,"journal-issue":{"issue":"12","published-print":{"date-parts":[[2005,6,15]]}},"URL":"https:\/\/doi.org\/10.1242\/jeb.01618","relation":{},"ISSN":["1477-9145","0022-0949"],"issn-type":[{"value":"1477-9145","type":"electronic"},{"value":"0022-0949","type":"print"}],"subject":[],"published":{"date-parts":[[2005,6,15]]}}}