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Deadly Strike Mechanism of a Mantis Shrimp This Shrimp Packs a Punch Powerful Enough to Smash Its Prey’S Shell Underwater

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Deadly Strike Mechanism of a Mantis Shrimp This Shrimp Packs a Punch Powerful Enough to Smash Its Prey’S Shell Underwater 22.4 Brief comms MH 15/4/04 5:40 pm Page 819 brief communications Deadly strike mechanism of a mantis shrimp This shrimp packs a punch powerful enough to smash its prey’s shell underwater. tomatopods (mantis shrimp) are well known for the feeding appendages they a b c Suse to smash shells and impale fish. Saddle Here we show that the peacock mantis shrimp (Odontodactylus scyllarus) generates c an extremely fast strike that requires major v energy storage and release, which we explain 0 ms m in terms of a saddle-shaped exoskeletal spring mechanism. High-speed images p reveal the formation and collapse of vapour bubbles next to the prey due to swift d movement of the appendage towards it, indi- 1 ms cating that O. scyllarus may use destructive cavitation forces to damage its prey. Stomatopod appendages were previously thought to be limited to a maximum speed of 10msǁ1 (ref. 1) but, by using new imaging technology and a faster species,we have mea- 2 ms sured the dactyl heel reaching peak speeds of 14–23 m sǁ1,peak angular speeds of 670–990 rad sǁ1, and peak acceleration of 65–104 km sǁ2 within an average period of 2.7ms,making O.scyllarusperhaps the fastest 3 ms appendicular striker in the animal kingdom. To generate these extreme movements in water, a large amount of energy must be released over a short period. Stomatopods Figure 1 The mechanics of a stomatopod strike. a, A high-speed image sequence illustrates the distal extension of the saddle (orange increase their power output through the use triangles) occurring simultaneously with the extension of the smashing heel of the dactyl and propodus (blue triangles). Scale bar, 1 cm. of a click mechanism,in which latches prevent Appendage kinematics were based on 6 individuals and 7–12 strikes per individual; measurement error, DŽ4%. Images were recorded at appendage movement until muscle contrac- 5,000 frames sǁ1. b, The compressed (top) and released (bottom) saddle on the raptorial appendage (raptorial segments: m, merus; tion is maximal1,2.When the latch is freed, v, meral–V; c, carpus; p, propodus; d, dactyl). c, The saddle modelled as a spring (orange) that stores elastic energy to drive the move- stored energy is released over a shorter time ment. Top, pre-strike phase. The lateral extensor muscle and apodeme (red) pull on the carpus to compress the saddle, while flexor than the duration of the original muscle con- muscles (not shown) engage a click mechanism to prevent extension of the appendage1. Bottom, the strike occurs when the latch is traction. Some extreme animal movements released1, enabling the saddle to extend and two pivot points to rotate in opposite directions. The meral–V forms a pivot point (black also require a specialized spring, because the circle) as it rotates distal–ventrally (anticlockwise here) and pushes the second pivot point, the carpal–meral fulcrum (white circle), distally. muscle fibres and tendon store insufficient The isometrically contracted extensor muscle maintains a constant distance between its carpal and meral attachment points (red circles), elastic strain energy3–5.We conservatively cal- forcing the carpus to rotate (clockwise here) and driving the dactyl heel towards the prey. culate that a minimum power requirement of 4.7ǂ105 watts per kilogram of muscle is nec- fraction of the strain energy underlying the occurs in fluids when areas of low pressure essary for a typical strike, which is orders of work requirements in an average strike (see form vapour bubbles that collapse and yield magnitude higher than that available in the supplementary information), which means considerable energy (in the form of heat, fastest-contracting muscles3,6 known. More- that stomatopods need a specialized spring. light and sound); these can be sufficient to over,the lateral extensor muscle and apodeme In our model for the stomatopod strike, destroy boat propellers and other hard sur- (arthropod tendon) could store only a small elastic energy is stored in a compressive, faces7.Cavitation has been noted in snapping saddle-shaped spring (Fig. 1) that is a stiff shrimp, which appear to ‘shoot’ cavitation exoskeletal structure located dorsally on the bubbles at prey items to stun them8,9. merus (the enlarged proximal segment) of all By contrast, cavitation in stomatopods stomatopods.Hyperbolic–paraboloid (saddle- occurs between the surface being struck and shaped or anticlastic) surfaces are used in the dactyl heel. Although the heel is highly engineering and architecture: their opposite mineralized10,the surface becomes pitted and and transverse curvatures reduce failure damaged over time; stomatopods moult fre- by distributing stresses across the three- quently and produce a new smashing surface dimensional surface. Likewise, the saddle every few months. In addition to their novel shape of the stomatopod’s spring minimizes energy-storage mechanism and remarkable the probability of local buckling while com- striking speed, stomatopods may be using pressing and extending. To our knowledge, cavitation forces to process their prey. the stomatopod’s saddle is the first biological S. N. Patek, W. L. Korff, R. L. Caldwell hyperbolic–paraboloid spring to be described. Department of Integrative Biology, University of Figure 2 The mantis shrimp Odontodactylus scyllarus strikes a Stomatopod smashing produces a loud California, Berkeley, California 94720-3140, USA snail. Cavitation bubbles (arrow) form and collapse between the pop, and high-speed video images reveal e-mail: [email protected] dactyl heel (red) and the snail (black). Scale bar, 1 cm. evidence of cavitation (Fig. 2). Cavitation 1. Burrows, M. Zeit. Vergl. Physiol. 62, 361–381 (1969). NATURE | VOL 428 | 22 APRIL 2004 | www.nature.com/nature 819 © 2004 Nature Publishing Group 22.4 Brief comms MH 15/4/04 5:40 pm Page 820 brief communications 2. Burrows, M. & Hoyle, G. J. Exp. Zool. 179, 379–394 (1972). troglodytes schweinfurthii) that lived about 3. Alexander, R. M. & Bennet-Clark, H. C. Nature 265, 114–117 a 800 km to the south-east in Gombe National (1977). 10,11 4. Bennet-Clark, H. C. in The Insect Integument Park in Tanzania .The new virus, which (ed. Hepburn, H. R.) 421–443 (Elsevier, Amsterdam, 1976). we designate SIVcpzDRC1, represents a 5. Gronenberg, W. J. Comp. Phys. A 178, 727–734 (1996). Kisangani o Riv third lineage within the well circumscribed ng er 6. Alexander, R. M. Comp. Biochem. Phys. A 133, 1001–1011 Ptt o C Pts P. t. schweinfurthii SIVcpz radiation, and is (2002). DRC 7. Brennen, C. E. Cavitation and Bubble Dynamics (Oxford clearly distinct from the P. t. troglodytes University Press, New York, 1995). Kinshasa SIVcpz clade that includes all known strains Gombe 8. Lohse, D., Schmitz, B. & Versluis, M. Nature 413, 477–478 400 km of HIV-1 (Fig. 1c, and see supplementary (2001). information). 9. Versluis, M., Schmitz, B., von der Heydt, A. & Lohse, D. Science 289, 2114–2117 (2000). These results indicate that chimpanzees 10.Currey, J. D., Nash, A. & Bonfield, W. J. Mater. Sci. 17, b in the vicinity of Kisangani are endemically 1939–1944 (1982). 1 234567891011 gp160 infected with SIVcpz that is highly divergent Supplementary information accompanies this communication on gp120 Nature’s website. from HIV-1, thereby ruling out these apes as Competing financial interests: declared none. p66 the source of HIV-1 and refuting the p51 gp41 OPV/AIDS theory.Instead,each of the many circulating HIV-1 variants comprising Origin of AIDS p31 groups M, N and O is linked to SIVcpz from p24 P. t. troglodytes(Fig.1c),the chimpanzee sub- Contaminated polio species native to west-central Africa1,12. p17 vaccine theory refuted Given that fears about the safety of polio vaccines are currently threatening the global espite strong evidence to the con- campaign to eradicate the disease8,our clear- trary1–5,speculation continues that cut evidence against one of the key sources of c 100 HIV1ELI the AIDS virus, human immunodefi- 100 HIV1LAI M concern is timely. The molecular epidemio- D SIVcpzPtt and HIV-1 ciency virus type 1 (HIV-1), may have HIV1U455 logical data presented here, together with crossed into humans as a result of contami- SIVcpzGAB1 data suggesting that HIV-1 group M origi- 100 HIV1YBF106 N nation of the oral polio vaccine (OPV)6–8. HIV1YBF30 nated 30 years before OPV trials were 65 82 SIVcpzUS 1,13 This ‘OPV/AIDS theory’ claims that chim- 98 SIVcpzCAM3 conducted and the absence of detectable panzees from the vicinity of Stanleyville — 58 66 SIVcpzCAM5 SIVcpz or chimpanzee DNA in archival now Kisangani in the Democratic Republic SIVcpzGAB2 stocks of OPV2,3, should finally lay the 100 HIV1MVP5180 O of Congo — were the source of a simian HIV1ANT70 OPV/AIDS theory to rest. SIVcpzANT immunodeficiency virus (SIVcpz) that was SIVcpzPts Michael Worobey*, Mario L. Santiago†, 99 SIVcpzDRC1 transmitted to humans when chimpanzee 99 76 SIVcpzTAN1 Brandon F. Keele†, Jean-Bosco N. Ndjango‡, 64 SIVcpzTAN5 tissues were allegedly used in the prepara- 100 Jeffrey B. Joy§, Bernard L. Labama||, 6,7 SIVcpzTAN3 tion of OPV .Here we show that SIVcpz 0.1 substitutions/site SIVcpzTAN2 Benoît D. Dhed’a‡, Andrew Rambaut¶, is indeed endemic in wild chimpanzees of Paul M. Sharp#, George M. Shaw†✩, this region but that the circulating virus is Beatrice H. Hahn† phylogenetically distinct from all strains of Figure 1 The Kisangani expeditions and refutation of the OPV/AIDS *Department of Ecology and Evolutionary Biology, HIV-1, providing direct evidence that these theory. a, Map of the Democratic Republic of Congo (DRC) and University of Arizona, Tucson, Arizona 85721, USA chimpanzees were not the source of the neighbouring countries showing the ranges of Pan troglodytes e-mail: [email protected] human AIDS pandemic.
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