CINXE.COM

<!DOCTYPE html><html lang="en-US"><head><title>Just a moment...</title><meta http-equiv="Content-Type" content="text/html; charset=UTF-8"><meta http-equiv="X-UA-Compatible" content="IE=Edge"><meta name="robots" content="noindex,nofollow"><meta name="viewport" content="width=device-width,initial-scale=1"><style>*{box-sizing:border-box;margin:0;padding:0}html{line-height:1.15;-webkit-text-size-adjust:100%;color:#313131;font-family:system-ui,-apple-system,BlinkMacSystemFont,Segoe UI,Roboto,Helvetica Neue,Arial,Noto Sans,sans-serif,Apple Color Emoji,Segoe UI Emoji,Segoe UI Symbol,Noto Color Emoji}body{display:flex;flex-direction:column;height:100vh;min-height:100vh}.main-content{margin:8rem auto;max-width:60rem;padding-left:1.5rem}@media (width <= 720px){.main-content{margin-top:4rem}}.h2{font-size:1.5rem;font-weight:500;line-height:2.25rem}@media (width <= 720px){.h2{font-size:1.25rem;line-height:1.5rem}}#challenge-error-text{background-image:url(data:image/svg+xml;base64,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);background-repeat:no-repeat;background-size:contain;padding-left:34px}@media (prefers-color-scheme:dark){body{background-color:#222;color:#d9d9d9}}</style><meta http-equiv="refresh" content="390"></head><body class="no-js"><div class="main-wrapper" role="main"><div class="main-content"><noscript><div class="h2"><span id="challenge-error-text">Enable JavaScript and cookies to continue</span></div></noscript></div></div><script>(function(){window._cf_chl_opt={cvId: '3',cZone: "academic.oup.com",cType: 'managed',cRay: '8f4855d9da5fce65',cH: '2rsckMBleGcIFXhT9tAYPGoLp8fehfhsLOgJalMM_Sw-1734621193-1.2.1.1-0tRssLTFDjr0VRrX6fmjGkfKvfQH5EVxNcqSeOEd0mGF3jlaG1L2p_o5Rb_SVuee',cUPMDTk: "\/treephys\/article-pdf\/39\/5\/717\/28871432\/tpy142.pdf?__cf_chl_tk=3XyVrtL8R0acy05DvQQPyzAfIOP8D7U5DQQ8AylcB7w-1734621193-1.0.1.1-WeUrn54lxVkoHJl8AvTiEDyU6kTxs7prOqyUKLFYdgw",cFPWv: 'b',cITimeS: '1734621193',cTTimeMs: '1000',cMTimeMs: '390000',cTplC: 0,cTplV: 5,cTplB: 'cf',cK: "",fa: "\/treephys\/article-pdf\/39\/5\/717\/28871432\/tpy142.pdf?__cf_chl_f_tk=3XyVrtL8R0acy05DvQQPyzAfIOP8D7U5DQQ8AylcB7w-1734621193-1.0.1.1-WeUrn54lxVkoHJl8AvTiEDyU6kTxs7prOqyUKLFYdgw",md: "qM7eHXnJoGYQjkJSOOp5eSmbC5GbpWmW94zcAy0mycE-1734621193-1.2.1.1-9l.2eFhH6tgL.Y2xvGZMf2IP8KbuHk0_SDll7YFgk1OPgg8yNwmVTwcISiSLOM_oRViLokAiVFokolGyczkv1hQVDCSEjbOmwE1_W79_yc0n_DD8WsJN6Oi1WnmQ3GqVaLLCv1DzLo31P84PynEdrHmUve0PUqC1_ZJfyR5yyjtP08qykPNbwnl2.PNyTvmYZyi_nyaNQpxC.TBzpnrruL7kKJH.eKwK.JqKeifmhHEYlbtmgQWn.7Tl8McFIFMK6Ze09jsqCTsr_6Mi5nePzjK_fAkQ_8DRJh93IiNAc6xu4wmMtdFHwx7eF1pJ9o.XWr9VOXSMdaPABFCgqDJ3mpArgHKcjugkEdlpFP4wv.Bs.1BHp_0ucvJfMAsWT3O6z4CMXGzm011g6GF0lzXizN0w086zx2A3SbKkwFBrw7nTtFU.TUHH6A_C8Ua1teKL0RKsxnjCiQE9ISwdvFmiIvEu4IjJtbCfqQU7jUBschrGKsjakmhFguNH59O2SdhpgYUWPQqTPENIFlzpoaeWKlP10ddkoaFQ6Wa2lV2S.pxxm.MSy.C.MbiAvSio9gzydc_C9ol4Z6wmiK6Vb4y9D9kzQe_zqPVE9o2HKjUkFfG9tfUJDEpM9SVXI7VYTJ8k1bpkaphmIcKJkaOPf1_fKjwqt.z_g7eoIY_rmiyNQAtuHEj9sT_amUoKPWQb3hkjDAdnZ1hO2.nUagETYN8bsjyK_7fEhv4HQLb4iCGfblyYdWkN58NZGntI91X9t6OZUr4p3P8CwMO2UIO6AzrCuxHgzjGoWLCVhB2etr.ABtZPvLJZa4gndtGiWyVNUngWPnRFHIo1yIuJx4m6pKuGjeSC.YUwgaT3H2YAseMIxbsnmHGeL54.fgow_ePj.FrHA.16Ai5g8vr1pyUbHzRx5T4VwVKX_.o0x2_GUxTiMXPMi7kLj1gF8MlY1M13MJXmOXBCouJxCo1RR.0ta1rWcKiKuVq_3XargWESokWyx5Ia9D82vDZkHMuq77NugVIWYHHHkeGrmyBj4B1_9E65I2kp6k8sAQ4GYme5YIAXbeQIWF.aAEB4nZ5tWEsloWgIr4s1ZigCUCIumm7jPQLyCZLkWJzHByGX2Ouaf1Tpl9lKFKlN1OK1GrLVRoBlTdCj7qJhoNih6xiGDOhUQMuAVrw4gDGFKXB0ahGdj.GExPQjNsSIWsKES0C_N4pmY1G3ptH5eJKKmIPmbakFVMdqW98.0CSvP20igXLvM1DNUOU5YTKvUQfBkcTK7j2pL7.6AS0lLdCbedg26SwdFZ3_rZY14St95zUL6L2JAPxij_yJ0aBPd6PlRgndFq_MGPzatGDbqCO4Fb6t4k..nKhIoFn1F9k6tZWT8P.vyjSZmDlS947Trw6SYY4QsuDmkXP0hsTMHEzp7kNVFX4On7FLfxcprmLNqCF2_XYMqNJzfSBnMjwmThtsLbCtkaO70ACkCtJtBQCy.RBZsjOE4AZs3NDnuRFGHse3hcjkJCwSgkaMDHr2MWp2sPt8o0hvc4TvgGzNhhnIInar2O1ym2lS3FRM3o1hKshlodDKRK3wwf1FCfcOrTQ9M.vi0Q6wTFFldFr.lnP7yhp2z7VrHiGH_YKB1XHRsLncZe4vjOA5nR2Fyph.CjXsHzTbmOZLwr9TNt6oQsKnvI70F8OQKYbHdqvnWIhdRPVNbrP7V7h_nMUh0Z9.iOWa5JKfC9zgPtNtBDMqY1CP9AEt1PjdGQ9sLnA.lkT1XIHIiNFh9b2RpHUByssNWrUkUidLrjPkBEd7e9Lll9hcss.qXTAcC6DQ7d0_6ZicGTxHDj1FO30klAsYcvzmrgZqXANfEpjRVg7gV0CWLifvrEyTfNQc76Kb2GSILztTo8Q99eShHINxeE8vqZlj23MS2PB4ZgT_II2iCXOkjv6tgabs9PwVi7uvu7Bnj8LtiSOLm23ceEPjWXUaji0UBlSVATZjDXmIXBynpzCXKk17pWmcTLsrndnyXpOl1Q8ANLyX1HPVSgSU1HLpXxS7Ar3Cd2I7JRCPbrrq3NMelsetRMrpRkYq04WvCFE8tnqt.aLX_ehMPcKaz0KCOY2cFA2erRgkeTqOCRujWjPWoSwvqv77DBNYCkklgn39lWJXfbdr1C48_wLsL8nv171Wbqnz6CSa.oGNR3t79n2ikpxtP0W3T_0sJWtcX7CaoEWjwsv3B6tyawrf0JaKwty7AT0dzsMG1Vu1OvLpVyM85ALWlscBnZSNA9DwrsQqzrlo.LirRHnrwmfTOloiwyNvxg7F2Rzc4Vo3opfysuVp7K90Y7AWP9bwIBvdHZSTfBT3.41SwDtd0i3lAywrnfarHBp1LwtWCmG9d.kT5wJHQIV4liSUHO4lIimaExVrEHo2NjEsQ3Mj9qOQdeVjk_bVg71rUeyWZdwNPeOLzBNkzROyfXO0c9ZGxI.COMkWbsSAcDqMHLTESJEBokflLU9I9iUNwTSfGLmaFQp7dp1esf1gWUIT5mrwAZ2yWpUd_T4thyasmfFD.8xx2Im8GxWtJpQL.hL1w5nowcooIDri.JzzcP1UTawn.hQuIa9hoswpK6PygznatLjPe.wd5b3J92w8F72ZD_IVSg5U_ohSl_NZiQ3oRHw4gqaoc0gIH.mbDRsL3zkWBPfCAFq5fNp2Ddh2u46Q42kqA1EpSIu9vH1GeW3t8ydBZT2vpguxL3097.1LQKSY8c8rFKDvQlOvFA96NXAWQpHrvJL06Nn0Nqaa0vrtt9xOWSNuRbx23id4iMiAhh9Rd8oZD9Y4jPnuVyXXlCN0rkw0W3cdz5ibTodoMTxspVMF9YfjXyKh_JLV09ohV15HNXbpGrZWhzlTVi1sgKcOIErDTw1mJbhJEEonE4qWEnMqt0ypBA8v_N.xdxRQViQMMrI2mbh2UtfzLtogeDylkO4BGGUTeZJcLPtRGFd1FfcMIlTSDtEFoDGoYOp4bcgAoIuPxF1HZ9U1TZIZG8lKTH1HspCPPRGUMU3GTclDuCQZqCIrIZs9u1WEf6cxzgL0XmTEX_k",mdrd: "XAbcPG7ZtkEfCAhEiDLs6cd5exBu7VTvcW3cWd8za1s-1734621193-1.2.1.1-ipur1xIfLSnKUQgs.qwktggUN2pz9lHaTEqq_I3hGi6fjPxOIU.acZtpwP1KvWw8GRRiLAe6bgzXSNxOfe.BE0bq5to_kq46QNuucBzB0dLw124GV.8PbVnee2OHOAI8ykTQiA0szD6ZNyMqzYw2aE.80z6DBwJODHahKmEcF4MREz3JLkfJ0fePKigTcbT9g7JB51CuS8iHtD5v2Qs34wiYhVUXTHe.ERrF0oVTiYdQKZZef96WQJHIM1x0ooXKTLIfqYeLj.FsTUOWmgUntKJRXnLNrAhAY3b6jzZGY6ApQZeokMcbLpPHg6Htg4trA6af3mWxojiSOQhnsovp2ALzUt4NQpVVgdckGg9dGxnmpu6svu04h1SdZtnSo_8tW1f3TKr50YZ.TAhUUxiErl299xW93PNKI3OOfY50f26bv.4VBLxeH.1LwyBbNVUNdxajb38OVbcMl1Aycz2yVV8X_sqpwU.Y2mes1TUP7EIEmAvStVtlaDML1LJMwt4x822yFTS2VCxe3WSZ4XA4aKBlbJA1GHjnR817.L2iDMsjSeQHkHMqJJNMUGducD.axdxvI6bpH5BPswFjAZomUXQIM1PI5qtet9pn12QEbi3U_NPcmpx0ZJ9CHMCaYnCBAHg3yP2_ukrXy.rFUdP3Jdhwu_0iBqmcRLQ8eWyqNtl8fVDhcDuRoJzkjnPQ8IudeuElYnf45hriTh23CEjMpB88XC5NuGSHVrbXfs1.z9HaNlUWkfpWjSDd_I410GZdLDLOhMqbT9iS07uro80GCfR3e.xCrBeLcosqNsVs6XdR8LnzXn880uv2661jtix95sSQ8VJGqswKRZajvuAkhD6ruzaupyS7SYeHr74j12VjBUkOobx4765t7VWSWQW_vyUqVtAVkcOBKd5F5U7FJZ28OQhkPI9kjcJArRxpMLAh40m82vzq0IPQcsciolPWeAVRWfzxn_Fzu0HSkk2csgkzuhNu8DuBInQ6M55zCJPSWCadYQ4llootYks7qX418tzQqr1uiZSWwFNrlKF.6LQcaC7Fb0NnCF78U4cuju.1iMGKjS3Dj_yc8MGqx8qCsqq7tnkB76twL0HqjOeKfqJVN1AqtMpT_jGX9slxZ3ywjU1cZ4ANngbVtjsImcKkjC0E6ZRRiihwN7DlPNWOfVH8kGY_ErRjM3z6zsyGiif2Ap5bOcytukuZz0IaQegT5BlWCYr7Bblr.K8DjHPs85f7pK0xxgVxgtzpSHx7otEEDNysXmYRiUbgABZNphtnt.wnL8r71fRCDEEV7BL_LFHAGplgoF6RfZpWexlmNQJxf3ftAh0XwV_QXTkWbxTIIIKZ9lgUpBTuHb9YuoHscF0_3XmmtTrTxtF9dst6Y1ISOUi7GQqwbI4PmmLLIx3Knzc2uiXVKh_2olUAq1ZIzgoQnh0k58ai2OWYMOZ7q6kFdxTsmbFEkwI8HjmRe_e2tZaT2vNGevBj393Q7BEDXWGL1d44sxuld8_NB7DHU920q6dK0U4JTJZ6qQu_PFGFsrleavBGPUlpopT0yT1jThl6f19NV1MHS.R6VnyuBZyK7sU1WzuxcrFCnuTiORiJ3Pyk.BFybf9Z7hhPdcMEQJsS0zQ5WbD6Kvltfcu1CNDTKOlRP9U7sc0tmNZ8ZEOQNWN9ad38UlJkN2g4Q7cCM9Gqe22nwzXqG5GwDDov1r0JyMrYeerc6kg2NB41c4CDRqkNyO2SNI52XBN2DYpPKQDftkk33AvSM8LcUZvQHKv4f9Y9nSxqkIMr6u0eKsBpfiwFtrhEbD7ZiusuI3MkOdhDGm6gTALdWhtqruEoVSfE8_80E34vKE59dHYgnrysDdl3.OlNo8pY.ZfCma0XbYOOQEDIUozMMt5mhj9Xa7eT2vsqWOJI5sGLV1O8lyWbaDOdsYFmFoVhQfIar_is_hmprF9kIN8ncT7jqIshGsUGAEY8MZMXFQLdpP3ejOlC"};var cpo = document.createElement('script');cpo.src = '/cdn-cgi/challenge-platform/h/b/orchestrate/chl_page/v1?ray=8f4855d9da5fce65';window._cf_chl_opt.cOgUHash = location.hash === '' && location.href.indexOf('#') !== -1 ? '#' : location.hash;window._cf_chl_opt.cOgUQuery = location.search === '' && location.href.slice(0, location.href.length - window._cf_chl_opt.cOgUHash.length).indexOf('?') !== -1 ? '?' : location.search;if (window.history && window.history.replaceState) {var ogU = location.pathname + window._cf_chl_opt.cOgUQuery + window._cf_chl_opt.cOgUHash;history.replaceState(null, null, "\/treephys\/article-pdf\/39\/5\/717\/28871432\/tpy142.pdf?__cf_chl_rt_tk=3XyVrtL8R0acy05DvQQPyzAfIOP8D7U5DQQ8AylcB7w-1734621193-1.0.1.1-WeUrn54lxVkoHJl8AvTiEDyU6kTxs7prOqyUKLFYdgw" + window._cf_chl_opt.cOgUHash);cpo.onload = function() {history.replaceState(null, null, ogU);}}document.getElementsByTagName('head')[0].appendChild(cpo);}());</script></body></html>