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: "royalsocietypublishing.org",cType: 'managed',cRay: '9149502a1aaa3e5f',cH: 'yviFjCyjb9CtxiFUy.clFpDQ1GyfXuESJ_qiQi1jCaM-1740000155-1.2.1.1-BY7WO5QEZ3NjuImjuj0pNsFB9fDwgmaNncrjnsSRx0BRoLSFRyoc9436q83I_m73',cUPMDTk: "\/search\/advanced?__cf_chl_tk=RwPiWTpF2j3eVODKYgCExtgA92pLA7SnRBPOPNdYiX0-1740000155-1.0.1.1-ESB..SHztQEuthRTPm.b8kwS39MTVSfTnSwrqMZ3eu0",cFPWv: 'g',cITimeS: '1740000155',cTTimeMs: '1000',cMTimeMs: '390000',cTplC: 0,cTplV: 5,cTplB: 'cf',cK: "unsupported-browser",fa: "\/search\/advanced?__cf_chl_f_tk=RwPiWTpF2j3eVODKYgCExtgA92pLA7SnRBPOPNdYiX0-1740000155-1.0.1.1-ESB..SHztQEuthRTPm.b8kwS39MTVSfTnSwrqMZ3eu0",md: "knZMW3AfdQQWUsraak0P2.vnYgulrBLdBdTV9NX2yRI-1740000155-1.2.1.1-FqZSt2y5SZRQJ9LzpLmTLP63V8QcsCTbZKuD2isxr6ZtAu70oDL4tWi0.rKCShnFik3rQsN4STD9jIzzWt04hQAtq3pJaEhJafS5oGfJR2BIRICUf9M3e_P6npH6U1nVNQkbCp9kLnQHSDdQXPdm8sgdtiQudrhT0GcgGktCl9YxlY9jzHdeczT_Eiu.Olzk5iUeMnJyjsOMAa8f52c0.50hkEfL0VTWByzhO3x.Va_R0j_i_Zw16GWvOttPt5S2cnlupUE1Vth5BzU1gm0jGe7Q4AbVQH74dPUJHqT6PlW5StIsAMJDhiGicpHdiXeNEB8dXCGnDFGRjq6fy3wCMJ9YiQOBF6cM9d1LxktlDfmWem8jZ_eTewyxx0jih5yyyDLajUlo5kYcU4atMrCMp1CPuGzp4LSP.PSIIgCvWa3dFkJQVjZXR6oBHyyC8VQWP847cwXNFMvkibKxh9bGZSFh58aKhwauf_Huf1GqXSwPktWlVJ29XNzziGzskkN36RkDPl.SLT03ymwJ2dXXlAJP7z3SetALNb7SlqfsH8W0_ZN.N79JBHPN6R2iHk_B.xZtwL8kZvUAM5RK4v7OtqJmhaVw3UhOt3mRMaWdNMY5KkrvMozh7TGNYJbhCI1siUclLVGzOk4ben3pwTMa5qgNt0gb.5OnQRaBV2MLs2yfjb4PQXWHxWuE2N9iDagTWAaGUtS3vubIXInHeLF9PXqa6JUc2Cb7twQgGVzI9TFw5MQXQNNAaK.p3L33Tm1vDWYmyg.92LfZqkqdo6FWFDBUdqpwAvT5iKOgbuRrI2caMkQQxe2T89okiy7WOI8mF38ZJo9.20i8IqZHaVvAVqn9LRB0m1qpJQBP20aS2aKJKqNturBokbGodpJqUBvw3KdZUr13gNfoTkB4136fPspAbL_mvUjhU30zYCWr11Yp0NZQB8nWEFhGIw_gy_oTD5SQfPalzBfZxHRkbB85rq3J08oaFilyHR_sPut57c_L9kaySw4KFEUhD78LOWo_j.ygM2EzKsPRMUs0nJgW4E_pMLa9Oj9lMQaBHobnv9aVh_BpX61Up6jFJ8uGlpaxNT4Yam1oGsmZEhymyTVTgY1NgThnsGLwfcXElH7Z7Cv5xexMyNOoNyLK9U9mt3JQyIYUVPqQ3DzP1aaIbVEMdPGUcjUuWntsUNumkgbE.RU.nO2BhoIPwLVk4_UDsj0d469fhOFvndHhKAZNSIPleR8sD9WJYfGhvVJhOBDaEGjyALOPWakiLXs6eYUG388OiK1w0sPhEcgsHeCrA8n_Ouc7f5xu4.McTGlrNAmUFxUBcL09gFFwv1oZyoEqDPIM0_l5W3stNQyvUZ70KqtdlOJWSwGrp6XMZglQOrHDRNuCc4PaE1ZVMW2pR5xw2SHvFrr8gnnEJa3.b8H_VP4U4lk_vQq99kjZEtT.W9FXXYly2155gKPbGcDK0g6RAxnA8ioWUDGOLKJDcDdCsKHqqcOUz9L7a8Sui9b.PBxvNGkLb.WhaaV8tCrfAty4S6mEuPjkMAHrO.r2RF9_EfXWdPo7q9htRB1liA.5khBR4t5SHVLbSwbpYe71CpFrpSZsDRj_3WRetMfo_uGziHor9WRIDVZrDoVgVLxfHoQGljOsEdDZTgjD8MjAsaAv2ZWd.JsmAr0_6KdfCDepPQst97RBXIqaHTcZgCHE4s4yYFQBuINpTdhXKvjMqb1AURBz6Na_wz6QuWNkYpeLKWrtfBaVtv4a2ypYPZSwR662O9glV.ilTqVpSo8M1LQ2T_g_MRAcs6Gmatt5RIbGe8hwSG4pUH4R22YRYqrL0WlL6pexaB75pIDB1Dy_7RbEq2CRastKG.6QjURMrqGyKWQICzd0N3sUdYBYQkUeQzUc4SqyCdu_gjzMs2a8g3snMnEVmQn5Xvl.hRhFW0o1Qq1MGo7.6tPG3l3BSOYvbghnc8HvFD6liIpTzr8Vpl90NdI0OGC8_P5j0niBwoG6uFE_YARfQOVoU._rIXQWEOitShqihcWSEbGqqLkc3kiCykqdd5QJ1THaC9iNmeT3SO622S4lhjq7MwgMwZyP1hXuXWkBM2ztGd6ad7V7FBBKgNK9OZZDsRGMFuXLeCHtk8xfTc0xJXh7oZYYeICjs.foCeks4E3c2sMVySLvGGQBp.Ujjn7AnYQI0DxJvJvc38wlgVYOKL1Lg0qu7c7aTHMQeLMOO_m9v39uoYAb5yobILxfcdeURs0ZXoXWwFIhXLGoWSK91KGmFucQio9KfockSMYY193Ueg4WR1Wy6JbH91gHDzZ4XbGGlqJFHxS4nDy.rZgOzs0w7NwD3oLPLf0xL7n5Yy1VdEZw3blWqywoiMakmWgAJNJdbcySm0DZmVJTljHLDqQEthUtQIa4t5aAWrGnzpJMno6vRIsv1.jn7ZPD.APYTpzfBcow6_oGYu3mdU4CPYM2uc2urukloA4nbRlujrtrrzSzA8p63rbn7w87iPgUP3TFIwHr0dor8u4ZOhS67_oPhZYHsN2WR7CVYQGDXAgHeBWPfP1kqaL_CQLbfvxsorssRvGlX3N6VU0CsQ",mdrd: "pyDTkUDmEcjOoZR.2V24lf5coNRBeKj.YHqU8wePRbo-1740000155-1.2.1.1-uzYtTDOn_rnJm_VyQ5oHvmU5xVw1QHMscRirEEuXnkjzA2f9aw8Gqwm163xAREok97GToU8rwlYl61mtowaMIANHqGp.vetn0MjYy5iBHLwvlTecZBz.gvjW3_aPvbslDwY8NvD_YJyuFtvASKbRTRSGE1tbLVIOp82tQ1SJ8Uo5p90SERP6dnBBRitEnulSPo2OGG6D0h.BwUNUXDLA2v_a_Z83hp_Vp4VlrR4AHcc2ZVhK7TcgfgvYmXlKNZrkpRgDwkhz3p73UGAoslCffJCJ..VmwOMm8m2itdsS6CuF0nuvKjruAcwoCkhHExLukLYZDuwgFf9Q7CF96z2vZUDXTsMwG25HcoIzkBUGUnuZ.FA9fG0FNkCbMPZVNcyRlYOsCY0Wkt.A.KrP6U9ZD29y6EtXcN9WGB577GH10MV.Wh6RYkbLBtfhvgpu9iL9DfPT1DyHK_vdjj3ighrVkK1W6Syehw5am_k2vRGwdsi7GuTbnmI4Ym8GuHyCXFkWV3q9pBukbSn5zpBCj5DcfOXcDt8q62zm5WVIQx__mPv7KHmGuOB_noq8qXaRXK4BSFGD3M5c6Ao22s0EjOrBGpPYvQe2WUarP13WrRI0YeSrkvb96HyzwGfKbfNQY0K2ieh.9sVny6texEp1ERRJhRPV1BJWb9HZlJQVv3NDUxJckWcRW4J4vFn3.ZX9K5iTWTA9NTWl12Cx5ITBLvfuyOcwvnqJ_k7ASa5Vorxqh1.ZTtDIMQ5b4fVIsETbU6XZMbIxsXiV5iaeyyZMA6qJoj_ycE88FK2IexOgsyYy3olUEqbOlujtmXxzizkZF7w3bHCfXjfMWqgTw.VMNBytw5VqRNt2naJrsPtRv4pZmkHrP0pIueR0GULljg2CQKTfG2C9GpbNMMpjPFyjRVYDiIutgrp2a126iYoLacN2mPqbgQQi0CXwMVYJ2nocq5JqQfRxIqp4Z1uzwS2unzRN5gXU7kD3Gt3YHGGvqJDXPwjIcskUm.f.OsTfQ9ZuS_0B6ckxDumBhZ5eCw_DGmSH9JR8_E7xmGuhdjUl2kTg5nf9sethnW1qBG.hkktvjBzuucgcnPZYMI5Hq7WREnpcUsC.y_Ab5c3io5uXYNkQoAFL55MH5dgpWVkcv.FtYZ3hnGz7HReCQflii3LsGePLcWWFg9AEhf5sqA0S.K0oaC9_5MbKot.Nc1tOFw1rxVYB4gpZsju7LCT.jhiQBb.0HOb_KBlHzemZWSgMFpHJ2IrRck8mFpljBcAOKFHQhGm1hHXAEqSBiIU67_7LZJcsYPAp1Iqr5dWS9Undr6udqY15DJHsPpX_ymaCECbVXh1AWKJmdGNkxWBe4cIqdxYvsm4QBCFdbQE5qGsT8is7FjecBTTNV6MmDRgTHdcCmbiB9m4gjpXVZf8xi8YX8v8rOE_GmU0sE7o4vjPvE3xQPsInKmiuQPFgtcd1vIHyT2mv2IpFiUasmDX9_FeDK9Rx4VYyXeISrZKyIJdubMuqBnVZHwzoA6PVsGtbHA0TIiW6.w5kVrPkvPWHWEXknOQIRi2jHuBnnxkQowJ4EGIzbRHdpdqRml3JAUF_r_tPyuBrr1QFrKGb7F99O7jkugqWpumLfvFwk6rZ_gnffr2zvSIC4QLJxd46DeYbBYlxR0TYr6YmjvJyGxt3r4hx5vNJuLaqnEOdga5Awj4w_fm3RuFjOE0LKQx66luSbKS5X1R75CB824upTqIBfEPlLZVE7z1CbHPTlrUUJDNPHlQUiHt14aAy2r.0H5y_nhyxPp8dOyZI2Q_34mfspmaT9PNpDCg32A9vj9eRg31x2fd3_uHzfmIPkUAd0LZhKxW6mkUq_fXlMw39Vuja8NmJ.xfQqJzf2yReFIObIOslhFUVMJ2AXyp_mk12vrS2U0ATu0A7DPLVcnRnReV39pkAxlrAASVMAW2uFMmF0FWq2WS9rez_EfNaZY8fKXqMTS6SQjYi_RrQBMjcJ3r0XAh5QRLd.w"};var cpo = document.createElement('script');cpo.src = '/cdn-cgi/challenge-platform/h/g/orchestrate/chl_page/v1?ray=9149502a1aaa3e5f';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, "\/search\/advanced?__cf_chl_rt_tk=RwPiWTpF2j3eVODKYgCExtgA92pLA7SnRBPOPNdYiX0-1740000155-1.0.1.1-ESB..SHztQEuthRTPm.b8kwS39MTVSfTnSwrqMZ3eu0" + window._cf_chl_opt.cOgUHash);cpo.onload = function() {history.replaceState(null, null, ogU);}}document.getElementsByTagName('head')[0].appendChild(cpo);}());</script></body></html>