CINXE.COM

Just a moment...

<!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: '913b3e68afb891ba',cH: '_N.21bFGFmvSomIwAbFajcgbAQ9bVwn4Urr2Wdcj_Bo-1739852627-1.2.1.1-dRXoNiHrunieSh47T.1TAAiWKOrjXQ.qOjDw.xLBGzRvxSv677VgcnzrXdzzGJhX',cUPMDTk: "\/sysbio\/article\/61\/1\/150\/1678583?__cf_chl_tk=xRP1xXVFNZyUKrhzinRc.ecoG1TkeeQfYxBq4r_1.es-1739852627-1.0.1.1-cXQE.sV.3NdUwAdRXAQwIqqwrQddztIWxsHf3YwlF9E",cFPWv: 'g',cITimeS: '1739852627',cTTimeMs: '1000',cMTimeMs: '390000',cTplC: 0,cTplV: 5,cTplB: 'cf',cK: "",fa: "\/sysbio\/article\/61\/1\/150\/1678583?__cf_chl_f_tk=xRP1xXVFNZyUKrhzinRc.ecoG1TkeeQfYxBq4r_1.es-1739852627-1.0.1.1-cXQE.sV.3NdUwAdRXAQwIqqwrQddztIWxsHf3YwlF9E",md: "IUM3u9xCXTVAAiHHTU8EXoNWiiPtJ1RnTVCnmH9RVuQ-1739852627-1.2.1.1-UJZdCiaucU0ZKIDbz.Zp6gHFo8HZyA9WiBWPnXEq42LHTOcVB68MuZWE8DKspA5Qp9sd3AiJjMOdx_uic2kGK8CaWNhYhM8xxE_DKCg2Kio_BzYrcDed2z.l85nUdwzC1qe_J2hcJ.n5_Fne.Xt8fu3.Jl0pljejCCs5xrxd9sgme9xFgvZLLgYn5fk8VdKAubDCrKWWBuuuGDBEB4tjS4ywvQPOEEcZB3HswVv5UM_pqd7MbQZrX.Z0YBbMVvsL3MVvKQIIZutS3bO1Nk9j2QT633sSLwuWhPhIuHXSDZz6Qv68uYUi7wrRujPPs_.bFPvjxWRBlSV.tPNUq7GsYlmQ0hjDH6zJqrUMpRgQSpVGVOOFmHvnl1IV2R9U01APQs9FQj4_NaSG2.sd3dgSnVf01CXDN4HBUiVFW66TVnpnAfrMHGAXGPuszp2_PYKkRR8QKO878LjT28pMaus57SuYScIzmVQnW6xAS3zi88FfpCoV.hVxrxs6zJ2kXD3LP3NmuM4yLjXspqiPuXnWSV76Jc4Qs2nGats9jDZli9I1i.Jidak8P.8hutOSQZuGeqyGLcI6SGUW0u_m9167blmAn_QT_zH8LePLD5T6prgSoz2qoFeTgsGb7XN_ns95LCmslcxUeATHbBCgxfJNfQps2JJNNOudDOf_8v.oix7cNTNcc.fBjWjITTgtS7nqhxmK_WpWm31BRlWuR8z4m2g0gsg_IDtD9UIs6MHqAXcFa.frU9plYJOBN2zPaQ_JHgO3J6pdjkshoBoZ3KGffOZ_EuAFY5plQ6eDPq1EyfVoebuvpLOllpAKM.b_jEJdvHp4j9ZTyGJcYOtxkH1u8xV4kWzsO_zfA6_RJs4mreSs8plo.JBU8ofz4eGsTPc3ZcdxSH_F_VqzgnMU9pHEL3j4ePNZXEe87_ipR73653Iqs34gaSi9Lng2m0aRTpPHMcW4fjcuNRzTohl6ROGMLqMslLMJl.cDHaD89Xx.t3EVoYG2QRiRRSkj7oXDtFhAo9DXL6iYsSyqsVvRz_wRyK1yozCE.I2pQGV_PS83S6htaZoZYhYmJHpcaKqsJVW4_MPlOJY.U750nx75pNo37IbntWG9Ny4KnmDFEtx0NGnOLv_zYATY.mjuytzL0dhp3T.Mz9jBPWvxkd6_Mm.LApDPkooAhsqpmvPQc.u005V3Cuacx57O5jG7zQxxlVLSW3ujKZ9.x9xwrXtUCuiX..cCgA3ivETyZMICeDzwo55FywAh40Y2AAQyJQo5YRf3uepzJLirPiVBNKtsTnulacsgUZqyQG4di_i.SANyK3SWaeUpsN5MBHMprf1lf72xvuckP6LvJ5qpTweCBrokNJxbHp8DRmhc.sLiYHUwfTNjZqcUWNSNVNa4Ca6hE9s12is5OHjaMFkt0eSgx1Mx04OF3diEyotkz5XVW1ngnh76VBuDwjOju6_kV4WagqkFFRgL5mDYyiRFJEBXrtF84pEdYGoSLhBpmr81Y2Z2xOeJeWK7UFnQH0R7YwHGGz3jV4slv7Egk1n3OVo3SlCzqwUQgzkRZP7npKiZ_sRX5N.2Ctg1NJEwbBuXdJLfZXZli6rgmxvHFRrIBXeSEIJeW77M0X74fGSEZQK8ojIlN4NgpbTPWItvMpBFEVwNVECMiPmICp6ypZyb67DRKPrqKAW56o5.BLHwovycf2cDDJ_VzbZOgRz4bWgQiG2M6QNfcbGoMwlgn_n5g3BMs.09FYu6T1u1q4tUvInXNlF90WR7.ReGK87yLZfD5aU5QrOk2BcTxeQ_9LU32GAP4iRwfOIKjN2jm0QGi48cFhMR2Wl3iMoWhtvBl742fIzi94Bx8h4uVFcfUP5QIM5OWdFRxlHd6oYRAPT82B1Yt3zV5APOU1z68N2Hu8KkeB7k0dRG0Lni3kn_00BbgTSxTvM.iQw3jL8i65aSpOIQe77JHOzzzT2hMkpuk7BcDvC4b2Z1ihDe78cuqgh4_XLWEhLTMLFUoOE0LQHmIafpsPZXdpFXwASVkihRq0E6jCiggUYATeWtJfY111JG4OvSJ3e18aWYr7CgQU6bnKwQKElJ3yx1JqogtL5iLGnzJjbun7U6bg_KkrSboCYPwPwIpaEjonpqyB77HLMfpmhFcSFVj6UHheJiUFAGwIEtFOOeynat_X8iaB.7w3sNBOZTJxJOgMYd7loYkTfDlXHKC51J69GkEQ92sGNVhaS4k1gTsKI2cuUIT9I_vaZ4jbnsVphBQD0siy3h0MMnrK_sbg3h6UMbZEQRjjRtP0Wjr4F5iqcjI7xPzeZDuJ8GBeRg.sd4FwjlhbU9NmTjOgT5_rLHQ31yXo2mGJZuwsXEPQkRh5d_NQQ0RLtfUleGiEjURw4XnIzA6._TtbTQIevxHwVDniVj_uHUvvqlLSqh7ORc0U4Tn9n_hvPdTN8UG0Uyg4DnuVDapkRtZUEgm2f_FCZtOd0CcmSFJcFG5Skwo14Znvd27kAQHQ29ZZnXwoGtCeEsVSn41ptcTw0FVE7kpIP4FbOdhroMYE0rpamvcBV8zZ8gcf1g14ennx6GdiNdIwbrVxSNtbJrRc3TR_bjyQOhnrU",mdrd: "WQq59qCQ23Tr6FdJ0h6hvSjOcUgm2DvlAPD8nvErung-1739852627-1.2.1.1-VRxpHU.k6iIbq9pbuKW4xrlaU7LCGj9A44dUMzoV23h_lb.GrohNPcB67WsE7VL27nTUecapusreY1BId9MP76dac52b8AW09K0ze5Ue55Oci7ICtD7s4dO.9kNJc6Hoa6UXob3VvO8zZkANgWnYTKblcfKQVNIQPJ0ddaUN1C9tQeG75l5Mlvpb2I31vFOv4tMUs62vnvT0w.WsIWlCkmlm0CnlWgRgPchiWPyQRWFxrN.b8xrFg8sbcagPfQmGS0ZMlO1ejn1nsNccGzAa_KTZCtsfHKOeNYIQdor4yAxnrfYPgfWYXtOb4Jb_0bVaBcPudf1tovVu3LQGP4iCEtv3xEdptMfx9umH07fo8JgwNt9labBCmw85jPFvwRwfleDZAhX9OsxITmKVE_itNSRiyeAV21H4dE.85upXFxHewne.LMII5lZ7tCvHTopyY4Ikwm2kIbcBCscYr6b39g1GQB83o.UIPxcR.INz.6hSwTVVMD6ffn9vJPphNcBJfydlg6NiEw8L319i9xwBsUeoOxokAw9QAwP.iooq6NbBBJkKM4Adi82HgKcKQVb2IjUWtZ1MDW11PC5UBhSYeYvKuLopLGkgxOlNpAeWpQXbkfy6VaYM40jEFVpNOduQJa9rfZs2niSSlQ32UobYbTkZA6gPZufZKqwSt2CeI0vixeQ4LMJa5hoZOma3_X2yv1.zzQsp6xTydJY_VRXD_g4Hsgu6b53NKQIMCFo6agqDVMinZxvdOhFrwi2Ff6V.wf_1aJbDD6nnji6pucHGzqC0aVpZ48I9hEZvykGJGfyPplmZRf1W60s078JDiC0t0X0zwUNyeYPs2Mt0TnypJtg_vaioI7fLUiQuWgMbNzgl4_UuxtDfUKHHjp5C4Wns.u6hF3G896quK_fkqvNc3Tkb99Xp6I2S0PYKbPoUUm7yY3SUw0LJUbiBWY6k5wbNa775AvbLMQMLpaT5ip94Rmot.fz9vvAkofeKWUduFNNr5fm2xC9rTD5F_dT28kttAIiLWkDN8BD.yUs_VBnFv_zGlQvV.wfQJvauLVyRjeX6LQh.qGIUIt4r7U29kjeQa_nqjL2jpJMlW42OaF6y.oTZPi.Wk1QGxIKcVCb5AjmA.CsK5gaeI7ZlpSCkOpv3_0cikWjMfzokPFOe3mIwP4._gk6nzSHMhyMvQdXL35ng.NPw2gFID1eUYizIqlA_Iq4_OYz91e2K55CpCgKvrTzaeHspUUYS2_ItqlVCFeEZ41h5jo0ZSEcDFfNanjfGqV.9vfV5R4272We0tvxpGYFXqZ8tzSvZwXYZ8eYD2rs_JueGx0XlH6q.Y1hWoQM3zMXzDZsKvJwSnHAYzdvZA_AdxEJmQZVL.gZv04CQHQCymXOSctf_p9HJBP4mYZyMig1ERTXOBIZ71ou6VUXrIHbDfVIpuOqgNF1xFYoB5knUVwiy9FPBw7j4DP.nnXN2OwXpKR.UgM2dBYfmFD7dSeQPaRFyjwuOLkg48LeDQAgDzskvCWX9EdaRqmaCnfkImTOpUPJxTEi6GrocEIb8UTtwDbEo1KZSiCyiQK.pOJvydeC7S2WqU4mNaIPdRM0F1sB89zqdhgud4JiaEWmefc4xGR9Gu2g591PiXnkshifDIAkHVUtEjvIVJJGNKHx4utsTA_5cLni68hxy_CC8vKRMkjVVl9NqcJOBOtupMpRHejoUUgkL6iOeu3hjRgMuBH.ISz1wpVNk8MR_kl6Pwru2iCMO90mjNGvFLujcUNQsaoIS_OonN2SlL.BWoZIuzOnvprHd8U.GJK0iJ276.7lg.UgqKzloksR0iGZrCGKF4_i4VCtg_C7BRyrcJ5tjToysBvMEvN7bV8SnHii0CU1zFuNvh23fEi1g7o9n3Lj1O9_DrXD99gRbMT6TaRNVO.0wIXcH4_tJ2Ci5xATNK1qpfH4GwJxKSiJqzY_W4xET0BCre9Yk4QsiDvDKxQVNRHcnRIQe6idHZDj919K.rQ"};var cpo = document.createElement('script');cpo.src = '/cdn-cgi/challenge-platform/h/g/orchestrate/chl_page/v1?ray=913b3e68afb891ba';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, "\/sysbio\/article\/61\/1\/150\/1678583?__cf_chl_rt_tk=xRP1xXVFNZyUKrhzinRc.ecoG1TkeeQfYxBq4r_1.es-1739852627-1.0.1.1-cXQE.sV.3NdUwAdRXAQwIqqwrQddztIWxsHf3YwlF9E" + window._cf_chl_opt.cOgUHash);cpo.onload = function() {history.replaceState(null, null, ogU);}}document.getElementsByTagName('head')[0].appendChild(cpo);}());</script></body></html>