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: '9135922d0d0a401a',cH: '.HvjP1WrKGBbPEyzjH7f9f2GuwxcpgySGaWp2E62M9Q-1739793143-1.2.1.1-C1piMTK13xNG1qFrrvfJ1OQ.Fca0YzGxW9eofUsJaWzNH.v8FM9ux8rhv7Ppqtan',cUPMDTk: "\/gigascience?__cf_chl_tk=OMtKdaki3EGEzb6LIdZ3.JnI2OKuZf6j9lTU7xSRAgQ-1739793143-1.0.1.1-MsO82aT13xEEBER1x3F8MtPdLhX1AIzUaM_wzcB.4D4",cFPWv: 'g',cITimeS: '1739793143',cTTimeMs: '1000',cMTimeMs: '390000',cTplC: 0,cTplV: 5,cTplB: 'cf',cK: "",fa: "\/gigascience?__cf_chl_f_tk=OMtKdaki3EGEzb6LIdZ3.JnI2OKuZf6j9lTU7xSRAgQ-1739793143-1.0.1.1-MsO82aT13xEEBER1x3F8MtPdLhX1AIzUaM_wzcB.4D4",md: "BOK8TBjKBswRWe676u685biuX6IiBaD.gUBgFEUADU4-1739793143-1.2.1.1-UdIt_5.qssF7y1xZRuGElCZTMjwAfNxx2WkQr.Rw7rXdNfcZ8oj3Ir272eBs9cmBHTOmzM0VYsA_ZL27Wnruk1ckNa1rPGegJVdIMeb81c.i43DNfVUAZQB_a6T2V9.pwZIp6Woad25WFyz7yV5dTr.gYoowk1ETwrni7eGpucOoL5wSL8cQcvnTagPjL6QxV8UhsWjUOiheS52qvlSJisoTccfmj9XlM0gDeZCISdplWqtrBMgkphWFxcJPPiopzqu9kiqeeH075SgyaMJ.3ZSNlR_sbG7HcWOlUyYQf5DfkEoEvzdqONo0cpaiz0zn8nhsAMKERi6ieaKuz66vXTceAT2V_HasP__fsgZDssMKBUf6JZxulpiaAX8cfWAbEFuDOkViXcqBun36AsfEIM8Ec_s2unHKDuGrqyJ7__FZO96ySqY3dqU0.oPRGhS8962DPcIqhYOaf80gf0yotIt91mXccPvXMX_29N7ufteirTRBDmhRK12Jl95EpHn5MqmZGQZhoUoaJNa6N4fnDSNrN6e5ljfsRtDPBjeh6Ky6Nmmoe_Xt5AyFvaX79ny93bXyFDK9IG.yJfxoOditE_i2nHX1rNc0W1_Ld1yH58sxTZgpfd1kQHNxvZVRs.woIA3hAyi_QF.L4o.zhZsqOL4PBppwG3_dQiNInJuqi9uLy19qSO0a0x_azc9yStOiLAHVwdu7997eta2aFA0BkLHC62xTn03s6vZ2IUYV1qSwjcY3BgC6epm.UPx.CznA39M93T4Xb4y4vaJGn39eGXchpZ4VdChTMuNOVMIt1Vpk5IGVoelWbNDIsTe5lX0i5dHC.uE_HetQkuDEp7DuA2jWrCM0FbNIWJUZxyslpKHoe6AlTC1aCvwGmwrhzp421yB0e1TxBIRvwv8668ydfV2U1s2z09_kNwsaFpf2iI3L4lo7pOTcGtTB.X98lXVpvAv2.0Y0sOsDCEAUVTZWMJO35QO46ZReI9_ShFdYKOVvmmQR1vpiyyG5MIZKymAlrTmpR8XhEzwttVvrJfxv_jQLfD7fhpwM22_ATDaOYmvuPkdz161I1C_WwR4NdcTHKxb5d_iI4qN7LGSTh31zjtxihJWcgbivkthI.deVMxOCRQtzvssxNRjFf624Sdr944ZL2cX2SAjvpWJ0bOVihbQUGWRYcmeJLQkqxAdfzqel7sIwq2xbPd9z_pQnozYPeE6ZtT7Q9yur5eB_QfUDb70P6ADUT5RBUWEcgvF4wMF5WdDsZEqjmZR81vVk3Gq6gPyWPBBQVo.wWJYcMUcnvf4o1Vvsbx386SA.P0pPxZdjtzaOIEikJiLcAMi3xjFEep.8NISnQFdsKMcrrVskf9oRlFn7QkpZ6HIctdkJYMYB0gFDjQWGrllLlFFqMiTbVrt9Yz09BzYHLfrxnB_5OraYNf_G3RSrVqvHz56uDhs0c0RnkE3y4kgwLMBjH3rmKrKhN.p2Kom_aac.CYQKcnLoM3M3a5FIyd.aZ8VqN7tVlLfPqgxECizE2sODcgM1F1yrrF.LK1BBENCAyQ8GIu1809oE1lX8Q.Qm4rIiDDrkBO6D.xvFoxSHxUZkLq14W_QIqCKqq57VFfobomAcaOaXUWbrfNBS06umO.mN8ix7XyjV1Oy48aLNiT7UGJrSrKOnog16R8S_18evkq8lQHVj4iFMhLpgrajDlQPsuaGO5xX4W4ySj.OZrQvHB5.M5MA591eB3HCfmwZGsdDMUnGdgiVRCYae4oEt1pFf3l55ruqRClewiXUZ5aBS7qqcnTbLB4GeQiaaVPRhgM_E_Q3y8fCa.IEhR0ZPbe.WjPoYFptlLd.v4ojrSqdZjsobN6G8u_6MJosOM5La2AglIAWCiG8kBYbZh6joPaH.akRR6n025w9GNL3hkcMC1vZjxUc94eWoW_1jgEjw1A0njDS9Jzq1ZELqIH2WyJ9PJ8FCeqo7bjkNALnW5VfMA87396Fyev3I3FnE.IxJEIEbm.n2APoA41HR.WWbjtSo75wXxENUFV8NRHVvKIW5hVIp4USCpqhKuJhxlvOxKJkYnD5QbuvQscsbm7QipgK12dpt6WDd5xb_CBehoXgYoH2M7clDgQ41N5Od2eTiLYlrmO0Q_3KiBWQQeE63C1FdIUMvuKzni8Ham.OAqup_tVqzWNONYS6Eh4dFGC8vv5eTUsMq4lbwF3.pbSglLilUonLNwM.0U_gGdICrwqeF_yXkyeu2al5_h9R_mp2Zyn1zAa24nIEP8YCGou3oiS7S_QTFB2eWNlUJH07mfqg8ghyCuYPtjrw39dnBTNF0tV4Fmp1gOwwKcisdQx8AJMvMd.xjBcqyFGy78OpNStdYTECe6vsorn8aqzW9Chm9mUF2wq2kXoRCqvMst3dlF1PexKeB9Ra8Dru4oJeJkR566y9sVijf6_o.OFLeLIYUjqh.ebuIyvKm5a2V1zUfgOS0QxGnKRLyK8n6.WRasZ70qWyIXQUmbrR.enq6Zwuwt8s2En2PI3hyaqfMBhXfMwRNPfg",mdrd: "5zz7JwQnEjJuXhKvk7ZA5mRdHSSGFGwW2h1nO1_wBAQ-1739793143-1.2.1.1-iENg6rHzRnpIJ3dcxGyEXjz8ew3Pb_g1ugLb4pDGd6WCjb9sz9tQf7OKbGLJkSDO674Osi9aus5gbCwQ1XXZDJqyYTMlNg4FRSCXStwSXbRDWX2yDe0ib_SroEZZVRIoWcV8AbaLFRnK0mN3cwZJYrP6MhOZfudkre_r9sl0sFXOls7OObIcOlA.RPpqI3CSXBoaSkMfZ6JsBpAC6qEKhmL1HmdX6qMKOrV.YbALe71V6xkoeRALePGK5q_kVavTvbbM2zthRHn9820oJlsYeaGQMYdRA8OdrTBgo6GXnwHZF48qHQKx8PKtjoLVQqrpnaNLiI1tiliJZym3r6UmWc9ztebMiaIFj07sR6I9SgCCog_p5Gew.eJ8X.FSBOi46d_PsNbAT1ZzECYoOE2IJO1AzRRfzHyjv1cG2NaygT8EAaXqku0_G7_CEFTMTMCjRbfA5O2t.k9XyWYqiEK6vJsPrPxRgon3g1XIdPaGuBz.tlwmHqx6wdwxYPVi4WSawtdd_iV_jwRh1UrBMdEAEW22aW9UMPtHlvYGKhWSKuBVCIP8BO.UnLrFCv1HR53LpDR1gO4CUYN8snuzGtXFDK.xxG5DVkSljPioOAlP9Jl6qKvyF6dUb7WCQCEMavcZ7gDQvli1TYT84tUaCw9Kd1hz1JKRMh7b0KbpwHXuxFMzpvK2Ll.lQ5Qr_Rm5uoOO6VGLIrwyi3bAq9SBufKZUbI3x9DhwoLmZyyq52I7KUiHeS_DLcp6yN.O8ZTF6bAV6dYNVIX82ilgQVjZkqj2NAIIpzx2_.GzssNCWZwLAQ4.4sLk6W8h3RRaTjAVSu.IpN0rbW3LEWZb7q0hFmq.jCcDW4W4aeLYnVcg03YvE9bLjmFBtmq.lwaWNHhf3klCDvNK8GWo0_OK_xdATT3V6edXMmu0w9.lXLpzpf8csXZkSamQV67JX8Zn11cMNXr..yiXVJuBZ5irXHFBpxLjqmv5hTZfCVEUYxApYID46XIHXcpeZ_vlzvVIoPYoDQALopi.tKglmNA5zUgcTZ_fmF8.Tg.I_lv1o1I9luUHqZcxmeZxoe6JU7tpCpwJ_MfU8m33Yqok_1__SA.uI.8KelDwjLxVetJMTjwCF8obJ3z_SddHjB7l3QXkVrtvuViAjxEk2uUSlO5xUpdE07ys_B7fk_Miyywib4ywi5iQa154kInL2Zg.tSTBpCYs0DjcebxhxrwR3aWd2K68vzXcOa4A9.a0Rmypwx6kWepSlBzsDj5tCYkBuPCKqiIOi_Yd6TvsMa43CS425Afv_oZJdsikWbrL7uwJsVanZNAZhm2_fNBoJPZZMniF248mpzYslzvTQVXQS.YQX8YtIX9H8D.Slaauvou5D6QinHlCIjpQzQhuxzkacrS_28tjcC525jbcuXXcS0f0fpVMM3UQ8TB0kP93tiDqAHxt.80VnREz3RUy2zvpICSLsGxAgkp52LbhncdhT1fGbffNcj61_wfOCmq77mpAdYEpIsiEZcuv2Gd8.rtwCInk22jaFfr.uh6GhUd8ONe0bDLROn7KRwp1lAO6Aw4QchgWi9_DKTxvvLvfSR0.E0t3tHQid_d8YEtxvnBx_X1YqAiSW9YuncSe22gOuzp6RUbjH5kszJcoi9eFb2n9wH5OUug7oyjMeQeDexWXB4udVZQfxktTvFB7GEzUkEhK.yR2n7kxWYOjvbScXSE6J9t1mVnwhMdVeBtEQ15xcWOJsfVS_ILraK8i.SbuDz7zEkh0uGw73u82ilV22JiduHlP7xfJ64pVXwB3dnZgd9jc7hedjDPfHtKryl_SV46ocIcevuXtD28HmTLq99yaLshHymSs93a6FU_I4FbWj6LbGhvunVG1jDX9NoQUgNaOs3VKiHA1XEWWpHy8bADTBO3a1q7K9CJBCNbRkmrYpVValWJEy1axZ.osieJfu_oq0SrkTbXB8SXjoHxX7ZLYBJDSQKE0yDop"};var cpo = document.createElement('script');cpo.src = '/cdn-cgi/challenge-platform/h/g/orchestrate/chl_page/v1?ray=9135922d0d0a401a';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, "\/gigascience?__cf_chl_rt_tk=OMtKdaki3EGEzb6LIdZ3.JnI2OKuZf6j9lTU7xSRAgQ-1739793143-1.0.1.1-MsO82aT13xEEBER1x3F8MtPdLhX1AIzUaM_wzcB.4D4" + window._cf_chl_opt.cOgUHash);cpo.onload = function() {history.replaceState(null, null, ogU);}}document.getElementsByTagName('head')[0].appendChild(cpo);}());</script></body></html>