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: '913547bd0f7ffd67',cH: 'IxP_4kfc2gjFUQBTz_B0UROYoWa4dUighTPWqOpuxr8-1739790094-1.2.1.1-YNoSlDDYPbyt.jiJhRn_ZrQlQGoLIzDrLLa6WTqFw5Ui6wBTFO3hxua8tO6xODG3',cUPMDTk: "\/gsajournals\/pages\/plants-genetics-and-genomics?__cf_chl_tk=QxR.XGuZpXPWrJFP1eb0lwZAsTkZs4Cfei055kcbIUE-1739790094-1.0.1.1-4LvRiCjesxsBJ_Etgvbk7_xPj9vLnNfGcZTfdb3OWaQ",cFPWv: 'b',cITimeS: '1739790094',cTTimeMs: '1000',cMTimeMs: '390000',cTplC: 0,cTplV: 5,cTplB: 'cf',cK: "",fa: "\/gsajournals\/pages\/plants-genetics-and-genomics?__cf_chl_f_tk=QxR.XGuZpXPWrJFP1eb0lwZAsTkZs4Cfei055kcbIUE-1739790094-1.0.1.1-4LvRiCjesxsBJ_Etgvbk7_xPj9vLnNfGcZTfdb3OWaQ",md: "c5lG3TXNT5b3xjYgLtFaM90JrZh.grSk_IMT6EKVpTo-1739790094-1.2.1.1-BsNDbXpPujQVYWp2R0PFxxcIah8O6cK2FihLCSUISW3PTMu0NTAXFKBBmqgK6f6Qt9R06BNVS5vKe5UfuoROAwy19YdO1jVzKMyDAauzCYFxw.cphYhDRMcxIjhzAgKu25RhZcE7xLm3fV3m_4lQRA8rrkNWXnleUsXTIoZf.zG_sCgfxdYZLexj3hPmz_TjA4PJ5.HR3Cxe3lAXt8H3Q95OtKA2jcEva3Mda2zlKFY715KDpFW.1eRt1P05oZvnDEpyeaj6RF9yYwu1BUqMDFbrgishWkny9XrFRnil.DOrRwkzfwlWpwl_rhyJW0Cb.jQYi9bZUq0gWuFEpodR8u3RwTCuap0mB.LIvRbAlzJ2aSQDu8dA4P6MgnV.WynCBNfPKWRxYW8ZFn01W4P7TfwPMiXEIBAD2wAdFCuWggQqwInoO_PoMyxsFGgqGSgWu5TWw2S6CqnAFzC8WgFWpMZXDDc52dLXPB3zbu6Utn3ldfh_2.UwDvjbaF3_w6Tm6LdyDj8KtYWILEV.JfHVR0OF0xaX7HcQMglXp0k7qjLj16euQxaSzWhvQTfoU.jOPNPvOmse4cAMO_x8f96g0s0SxqtAb9iqds4SDh4PHTruzc1LOstCrrJbHw8EELE_dCcH5pJzaP3x0Zc3HDyqLMGEujktVvoNUyyMPuwGzucubhhm.cG4lsryrcNAnNVXeAE0eaFsMYggIQZg3wrSX2H2Y50zgm7ypUowFDEHy35HGiBdUkflqPGA_3mGo3E2F.ajM4Wu8TL9ZcXGxEZZ4vBC2xJBxUsm6jS5jWj.RuFbPS7gz1Vw1J_86VQ3lXMZTLU9vaCUoOzUNMcNsbhbgykvKVm4dxScB.EdzUnpEUgX3MpJyEnwT2cOhJ574p4qnHWKpjMRrbiFYllKI6ypcKZ07_WcYBWBOZWb6E0qowjlwDRuBmlnmzPG8EsUm3IaBcBcHSHOp.kEObAIJYxrSElx8NmBfFeCrBbaPY5dsbvsS9Q6ibvYC_aCZC_MD417vvqmae9KlSJ4rMLLc6xf13pf6Bcf_RQwk1JPc0ubqts5pnodcnuTXAYCyJdX7WqlcfYNcN0JB_zb6t7HCNYgkVXdawzRTvIwuP5ijRo4e.cpr0r1dvi2jY12oa6jTC46I_HfiauMnlBhKU1fAQuAAcdddAUJoibr795Ee7zwedO.WNZEUIU01YmSOKOlGPjuxmmSzo8_bUETM7RVqD4LDXxHcuZnPVnzEfrHsziaXrWopDF02F7Y4RSmS6zT3OEXbBYEh._j9pCCugfa9rYqO_1K9RiHKifbT2uE_kVoUoCWXyIMtc7Ld2WSxYi.mmqNSDRcHkPFNLGZM9KzadT1EHlLhY5iXVAhxOD2J_2O6XOE0t4fmsgwVtw8_I8vAKcTe3S6M9pzY.PbPhFO1BECi8hVvo.O_it8rHSeKDxUZhbedYkxci47Ob24ve_Pk3QGmfGV6pAKFBRcVxa__91ecUNH9673xB9ZihLoc6ef04Cct9w4shTk3Bz6h58Qux.e5YBlA_tKvor9tiemk4CNYaaijH4b.NGNM4yCLEjjohhy68CPOG8Y8kFPBRAmOvDRAUjjI0jDtbcdSwGBGFbNTUE1ymKJn38EPmyOMZRseHqZab_2sqcYlCymVFivWVnKZm83Mz6huxkTGuGfpS17p6PEN1PLimgXwXLy93guX2gHoUwLWB6uBKZzcqMfJYwyKS62aLQvooapAxm.NNIEdOVBLdeLvz7VjE27mCxgVJmUiNpH0dP9PBRyHUJcMTLZkjuU_QNmmoZyoWXXbWc66Oo.UV8zXTbzZh6pwKXRRqrAOFk9m.Mvl28PuLEaIxXLUs4lv9asUcPD6an9EPOQlmmWEfjlMAmIwHHuz9Oy1QyORVvIcX258MLalCc1KP7.McJO8C56PPnTMnBodNuvUhaLm0yxw35t5IGN0stlcOkyOgp2lKhkXHKAi_l5wNuPNfoLZIBRPpuJ2cV0ebK86Q_O_ETT3cehceza8_YNph1_fI4V48OMrJm9ZdrOFoRHN34q.nLLCp6n0Q3HsOliImc5a_ZlMTucKGifaj88qi3PHWoOawdOW3CqViqYS_d5vb9ybmxGOK43MVjy_HyZ1xcQugyOYeMLQEYrNOLwdMpFpzhrb6s6s_QJG7mRLtX1YYc4x2hDyCvLOvihSz7tWj0F8IMF8MPe0SBPam1Ii7lVvtErivDbcBNekEJ5uUxlRasFMz0cDteg33NNcoCTqVfaLXRHuDwSRv9bmPfLKb93fO7QA6aI_8FsGzz9ID.lHILX__V_F59wLacPxT9Oy6L.LFxt0_im7ScA0F_fEAWe5Imi2y5zpqevVgAXJgcaqasjN8wnI0w0ngI6lIVtBJzO3rQTn3gjrFrZm3i0L3_Q8JDpj4aFmzyxieeoKk3iYcnDDxv6_TqDIut1Fh2vxtFuE571Y9lLRDYas6srM6sXm3SJUquRtfV3a5pK.aBDVKNzIjL0mOhF95TGH78zrjdCMGHPcUqqHljziwxmQkgVwRqiXVJ.1Zh_gbJvrdxXIeGAFGI0H5JXcQHFTvWV16SkyRYi7DM5gvFTULCHfceDiVjH4jVdGLznEEwi_B2wQM6AerL.QJzHO5ZE72WcsA",mdrd: ".rh0PsiEq5bj6m82NpK2v0gwVvXs4q1pwQnhSOkpx1Q-1739790094-1.2.1.1-VTwT2MGT04dtbx8BUytO1xUFK8fUDOyUp1xuaj7HuhzQlI91GL.ATs5HhpwYrN4P9RyXNgyE2Jp28yfeJJR54gDkti72USZDtvFIC2JYXeBqtF1ZITCHGgvnucBwJepC13NLX58sU04fUqNdd0lkkg06zN2T3s3krSez4ZoaYupqqrvxV3heyKdL80iXrg3osFOqHNVyT9WOa1x3FcOodcIf_1q6B0Cs9P6CJxsCdKeCcAOeNacNbhAHLVWtdwbG_CqN0iGRQaYt4I8FGm6k0t9vV.jLHMqU.cuRR6tB_MbhXKStsWWACeSbf7GQKGdu4VUck1OfdJjce9i.DYxAAOCO8yxuoExMECewrb_cLsPa.qhog5rMYen5lw00erd9HSMjLbnqbyMl6JaaKtCwQD72Gq6dgP4zUYgN6rMxzyhFnzCJ6jmz4OQTrTu7zWQoDZze5Kja6wTbSSPb5ucx2LkTwREvetnh1etPYly_0Ii4tDPkuusz65IWNOlmGyXicx0Q30Qt7x5yzPi7thlhNWUOHnzG.ddAAmloSvCaxli4wbyRFhbK9xLI_J1OW_pjAdX7FQvtczHeovbN4pd95BQhtAujVyOdqAlQoCO7gTqbKFeHWWL4ux4iejlMJpXNMkjhsO8MXxzLRZAj1T52AALF6qRb3WKqm21hMP.2Oyw9K41R6C_n2VAB6abDffGZESKKkQRKrFYRC34fyjz3LGpyoZHSKb2UE8ymyqDem1EkZL6t2OwtIbkLpXqxcLOFt9JhEMrqZ3M_qBp81bd1wGZScsNOQ3AvGkqB1pQVFFmgZgs3mqt1_O0chapAukzXCVRr6lFPJPoIw80TFn4wq00znrnGBXc3z5n8v0741tjmLuUkZJXOONEiTvvGRVZWvifyyIzVYqC0x8O9rBlHp02LRw8kI3Mlnb6VVgygYswZPuhrDntiTkSEkj.2HPYdBBvLZ5bBxyFdLhJuvMA5xkHnzn_BBzV_2jfHE3BmioXzTlv3iEMsjhmWTC5WsdYDRIYIS.5Gi1On.jurGTEeTQU0G6kR108pr9k86FaQdNHRNz1tBgYLwg5kJL4hBOw7CQthZ5pv5wcpzbdeoZzlpRTErrTwkvNwaAscqvNfUtt2y5rCvQHS3DqMPkdknW4J0ypZ4SPmqM_NCjvsNrpj.qoHZe0U5WK5dfcXMQp.JVt.Jww9E_S1d653Mi8boN.0ELyZpphs8Nsf_xCKQIc_R7dGkD22x10SKGPT4E7oscRoSVqcuspNYLu_GqoqkDFNd6uzDkJWzZd.jEbuY3mCOm.kosToc5gmPvoUXiNCdylPvP7xydDfQ81rBReYnGArm.ZyGk5Df.6xpRcwgHByDRZqAR0aSt72iN71N1GSclwrGEC0.5lLRIZoXDtnSop0fMCM.6phCOijXgqHeQYQrQ3CWOtDx0trvuc7BiL6VMm8v7DUCCSpFbKzaBIV3Up6n7yWflDluVQSKvDGc9SsBU4Kr_Iai0xs67tCPwskD5eeKSK0Eha8XxTMtGYgf0H9uKTYKLHYUMOsz92EBixJzDOHXPAMs6QrSSgYCimcdiddkQlm1TFEzAsCn5cs0x0q8ZjO9iJd2GyrtxecjC5hPmuoP8kZrSAJ.e6LnbR3GhYsUduyFjKgRsTf1xDixC_HM0_Orh06iaHj1mVdOHqltflyaY0dR60pK2aP4_nhdq3Bf3yohm7WZpfcFjvKAHCczoQ2D9p6Tu12ShcVibab7XW8RIOe0HxMmZOIcLhKae0wFeKU5jzEuySpsIDgI9QlKCay5KmjYt.AQ_Uo8B2GzYZS36OfTdXOA3MA__reEFM_98SJm1tXIrCm9aLWY3USEldQibV4FUihJt1dAoWNWYg2SHMFW3v5WiNGdZI_tJ8z38K4lc.Kuuw49aFYB4DXEZTz6bqMHYh58jkDqIdZ26UAt2cD4TzWFKSo1d9iO9LQfh3_QYAnASoxz3Rbnc62"};var cpo = document.createElement('script');cpo.src = '/cdn-cgi/challenge-platform/h/b/orchestrate/chl_page/v1?ray=913547bd0f7ffd67';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, "\/gsajournals\/pages\/plants-genetics-and-genomics?__cf_chl_rt_tk=QxR.XGuZpXPWrJFP1eb0lwZAsTkZs4Cfei055kcbIUE-1739790094-1.0.1.1-4LvRiCjesxsBJ_Etgvbk7_xPj9vLnNfGcZTfdb3OWaQ" + window._cf_chl_opt.cOgUHash);cpo.onload = function() {history.replaceState(null, null, ogU);}}document.getElementsByTagName('head')[0].appendChild(cpo);}());</script></body></html>