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However, the molecular mechanisms regulating cone enlargement after fertilization remain poorly understood. Here, using\n                          <jats:italic>Torreya grandis<\/jats:italic>\n                          , a representative gymnosperm species with persistently low cone enlargement efficiency, we elucidated the molecular regulation of cone enlargement and identified sucrose non\u2010fermenting 1\u2010related protein kinase 1 (TgSnRK1) as a novel negative regulator of this process.\n                        <\/jats:p>\n                      <\/jats:list-item>\n                      <jats:list-item>\n                        <jats:p>\n                          TgSnRK1 physically interacted with the WRKY transcription factor TgWRKY47 both\n                          <jats:italic>in vitro<\/jats:italic>\n                          and\n                          <jats:italic>in vivo<\/jats:italic>\n                          . Functional analyses revealed that TgSnRK1 acted upstream of TgWRKY47 and suppressed cone enlargement in\n                          <jats:italic>T. grandis<\/jats:italic>\n                          .\n                        <\/jats:p>\n                      <\/jats:list-item>\n                      <jats:list-item>\n                        <jats:p>\n                          Phosphorylation assays revealed that TgSnRK1 directly phosphorylated TgWRKY47 at Thr\u201075. This phosphorylation event markedly reduced the transactivation activity of TgWRKY47, thereby attenuating its activation of the cell wall\u2010remodeling gene\n                          <jats:italic>TgEXPA2<\/jats:italic>\n                          , a key regulator of cell expansion during cone enlargement.\n                        <\/jats:p>\n                      <\/jats:list-item>\n                      <jats:list-item>\n                        <jats:p>Together, these findings establish a phosphorylation\u2010dependent TgSnRK1\u2010TgWRKY47\u2010TgEXPA2 regulatory pathway in which developmental attenuation of TgSnRK1\u2010mediated phosphorylation of TgWRKY47 at Thr\u201075 enhances TgWRKY47 transactivation activity, thereby promoting TgEXPA2\u2010dependent cone enlargement. This study advances our understanding of reproductive development in gymnosperms and identifies potential molecular targets for improving reproductive efficiency and seed yield.<\/jats:p>\n                      <\/jats:list-item>\n                    <\/jats:list>\n                  <\/jats:p>","DOI":"10.1111\/nph.71525","type":"journal-article","created":{"date-parts":[[2026,8,28]],"date-time":"2026-08-28T10:56:06Z","timestamp":1787914566000},"page":"892-908","update-policy":"https:\/\/doi.org\/10.1002\/crossmark_policy","source":"Crossref","is-referenced-by-count":0,"title":["Phosphorylation of the transcription factor\n                    <scp>TgWRKY47<\/scp>\n                    by\n                    <scp>TgSnRK1<\/scp>\n                    modulates cone enlargement in the gymnosperm\n                    <scp>\n                      <i>Torreya grandis<\/i>\n                    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