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We combined transmission electron microscope and confocal microscopy to reconstruct three\u2010dimensional internal leaf structures of C\n                          <jats:sub>3<\/jats:sub>\n                          , C\n                          <jats:sub>2<\/jats:sub>\n                          , and C\n                          <jats:sub>4<\/jats:sub>\n                          <jats:italic>Alternanthera<\/jats:italic>\n                          species to reveal differences in the M\u2013BS interface and cell\u2010to\u2010cell communication capacity.\n                        <\/jats:p>\n                      <\/jats:list-item>\n                      <jats:list-item>\n                        <jats:p>\n                          We found that M\u2013BS connectivity increases stepwise from C\n                          <jats:sub>3<\/jats:sub>\n                          to C\n                          <jats:sub>2<\/jats:sub>\n                          and to C\n                          <jats:sub>4<\/jats:sub>\n                          species, but these shifts are achieved via different combinations of underlying traits in C\n                          <jats:sub>2<\/jats:sub>\n                          vs C\n                          <jats:sub>4<\/jats:sub>\n                          species. Specifically, increases to vein density were unique to C\n                          <jats:sub>2<\/jats:sub>\n                          whilst enhancements to BS cell, pit field, and plasmodesmata densities were unique to C\n                          <jats:sub>4<\/jats:sub>\n                          <jats:italic>Alternanthera<\/jats:italic>\n                          species. Both C\n                          <jats:sub>2<\/jats:sub>\n                          and C\n                          <jats:sub>4<\/jats:sub>\n                          <jats:italic>Alternanthera<\/jats:italic>\n                          shorten BS cell length, which may facilitate tighter BS packing to enhance M\u2013BS connectivity.\n                        <\/jats:p>\n                      <\/jats:list-item>\n                      <jats:list-item>\n                        <jats:p>Our findings highlight the diverse mechanisms through which photosynthetic innovation can emerge, providing valuable insight for evolutionary biology and agricultural bioengineering efforts.<\/jats:p>\n                      <\/jats:list-item>\n                    <\/jats:list>\n                  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