摘要
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Photosynthetic pathway characteristics were studied in nine species of <i>Heliotropium</i> (<i>sensu lato</i>, including <i>Euploca</i>), using assessments of leaf anatomy and ultrastructure, activities of PEP carboxylase and C<su...
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Photosynthetic pathway characteristics were studied in nine species of <i>Heliotropium</i> (<i>sensu lato</i>, including <i>Euploca</i>), using assessments of leaf anatomy and ultrastructure, activities of PEP carboxylase and C<sub>4</sub> acid decarboxylases, and immunolocalization of ribulose 1.5-bisphosphate carboxylase/oxygenase (Rubisco) and the P-subunit of glycine decarboxylase (GDC). <i>Heliotropium europaeum</i>, <i>Heliotropium calcicola</i> and <i>Heliotropium tenellum</i> are C<sub>3</sub> plants, while <i>Heliotropium texanum</i> and <i>Heliotropium polyphyllum</i> are C<sub>4</sub> species. <i>Heliotropium procumbens</i> and <i>Heliotropium karwinskyi</i> are functionally C<sub>3</sub>, but exhibit 'proto-Kranz' anatomy where bundle sheath (BS) cells are enlarged and mitochondria primarily occur along the centripetal (inner) wall of the BS cells; GDC is present throughout the leaf. <i>Heliotropium convolvulaceum</i> and <i>Heliotropium greggii</i> are C<sub>3</sub>-C<sub>4</sub> intermediates, with Kranz-like enlargement of the BS cells, localization of mitochondria along the inner BS wall and a loss of GDC in the mesophyll (M) tissue. These C<sub>3</sub>-C<sub>4</sub> species of <i>Heliotropium</i> probably shuttle photorespiratory glycine from the M to the BS tissue for decarboxylation. <i>Heliotropium</i> represents an important new model for studying C<sub>4</sub> evolution. Where existing models such as <i>Flaveria</i> emphasize diversification of C<sub>3</sub>-C<sub>4</sub> intermediates, <i>Heliotropium</i> has numerous C<sub>3</sub> species expressing proto-Kranz traits that could represent a critical initial phase in the evolutionary origin of C<sub>4</sub> photosynthesis.
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