Physiological integration in the clonal perennial herb Trifolium repens L.

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Abstract
Translocation of 14C-labelled carbohydrates between the parent stolon and branches, and among branches, of Trifolium repens plants was investigated in two glasshouse experiments to determine patterns of physiological organisation in this clonal species. Differential defoliation treatments were applied to the parent stolon and/or branches to test the sensitivity of translocation to the short-term carbon needs of defoliated sinks. Strong reciprocal exchange of carbohydrate between the parent stolon and branches was observed, with 18 41% of the 14C exported from leaves on the parent stolon moving to branches, while branches simulta-neously exported 25% (for old source branches) to 54% (for young source branches) of the 14C they assimilated to the parent plant, including translocation to other branches. Branch-to-branch translocation occurred both acropetally and basipetally. Parent-to-branch, branch-to-parent and branch-to-branch carbon fluxes all increased in response to defoliation of the sink, at the expense of carbon supply to stolon tissue or roots of the source module. Reduced export to stolon tissue of the parent axis played a major role in facilitating C reallocation from leaves on the parent stolon to defoliated branches. The observed patterns of C allocation and translocation could be adequately explained by accepted source-sink theory, and are consistent with a high degree of intra-plant physiological integration in resource supply and utilisation. This information provides mechanistic explanations for aspects of the growth dynamics and ecological interactions of T. repens in the patchy environment of a grazed pasture.