Highways for internal virus spread: patterns of virus movement in the stoloniferous herb Trifolium repens

Resource sharing between connected ramets can be advantageous for clonal plants. However, widespread vascular integration bears the risks of enhanced internal pathogen spread, which may be one of the most prominent disadvantages of clonal growth. In this paper we analyzed patterns of internal virus spread in ramet groups of the stoloniferous herb Trifolium repens L. (white clover) and investigated the effect of leaf ontogeny on intraclonal disease development. We inoculated single leaves of T. repens with White clover mosaic virus and analyzed the infection status of ramets at different distances from the point of infection and in leaves from different developmental stages. White clover mosaic virus infected all young developing plant parts positioned on basal branches or on the main stolon. Leaf ontogeny strongly affected disease development, and fully mature leaves on the main stolon remained virus free. The pattern of plant-internal virus spread was not affected by heterogeneous light conditions. Despite the well-described advantages of physiological integration, our data suggest that clonal integration may lead to negative selection pressures on clonal growth in pathogen-prone environments.