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The costs and benefits, measured in terms of dry weight, of physiological integration between clonal ramets, were analysed in two experiments conducted on the clonal herb Glechoma hederacea. Firstly, integration between consecutively-produced ramets was examined in an experiment in which stolons grew from one set of growing conditions (either unshaded or shaded and either nutrient-rich or nutrient-poor) into conditions in which light or nutrient level was altered. Comparisons were made between the dry weight of the parts of the clones produced before and after growing conditions were changed, and the dry weights of the corresponding part of control clones subjected to constant growing conditions. In a second experiment, integration between two distinct parts of G. hederacea clones was investigated. In this experiment clones were grown from two connected parent ramets and the parts of the clone produced by each parent ramet were subjected independently to either nutrient-rich or nutrient-poor conditions. Ramets in resource-rich conditions provided considerable physiological support to those in resource-poor conditions. This was measured as a dry weight gain compared with the weight of the corresponding part of the control clones growing in resource-poor conditions. However, when stolons grew from resource-poor conditions into resource-rich conditions, there was no similar evidence of the resourcepoor ramtes receiving support from resource-rich ramets. Physiological integration did not result in dry weight gains when this would have necessitated basipetal translocation of resources.Because of the predominantly acropedal direction of movement of translocates in G. hederacea, the structure of the clone was important in determining the effectiveness of integration between ramets. Where physiological integration was effective, the cost to the supporting ramets in terms of dry weight was insignificant. Physiological integration allows clones to maintain a presence in less favourable sites with insignificant cost to ramets in favourable sites, thereby reducing the probability of invasion by other plants, and providing the potential for rapid clonal growth if conditions improve. Integrated support of ramets in unfavourable conditions also enables the clone to grow through unfavourable sites, thus increasing the probability of encountering more favourable conditions by wider foraging.

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Clones of the gynodioecious herb Glechoma hederacea L. produce ramets which flower and bear seed, ramets which flower but fail to bear seed, and sterile ramets. Performance and survivorship of these subsets of ramets were compared for hermaphrodite and male sterile clones in contrasting grassland and woodland habitats for 2 consecutive years. For all clones, ramet densities on given dates were comparable between years. Winter survival of ramets exceeded 95% and mortality during the growing season was exponential. No genet recruitment was recorded throughout the study. For all clones, flowering ramets were larger than nonflowering ramets. Both flowering and nonflowering ramets of hermaphrodite clones, from grassland and woodland, were significantly larger than corresponding types of ramets from male sterile clones. A higher proportion of ramets of hermaphrodite clones flowered and set seed than ramets of male sterile clones. Flowering ramets of hermaphrodite clones produced more flowers than flowering ramets of male sterile clones. For all clones, half-lives of subsets of ramets declined in the order: seed-bearing > flowering but not seed-bearing > nonflowering. Within the grassland habitat, ramet subsets from hermaphrodite clones had longer half-lives than corresponding subsets from the male sterile clones. In the absence of greater flowering and seed set by male sterile clones, self-incompatibility of hermaphrodite flowers, which has been reported previously for G. hederacea, is crucial for the maintenance of clones of both sexes in populations.