Pollinator directionality as a response to nectar gradient: promoting outcrossing while avoiding geitonogamy.

Plants with multiple flowers could be prone to autonomous self-pollination and insect-mediated geitonogamy, but physiological and ecological features have evolved preventing costs related to autogamy. We studied the rare perennial herb Dictamnus albus as a model plant, with the aim of describing the plant-pollinator system from both plant and pollinator perspectives and analysing features that promote outcrossing in an entomophilous species. The breeding system and reproductive success of D. albus were investigated in experimental and natural conditions, showing that it is potentially self-compatible, but only intra-inflorescence insect-mediated selfing is possible. Nectar analysis showed gender-biased production towards the female phase, which follows the male phase, and during flowering, full blooming is found in flowers at the bottom of the raceme. Among a wide spectrum of insect visitors, three genera (Bombus, Apis, Megachile) were found to be principal pollinators. A study of insect behaviour showed a tendency towards bottom-to-top flights for the most important pollinators Bombus spp. and Apis mellifera: upward movements on the racemes could be explained by foraging behaviour, from more to less rewarding flowers. In accordance with the 'declining reward hypothesis', bumblebees and honeybees leave the plant when gain of reward is low, after which few flowers are visited, reducing the chance of self-pollen transfer among flowers. Intra-flower self-pollination is prevented in D. albus by protandry and herkogamy, while the nectar-induced sequential pattern of pollinator visits avoids geitonogamy and tends to maximise pollen export, promoting outcrossing. All these features for preventing selfing benefit plant fitness and population genetic structure.