In angiosperms selection has led to the utilization of numerous substances other than pollen and nectar that help to insure repeated visitation by pollinating animals. Here, we group the various substances into nonnutritive and nutritive rewards and discuss within each group the specific kinds that occur. In our discussion of nutritive rewards, we emphasize floral oils, lipids produced by one of two types of specialized secretory organs called elaiophores and which serve as nutritive rewards for certain New World anthophorine bees. Although discovered only within the last 15 years, the syndrome of oil production now appears to be one of the most widespread kinds of floral rewards. We report here for the first time the occurrence of oil production in the Solanaceae (Nierembergia). It is apparent that oil production has evolved independently many times, but plants which produce oils that are collected by female anthophorine bees show similarities in the chemistry of the oils and the types of structures that produce them. It is not clear whether other groups of plants reported to produce oils but which are not pollinated by anthophorine bees possess an analogous system or not. Floral rewards can be considered any component of a flower or inflorescence that is used by animals and, because of this use, insures repeated visitation that will lead to pollination. Without doubt, pollen and nectar are the primary rewards offered by flowers to visiting animals in order to buy their services as pollinating agents. Of the two, nectar is sought by a wider array of animals than pollen. On the other hand, pollen is the primary reward for which bees, probably the single most important group of pollinators, visit flowers. The role of pollen and nectar in the attraction of potential pollinators has been appreciated for hundreds of years, but we have only recently begun to realize the complex nature of these two rewards. Nectar, for example, formerly considered to be a simple sugar solution, has been shown to consist of a variety of chemicals dissolved, or suspended, in an aqueous solution. These range from mixtures of one to three common sugars (glucose, sucrose and fructose) to more complex sugar solutions (Percival, 1961) or combinations of sugars, free amino acids, vitamins, lipids, and other compounds (Baker & Baker, 1975; Baker, 1978). The complex chemical nature of pollen has been realized for a century (refs. in Barbier, 1971), but only in the last twenty years have researchers begun to explore the varied nature of specific enzymes contained in the pollen walls and their possible roles in incompatibility reactions (Stanley & Linskens, 1974). These same enzymes may play a role in pollen recognition by specific pollinators. The chemistry of pollen is in fact so complex that it has been impossible to provide a precise description of pollen chemistry that is all-inclusive. The continued elucidations of the intricate nature of these common rewards has spurred studies of pollination biology and provided an impetus for the investigation or reinvestigation of other floral rewards.
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