A lever action hypothesis for pendulous hummingbird flowers: experimental evidence from a columbine.

BACKGROUND AND AIMS: Pendulous flowers (due to a flexible pedicel) are a common, convergent trait of hummingbird-pollinated flowers. However, the role of flexible pedicels remains uncertain despite several functional hypotheses. Here we present and test the 'lever action hypothesis': flexible pedicels allow pendulous flowers to move upwards from all sides, pushing the stigma and anthers against the underside of the feeding hummingbird regardless of which nectary is being visited. METHODS: To test whether this lever action increased pollination success, we wired emasculated flowers of serpentine columbine, Aquilegia eximia, to prevent levering and compared pollination success of immobilized flowers with emasculated unwired and wire controls. KEY RESULTS: Seed set was significantly lower in wire-immobilized flowers than unwired control and wire control flowers. Video analysis of visits to wire-immobilized and unwired flowers demonstrated that birds contacted the stigmas and anthers of immobilized flowers less often than those of flowers with flexible pedicels. CONCLUSIONS: We conclude that flexible pedicels permit the levering of reproductive structures onto a hovering bird. Hummingbirds, as uniquely large, hovering pollinators, differ from flies or bees which are too small to cause levering of flowers while hovering. Thus, flexible pedicels may be an adaptation to hummingbird pollination, in particular due to hummingbird size. We further speculate that this mechanism is effective only in radially symmetric flowers; in contrast, zygomorphic hummingbird-pollinated flowers are usually more or less horizontally oriented rather than having pendulous flowers and flexible pedicels.

Mucilage-bound sand reduces seed predation by ants but not by reducing apparency: a field test of 53 plant species.

Seed mucilage, a coating on seeds or fruit that becomes slimy and sticky when wet, has evolved convergently many times across plants. One common consequence of having seed mucilage is that sand and dirt particles stick to wet seeds and remain tightly bound to the seed surface after the mucilage dries. Here, we test the hypothesis that a mucilage-bound sand coating protects the seed from seed predators; either as a physical barrier or by reducing apparency of the seed (i.e., camouflage). We experimentally manipulated the sand coating on seeds of 53 plant species of 13 families and assayed the defensive benefit of the sand coating in feeding "depots" near harvester ant nests in California's Central Valley. Consistent with a defensive function, sand coating reduced ant predation on seeds in 48 of the 53 species examined. To test whether this striking benefit was due to reduced apparency, we conducted an addition experiment using flax seeds in which we factorially manipulated the color of both the background substrate and the sand coating, creating visually apparent and unapparent seeds. Our results did not support the reduced apparency hypothesis; seeds coated in background-matched sand were removed at the same rate as seeds coated in unmatched sand. The defensive benefit of a sand coating was not well-predicted by seed mass, entrapped sand mass, or sand mass scaled by seed mass. Together, our results demonstrate that seed mucilage is a phylogenetically widespread and effective seed defensive trait and point to the physical barrier, not reduced apparency, as a mechanism.

A chloride- and calcium-dependent glutamate-binding protein from rat brain. Identification as a ubiquitous constituent of the inner mitochondrial membrane.

We have recently solubilized and enriched a chloride- and calcium-dependent glutamate-binding protein from rat brain (Brose, N., Halpain, S., Suchanek, C., and Jahn, R. (1989) J. Biol. Chem. 264, 9619-9625). The partially purified protein fraction, containing two major protein components of 51,000 Da and 105,000 Da, was used to generate a rabbit antiserum. This serum quantitatively precipitated the binding activity from membrane extracts. Small amounts of the antiserum inhibited glutamate binding when chloride was absent from the incubation medium. Three protein bands were labeled by the serum on immunoblots. From the affinity purified antibody fractions contained in the serum, only the antibodies directed against a 51,000-Da protein were able to immunoprecipitate the binding activity, indicating that this protein is an essential component of the binding site. A survey of a variety of rat tissues by immunoblot analysis revealed a ubiquitous distribution of the protein. After subcellular fractionation of liver and brain, the 51,000-Da protein copurified with mitochondrial markers. Furthermore, exclusive labeling of mitochondria was observed by light and electron microscopy immunocytochemistry. Subfractionation of purified liver mitochondria resulted in a selective association of the protein with inner mitochondrial membranes. Pharmacological characterization of glutamate binding to liver mitochondrial membranes revealed a pattern almost identical to that of the chloride- and calcium-dependent glutamate-binding site in rat brain.