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A lever action hypothesis for pendulous hummingbird flowers: experimental evidence from a columbine.
Background and aimsPendulous 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.MethodsTo 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 resultsSeed 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.ConclusionsWe 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
Effective regulations with little effect? The antecedents of the perceptions of environmental officials on enforcement effectiveness in China
Enforcement gaps are an especially vexing problem in China due to pervasive "pro-growth" local government priorities, the weak administrative capacity of environmental agencies themselves, and relatively weak levels of societal support for a cleaner environment. This study seeks to examine this problem from the perspective of the local enforcement officials by empirically examining the relationships between these and other influences on their perceptions of enforcement effectiveness. Using samples of enforcement officials from the fast-growing cities of Chengdu, Dalian, and Guangzhou, this study finds that many of the more commonly cited problems related to regulatory enforcement were not as influential in accounting for variations in perceptions of enforcement effectiveness than the current literature might suggest. Moreover, this study also finds that the pattern of influences varies greatly across jurisdictions, with only the officials' belief in the legitimacy of the governments' policies being significant in all three samples. Strong influences on perceptions of enforcement effectiveness in two of the three samples include the environmental values of enforcement officials, their perceptions of organizational capacity for enforcement, and their assessment of government support for environmental protection. Although government support was found to be a strong predictor of enforcement effectiveness in two of the three samples, the influence of societal support had a somewhat mixed and more complicated effect. In addition, this study suggests that further improvements in enforcement effectiveness may be possible by cultivating or selecting enforcement officials with strong environmental values and beliefs in the legitimacy of the government's environmental policy to take charge of enforcement. Because it is generally accepted that local environmental protection bureaus are generally upgrading their organizational capacity for improvement as the result of increasing central government support for environmental protection and institutional restructuring, and because environmental awareness in China is growing, this study suggests that some incremental progress is likely in China's efforts to close the enforcement gap. Unfortunately, such improvements are likely to be masked by the steep trajectory of economic growth, the narrow scope of regulatory control (i.e., with scant attention paid to nonindustrial sources), and the migration of industrial pollution to rural areas