Abstract
A nectar diet is simple in nutritional composition and easily digested, but may vary greatly in its proportions of sugar and water. Here, we apply the geometric framework, a modelling approach for investigating how animals balance nutrient needs in multidimensional and dynamic nutritional environments, to captive whitebellied sunbirds (Cinnyris talatala). We address the question of how these small birds (~8 g) prioritise sugar and water intake, and how dietary salt content interacts with sugar and water intake. Sunbirds kept at 20°C and provided with moderate to high sucrose concentrations (≥1 M), together with supplementary water, converge on an intake target of 2.79 g day−1 of sucrose and 7.72 g day−1 of water: equivalent to 0.85 M sucrose. When the birds are given more dilute sucrose concentrations, they defend their sugar intake by over-ingesting water, up to a ceiling of 47 g day−1. Sugar intake thus gets priority over water intake, but the birds have a finite capacity to over-ingest water to gain the target level of sugar. Regulation appears to be less precise when birds are given a choice between two sucrose solutions than when they choose between a sugar solution and supplementary water. Intake targets vary in response to internal and external factors, and sunbirds increase their sugar intake in response to increased activity and cold, irrespective of nectar concentration. They also compensate for interruptions in foraging activity, whether overnight or during the day. Interactive effects become evident when sodium is included as a third nutrient: on very dilute nectar (≤0.1 M), where sunbirds lose body mass, the addition of sodium to the diet helps to achieve the carbohydrate intake target, while raising the ceiling on water intake. This analysis provides a new perspective on nectarivory, while adding to the comparative database on nutrient regulation and emphasising water as a nutrient.





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Acknowledgments
This work was supported by the University of Pretoria, the South African National Research Foundation and the Claude Leon Foundation. DR received support from the Massey University Research Fund.
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Communicated by I.D. Hume.
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Köhler, A., Raubenheimer, D. & Nicolson, S.W. Regulation of nutrient intake in nectar-feeding birds: insights from the geometric framework. J Comp Physiol B 182, 603–611 (2012). https://doi.org/10.1007/s00360-011-0639-2
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DOI: https://doi.org/10.1007/s00360-011-0639-2