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SOYBEAN and BEES

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honeybees can discriminate between sucrose concentrations differing by only 5%. They<br />

showed that bees can distinguish between 50% <strong>and</strong> 45%, but not between 50% <strong>and</strong> 47.5% or<br />

between 47.5% <strong>and</strong> 45% sucrose.<br />

Carbohydrates <strong>and</strong> free amino acids in the nectar are paramount for attraction, but<br />

because animals differ in their nutritive preferences, the composition of the nectar<br />

determines the spectrum of its consumers. For example, hummingbirds, butterflies,<br />

moths <strong>and</strong> long-tongued bees usually prefer sucrose-rich FNs, as do most ant species<br />

that feed on EFN, whereas short-tongued bees <strong>and</strong> flies prefer FN rich in hexoses<br />

(Gonzales-Teuber <strong>and</strong> Heil, 2009a; Nepi <strong>and</strong> Stpiczynska, 2008; Nepi et al., 2009;<br />

Blütgen <strong>and</strong> Fiedler, 2004). However, some nectarivorous birds <strong>and</strong> ants lack the<br />

enzyme invertase, being not able to process sucrose, thus preferring sucrose-free nectars,<br />

according to Heil et al. (2005) <strong>and</strong> Martínez del Río (1990).<br />

Previous studies reached quite different conclusions. Whitehead <strong>and</strong> Larsen (1976) determined<br />

that the maximum response of honeybee galeal chemoreceptors occurs with<br />

sucrose concentrations of about 1.5 molar (50% w/w) <strong>and</strong> glucose or fructose concentrations<br />

of about 3.0 molar (50% w/w). Honeybee carbohydrate preferences, based on nectar<br />

contents of plant species preferred by bees, indicated that honeybees might prefer<br />

nectar with relatively equal quantities of fructose, glucose, <strong>and</strong> sucrose (Furgala et<br />

al., 1958; Kropacova, 1965). However, Wykes (1952) examined the gustatory response<br />

of honeybees to sugar solutions varying in composition but having the same total sugar<br />

concentration, showing that honeybees prefer solutions of sugars in this descending<br />

order: sucrose, glucose, maltose, <strong>and</strong> fructose. Bachman <strong>and</strong> Waller (1977) <strong>and</strong> Waller<br />

(1972) showed that honeybees prefer sugar solutions in which sucrose is both the main<br />

constituent <strong>and</strong> is near a concentration of 50%.<br />

Besides carbohydrates <strong>and</strong> amino acids, which are present in high proportion in nectar, other<br />

compounds are involved in the nectar capacity of attraction. Volatile organic compounds<br />

(VOCs), like benzyl acetone, have been related to pollinator attraction, <strong>and</strong> scented petals<br />

have been known for centuries. In line with this information, nectar odors are considered a<br />

relevant signal for pollinators (Raguso, 2004). On the other side, gelsemine <strong>and</strong> iridoid glycosides<br />

exhibit repellent properties (Heil, 2011).<br />

Weiss (2001) demonstrated that butterflies <strong>and</strong> moths prefer artificial flowers containing<br />

scented nectar in contrast with ones containing pure sugar solutions, while Röse et al. (2006)<br />

mentioned that parasitoid wasps localize cotton (Gossypium hirsutum) EFN using only its<br />

odors, the same for mites that use nectar odors to distinguish between host <strong>and</strong> non-host<br />

plants, according to Reyneman et al. (1991).<br />

70 SoybeAn <strong>and</strong> bees

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