613
INTRODUCTION
Many animal species show preferences for certain food types (Ayala-
Berdon et al., 2008). The amount of food consumed by the animal
is usually reduced as levels of available energy are increased
(Collins, 1981; Slansky and Wheeler, 1992). Studies of this ‘intake
response’ (Castle and Wunder, 1995) have mostly focused on
nectarivorous birds (Collins, 1981; López-Calleja et al., 1997;
McWhorter and Martínez del Rio, 2000) but these issues have
seldom been addressed in frugivores (see Levey and Martínez del
Rio, 1999).
According to Baker and Baker, hummingbird-pollinated plants
produce nectar that is rich in sucrose, whereas passerine-pollinated
plants produce nectar in which hexose sugars are dominant (Baker
and Baker, 1983). Some choice tests have shown that several
hummingbird species prefer sucrose to glucose or fructose
(Hainsworth and Wolf, 1976; Martínez del Rio, 1990a), whereas
some passerine species prefer glucose and fructose to sucrose
(Martínez del Rio and Stevens, 1989; Martínez del Rio et al., 1989).
However, Lotz and Schondube suggested that birds will generally
prefer hexose sugars to sucrose at low equicaloric concentrations
but will prefer sucrose to hexose at high equicaloric concentrations
(Lotz and Schondube, 2006). A possible explanation for this switch
at high concentrations is that sucrose nectar contains double the
energy content per unit of osmotic concentration than hexose nectar
(Beuchat et al., 1990). This reduced osmotic concentration may
increase post-ingestional intestinal energy delivery rate by
increasing gastric emptying rate (Karasov and Cork, 1994;
Schondube and Martínez del Rio, 2003). If the delivery of high
osmotic, concentrated solutions is not regulated birds may
experience osmotic diarrhoea (Lotz and Schondube, 2006). Sucrotic
birds may therefore lessen their osmotic stress by choosing sucrose
rather than hexose at high concentrations but at low concentrations
this effect may have less consequence (Lotz and Schondube, 2006).
However, Johnson and Nicolson recommend that a more useful
distinction would be between specialist and generalist bird
pollination systems (Johnson and Nicolson, 2008). They suggest
that generalist bird-pollinated plants are characterised by large
volumes of nectar (probably as a result of occasional avian
nectarivores having a much large body size) with low
concentrations and low sucrose proportions whereas specialist bird-
pollinated plants are characterised by smaller volumes of nectar
with higher concentrations and high sucrose proportions. It has also
been suggested that preferences are based on energy yield for
relatively small specialist nectarivores (Downs and Perrin, 1996).
However, experimental procedure may affect the results of
choice tests (Brown et al., 2008). Previous studies have used
solutions that are equicaloric (Fleming et al., 2004), equimolar
(Downs, 1997; Downs, 2000) or solutions that are equivalent by
mass (Lotz and Nicolson, 1996; Blem et al., 2000). Equimolar
solutions offer equal amounts of molecules per solution, but a
sucrose solution at a particular concentration has approximately
double the energy of an equimolar hexose solution (Hixon, 1980;
Schoener, 1983; Downs and Perrin, 1996; Downs, 1997). Sucrose
solutions that are equivalent in sugar mass to a hexose sugar solution
have approximately 5% more available energy (Fleming et al., 2004).
It has therefore been suggested that in choice tests sugar solutions
that are energetically equivalent should be used (Fleming et al.,
2004).
The Journal of Experimental Biology 214, 613-618
© 2011. Published by The Company of Biologists Ltd
doi:10.1242/jeb.047217
Food preferences of Knysna and purple-crested turacos fed varying concentrations
of equicaloric and equimolar artificial fruit
Amy-Leigh Wilson and Colleen T. Downs*
School of Biological and Conservation Sciences, University of KwaZulu-Natal, Private Bag X01, Pietermaritzburg, 3209, South Africa
*Author for correspondence (downs@ukzn.ac.za)
Accepted 27 October 2010
SUMMARY
The effects that different fruit sugar types and concentrations have on food preferences of avian frugivores have been relatively
poorly studied. Although it has been recently advocated that preference is based on equicaloric energy it is also important to note
whether preferences change as energy content changes. Therefore, sugar preferences of equicaloric and equimolar artificial fruit
of different sugar types at varying concentrations and molarities were investigated in two relatively large South African frugivores,
Knysna (Tauraco corythaix) and purple-crested (Gallirex porphyreolophus) turacos. Artificial fruits containing 6.6, 12.4 or 22%
sucrose or glucose, and artificial fruits containing 0.42, 0.83 or 1.66 mol l
–1
sucrose or glucose, were used to determine sugar
preferences. Knysna turacos preferred the sucrose to the glucose equicaloric artificial fruit diet at low concentrations whereas
purple-crested turacos showed no preference for either diet. Both turacos species preferred the sucrose equimolar artificial fruit
diet to the glucose at low concentrations. At high concentrations neither species showed a preference for either equicaloric or
equimolar artificial fruit diets. This suggests that energy requirements influence food preferences more than sugar type and that
birds will select fruit that is higher in energy irrespective of sugar type. This complements an earlier study on digestion of
differing equicaloric and equimolar artificial fruit sugar types. It again emphasizes the need for future studies looking at the
composition of indigenous forest fruit sugars in order to obtain insight into the role of these avian frugivores as potential seed
dispersers of fruiting tree species.
Key words: sugar type, sugar preference, energy requirement, frugivory, Knysna turaco, purple-crested turaco, artificial fruit.
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