Use of a sound-based vibratome by leaf-cutting ants.

Leaf-cutting ants harvest fresh vegetation that they then use as food for symbiotic fungi. When cutting leaf fragments, the ants produce high-frequency vibrations with a specialized organ located on the gaster. This stridulation behavior is synchronized with movements of the mandible, generating complex vibrations of the mandibles. The high vibrational acceleration of the mandible (up to three times the gravitational force at peak acceleration at about 1000 hertz) appears to stiffen the material to be cut. An identical effect is achieved when soft material is sectioned with a vibratome. This hypothesis is supported by experiments simulating the cutting process with vibrating isolated mandibles: When tender leaves were cut, the vibration of the mandible reduced force fluctuations and thus permitted a smoother cut to be made.

Perceptual differences in trail-following leaf-cutting ants relate to body size.

Leaf-cutting ants of the genus Atta have highly size-polymorphic workers, and size is related to division of labor. We studied trail-following behavior of different-sized workers in a laboratory colony of Atta vollenweideri. For small and large workers, we measured responsiveness and preference to artificial conspecific and heterospecific pheromone trails made from poison gland extracts of A. vollenweideri and A. sexdens. Responsiveness was measured as the probability of trail-following, and preference was measured by testing the discrimination between one conspecific and one heterospecific trail. Minute amounts of the releaser component methyl-4-methylpyrrole-2-carboxylate (0.4pg/1m), present in both, conspecific and heterospecific trails, suffice to elicit trail-following behavior. Workers followed heterospecific trails, and these trails (after normalizing their concentration) were as effective as conspecific trails. Small workers were less likely to follow a trail of a given concentration than large workers. In the discrimination test, small workers preferred the conspecific trail over the heterospecific trail, whereas large workers showed no significant preference. It is suggested that large workers primarily respond to the releaser component present in both trails, whereas small workers focus more on the conspecific traits provided by the blend of components contained in the trail pheromone.

Thermal radiation as a learned orientation cue in leaf-cutting ants (Atta vollenweideri).

We explored the ability of leaf-cutting ants (Atta vollenweideri) to learn the location of a food reward by using thermal information as an orientation cue. During training of single workers, the conditioned stimulus was a distant thermal source placed frontally, 15 mm away from a platform having a leaf fragment as reward. After training, single workers were confronted with the choice between two sides, one being coupled, in a pseudo-randomized design, with a thermal stimulus heated 5 degrees C above environmental temperature. After 10 learning trials, workers significantly chose the side with the thermal stimulus. This showed that workers can use thermal information for spatial orientation in the context of foraging, which may help them to locate, for instance, highly attractive sun-exposed leaves. Thermal radiation alone as orientation cue was sufficient to allow learning, since preclusion of thermal convection during training and test did not impair workers' response. Shielding of both thorax and gaster from the thermal source did not weaken learning, suggesting the sole participation of head and antennae in temperature reception. A thermal stimulus heated 1 degrees C above environmental temperature could not be used as a learned orientation cue, even when foragers were allowed to directly contact the thermal source.

Fluid intake rates in ants correlate with their feeding habits.

This study investigates the techniques of nectar feeding in 11 different ant species, and quantitatively compares fluid intake rates over a wide range of nectar concentrations in four species that largely differ in their feeding habits. Ants were observed to employ two different techniques for liquid food intake, in which the glossa works either as a passive duct-like structure (sucking), or as an up- and downwards moving shovel (licking). The technique employed for collecting fluids at ad libitum food sources was observed to be species-specific and to correlate with the presence or absence of a well-developed crop in the species under scrutiny. Workers of ponerine ants licked fluid food during foraging and transported it as a droplet between their mandibles, whereas workers of species belonging to phylogenetically more advanced subfamilies, with a crop capable of storing liquids, sucked the fluid food, such as formicine ants of the genus Camponotus. In order to evaluate the performance of fluid collection during foraging, intake rates for sucrose solutions of different concentrations were measured in four ant species that differ in their foraging ecology. Scaling functions between fluid intake rates and ant size were first established for the polymorphic species, so as to compare ants of different size across species. Results showed that fluid intake rate depended, as expected and previously reported in the literature, on sugar concentration and the associated fluid viscosity. It also depended on both the species-specific feeding technique and the extent of specialization on foraging on liquid food. For similarly-sized ants, workers of two nectar-feeding ant species, Camponotus rufipes (Formicinae) and Pachycondyla villosa (Ponerinae), collected fluids with the highest intake rates, while workers of the leaf-cutting ant Atta sexdens (Myrmicinae) and a predatory ant from the Rhytidoponera impressa-complex (Ponerinae) did so with the lowest rate. Calculating the energy intake rates in mg sucrose per unit time, licking was shown to be a more advantageous technique at higher sugar concentrations than sucking, whereas sucking provided a higher energy intake rate at lower sugar concentrations.

Olfactory conditioning during the recruitment process in a leaf-cutting ant.

During recruitment, workers of Acromyrmex lundi are conditioned to the odor of the food fragment initially carried by the scout worker. The learned odor cue is used as a decision criterion during food collection.

Haemolymph sugars and the control of the proventriculus in the honey bee Apis mellifera.

Crop emptying and rectal filling rates were investigated in bees trained to collect defined amounts of sucrose solution. Crop emptying rates strongly depended on the sucrose concentration of the collected solution. There was a close match between the energy expenditure of the bees and the amount of sucrose transported through the proventriculus, irrespective of the fluid dilution. Results indicated that the controlling variable was the amount of sucrose flowing through the proventriculus rather than the volume flow. In order to distinguish between haemolymph osmolality and haemolymph carbohydrate levels as factors controlling the activity of the proventriculus, bees were injected with either metabolizable or non-metabolizable carbohydrates. Only the injection of metabolizable carbohydrates modulated the activity of the proventriculus, indicating that the titers of metabolizable carbohydrates are involved in the feedback loop controlling crop emptying, and that haemolymph osmolality alone does not influence the activity of the proventriculus.

Haemolymph sugar levels in foraging honeybees (Apis mellifera carnica): dependence on metabolic rate and in vivo measurement of maximal rates of trehalose synthesis.

Previous investigations of haemolymph sugar levels in honeybees have reported very different results, probably because different experimental conditions affected the activity levels of the animals. The present study investigated the dependence of haemolymph sugar levels in foraging honeybees on metabolic rate and whether the haemolymph sugar level is regulated. Free-flying foraging bees were trained to collect controlled amounts of sucrose solution of different concentrations (15%, 30% or 50% sucrose w/w). Immediately after feeding, metabolic rate was measured over a given time depending on the sucrose concentration, then crop-emptying rate and haemolymph sugar levels were measured. Bees exhibiting a wide range of metabolic rates were compared to establish whether the observed differences in haemolymph sugar levels were due to limits in the supply of sugars from the crop or in the rate of trehalose synthesis in the fat bodies. Independent of the concentration of the sucrose solution supplied, haemolymph trehalose, glucose and fructose levels were constant for metabolic rates from 0 to 4.5 ml CO(2)h(-1). At higher metabolic rates, trehalose concentration decreased while that of glucose and fructose increased, with the exception of bees fed 15% sucrose solution. As the supply of sugar from the crop via the proventriculus was sufficient to support even the highest metabolic rates, the observed pattern must result from an upper limit in the capacity of the fat body to synthesise trehalose. The maximal rate of conversion of glucose to trehalose in the fat body was therefore calculated to average 92.4 microg glucosemin(-1). However, for bees fed 15% sucrose solution both the rate of conversion of glucose to trehalose and the rate of sugar transport from the crop to the ventricle were limited, together resulting in a decrease in total haemolymph sugar levels for metabolic rates higher than 5 ml CO(2)h(-1).

Wind-induced ventilation of the giant nests of the leaf-cutting ant Atta vollenweideri.

To understand the significance of elaborate nest architecture for the control of nest climate, we investigated the mechanisms governing nest ventilation in a large field nest of Atta vollenweideri. Surface wind, drawing air from the central tunnels of the nest mound, was observed to be the main driving force for nest ventilation during summer. This mechanism of wind-induced ventilation has so far not been described for social insect colonies. Thermal convection, another possible force driving ventilation, contributed very little. According to their predominant airflow direction, two functionally distinct tunnel groups were identified: outflow tunnels in the upper, central region, and inflow tunnels in the lower, peripheral region of the nest mound. The function of the tunnels was independent of wind direction. Outflow of air through the central tunnels was followed by a delayed inflow through the peripheral tunnels. Leaf-cutting ants design the tunnel openings on the top of the nest with turrets which may reinforce wind-induced nest ventilation.

Assessment of food source profitability in honeybees (Apis mellifera): how does disturbance of foraging activity affect trophallactic behaviour?

When forager honeybees (Apis mellifera) return to the hive after a successful foraging trip, they unload the collected liquid to recipient hive mates through mouth-to-mouth contacts (trophallaxis). The speed at which the liquid is transferred (unloading rate) from donor to recipient is related to the profitability of the recently visited food source. Two main characteristics that define this profitability are the flow of solution delivered by the feeder and the time invested by the forager at the source (visit time). To investigate the effect of visit time on trophallactic behaviour, donor foragers were trained to a rate feeder that could deliver different flows of solution. We dissociated visit time and flow of solution by introducing pauses in the solution's deliverance at different moments of the foraging visit. We analysed whether timing of the non-deliverance period within the visit is important for the forager's assessment of resource profitability. During the subsequent trophallactic encounter with a hive mate, unloading rate was related to the total time invested by the forager at the food source only if the ingestion process had already been started. These results together with previous ones suggest that foragers integrate an overall flow rate of solution of the feeder throughout the entire foraging visit.

Nectar feeding by the ant Camponotus mus: intake rate and crop filling as a function of sucrose concentration.

In independent assays, workers of the ant Camponotus mus were conditioned to visit an arena where they found a large drop of sucrose solution of different concentrations, from 5 to 70% weight on weight (w/w). Single ants were allowed to collect the sucrose solution ad libitum, and feeding time, feeding interruptions, crop load, and intake rates were recorded. Feeding time increased exponentially with sucrose concentration, and this relationship was quantitatively described by the increase in viscosity with concentration corresponding to pure sucrose solutions. Ants collecting dilute solutions (5 to 15% w/w) returned to the nest with partial crop loads. Crop filling increased with increasing sucrose concentration, and reached a maximum at 42.6% w/w. Workers collecting highly concentrated solutions (70% w/w) also returned to the nest with a partially-filled crop, as observed for dilute solutions. Nectar intake rate was observed to increase with increasing sucrose concentration in the range 5 to 30% sucrose. It reached a maximum at 30.8%, and declined with increasing sucrose concentration. Results suggest that both sucrose concentration and viscosity of the ingested solution modulate feeding mechanics as well as the worker's decision about the load size to be collected before leaving the source.

Foraging in the ant Camponotus mus: nectar-intake rate and crop filling depend on colony starvation.

The effects of colony starvation on the dynamics of nectar collection were studied in individual workers of the ant Camponotus mus. A laboratory colony was first deprived of carbohydrates for 15days, and thereafter fed daily ad libitum with diluted honey until satiation. During these two successive experimental phases, the probability of feeding, crop filling and fluid-intake rates were recorded daily for individual foragers collecting a 10% (w/w) sucrose solution. The feeding responses of individuals varied with the nutritional state of the colony. When the colony was deprived of sugar, acceptance of the sucrose solution was higher than under satiation. Feeding time increased with increasing starvation. During deprivation workers fed nearly continuously on the solution, whereas a number of feeding interruptions occurred under satiation. Crop filling also increased with increasing starvation, and showed a marked decrease when the colony was satiated. Fluid-intake rate during the deprivation phase was roughly twice that during the satiation phase. This matched well with the difference in sucking frequency recorded during ingestion in satiated and starved workers, which was also higher during starvation. Results indicate that the responsiveness of foragers, determined by the nutritional state of the colony, influenced both foraging decisions and the dynamics of fluid intake.