Effects of harvester ants on plant species distribution and abundance in a serpentine grassland

Seed harvesting ants can have important effects on the composition and structure of plant communities. We investigated two effects of Messor andrei, the black seed-harvesting ant, on a serpentine grassland plant community in northern California. First, to determine if selective seed predation by ants affects plant community composition, we excluded harvester ants from 1-mediameter circular plots of grassland. Abundances of all species on these plots and on control plots were measured before and after exclosure. Second, to determine if M. andrei nest mounds affect plant community composition, we compared plant species abundances on and off nest mounds. M. andrei deposit large amounts of organic matter on their nest mounds over a foraging season, so mounds may alter the edaphic environment. The exclusion of seed-harvesting activity did not cause changes in the plant community. Nest mounds had a strong effect on plant communities: there were many more grasses and fewer forbs on ant mounds, although at least one forb, Lepidium nitidum, produced twice as many seeds when it grew on nest mounds. We found that nest mounds formed islands of higher-temperature soil in the serpentine grassland. Received: 31 March 1997 / Accepted: 6 May 1997     

Effects of experimental manipulation of inflorescence size on pollination and pre-dispersal seed predation in the hummingbird-pollinated plant Ipomopsis aggregata

Large floral displays should theoretically provide advantages to plants through increased pollinator visitation and resulting fruit and seed set. However empirical tests of the response of pollinators to floral display size have been limited by a lack of direct experimentation, and the results of such studies have been equivocal. In addition, other selective agents such as pre-dispersal seed predators might modulate effects of floral display on pollination. By artificially altering flower number, we examined the direct effects of floral display in the monocarpic herb, Ipomopsis aggregata (Polemoniaceae), on visitation rates by broad-tailed and rufous hummingbird pollinators, as well destruction of fruits by a pre-dispersal seed predator (Hylemya: Anthomyiidae). In addition, we quantified the ultimate effects of flower number on female reproductive success. Plants with larger floral displays were most likely to be visited first in any given foraging bout (P < 0.01). As expected, plants with more flowers received more total flower visits. However, we found no gain in the proportion of flowers visited for many- versus few-flowered plants, or the total number of approaches/hour. In fact, a significantly greater percentage of flowers were visited on few-flowered plants. Plants did not compensate for our reduction in flowers by increasing investment in the number or proportion of flowers that set fruit, the number of seeds/fruit, or seed weight. Pre-dispersal seed predation was greater for many- than for few-flowered plants (P < 0.001), but this did not offset the potential fitness gains of producing large displays. Our data support the hypothesis that large floral displays function primarily in long-distance attraction of pollinators, and enhance maternal success. Received: 21 March 1996 / Accepted: 24 October 1996     

The exploitation of an ant-defended host plant by a shelter-building herbivore

Larvae of a Polyhymno species (Lepidoptera: Gelechiidae) feed on the ant-defended acacia, Acacia cornigera, in the tropical lowlands of Veracruz, Mexico. Polyhymno larvae construct sealed shelters by silking together the pinna or pinnules of acacia leaves. Although larval density and larval survival are higher on acacias not occupied by ants, shelters serve as a partial refuge from the ant Pseudomyrmex ferruginea (Hymenoptera: Formicidae), which defends A. cornigera plants; thus, shelters provide Polyhymno larvae access to an ant-defended host plant. P. ferruginea ants act as the primary antiherbivore defense of A. cornigera plants, which lack the chemical and mechanical defenses of non-ant-defended acacias. Thus, defeating the ant defense of A. cornigera provides Polyhymno larvae access to an otherwise poorly defended host plant. Damage caused by Polyhymno larval feeding reaches levels which can kill A. cornigera plants. Received: 6 June 1996 / Accepted: 6 September 1996     

Effects of aerobic exercise on the torque-velocity relationship in cycling

The kinetics of the torque-velocity (T-ω) relationship after aerobic exercise was studied to assess the effect of fatigue on the contractile properties of muscle. A group of 13 subjects exercised until fatigued on a cycle ergometer, at an intensity which corresponded to 60% of their maximal aerobic power for 50 min (MAP60%); ten subjects exercised until fatigued at 80% of their maximal aerobic power for 15 min (MAP80%). Of the subjects 7 exercised at both intensities with at least a 1-week interval between sessions. Pedalling rate was set at 60 rpm. The T-ω relationship was determined from the velocity data collected during all-out sprints against a 19 N · m braking torque on the same ergometer, according to a method proposed previously. Maximal theoretical velocity (ω₀) and maximal theoretical torque (T ₀) were estimated by extrapolation of the linear T-ω relationship. Maximal power (P _max) was calculated from the values of T ₀ and ω₀ (P _max = 0.25 ω₀ T ₀). The T-ω relationships were determined before, immediately after and 5 and 10 min after the aerobic exercise. The kinetics of ω₀, T ₀ and P _max was assumed to express the effects of fatigue on the muscle contractile properties (maximal shortening velocity, maximal muscle strength and maximal power). Immediately after exercise at MAP60% a 7.8% decrease in T ₀ and 8.8% decrease in P _max was seen while the decrease in ω₀ was nonsignificant, which suggested that P _max decreased in the main because of a loss in maximal muscle strength. In contrast, MAP80% induced a 8.1% decrease in ω₀ and 12.8% decrease in P _max while the decrease in T ₀ was nonsignificant, which suggested that the main cause of the decrease in P _max was probably a slowing of maximal shortening velocity. The short recovery time of the T-ω relationship suggests that the causes of the decrease of torque and velocity are processes which recover rapidly. Accepted: 25 November 1996     

Nitrogen storage forms in nine boreal understorey plant species

Storage forms of N were studied in below-ground structures of nine boreal forest understorey plants. The ericaceous shrubs Vacciniumvitis-idaea and V.myrtillus, the fern Gymnocarpium dryopteris, the grass Deschampsia flexuosa, and the herbs Epilobium angustifolium, Maianthemum bifolium, Solidago virgaurea, Geranium sylvaticum and Trientalis europaea were sampled in early summer and late autumn from plots fertilised with a complete mixture of nutrients and from non-fertilised control plots. Concentrations of total nitrogen, insoluble and soluble proteins, free amino acids and nitrate were measured, and changes in absolute and relative concentrations of these N fractions between early summer and late autumn were used to identify the forms in which the plants store N. In all species studied, the concentration of free amino acids increased both between summer and autumn and in response to fertilisation, while the concentration of protein N increased only in response to fertilisation. Thus, free amino acids appear to have a central role in N storage. In all of the species except G. dryopteris, D. flexuosa and S. virgaurea, arginine dominated the pool of free amino acids and thus arginine was the major form of stored N in most species. In D. flexuosa and S. virgurea, however, asparagine and arginine together were the major forms of stored N, while glutamine was the major free amino acid, and N storage form, in G. dryopteris. Received: 10 March 1996 / Accepted: 22 December 1996     

Interspecific interactions and co-existence in dabbling ducks: observations and an experiment

We studied the possible role of resource limitation and interspecific competition in assemblages of dabbling ducks on breeding lakes in Finland and Sweden with observational and experimental data. After initial vegetation mapping and yearly censuses of ducks in 1985–1990, we collected observational data in 1991–1994 from 28 lakes with natural populations of mallard Anas platyrhynchos and teal A. crecca. Mallard and teal co-occur over vast areas in the Holarctic and they are the only breeding dabbling ducks on many oligotrophic lakes. Both species are migratory in our study regions, teal arriving later in spring than mallards. Log-linear model analysis of observational presence/absence data revealed a positive, not a negative, association between the species. This association was independent of habitat diversity as well as of lake size. Mallard-teal interaction was also studied in a cross-over introduction experiment in 32 other lakes in two years. Wing-clipped mallards were introduced to breeding lakes before the arrival of teal to induce resource limitation and interspecific competition, hypothesized to reduce lake use by teal. The density of mallard pairs on experimental lakes was 2.9–8.0 times higher than on controls, but there was no negative response by teal to the treatment. This is the first combined observational-experimental demonstration of lack of interspecific competition in waterfowl. Our results indicate that heterospecific attraction may affect species co-existence in dabbling ducks. Received: 28 October 1996 / Accepted: 13 January 1997     

Effects of 3 days of carbohydrate supplementation on muscle glycogen content and utilisation during a 1-h cycling performance

This study compared the effects of supplementing the normal diets of six trained cyclists [maximal oxygen uptake O_2max) 4.5 (0.36)l · min⁻¹; values are mean (SD)] with additional carbohydrate (CHO) on muscle glycogen utilisation during a 1-h cycle time-trial (TT). Using a randomised crossover design, subjects consumed either their normal diet (NORM) for 3 days, which consisted of 426 (137) g · day⁻¹ CHO [5.9 (1.4) g · kg⁻¹ body mass (BM)], or additional CHO (SUPP) to increase their intake to 661 (76) g · day⁻¹ [9.3 (0.7) g · kg⁻¹ BM]. The SUPP diet elevated muscle glycogen content from 459 (83) to 565 (62) mmol · kg⁻¹ dry weight (d.w.) (P < 0.05). However, despite the increased pre-exercise muscle glycogen stores, there was no difference in the distance cycled during the TT [40.41 (1.44) vs 40.18 (1.76) km for NORM and SUPP, respectively]. With NORM, muscle glycogen declined from 459 (83) to 175 (64) mmol · kg⁻¹ d.w., whereas with SUPP the corresponding values were 565 (62) and 292 (113) mmol · kg⁻¹ d.w. Accordingly, both muscle glycogen utilisation [277 (64) vs 273 (114) mmol · kg⁻¹ d.w.] and total CHO oxidation [169 (20) vs 165 (30) g · h⁻¹ for NORM and SUPP, respectively] were similar. Neither were there any differences in plasma glucose or lactate concentrations during the two experimental trials. Plasma glucose concentration averaged 5.5 (0.5) and 5.6 (0.6) mmol · l⁻¹, while plasma lactate concentration averaged 4.4 (1.9) and 4.4 (2.3) mmol · l⁻¹ for NORM and SUPP, respectively. The results of this study show that when well-trained subjects increase the CHO content of their diet for 3 days from 6 to 9 g · kg⁻¹ BM there is only a modest increase in muscle glycogen content. Since supplementary CHO did not improve TT performance, we conclude that additional CHO provides no benefit to performance for athletes who compete in intense, continuous events lasting 1 h. Furthermore, the substantial muscle CHO reserves observed at the termination of exercise indicate that whole-muscle glycogen depletion does not determine fatigue at this exercise intensity and duration. Accepted: 25 November 1996     

Lack of evidence for local adaptation to individual plant clones or site by a mobile specialist herbivore

Several studies have documented local adaptation by sedentary insects to individual phenotypes of their host plants. Here, I examined whether a similar phenomenon could be found in a mobile, specialized insect, the sumac flea beetle. Previous work has shown that sumac individuals differ in their suitability as hosts for these beetles and that differences have both an environmental and a genetic basis. Using beetle populations collected as eggs from eight different sumac clones along an east-west transect, a reciprocal transfer experiment was conducted to determine whether there was any evidence for local adaptation by beetles to individual plant clones or to site. Variables examined were larval survivorship past first instar, development time, weight at pupation and patterns of predation by enemies. While no evidence for local adaptation was found, there were significant effects of plant clone on which larvae developed, origin of the larval population and the interaction of these effects on larval performance. For larval weight at pupation, there was also some indication that trade-offs may exist in ability of larvae to use different host plant clones. In addition, there were significant environmental effects on several measures of larval performance. Predation rates differed by plant clone, but not by site or with respect to origin of larvae. While no evidence for local adaptation was found in this study, prerequisites for finding such patterns may exist in this system. Received: 23 May 1996 / Accepted: 26 September 1996     

Effects of changes in atmospheric carbon dioxide on the location of hosts by the moth, Cactoblastis cactorum

Sensory organs that detect CO₂ are common in herbivorous moths and butterflies, but their function has been unclear until now. As the CO₂ gradients in the vicinity of a host plant depend on its physiological condition, CO₂ could provide a sensory cue for the suitability of the plant as a larval food source. This study investigated whether changing the atmospheric CO₂ concentration affected oviposition by Cactoblastis cactorum on its host, the cactus Opuntia stricta. On host plants exposed to rapid fluctuations in CO₂ concentration, the frequency of oviposition was reduced by a factor of 3.2 compared to the control. As the fluctuations mask the much smaller CO₂ signals generated by the plants, this suggests that those signals constitute an important component of the host identification process. On host plants exposed to a constant background of doubled CO₂, oviposition was also reduced, by a factor of 1.8. An increased background reduces host signal detectability, partially as a consequence of a general principle of sensory physiology (Weber-Fechner's law), and partially due to other factors specific to CO₂-receptor neurons. Received: 4 October 1996 / Accepted: 16 January 1997     

Effects of hydration state on hormonal and renal responses during moderate exercise in the heat

The effects of hydromineral hormones and catecholamines on renal concentrating ability at different hydration states were examined in five male volunteers while they performed three trials. Each of these trials comprised a 60-min exercise bout on a treadmill (at 50% of maximal oxygen uptake) in a warm environment (dry bulb temperature, 35°C; relative humidity, 20–30%). In one session, subjects were euhydrated before exercise (C). In the two other sessions, after thermal dehydration (loss of 3% body mass) which markedly reduced plasma volume (PV) and increased plasma osmolality (osm_pl), the subjects exercised either not rehydrated (Dh) or rehydrated (Rh) by drinking 600 ml of mineral water before and 40 min after the onset of exercise. During exercise in the Dh compared to C state, plasma renin, aldosterone, arginine vasopressin (AVP), noradrenaline and adrenaline concentrations were increased (P < 0.05). A reduction in creatinine clearance and urine flow was also observed (P < 0.05) together with a decrease in urine osmolality, osmolar clearance and sodium excretion, while free water clearance increased (P < 0.05). However, compared to Dh, Rh partially restored PV and osm_pl and induced a marked reduction in the time courses of both the plasma AVP and catecholamine responses (P < 0.05). Values for renal water and electrolyte excretion were intermediate between those of Dh and C. Plasma atrial natriuretic peptide presented similar changes whatever the hydration state. These results demonstrate that during moderate exercise in the heat, renal concentrating ability is paradoxically reduced by prior dehydration in spite of high plasma AVP levels, and might be the result of marked activation of the sympatho-adrenal system. Rehydration, by reducing this activation, could partially restore the renal concentrating ability despite the lowered plasma AVP. Accepted: 23 April 1997     

Effects of emergence date and maternal size on egg development and sizes of eggs and first-instar nymphs of a semelparous aquatic insect

We examined whether or not sizes of eggs and offspring were related to emergence date or maternal size in a semelparous aquatic insect (the burrowing mayfly, Hexagenia) in which parental care is lacking and oviposited eggs are passively dispersed. We quantified the size of males and female imagos over the emergence span at a site on the Detroit River, Canada, and investigated relationships between emergence date and female size and (1) egg size and (2) size of first-instar nymphs. Although size of female imagos (H. limbata and H. rigida combined) declined significantly (P<0.025) over the emergence season, there was no significant relationship between body length and emergence date for males of either species. Males were significantly (P<0.001) smaller than females. H. limbata eggs, subsampled from three individuals from each of three size classes of female imagos collected on seven sampling dates, were measured using video image analysis. Eggs (n=100) oviposited by each of 63 H. limbata imagos were inspected daily for hatching. Newly hatched nymphs were removed, counted and measured. Egg size (P<0.001) and size of first-instar nymphs (P<0.001) varied significantly with emergence date, but not maternal size. The largest eggs and newly hatched nymphs occurred at peak emergence of adults. The synchronous release of larger (faster-sinking) eggs may result in reduced predation. Plasticity in egg development time and egg and nymph size may account for the ability of this taxon to recover from episodes of massive population reduction. Received: 12 March 1996 / Accepted: 24 February 1997     

Effects of haemoglobin O2 saturation on volume regulation in adrenergically stimulated red blood cells from the trout, Oncorhynchus mykiss

Adrenergic stimulation of trout red blood cells activates a Na⁺/H⁺-exchange. If unopposed, the ensuing increase in cell Na⁺ leads to an isosmotic cell swelling. In this study the effect of the level of haemoglobin O₂ saturation on volume regulation has been investigated in adrenergically stimulated red blood cells from trout: at full haemoglobin O₂ saturation, net influx of Na⁺ through the Na⁺/H⁺-exchanger was balanced by net efflux of K⁺ and no increases in cell volume took place. In contrast, at low O₂ saturation (8–14%) adrenergic stimulation led to a substantial increase in cell Na⁺, K⁺ and volume. Moreover, cell volume recovery after adrenergic swelling was incomplete at low O₂ saturation, whereas cells at high O₂ saturation exhibited a fast and complete cell volume recovery. In cells exposed to alternating high and low O₂ saturation, volume regulation was similar to the regulation found in cells maintained at high O₂ saturation. In cells at high O₂ saturation, extrusion of cellular Na⁺ by the Na⁺/K⁺-pump significantly contributed to the volume decrease. It is concluded that trout red blood cells at high or alternating O₂ saturations possess a powerful regulatory volume decrease response that is shut off at low O₂ saturation. The physiological implications of this regulation is discussed. Accepted: 30 September 1996     

Acorn selection by Mexican jays: a test of a tri-trophic symbiotic relationship hypothesis

By caching acorns, jays serve as important dispersal agents for oak (Quercus) species. Yet little is known about which acorn characteristics affect selection by jays. In the traditional model of jay/oak symbiosis, large, brown, ripe acorns free of invertebrate parasites (e.g., Curculio acorn weevils) are selected by jays. Recently, it has been suggested that a tri-trophic relationship between oaks, jays, and weevils may have evolved to counter the negative dietary effects of acorn tannins. Under the tri-trophic model, jays would preferentially select acorns containing weevil larvae. We tested the assumptions that (1) acorns containing curculionid larvae exist in sufficient quantities to support jay populations and (2) jays can detect, and preferentially select, acorns containing weevil larvae, and investigated the cues by which jays select acorns. Captive Mexican jays (Aphelocomaultramarina) were presented Emory oak (Quercusemoryi) acorns in aviary feeding trials. Large, dense, viable acorns free of curculionid larvae were preferentially selected. Contrary to results of previous research, color did not affect selection. Acorn viability increased and curculionid larval occupancy decreased in adjacent savannas and isolated stands relative to existing oak woodland, perhaps favoring oak recruitment into adjacent lower-elevation grasslands. Our results compel us to reject the tri-trophic model for this system, and are consistent with the traditional jay/oak symbiosis model. Relatively long-distance dispersal of viable acorns favors Emory oak replacement, and spatial patterns of acorn viability and curculionid parasitism suggest expansion of Emory oak into adjacent low-elevation semi-arid grasslands. Received: 29 February 1996 / Accepted: 26 September 1996     

Somatic variation during long term subculturing of plant cells caused by insertion of a transposable element in a phenylalanine ammonia-lyase (PAL) gene

We have identified a new En/Spm-like transposable element, Tdc1, in the 5′ flanking region of a phenylalanine ammonia-lyase gene (gDcPAL1) that is normally induced by transferring cells of carrot suspension cultures to fresh liquid medium (transfer or dilution effect). The initial integration into gDcPAL1 occurred more than 4 years after culture initiation. Tdc1 was first detected in gDcPAL1 genomic clones of a genomic library made from cells of the same cultured cell line 7 years after its initiation and thus following repeated subculturing. Twelve years after initiation, about 5–10% of the cells had Tdc1 inserted into the gDcPAL1 gene, indicating that Tdc1 insertion into gDcPAL1 occurred in one (or more) cell(s) during the first 4–7 years of subculturing. These mutant cells did not disappear during numerous passages; instead the proportion of cells having this Tdc1 inserted into gDcPAL1 has been increasing over the last 5 years. The promoter activity and the inducibility by transfer/dilution of the gDcPAL1 gene harboring Tdc1 is reduced relative to wild type. Finally, we show that insertion of a transposable element is one of the mechanisms that can cause variation of plant cell cultures during repeated subculture. Received: 11 October 1996 / Accepted: 27 December 1996     

Shade-induced changes in the branching pattern of a stoloniferous herb: functional response or allometric effect?

Shade-induced changes in the branching pattern of clonal plants can lead to conspicuous modifications of their growth form and architecture. It has been hypothesized that reduced branching in shade may be an adaptive trait, enabling clonal plants to escape from unfavourable patches in a heterogeneous environment by allocating resources preferentially to the growth of the main axis (i.e. linear expansion), rather than to local proliferation by branching. However, such an adaptionist interpretation may be unjustified if (1) branching frequency is a function of the ontogenetic stage of plants, and if (2) shading slows down the ontogenetic development of plants, thereby delaying branch formation. In this case, architectural differences between sun- and shade-grown individuals, harvested at the same chronological age, may not represent a functional response to changes in light conditions, but may be a by-product of effects of shade on the rate of plant development. To distinguish between these two alternatives, individuals of the stoloniferous herb Potentilla reptans were subjected to three experimental light conditions: a control treatment providing full daylight, and two shade treatments: neutral shade (13% of ambient PPFD; no changes in light spectral composition) and simulated canopy shade (13% PPFD and a reduced red:far-red ratio). Plant development was followed throughout the experiment by daily monitoring primary stolon growth as well as branch and leaf initiation. Biomass and clonal offspring production were measured when plants were harvested. At the end of the experiment shaded plants had produced significantly fewer branches than clones grown in full daylight. In all three treatments, however, initiation of secondary stolons occurred at the same developmental stage of individual ramets. Shading significantly slowed down the ontogenetic development of plants and this resulted in the observed differences in branching patterns between sun- and shade-grown individuals, when compared at the same chronological age. These results hence provide evidence that shade-induced changes in the branching pattern of clonal plants can be due to purely allometric effects. Implications for interpreting architectural changes in terms of functional shade-avoidance responses are discussed. Received: 16 August 1996 / Accepted: December 1996     

Partial purification of mitochondrial ribosomes from broad bean and identification of proteins encoded by the mitochondrial genome

An approach towards the identification at the protein level of the ribosomal proteins encoded by the mitochondrial genome of broad bean (Vicia faba) has been developed. After Triton X-100 treatment of isolated mitochondria, a fraction enriched in mitochondrial ribosomes was obtained by successive centrifugation, first onto a sucrose cushion, and then in a sucrose gradient. Mitochondrial translation products were labelled in isolated mitochondria with [³⁵S]methionine and added to the enriched mitochondrial ribosomal proteins before separation by two-dimensional gel electrophoresis. Six spots, identified both by Coomassie blue staining and autoradiography, were analysed by protein micro-sequencing. Two of these were shown to correspond to ribosomal proteins S10 and S12. We conclude that these two proteins are encoded by the mitochondrial genome of broad bean and that the method described here can be used to identify other proteins encoded by the mitochondrial genome. Received: 4 September 1996 / Accepted: 30 November 1996     

The effect of a thiamin derivative on exercise performance

The purpose of this study was to investigate the effect of a thiamin derivative, thiamin tetrahydrofurfuryl disulfide (TTFD), on oxygen uptake (˙VO₂), lactate accumulation and cycling performance during exercise to exhaustion. Using a randomized, double-blind, cross-over design with a 10-day washout between trials, 14 subjects ingested either 1 g · day⁻¹ of TTFD or a placebo (PL) for 4 days. On day 3, subjects performed a progressive exercise test to exhaustion on a cycle ergometer for the determination of ˙VO_2submax, ˙VO_2peak, lactate concentration ([La⁻ ]), lactate threshold (Th_La) and heart rate ( f _c). On day 4, subjects performed a maximal 2000-m time trial on a cycle ergometer. A one-way analysis of variance (ANOVA) with repeated measures was used to determine significant differences between trials. There were no significant differences detected between trials for serial measures of ˙VO_2submax, [La⁻] or f _c. Likewise, ˙VO_2peak [PL 4.06 (0.19) TTFD 4.12 (0.19) l · min⁻¹, P = 0.83], Th_La [PL 2.47 (0.17), TTFD 2.43 (0.16) l · min⁻¹, P = 0.86] and 2000-m performance time [PL 204.5 (5.5), TTFD 200.9 (4.3) s, P = 0.61] were not significantly different between trials. The results of this study suggest that thiamin derivative supplementation does not influence high-intensity exercise performance. Accepted: 19 December 1996     

Oxygen consumption in the foraging honeybee depends on the reward rate at the food source

Oxygen consumption of the honeybee Apis mellifera ligustica was measured as a function of the flow rate supply of sucrose solution at an automatic feeder located inside a respirometric chamber. Trained bees freely entered the respirometric chamber and collected the sucrose solution supplied. The mean value of the O₂ consumption rate per visit increased with the sucrose flow rate, and for a given flow rate, with increasing locomotor activity. However, when no locomotor activity was displayed, O₂ consumption also increased with increasing nectar flow rate. Crop load attained at the end of the visit showed a positive relationship with the nectar flow rate; however, for a given flow rate, O₂ consumption showed either no correlation or a negative one with the final crop load attained. It is concluded that the energy expenditure of the foraging bee is controlled by a motivational drive whose intensity depends on the reward rate at the food source. Accepted: 30 July 1996