Flight of the honey bee VI: energetics of wind tunnel exhaustion flights at defined fuel content,speed adaptation and aerodynamics

To gain information on extended flight energetics, quasi-natural flight conditions imitating steady horizontal flight were set by combining the tetheredflight wind-tunnel method with the exhaustion-flight method. The bees were suspended from a two-component aerodynamic balance at different, near optimum body angle of attack and were allowed to choose their own speed: their body mass and body weight was determined before and after a flight; their speed, lift, wingbeat frequency and total flight time were measured throughout a flight. These values were used to determine thrust, resultant aerodynamic force (magnitude and tilting angle), Reynolds number, total flight distance and total flight impulse. Flights in which lift was body weight were mostly obtained. Bees, flown to complete exhausion, were refed with 5, 10, 15 or 20 l of a 1.28-mol·l^-1 glucose solution (energy content w=18.5, 37.0, 55.5 or 74.0 J) and again flown to complete exhaustion at an ambient temperature of 25±1.5°C by a flight of known duration such that the calculation of absolute and relative metabolic power was possible. Mean body mass after exhaustion was 76.49±3.52 mg. During long term flights of 7.47–31.30 min similar changes in flight velocity, lift, thrust, aerodynamic force, wingbeat frequency and tilting angle took place, independent of the volume of feeding solution. After increasing rapidly within 15 s a more or less steady phase of 60–80% of total flight time, showing only a slight decrease, was followed by a steeper, more irregular decrease, finally reaching 0 within 20–30 s. In steady phases lift was nearly equal to resultant aerodynamic force; tilting angle was 79.8±4.0°, thrust to lift radio did not vary, thrust was 18.0±7.4% of lift, lift was somewhat higher/equal/lower than body mass in 61.3%, 16.1%, 22.6% of all totally analysable flights (n=31). The following parameters were varied as functions of volume of feeding solution (5–20 l in steps of 5 l) and energy content. (18.5–74.0 J in steps of 18.5 J): total flight time, velocity, total flight distance, mean lift, thrust, mean resultant aerodynamic force, tilting angle, total flight impulse, wingbeat frequency, metabolic power and metabolic power related to body mass, the latter related to empty, full and mean (=100 mg) body mass. The following positive correlations were found: L=1.069·10^-9 f ^2.538; R=1.629·10^-9 f ^2.464; P _m=7.079·10^-8 f ^2.456; P _m=0.008v+0.008; P _m=18.996L+0.022; P _m=19.782R+0.021; P _m=82.143T+0.028; P _m=1.245·bm _f ^1.424 ; P _{mrel e}=6.471·bm _f ^1.040 ; =83.248+0.385. The following negative correlations were found: V=3.939–0.032; T=1.324·10^-4–0.038·10^-4. Statistically significant correlations were not found in T(f), L(), R(), f(), P _m(bm _e), P _{m rel e}(bm _e), P _{m rel f}(bm _e), P _{m rel f}(bm _f).Abbreviations A(m²) frontal area - bl(m) body length - bm(mg) body mass - c(mol·1^-1) glucose concentration of feeding solution - c _D (dimensionless) drag coefficient, related to A - D(N) drag - F _w(N) body weight - F _wp weight of paper fragment lost at flight start - f wingbeat frequency (s^-1) - g(=9.81 m·s^-2) gravitational acceleration - I(Ns)=R(t) dt total impulse of a flight - L(N) lift vertical sustaining force component - P _m(J·s^-1=W) metabolic power - P_{m ret} (W·g^-1) metabolic power, related to body mass - R(N) resultant aerodynamic force - Re v·bl·v ^-1 (dimensionless) Reynolds number, related to body length - s(m) v(t) dt virtual flight distance of a flight - s(km) total virtual flight distance - T (N) thrust horizontal force component of horizontal flight - T _a (°C) ambient temperature - t(s) time - t _tot (s or min) total flight time - v(m·s^-1) flight velocity - v(l) volume of feeding solution - W (J) energy and energy content of V - ( °) body angle of attack between body longitudinal axis and flow direction - ( °) tilting angle ( 90°) between R and the horizont in horizontal flight v(=1.53·10^-5m²·s^-1 for air at 25°) kinematic viscosity - (=1.2 kg·m^-3 at 25°C) air density     

Floral Lipid Chemistry of Byrsonima crassifolia (Malpigheaceae) and a Use of Floral Lipids by Centris Bees (Hymenoptera: Apidae)

Byrsonima rrassifolia (L.), a Neotropical malpighiaceous tree or treelet, has flowers that produce abundant lipids instead of nectar. Several species of Centris bees collect this oil. The floral oil shows variation between trees and can be separated into two types. One type is most common (11 of 14 samples) and consists of mono- and di-glycerides, some free fatty acids, a small amount of tri-glycerides and a trace of carbohydrate. The composition of the glycerides is predominantly (53–63%) esters of unsaturated 18C–22C fatty acids with the rest made up of 14C–18C saturated fatty acids. The second type of floral oil is similar to type one except that the samples contain large amounts of an unknown lipid more polar than the standards. Oil collected from the scopae of two representative Centris species, C. adanae and C. flavofasciata, was the same as the type one oil collected from B. rrassifolia floral elaiophores. The liquid provisions from the cells of several groundnesting Centris species was also found to be identical to B. crassifolia type one floral elaiophore oil with the exception of several minor ninhydrin positive compounds which may have been derived from the pollen which makes up part of the larval food provisions.     

Trapline foraging by bumblebees: I. Persistence of flight-path geometry

By setting out arrays of potted plants of Penstemon strictus,I tested whether freely foraging bumblebee (Bombus spp. ) workerswould establish regular foraging routes that reflected the geometryof the array. They did, passing through an asymmetrical arrayin a pattern that minimized interplant flight distances. Afterthe array was changed to a symmetrical pattern, however, theexperienced bees continued to show their previous asymmetricalflight patterns. New bees without experience on the asymmetricalarray showed no asymmetry on the symmetrical array. I term thispersistence of flight-path geometry "trapline holdover, " anddiscuss its implications for the study of animals' learningand foraging behavior.     

Local bumble bee decline linked to recovery of honey bees,drought effects on floral resources

Time series of abundances are critical for understanding how abiotic factors and species interactions affect population dynamics, but are rarely linked with experiments and also scarce for bee pollinators. This gap is important given concerns about declines in some bee species. I monitored honey bee (Apis mellifera) and bumble bee (Bombus spp.) foragers in coastal California from 1999, when feral A. mellifera populations were low due to Varroa destructor, until 2014. Apis mellifera increased substantially, except between 2006 and 2011, coinciding with declines in managed populations. Increases in A. mellifera strongly correlated with declines in Bombus and reduced diet overlap between them, suggesting resource competition consistent with past experimental results. Lower Bombus numbers also correlated with diminished floral resources. Declines in floral abundances were associated with drought and reduced spring rainfall. These results illustrate how competition with an introduced species may interact with climate to drive local decline of native pollinators.     

Bumble bee behavior and selection on flower size in the sky pilot,Polemonium viscosum

Summary In alpine Polemonium viscosum, plants having sweet-scented flowers are primarily pollinated by queens of the bumble bee species, Bombus kirbyellus. In this paper we ask whether two aspects of the pollination effectiveness of bumble bees, visitation rate and pollination efficiency, vary significantly with flower size in sweet-flowered P. viscosum.(i) Bumble bees visited plants with large flowers on 80–90% of encounters, but visited those with smaller flowers on only 49% of encounters. (ii) However, the gain in pollination that large-flowered plants obtained via increased visitation was countered in part because bumble bees deposited fewer outcross pollen grains per visit on stigmas of large flowers than on those of small ones. When both visitation rate and pollination efficiency are taken into account, the predicted value of a single bumble bee encounter declines from 1.06 seeds for flowers larger than 18 mm in diameter to 0.55 seeds for flowers smaller than 12 mm in diameter. Our results suggest that bumble bee pollinators of P. viscosum prefer flower morphologies that are poorly suited for precise pollination. Such behavioral complexities are likely to place constraints on the evolution of optimal floral design.     

Sterol utilization in honey bees fed a synthetic diet: Analysis of prepupal sterols

The sterols of prepupal honey bees, Apis mellifera L., from brood reared by workers fed chemically-defined synthetic diets containing cholesterol, campesterol, sitosterol, stigmasterol, 24-methylenecholesterol, or no sterol over a 12-week period were isolated, identified, and quantified. The major sterol present in each prepupal sample was 24-methylenecholesterol, but significant levels of sitosterol and isofucosterol were also present in every case, as was a very small percentage of desmosterol (usually < 1%). This is the first report of isofucosterol being identified in the sterols of the honey bee. A considerably larger percentage of each dietary sterol was found in prepupae reared by workers fed that particular sterol in the diet. This was most dramatic in the case of the cholesterol diet in which case cholesterol content increased to as much as 17.2% of the prepupal sterols, whereas cholesterol had not exceeded 2.2% in samples from other diet regimens. However, stigmasterol comprised no more than 6.3% of the total sterols in any sample from prepupae fed the stigmasterol diet. The preponderance of 24-methylenecholesterol in all prepupae, regardless of the dietary sterol provided to the workers, as well as the lesser quantities of sitosterol and isofucosterol present in all samples, suggest a unique system of utilization and metabolism of these dietary sterols by the worker bees. Apparently they make available to the brood varying amounts of unchanged dietary sterol plus considerable and fairly constant portions of 24-methylenecholesterol, sitosterol, and isofucosterol drawn from their own sterol pools.     

Nonrandom visitation of brood cells by worker honey bees (Hymenoptera: Apidae)

The visitation pattern by worker honey bees to cells in the brood nest was monitored on an artificially created brood pattern consisting of about one-fourth brood cells evenly distributed among empty cells. The majority (63 %) of the observed workers selectively entered larval cells. In contrast, some workers avoided egg cells when presented a choice of egg vs empty cells. The results suggest that larvae produce a general signal indicating their presence to worker bees. Eggs also seem to produce a signal, which is perceived to be different from the one from larvae.     

Oxygen consumption and flight muscle activity during heating in workers and drones of Apis mellifera

Summary Instantaneous oxygen consumption, muscle potential frequency, thoracic and ambient temperature were simultaneously measured during heating in individual workers and drones of honey bees. Relationships between these parameters and effects of thoracic temperature on power input and temperature elevation were studied. Oxygen consumption increased above basal levels only when flight muscles became active. Increasing muscle potential frequencies correlated with elevated oxygen consumption and raised thoracic temperature. The difference between thoracic and ambient temperature and oxygen consumption were linearly related. Oxygen consumption per muscle potential (l O₂ · g _–1 ^thorax · MP^–1) was two-fold higher in drones than in workers. However, oxygen consumption for heating the thorax (l O₂ · g _–1 ^thorax · (T_th-T_a) · °C^–1) was nearly the same in workers and drones. Thoracic temperature affected the amount of oxygen consumed per muscle potential (R₁₀=1.5). Achieved temperature elevation per 100 MP was more temperature sensitive in drones (R₁₀=6–10) than in workers (R₁₀=3.6). Q₁₀ values for oxygen consumption were 3 in workers and 4.5–6 in drones. Muscle potential frequency decreased with a Q₁₀=1.8 in workers and 2.7 in drones. Heating behaviour of workers and drones was different. Drones generated heat less continuously than workers, and showed greater interindividual variability in predilection to heat. However, the maximal difference between ambient and thoracic temperature observed was 22 °C in drones and 14 °C in workers, indicating greater potential for drones.Abbreviations DL dorsal-longitudinal muscle - DV dorsoventral muscle - MP muscle potential - T _a ambient temperature - T _th thoracic temperature     

The spatial precision of the honey bees' dance communication

Many social insect species that employ directional recruitment to food finds communicate the location of their targets with seemingly needless imprecision. Here we present evidence that the spatial precision of the honey bees' dance communication has been tuned by selection so that recruits are neither so accurate that they usually find areas which have already been depleted nor so inaccurate that they usually fail to find the advertised resources altogether. First, the bees' distance errors are similar in magnitude to their directional errors, and their angular errors decrease greatly with increasing distance to the target (decreasing more than fourfold between 100 and 700 m). As a result, the absolute scatter of recruits remains relatively constant with changing distance to the target (increasing by less than 50% between 100 and 700 m). This is to be expected if the optimal level of imprecision is the same for distance and direction and does not change with the distance of the target from the nest. Second, comparative studies involving three tropical- and one temperate-zone species (all Apis)suggest that the precision of the bees' languages may have been tuned in accordance with the spatial characteristics of the resources each species uses. We suggest that both the round dance, which conveys no directional information for nearby targets, and the high angular divergence in waggle dances indicating targets within several hundred meters of the colony are both understandable in this context.