The functional organization of resin work in honeybee colonies

Resin is an important building material in the nests of honeybees, but little is known about how it is handled within the nest and how its collection is controlled. We studied the functional organization of resin work to better understand how a colony adaptively controls its intake of resin. Two hypotheses have been proposed for how resin collectors stay informed of the need for additional resin: (1) the unloading difficulty hypothesis (resin need is sensed indirectly by the unloading delay) and (2) the caulking activity hypothesis (resin need is sensed directly while engaged in using resin). A falsifiable prediction of the latter hypothesis, but not of the former, is that resin collectors not only gather resin outside the hive but also regularly handle resin inside the hive (taking it from other bees and using it to caulk crevices). Consistent with this prediction are our findings that in the resin sector of a colony’s economy, unlike in the pollen, nectar, and water sectors, there is no strict division of labor between the collectors and the users of a material. Over the course of a day, bees seen collecting resin were also commonly seen using resin. Moreover, we found that the unloading locations of resin collectors are unlike those of water and nectar collectors, being deep inside the hive (at the sites of resin use) rather than at the hive entrance. This arrangement facilitates the engagement in resin use by resin collectors. We conclude that the caulking activity hypothesis is well-supported, but that the unloading difficulty hypothesis also remains viable, for we found that resin collectors experience variable delays in getting rid of their loads, from less than 15 min to more than an hour, consistent with this hypothesis. The stage is now set for experimental tests of these two hypotheses. Both may be correct, which if true will imply that social insect workers, despite their small brains, can acquire and integrate information from multiple sources to improve their knowledge of conditions within the colony.     

Honey bee foragers as sensory units of their colonies

Forager honey bees function not only as gatherers of food for their colonies, but also as sensory units shaped by natural selection to gather information regarding the location and profitability of forage sites. They transmit this information to colony members by means of waggle dances. To investigate the way bees transduce the stimulus of nectar-source profitability into the response of number of waggle runs, I performed experiments in which bees were stimulated with a sucrose solution feeder of known profitability and their dance responses were videorecorded. The results suggest that several attributes of this transduction process are adaptations to enhance a bee's effectiveness in reporting on a forage site. (1) Bees register the profitability of a nectar source not by sensing the energy gain per foraging trip or the rate of energy gain per trip, but evidently by sensing the energetic efficiency of their foraging. Perhaps this criterion of nectar-source profitability has been favored by natural selection because the foraging gains of honey bees are typically limited by energy expenditure rather than time availability. (2) There is a linear relationship between the stimulus of energetic efficiency of foraging and the response of number of waggle runs per dance. Such a simple stimulus-response function appears adequate because the range of suprathreshold stimuli (max/min ratio of about 10) is far smaller than the range of responses (max/min ratio of about 100). Although all bees show a linear stimulus-response function, there are large differences among individuals in both the response threshold and the slope of the stimulus-response function. This variation gives the colony a broader dynamic range in responding to food sources than if all bees had identical thresholds of dance response. (3) There is little or no adaptation in the dance response to a strong stimulus (tonic response). Thus each dancing bee reports on the current level of profitability of her forage site rather than the changes in its profitability. This seems appropriate since presumably it is the current profitability of a forage site, not the change in its profitability, which determines a site's attractiveness to other bees. (4) The level of forage-site quality that is the threshold for dancing is tuned by the bees in relation to forage availability. Bees operate with a lower dance threshold when forage is sparse than when it is abundant. Thus a colony utilizes input about a wide range of forage sites when food is scarce, but filters out input about low-reward sites when food is plentiful. (5) A dancing bee does not present her information in one spot within the hive but instead distributes it over much of the dance floor. Consequently, the dances for different forage sites are mixed together on the dance floor. This helps each bee following the dances to take a random sample of the dance information, which is appropriate for the foraging strategy of a honey bee colony since it is evidently designed to allocate foragers among forage sites in proportion to their profitability.     

Use of the ouput of a lead risk assessment model to establish soil lead cleanup levels

Three general methods to calculate soil contaminant cleanup levels are assessed: the truncated lognormal approach, Monte Carlo analysis, and the house-by-house approach. When these methods are used together with a lead risk assessment model, they yield estimated soil lead cleanup levels that may be required in an attempt to achieve specified target blood lead levels for a community. The truncated lognormal approach is exemplified by the Society for Environmental Geochemistry and Health (SEGH) model, Monte Carlo analysis is exemplified by the US EPA's LEAD Model, and the house-by-house approach is used with a structural equation model to calculate site-specific soil lead cleanup levels. The various cleanup methods can each be used with any type of lead risk assessment model. Although all examples given here are for lead, the cleanup methods can, in principle, be used as well with risk assessment models for other chemical contaminants to derive contaminant-specific soil cleanup levels.     

Uptake and metabolism of [<Superscript>14</Superscript>C]rinderine and [<Superscript>14</Superscript>C]retronecine in leaf-beetles of the genus <Emphasis Type="Italic"> Platyphora</Emphasis> and alkaloid accumulation in the exocrine defensive secretions

Summary. Sequestration and processing of pyrrolizidine alkaloids (PAs) by leaf beetles of the genus Platyphora were investigated. Tracer experiments with labeled alkaloids were performed with P. eucosma feeding on Koanophyllon panamense (Asteraceae, tribe Eupatorieae). P. eucosma catalyzes the same reactions previously demonstrated for P. boucardi specialized to Prestonia portobellensis (Apocynaceae): (i) epimerization of rinderine to intermedine; (ii) esterification of retronecine yielding insect-specific PAs; (iii) efficient transport of the PAs as free bases into the defensive secretions. P. bella feeding on Tournefortia cuspidata (Boraginaceae) shows the same sequestration behavior and ability to synthesize the specific retronecine esters. P. ligata, a species phylogenetically closely related to the PA adapted species and clustering in the same clade, but feeding on a host plant devoid of PAs, feeds easily on PA treated host-plant leaves, but does not sequester or metabolize PAs. P. kollari a species clustering outside the PA clade refused to feed on its food-plant leaves painted with PAs. The results are discussed in relation to host-plant selection of the PA adapted species and the role of PAs in chemical defense. Received 20 September 2002; accepted 18 November 2002.     

Social foraging in honey bees: how nectar foragers assess their colony's nutritional status

Summary A honey bee colony operates as a tightly integrated unit of behavioral action. One manifestation of this in the context of foraging is a colony's ability to adjust its selectivity among nectar sources in relation to its nutritional status. When a colony's food situation is good, it exploits only highly profitable patches of flowers, but when its situation is poor, a colony's foragers will exploit both highly profitable and less profitable flower patches. The nectar foragers in a colony acquire information about their colony's nutritional status by noting the difficulty of finding food storer bees to receive their nectar, rather than by evaluating directly the variables determining their colony's food situation: rate of nectar intake and amount of empty storage comb. (The food storer bees in a colony are the bees that collect nectar from returning foragers and store it in the honey combs. They are the age group (generally 12–18 day old bees) that is older than the nurse bees but younger than the foragers. Food storers make up approximately 20% of a colony members.) The mathematical theory for the behavior of queues indicates that the waiting time experienced by nectar foragers before unloading to food storers (queue length) is a reliable and sensitive indicator of a colony's nutritional status. Queue length is automatically determined by the ratio of two rates which are directly related to a colony's nutritional condition: the rate of arrival of loaded nectar foragers at the hive (arrival rate) and the rate of arrival of empty food storers at the nectar delivery area (service rate). These two rates are a function of the colony's nectar intake rate and its empty comb area, respectively. Although waiting time conveys crucial information about the colony's nutritional status, it has not been molded by natural selection to serve this purpose. Unlike signals, which are evolved specifically to convey information, this cue conveys information as an automatic by-product. Such cues may prove more important than signals in colony integration.     

Social foraging by honeybees: how colonies allocate foragers among patches of flowers

Summary To understand how a colony of honeybees keeps its forager force focussed on rich sources of food, and analysis was made of how the individual foragers within a colony decide to abandon or continue working (and perhaps even recruit to) patches of flowers. A nectar forager grades her behavior toward a patch in response to both the nectar intake rate of her colony and the quality of her patch. This results in the threshold in patch quality for acceptance of a patch being higher when the colonial intake rate of nectar is high than when it is low. Thus colonies can adjust their patch selectivity so that they focus on rich sources when forage is abundant, but spread their workers among a wider range of sources when forage is scarce. Foragers assess their colony's rate of nectar intake while in the nest, unloading nectar to receiver bees. The ease of unloading varies inversely with the colonial intake rate of nectar. Foragers assess patch quality while in the field, collecting nectar. By grading their behavior steeply in relation to such patch variables as distance from the nest and nectar sweetness, foragers give their colony high sensitivity to differences in profitability among patches. When a patch's quality declines, its foragers reduce their rate of visits to the patch. This diminishes the flow of nectar from the poor patch which in turn stimulates recruitment to rich patches. Thus a colony can swiftly redistribute its forager force following changes in the spatial distribution of rich food sources. The fundamental currency of nectar patch quality is not net rate of energy intake, (Gain-Cost)/Time, but may be net energy efficiency, (Gain-Cost)/Cost.     

The use of waggle dance information by honey bees throughout their foraging careers

We studied the extent to which worker honey bees acquire information from waggle dances throughout their careers as foragers. Small groups of foragers were monitored from time of orientation flights to time of death and all in-hive behaviors relating to foraging were recorded. In the context of a novice forager finding her first food source, 60% of the bees relied, at least in part, on acquiring information from waggle dances (being recruited) rather than searching independently (scouting). In the context of an experienced forager whose foraging has been interrupted, 37% of the time the bees resumed foraging by following waggle dances (being reactivated) rather than examining the food source on their own (inspecting). And in the context of an experienced forager engaged in foraging, 17% of the time the bees initiated a foraging trip by following a waggle dance. Such dance following was observed much more often after an unsuccessful than after a successful foraging trip. Successful foragers often followed dances just briefly, perhaps to confirm that the kind of flowers they had been visiting were still yielding forage. Overall, waggle dance following for food discovery accounted for 12–25% of all interactions with dancers (9% by novice foragers and 3–16% by experienced foragers) whereas dance following for reactivation and confirmation accounted for the other 75–88% (26% for reactivation and 49–62% for confirmation). We conclude that foragers make extensive use of the waggle dance not only to start work at new, unfamiliar food sources but also to resume work at old, familiar food sources.     

Potential effects of forest policies on terrestrial biodiversity in a multi-ownership province.

We used spatial simulation models to evaluate how current and two alternative policies might affect potential biodiversity over 100 years in the Coast Ranges Physiographic Province of Oregon. This 2.3-million-ha province is characterized by a diversity of public and private forest owners, and a wide range of forest policy and management objectives. We evaluated habitat availability for seven focal species representing different life histories. We also examined how policies affected old-growth stand structure, age distributions relative to the historical range of variability, and landscape patterns of forest types. Under the current policy scenario, the area of habitat for old-growth forest structure and associated species increased over time, the habitat for some early-successional associates remained stable, and the area of hardwood vegetation and diverse early-successional stages declined. The province is projected to move toward but not reach the historical range of variation of forest age classes that may have occurred under the wildfire regimes of the pre-Euroamerican settlement period. Ownership explained much of the pattern of biodiversity in the province, and under the current policy scenario, its effect increased over time as the landscape diverged into highly contrasting forest structures and ages. Patch type diversity declined slightly overall but declined strongly within ownerships. Most of the modeled change in biodiversity over time resulted from policies on public forest lands that were intended to increase the area of late-successional forests and species. One of the alternative policies, increased retention of wildlife trees on private lands, reduced the contrast between ownerships and increased habitat availability over time for both early- and late-successional species. Analysis of another alternative, stopping thinning of plantations on federal lands, indicated that current thinning regimes improve habitat for the Olive-sided Flycatcher, but the no-thinning alternative had no effect on the habitat scores for the late-successional species in the 100-year simulation. A comparison of indicators of biological diversity suggests that using focal species and forest structural measures can provide complementary information on biodiversity. The multi-ownership perspective provided a more complete synthesis of province-wide biodiversity patterns than assessments based on single ownerships.     

Influence of environment, disturbance, and ownership on forest vegetation of coastal Oregon.

Information about how vegetation composition and structure vary quantitatively and spatially with physical environment, disturbance history, and land ownership is fundamental to regional conservation planning. However, current knowledge about patterns of vegetation variability across large regions that is spatially explicit (i.e., mapped) tends to be general and qualitative. We used spatial predictions from gradient models to examine the influence of environment, disturbance, and ownership on patterns of forest vegetation biodiversity across a large forested region, the 3-million-ha Oregon Coast Range (USA). Gradients in tree species composition were strongly associated with environment, especially climate, and insensitive to disturbance, probably because many dominant tree species are long-lived and persist throughout forest succession. In contrast, forest structure was strongly correlated with disturbance and only weakly with environmental gradients. Although forest structure differed among ownerships, differences were blurred by the presence of legacy trees that originated prior to current forest management regimes. Our multi-ownership perspective revealed biodiversity concerns and benefits not readily visible in single-ownership analyses, and all ownerships contributed to regional biodiversity values. Federal lands provided most of the late-successional and old-growth forest. State lands contained a range of forest ages and structures, including diverse young forest, abundant legacy dead wood, and much of the high-elevation true fir forest. Nonindustrial private lands provided diverse young forest and the greatest abundance of hardwood trees, including almost all of the foothill oak woodlands. Forest industry lands encompassed much early-successional forest, most of the mixed hardwood-conifer forest, and large amounts of legacy down wood. The detailed tree- and species-level data in the maps revealed regional trends that would be masked in traditional coarse-filter assessment. Although abundant, most early-successional forests originated after timber harvest and lacked legacy live and dead trees important as habitat and for other ecological functions. Many large-conifer forests that might be classified as old growth using a generalized forest cover map lacked structural features of old growth such as multilayered canopies or dead wood. Our findings suggest that regional conservation planning include all ownerships and land allocations, as well as fine-scale elements of vegetation composition and structure.