Experimental evidence of a tripartite mutualism: bacteria protect ant fungus gardens from specialized parasites

Symbioses shape all levels of biological organization. Although symbiotic interactions are typically viewed as bipartite associations, with two organisms interacting largely in isolation from other organisms, the presence and importance of additional symbionts is becoming increasingly more apparent. This study examines the importance of a third mutualist within the ancient symbiosis between leaf-cutting ants and their fungal cultivars. Specifically, we experimentally examine the role of a filamentous bacterium (actinomycete), which is typically carried on the cuticle of fungus-growing ants, in suppressing the growth of a specialized microfungal parasite ( Escovopsis ) of the fungus garden. We conducted two-by-two factorial design experiments crossing the presence/absence of actinomycete with the presence/absence of Escovopsis within small sub-colonies of Acromyrmex octospinosus . In these experiments, infection by Escovopsis became much more extensive within fungus gardens and had a greater impact on the health of gardens in those sub-colonies with the bacterium removed from workers as compared to gardens with the bacterium still present on the ants. We establish that the actinomycete bacterium is most abundant on those major workers tending the garden, providing further support that the bacterium is involved in garden hygiene. We also found a significantly higher abundance of actinomycete on workers in colonies experimentally infected with Escovopsis as compared to uninfected control colonies. We suggest that mutualisms between antibiotic-producing microbes and higher organisms may be common associations that are mostly overlooked and that the role of symbionts in reducing the impact of parasites is likely an important aspect in the cost-benefit assessment of mutualisms.     

Foraging decisions of individual workers vary with colony size in the greenhead ant Rhytidoponera metallica (Formicidae, Ectatomminae)

Summary. The ability of worker ants to adapt their behaviour depending on the social environment of the colony is imperative for colony growth and survival. In this study we use the greenhead ant Rhytidoponera metallica to test for a relationship between colony size and foraging behaviour. We controlled for possible confounding ontogenetic and age effects by splitting large colonies into small and large colony fragments. Large and small colonies differed in worker number but not worker relatedness or worker/brood ratios. Differences in foraging activity were tested in the context of single foraging cycles with and without the opportunity to retrieve food. We found that workers from large colonies foraged for longer distances and spent more time outside the nest than foragers from small colonies. However, foragers from large and small colonies retrieved the first prey item they contacted, irrespective of prey size. Our results show that in R. metallica, foraging decisions made outside the nest by individual workers are related to the size of their colony.Received 23 March 2004; revised 3 June 2004; accepted 4 June 2004.     

The allocation of foragers in red wood ants

Abstract. 1. We studied how colonies of the red wood ant, Formica polyctena , adjust the numbers of foragers allocated to different foraging trails. In a series of field experiments, foragers were marked and transferred from one nest to another, related nest, where they joined the foraging force. Transferred workers acted as a reserve of uncommitted, available foragers.
2. Previous work shows that each individual forager habitually uses one trail. We found that for an uncommitted forager, the influence of recruitment initially is stronger than that of directional fidelity. Transferred workers were likely to use trails leading to new food sources. When transferred to a new nest, foragers were not likely to use a trail in the same direction as their original trail in the donor nest.
3. After a week, transferred foragers tended to develop route fidelity. Even after bait was no longer present, they continued to use the trail that had formerly led to a bait source.
4. We examined how colonies adjust numbers on a trail by experimentally depleting some trails. Colonies usually did not compensate for depletion: foragers were not recruited to depleted trails.
5. In general, the dynamics of foraging in this species facilitate a consistent foraging effort rather than rapid adjustments of forager allocation.     

Energetics of newly-mated queens and colony founding in the fungus-gardening ants Cyphomyrmex rimosus and Trachymyrmex septentrionalis (Hymenoptera: Formicidae)

Abstract.  The energetics of colony founding is investigated in the fungus gardening ants (Attini) Trachymyrmex septentrionalis and Cyphomyrmex rimosus . Similar to most ants, inseminated queens of these two species found nests independently unaccompanied by workers (haplometrosis). Whereas most ant founding queens seal themselves in a chamber and do not feed when producing a brood entirely from metabolic stores (claustral founding), the majority of fungus gardening ants must forage during the founding phase (semiclaustral founding). Laboratory-reared T. septentrionalis individuals comprise 84 dealate females collected after mating flights in June 2004. Twenty are immediately killed to obtain values for queen traits and another 20 after worker emergence for queen, fungus garden and worker traits. Cyphomyrmex rimosus comprise 22 dealate females collected in June 2005; ten of which are immediately killed and similarly prepared. Newly-mated T. septentrionalis queens have 25% of their dry weight as fat; whereas newly-mated C. rimosus queens contain 11% fat. These amounts are 50–75% less than most independently founding ant species. Trachymyrmex septentrionalis queens lose merely 5% of their energetic content during colony founding, whereas the total energetic content of their brood is more than three-fold the amount lost by the queen. Incipient T. septentrionalis colonies produce approximately half as much ant biomass per gram of fungus garden as do mature colonies. Similar to most ants, T. septentrionalis produces minim workers that are approximately 40% lighter than workers from mature colonies. Regardless of their size, T. septentrionalis workers contain much lower fat than do workers of claustral species. These data indicate that fungus gardening is adaptive because colonies can produce much cheaper offspring, making colony investment much lower.     

Fitness consequences of nest infiltration by the mutualist‐exploiter Megalomyrmex adamsae

1. Fungus‐growing ants are obligate mutualists. Their nutrient‐rich fungus garden provides a valuable food store that sustains the ant hosts, but can also attract social parasites. 2. The ‘guest ant' Megalomyrmex adamsae Longino parasitises the fungus‐growing Trachymyrmex zeteki Weber queen just after nest founding. The parasitic queen infiltrates the incipient nest, builds a cavity in the fungal garden, and lays eggs that develop into workers and reproductive males and females. 3. This study compared young parasitised and non‐parasitised laboratory colonies by measuring garden growth and biomass, and the number of host workers and reproductives. Host queen survival and parasite colony growth were also monitored. 4. Parasitised Trachymyrmex colonies had reduced host worker and alate numbers, as well as lower garden biomass, compared with non‐parasitised control colonies, confirming that M. adamsae is a xenobiotic social parasite. Host queen survival was not significantly different between parasitised and control colonies. 5. This is the first study that experimentally infects host colonies with a xenobiotic social parasite to measure fitness cost to the host. The natural history of M. adamsae and the fungus‐growing ant mutualism are evaluated in the context of three general predictions of (Bronstein, Ecology Letters, 4 , 277–287, 2001a) regarding the cost of mutualism exploiters.     

Caste specialization in behavioral defenses against fungus garden parasites in <Emphasis Type="Italic">Acromyrmex octospinosus</Emphasis> leaf-cutting ants

Division of labor and caste specialization plays an important role in many aspects of social insect colony organization, including parasite defense. Within leaf-cutting ant colonies, worker caste specialization permeates colony tasks ranging from foraging, substrate incorporation, brood care, and chemical defenses via glandular secretions and mutualistic bacteria. Leaf-cutting ants rely on protecting a mutualistic fungus they grow for food from microfungi in the genus Escovopsis that parasitizes the ant–fungus relationship. Here, we examine whether Acromyrmex octospinosus leaf-cutter ant castes (minors and majors) display task specialization in two behavioral defenses against Escovopsis: fungus grooming (the removal of Escovopsis spores) and weeding (the removal of large pieces of Escovopsis-infected fungus garden). Using behavioral observations, we show that minors are the primary caste that performs fungus grooming, while weeding is almost exclusively performed by majors. In addition, using a sub-colony infection experimental setup, we show that at the early stages of infection, minors more efficiently remove Escovopsis spores from the fungus garden, thereby restricting Escovopsis spore germination and growth. At later stages of infection, after Escovopsis spore germination, we find that major workers are as efficient as minors in defending the fungus garden, likely due to the increased importance of weeding. Finally, we show, using SEM imaging, that the number of sensory structures is similar between minor and major workers. If these structures are invoked in recognition of the parasites, this finding suggests that both castes are able to sense Escovopsis. Our findings support that leaf-cutter ant behavioral defense tasks against Escovopsis are subject to caste specialization, likely facilitated by worker sizes being optimal for grooming and weeding by minors and majors, respectively, with important consequences for cultivar defense.     

The effects of colony characteristics on life span and foraging behavior of individual wasps (Polybia occidentalis,Hymenoptera: Vespidae)

Summary We studied the effects of intrinsic colony characteristics and an imposed contingency on the life span and behavior of foragers in the swarm-founding social waspPolybia occidentalis. Data were collected on marked, known-age workers introduced into four observation colonies.To test the hypothesis that colony demographic features affect worker life span, we examined the relationships of colony age and size with worker life span using survivorship analysis. Colony age and size had positive relationships with life span; marked workers from two larger, older colonies had longer life spans (¯X = 24.7 days) than those from two smaller, younger colonies (¯X = 20.1 days).We quantified the effects of experimentally imposed nest damage on forager behavior, to determine which of three predicted behavioral responses by foragers to this contingency (increased probability of foraging for building material, increased rate of foraging, or decrease in age of onset of foraging) would be employed. Increasing the colony level of need for materials used in nest construction (wood pulp and water) by damaging the nests of two colonies did not cause an increase in either the proportion of marked workers that gathered nest materials or in foraging rates of marked individuals, when compared with introduced workers in two simultaneously observed control colonies. Instead, nest damage caused a decrease in the age at which marked workers first foraged for pulp and water. The response to an increase in the need for building materials was an acceleration of behavioral development in some workers.     

Colony Dehydration and Water Collection by Specialized Caste in the Leaf-Cutting Ant <Emphasis Type="Italic">Atta Sexdens Rubropilosa</Emphasis>

Social organization enables leaf-cutting ants to keep appropriate micro-ecological nest conditions for the fungus garden (their main food), eggs, larvae and adults. To maintain stability while facing changing conditions, individual ants must perceive destabilising factors and produce a proper behavioral response. We investigated behavioral responses to experimental dehydration in leaf-cutting ants to verify if task specialization exists, and to quantify the ability of ant sub-colonies for water management. Our setup consisted of fourteen sub-colonies, ten of which were randomly assigned to different levels of experimental dehydration with silica gel, whereas the remaining four were controls. The ten experimental sub-colonies were split into two groups, so that five of them had access to water. Diverse ant morphs searched for water in dehydrated colonies, but mainly a caste of small ants collected water after sources had been discovered. Size specialization for water collection was replicable in shorter experiments with three additional colonies. Ants of dehydrated colonies accumulated leaf-fragments on the nest entrance, and covering the fungus garden. Behaviors that may enhance humidity within the nests were common to all dehydration treatments. Water availability increased the life span of dehydrated colonies.     

Growth-related changes in predation behavior in incipient colonies of the ponerine antEctatomma tuberculatum (Olivier)

Summary We traced the development in the laboratory of 18 young colonies of the arboricolous ponerine antEctatomma tuberculatum. Colony foundation is of the partially-claustral type. During the early stages, when the colony is entirely dependent on the queen's behavior, the growth of the colony in terms of number of workers produced over time was relatively predictable. Afterwards, divergence in colony growth in function of the time increases as fast as the number of workers influences the efficiency of colony provisioning.Comparative analysis indicated clear changes in the predation behavior of foundresses and workers as colonies developed. For any stage of colony growth, all individuals provisioned the nest with dead prey or sugar-rich substances in the same way. However, prey hunting involves two different strategies. Foundresses and nanitic workers (originating from colonies with 9–15 workers) foraged actively, catching prey as the result of random encounters. Post-nanitic foragers (originating from colonies with 20–30 workers) and those from nature colonies developed an ambush strategy. Workers in these colonies gained experience at catching and handling prey during a period when they acted as nest guards, and so tended to be more efficient hunters than poorly experienced foundresses or nanitic foragers. The change in strategy was also positively correlated with an increase in the size of workers as the colony matured. A stable maximum in workers size is apparently reached only after the appearance of efficiently hunting foragers, presumably in numbers sufficient to provide adequate quantity and quality of larval food. Such a correlation between worker size and colony growth, assumed general for all ants, has not been demonstrated for Ponerinae before this work.     

Task-Related Variation of Cuticular Hydrocarbon Profiles Affect Nestmate Recognition in the Giant ant Dinoponera quadriceps

Recognition seems to be one of the more remarkable characteristics of social groups. In social insects, cuticular hydrocarbons are important for colonial recognition by providing a chemical signature for colony members. The acceptance threshold model predicts that colony members will accept conspecifics when the levels of nest mate cues dissimilarities are below the acceptance threshold. We tested the hypothesis that the encounter of a guard ant worker with a nurse may cause a delay in the process of recognition, because nurses from distinct colonies may share greater amount of chemical compounds. Dinoponera quadriceps guard workers decreased their effectiveness to recognize nurses rather than did to foragers. Alien foragers received significantly more bites and other stronger acts than non-nestmate nurses when they were experimentally introduced. In addition, guards took significantly more time to react against non-nestmate nurses than against alien foragers. Analysis of the cuticular hydrocarbon profiles corroborated our behavioural analysis that nurses from distinct colonies overlap greater amount of cuticular hydrocarbons.     

Influence of physical and chemical factors during foraging and culture of the symbiont fungus in Atta sexdens rubropilosa （Hymenoptera： Formicidae）

The aim of this paper is to investigate the influence of physical and chemical factors on transport and use of substrate for Atta sexdens rubropilosa workers. Three types of rectangular fragments were used to study the physical influence factors： filter paper with paraffin, filter paper without paraffin and polyester film. To study the chemical factors, some fragments were impregnated with organic extract of orange albedo, others were soaked with soybean oil and for the remaining ones nothing was applied. The fonowing parameters were evaluated： （i） attractiveness of substrate for transport and number of loading workers per treatment; （ii） foraged material incorporation; （iii） rejection by numbers of fragments deposited in the garbage or beside the fungus garden. All the polyester film fragments carried out to the fungus garden were subsequently rejected. We verified that chemical factors of the substrate were more quickly detected by the workers, whereas physical factors were used as a criterion in the decision-making to reject or accept the substrate collected.     

Workers’ Extra-Nest Behavioral Changes During Colony Fission in Dinoponera quadriceps (Santschi)

Ant colonies can reproduce by two strategies: independent foundation, wherein the queen starts a new colony alone, and dependent foundation, in which workers assist the queen. In the queenless species Dinoponera quadriceps (Santschi), the colony reproduces obligatorily by fission, a type of dependent foundation, but this process is not well understood. This study describes a colony fission event of D. quadriceps in the field and analyzes the influence of the fission process on workers’ extra-nest behavior. Based on observations of workers outside the nest, five distinct stages were identified: monodomic stage, polydomic stage, split stage, conflict stage, and post-conflict stage. The colony was initially monodomic and then occupied a second nest before it split into two independent colonies, indicating a gradual and opportunistic dependent foundation. After the fission event, the daughter colony had aggressive conflicts with the parental colony, resulting in the latter’s disappearance. Colony fission affected workers’ extra-nest behavior by increasing the frequency of rubbing the gaster against the substrate (which probably has a chemical marking function) and by decreasing the frequency of foraging during the split stage. After the fission event, the number of foragers was halved and foragers remained nearer to the nest during extra-nest activity. The spatial closeness of the parental and daughter colonies led to competition that caused the extinction or migration of the parental colony. Intraspecific competition was indicated by foraging directionality at the colony level, whereby areas of neighbor colonies were avoided; this directionality was stronger while both colonies coexisted.     

Task-Matching and Short-Term Size Shifts in Foragers of the Harvester Ant, Messor pergandei (Hymenoptera: Formicidae)

Some social insects produce workers of variable size and/or shape. One hypothesis to explain such variation is that it increases colony efficiency in performing diverse tasks, with greater efficiency leading to greater colony-level fitness. Using the granivorous ant Messor pergandei, which produces a unimodal distribution of worker sizes, I explored two corollaries of this ergonomic hypothesis: that there should be detectable task-matching and that the mean size should shift predictably with a change in task. In five experiments conducted in different years and sites in California deserts, colonies were baited with fragments of wheat or millet ground and sieved to different sizes. In six of eight colonies so baited the minimum linear dimension or mass of seeds carried by foragers was significantly related to forager size, with r² values ranging from 3 to 30%. In four of the five experiments, colonies were subsequently baited with either large seed fragments alone or small fragments alone. In three of these four experiments I detected a rapid response, over a few hours or days, in which foragers deployed by colonies shifted to larger or smaller mean size in directions predicted by size of the bait. These results suggest that an ergonomic function is one contributor to worker size polymorphism in M. pergandei. Although task-matching did not explain a substantial amount of the variance in forager size in all experiments, the rapid size shifts provide behavioral evidence that task-matching is important to colonies.     

The relationship between population size,amount of brood,and individual foraging behaviour in the honey bee,Apis mellifera L.

This study experimentally examines the relationship between colony state and the behaviour of individual pollen and nectar foragers in the honey bee, Apis mellifera L. In the first experiment we test the prediction that individual pollen foragers from colonies with higher brood quantities should exhibit a greater work effort for pollen resources than individual pollen foragers from colonies with low brood quantities. Eight colonies were assigned into two treatment groups; HIGH brood colonies were manipulated to contain 9600±480 cm² brood area; LOW brood colonies were manipulated to contain 1600±80 cm² brood area. We measured colony brood levels over the course of the experiment and collected individual pollen loads from returning pollen foragers. We found that, while colonies remained significantly different in brood levels, individual pollen foragers from HIGH brood colonies collected larger loads than individuals from LOW brood colonies. In the second experiment we investigated the influence of colony size on the behaviour of individual nectar foragers. We assigned eight colonies to two treatment groups; LARGE colonies were manipulated to contain 35000±1700 adult workers with 3500±175 cm² brood area, and SMALL colonies were manipulated to contain 10000±500 adult workers with 1000±50 cm² brood area. We observed foraging trips of individually marked workers and found that individuals from LARGE colonies made longer foraging trips than those from SMALL colonies (LARGE: 1666.7±126.4 seconds, SMALL: 1210.8±157.6 seconds), and collected larter nectar loads (LARGE: 19.2±1.0 l, SMALL: 14.6±0.8 l). These results indicate that individual nectar foragers from LARGE colonies tend to work harder than individuals from SMALL colonies. Both experiments indicate that the values of nectar and pollen resources to a colony change depend on colony state, and that individual foragers modify their behaviour accordingly.     

The effects of leaf deprivation on leaf-cutting ants and their mutualistic fungus

1. When leaf-cutting ants were deprived of leaves for 5 days, they increased their consumption rates of 'staphylae', the nutritive bodies produced by their mutualistic fungus. They also pruned their fungus garden more intensively, presumably stimulating increased staphyla production.
2. Depriving fungus gardens of leaves for 5 days led to greater rates of staphyla production compared with control gardens. The rate of substrate exhaustion also increased, possibly because of this increased staphyla production, which led to greater removal of fungal resources by the ants.
3. The availability of leaf substrate directly affected staphyla production by the fungus garden. When leaf fragments were inserted into fungus garden samples, staphyla production was depressed, suggesting that the fungus can allocate resources either to hyphal colonization of fresh substrate or to staphyla production.
4. Under conditions of leaf deprivation, food production for the ants effectively increased, due to the combined effects of increased pruning by the ants and the response of the fungus to lack of substrate. This will allow nests to survive for long periods in the field when forage availability is reduced.
5. Exposing parts of the colony to the stress of leaf deprivation showed that there is a high level of flexibility in the fungus garden's ability to produce staphylae. This may be important during the cyclic production of broods, when demands for larval food will fluctuate widely. Under normal conditions, the optimal strategy may be for the fungus garden to have a moderate rate of production with a low rate of turnover, unless demand for staphylae increases dramatically.     

Genotypic variability for rates of behavioral development in worker honeybees (Apis mellifera L.)

We demonstrate the effects of age and genotype on the likelihoodthat an individual worker honeybee (Apis mellifera L.) willbecome a forager. We established experimental colonies thatwere each initially composed of identifiable, nonforaging workersof similar ages (1–5 days old). Workers in each test colonywere the progeny of two queens, providing age and genotypicgroups of workers for comparisons. We then recorded the daythat each worker was first observed foraging. Older workerswere more likely to become foragers under our experimental conditions,even when age differences were just 1 day. At a given age, workersfrom different queens varied in their likelihood of becomingforagers. However, we found that neither age nor genotype (queensource) directly affected the likelihood that a forager wouldrevert to within-nest, larval care activities after the removalof nonforaging bees from colonies. The likelihood of revertingwas only dependent on how long a worker was a forager beforeremoving the nonforagers of the colony.     

The influence of colony population and brood rearing intensity on the activity of detoxifying enzymes in worker honey bees

Abstract. Cohorts of worker honey bees from a single parental hive, cross-fostered into colonies which differed in population and other colony parameters, were assessed for activities of enzymes involved in metabolic detoxication. Activities of glutathione S-transferase and mixed-function oxidase enzymes were negatively correlated with foster colony population, but positively correlated with the ratio of larvae (brood)/adult workers. Worker bees which had begun foraging had enzyme activity levels higher than any found in bees which were still performing in-hive duties. Elevated levels of detoxifying enzymes in colonies with low populations and high ratios of larvae/adults may be a protective mechanism to prevent poisoning of larvae by toxins brought to the colony by foragers.     

Regulation of worker activity in a primitively eusocial wasp, Ropalidia marginata

Ropalidia marginala, a tropical, primitively eusocial, polistinewasp, is unusual in that the queen (the sole egg-layer) is neitherthe most behaviorally dominant nor the most active individualin the colony. The queen by herself rarely ever initiates interactionstoward her nest mates or unloads returning foragers. There arealways a few workers in the colony who are more dominant andactive than the queen. Absence of the queen from her colonydoes not affect colony maintenance activities such as foragingor brood care, but it always results in one individual becomingvery aggressive and dominant. The dominant worker becomes thenext queen if the original queen does not return. The queendoes not appear to play any significant role in colony activityregulation. Instead, colony activities appear to be regulatedby several mechanisms including dominance behavior toward foragers,feeding of larvae, and the unloading of returning foragers,all mediated by workers themselves. Regulation of colony maintenanceappears to be based on direct evaluation of the needs of thecolony by the workers themselves. The queen however has perfectreproductive control over all workers; workers never lay eggsin the presence of the queen. It appears therefore that themechanisms involved in regulation of worker activity and workerreproduction are separate in R marginata. These findings contrastwith other primitively eusocial species where the queen actsas a "central pacemaker" and controls both worker activity andworker reproduction.     

Division of labour and seasonality in the ant Leptothorax albipennis: worker corpulence and its influence on behaviour

We address the organization of workers in social insect societies. We distinguish between changes in behavioural role over the nurse to forager role sequence, which may depend on changes in physiology, and potentially more rapid changes of task within role. We investigated the association between role and nutrient status in the ant Leptothorax albipennis. Worker lipid stores were quantified using a new body size-controlled method, and were related to worker behaviour. Worker lipid stores were evenly distributed amongst colony members at the end of winter, splitting rapidly into two distinct modes (replete nurses and lean foragers) in spring. The proportion of lean foragers increased throughout spring and summer, until most colonies contained only workers of this type. Callow workers then eclosed with intermediate lipid stores. We developed a computer vision system that tracks all nest ants to extract detailed behaviour of individuals of known lipid stores. Lipid storage was negatively correlated with a worker's foraging propensity, and with measures of spatial occupation in the nest and of activity. Different colonies showed a similar quantitative correlation between lipid stores and behavioural role, suggesting that lipid stores were not only correlated with the relative organization of individuals within each nest, but may also have influenced their absolute role. We reviewed the literature and found evidence that nutrient status influences role predisposition in social insect workers. We conclude that the distribution of worker roles may be linked to the balance between foraging income and energetic consumption within the colony directly via worker nutrient status. Copyright 2000 The Association for the Study of Animal Behaviour.     

Age and Behavior of Honey Bees,Apis mellifera (Hymenoptera: Apidae), that Perform Vibration Signals on Workers

The vibration signal is one of the most commonly occurring communication displays in honey bee (Apis mellifera) colonies. It may function in a ‘modulatory’ manner, because it causes a nonspecific increase in activity that enhances a variety of behaviors depending upon the age and caste of the recipient. We examined honey bee workers that performed vibration signals on other workers in three observation hives, each containing a population of marked bees of known age. In all three colonies, the mean age of the first performance of the vibration signal was significantly different from the mean age at which workers first performed waggle dances, carried pollen loads, or attended the queen. However, workers of all ages, except those less than 3 d old, could perform vibration signals. In older workers of foraging age, signal performance was most closely associated with recent foraging success. Younger workers that vibrated did not appear to be early-maturing foragers and thus their signals were probably not influenced by food collection. Rather, for these preforaging-age workers, signal performance was associated more with periods of orientation flight, during which younger bees learn the location of the nest and surrounding landmarks. Thus, the vibration signal may be triggered by different stimuli in different worker age classes. Because it elicits a general increase in activity in all recipients, the signal may help adjust many different colony behaviors simultancously to changes in foraging success and colony development.