Decay rates of attractive and repellent pheromones in an ant foraging trail network

Pharaoh’s ants (Monomorium pharaonis) use at least three types of foraging trail pheromone: a long-lasting attractive pheromone and two short-lived pheromones, one attractive and one repellent. We measured the decay rates of the behavioural response of ant workers at a trail bifurcation to trail substrate marked with either repellent or attractive short-lived pheromones. Our results show that the repellent pheromone effect lasts more than twice as long as the attractive pheromone effect (78 min versus 33 min). Although the effects of these two pheromones decay at approximately the same rate, the initial effect of the repellent pheromone on branch choice is almost twice that of the attractive pheromone (48% versus 25% above control). We hypothesise that the two pheromones have complementary but distinct roles, with the repellent pheromone specifically directing ants at bifurcations, while the attractive pheromone guides ants along the entire trail. Received 15 November 2007; revised 7 March 2008; accepted 18 March 2008.     

Ant navigation: reading geometrical signposts

Ants often travel along complex pheromone trail systems between their nest and foraging areas. A new and surprising discovery is that Pharaoh's ants can work out from the geometry of individual branch points on the trail whether they are heading towards or away from the nest.     

From nonlinearity to optimality: pheromone trail foraging by ants

Pheromone trails laid by foraging ants serve as a positive feedback mechanism for the sharing of information about food sources. This feedback is nonlinear, in that ants do not react in a proportionate manner to the amount of pheromone deposited. Instead, strong trails elicit disproportionately stronger responses than weak trails. Such nonlinearity has important implications for how a colony distributes its workforce, when confronted with a choice of food sources. We investigated how colonies of the Pharaoh's ant, Monomorium pharaonis, distribute their workforce when offered a choice of two food sources of differing energetic value. By developing a nonlinear differential equation model of trail foraging, and comparing model with experiments, we examined how the ants allocate their workforce between the two food sources. In this allocation, the most profitable feeder (i.e. the feeder with the highest concentration of sugar syrup) was usually exploited by the majority of ants. The particular form of the nonlinear feedback in trail foraging means that when we offered the ants a choice between two feeders of equal profitability, foraging was biased to the feeder with the highest initial number of visitors. Taken together, our experiments illuminate how pheromones provide a mechanism whereby ants can efficiently allocate their workforce among the available food sources without centralized control.     

Ants adjust their pheromone deposition to a changing environment and their probability of making errors

Animals must contend with an ever-changing environment. Social animals, especially eusocial insects such as ants and bees, rely heavily on communication for their success. However, in a changing environment, communicated information can become rapidly outdated. This is a particular problem for pheromone trail using ants, as once deposited pheromones cannot be removed. Here, we study the response of ant foragers to an environmental change. Ants were trained to one feeder location, and the feeder was then moved to a different location. We found that ants responded to an environmental change by strongly upregulating pheromone deposition immediately after experiencing the change. This may help maintain the colony''s foraging flexibility, and allow multiple food locations to be exploited simultaneously. Our treatment also caused uncertainty in the foragers, by making their memories less reliable. Ants which had made an error but eventually found the food source upregulated pheromone deposition when returning to the nest. Intriguingly, ants on their way towards the food source downregulated pheromone deposition if they were going to make an error. This may suggest that individual ants can measure the reliability of their own memories and respond appropriately.     

Communication in the migratory termite-hunting ant Pachycondyla (= Termitopone) marginata (Formicidae, Ponerinae)

The Neotropical species Pachycondyla marginata conducts well-organized predatory raids on the termite species Neocapritermes opacus and frequently emigrates to new nest sites. During both activities the ants employ chemical trail communication. The trail pheromone orginates from the pygidial gland. Among the substances identified in the pygidial gland secretions, only citronellal was effective as a trail pheromone. Isopulegol elicited an increase in locomotory activity in the ants and may function as a synergist recruitment signal. The chemical signal is enhanced by a shaking display performed by the recruiting ant.     

Effect of Trail Bifurcation Asymmetry and Pheromone Presence or Absence on Trail Choice by Lasius niger Ants

During foraging, ant workers are known to make use of multiple information sources, such as private information (personal memory) and social information (trail pheromones). Environmental effects on foraging, and how these interact with other information sources, have, however, been little studied. One environmental effect is trail bifurcation asymmetry. Ants forage on branching trail networks and must often decide which branch to take at a junction (bifurcation). This is an important decision, as finding food sources relies on making the correct choices at bifurcations. Bifurcation angle may provide important information when making this choice. We used a Y‐maze with a pivoting 90° bifurcation to study trail choice of Lasius niger foragers at varying branch asymmetries (0°, [both branches 45° from straight ahead], 30° [branches at 30° and 60° from straight ahead], 45°, 60° and 90° [one branch straight ahead, the other at 90°]). The experiment was carried out either with equal amounts of trail pheromone on both branches of the bifurcation or with pheromone present on only one branch. Our results show that with equal pheromone, trail asymmetry has a significant effect on trail choice. Ants preferentially follow the branch deviating least from straight, and this effect increases as asymmetry increases (47% at 0°, 54% at 30°, 57% at 45°, 66% at 60° and 73% at 90°). However, when pheromone is only present on one branch, the graded effect of asymmetry disappears. Overall, however, there is an effect of asymmetry as the preference of ants for the pheromone‐marked branch over the unmarked branch is reduced from 65%, when it is the less deviating branch, to 53%, when it is the more deviating branch. These results demonstrate that trail asymmetry influences ant decision‐making at bifurcations and that this information interacts with trail pheromone presence in a non‐hierarchical manner.     

Trail following and stowaway behaviour of the myrmecophilous staphylinid beetle,Homoeusa acuminata,during foraging trips of its hostLasius fuliginosus (Hymenoptera: Formicidae)

Summary The behaviour of adultHomoeusa acuminata on trails of its hostLasius fuliginosus was investigated both in the field and in the laboratory. The beetles were active from May to September, accurately following the foraging trails of their hosts up to 20 metres from the nest. Most of the time, they were ignored by the ants, but if attacked they raised their abdomen as a possible appeasement or defensive behaviour. On trails the beetles most probably act as food robbers, feeding on prey collected by ants. The following method, called stowaway behaviour, was used by the beetles: when a beetle encountered an ant carrying a prey back to the nest it jumped on the prey, probably feeding on it while being transported.Laboratory experiments on circular artificial trails demonstrated thatH. acuminata follows a water extract of hindguts of the ants, the source of the trail pheromone. Both beetles and ants responded to an artificial trail of 0.03 hindgut equivalent per cm, but the mean distance followed by the beetles was about twelve times higher than that covered by the ants themselves. In contrast, experiments with solutions of the six fatty acids reported as the active components of the trail pheromone showed that the beetles did not respond at all, and that the ants only respond to the fatty acids at a very high concentration.     

Just follow your nose: homing by olfactory cues in ants

How is an ant-equipped with a brain that barely exceeds the size of a pinhead-capable of achieving navigational marvels? Even though evidences suggest that navigation is a multimodal process, ants heavily depend on olfactory cues-of pheromonal and non-pheromonal nature-for foraging and orientation. Recent studies have directed their attention to the efficiency of pheromone trail networks. Advances in neurophysiological techniques make it possible to investigate trail pheromone processing in the ant's brain. In addition to relying on pheromone odours, ants also make use of volatiles emanating from the nest surroundings. Deposited in the vicinity of the nest, these home-range markings help the ants to home after a foraging run. Furthermore, olfactory landmarks associated with the nest enhance ants' homing abilities.     

Coordination of Raiding and Emigration in the Ponerine Army Ant Leptogenys distinguenda (Hymenoptera: Formicidae: Ponerinae): A Signal Analysis

Several glandular sources of trail pheromones have been discovered in army ants in general. Nevertheless, at present the understanding of the highly coordinated behavior of these ants is far from complete. The importance of trail pheromone communication for the coordination of raids and emigrations in the ponerine army ant Leptogenys distinguenda was examined, and its ecological function is discussed. The secretions of at least two glands organize the swarming activities of L. distinguenda. The pygidial gland is the source of an orientation pheromone holding the group of raiding workers together. The same pheromone guides emigrations to new nest sites. In addition, the poison sac contains two further components: one with a weak orientation effect and another which produces strong, but short-term attraction and excitement. The latter component is important in prey recruitment and characterizes raid trails. This highly volatile recruitment pheromone allows the extreme swarm dynamic characteristic of this species. Emigration trails lack the poison gland secretion. Due to their different chemical compositions, the ants are thus able to distinguish between raid and emigration trails. Nest emigration is not induced chemically, but mechanically, by the jerking movements of stimulating workers.     

Orientation by magnetic field in leaf-cutter ants, Atta colombica (Hymenoptera: Formicidae)

Leaf‐cutter ants (Atta colombica) use trail following to travel between foraging sites and the home nest. However, this combination of pheromone and visual cues is likely to be complemented by a directional reference system such as a compass, used not only when foraging but also during colony formation, where foraging trails degrade or where ants become displaced. One candidate system is the magnetic polarity compass. We tested the orientation of leaf‐cutter ants under a magnetic field of reversed‐polarity, with the prediction that the ants would show 180° deflection compared with control ants in an unchanged geomagnetic field. When the sun's disc was unobstructed by clouds, orientation was the same as that of control ants, implying that magnetic cues were not used to orient. However, when the sky was overcast, ants in the experimental treatment significantly shifted their mean orientation both in comparison with controls and reversed‐polarity ants under the sun. Although a total reversal in orientation was not induced, the results demonstrate that Atta respond to magnetic reversal in the absence of sunlight cues, and suggest a role for magnetic cues in determining direction during orientation.     

Food collection and response to pheromones in an ant species exposed to electromagnetic radiation

We used the ant species Myrmica sabuleti as a model to study the impact of electromagnetic waves on social insects' response to their pheromones and their food collection. We quantified M. sabuleti workers' response to their trail, area marking and alarm pheromone under normal conditions. Then, we quantified the same responses while under the influence of electromagnetic waves. Under such an influence, ants followed trails for only short distances, no longer arrived at marked areas and no longer orientated themselves to a source of alarm pheromone. Also when exposed to electromagnetic waves, ants became unable to return to their nest and recruit congeners; therefore, the number of ants collecting food increases only slightly and slowly. After 180 h of exposure, their colonies deteriorated. Electromagnetic radiation obviously affects social insects' behavior and physiology.     

Leaf-cutting ants as road engineers: the width of trails at branching points in <Emphasis Type="Italic">Atta cephalotes</Emphasis>

We used a simple engineering principle, which suggests that the width of a road needed for a smooth traffic flow is proportional to the peak traffic volume (“engineering hypothesis”), to analyze the adaptive significance of trail width at branching points in the leaf-cutting ant Atta cephalotes. Since the flow of outgoing ants splits at trail bifurcations and merges when ants return to the nest through the same paths, the sum of branch widths should equal the width of the trail section upstream of the bifurcation. We measured the width of branches and their preceding trail section and also performed field measurements and manipulations to analyze ant flow, number of collisions, and ant speed in different trail sectors. Contrary to the prediction of the “engineering hypothesis”, the sum of branch widths was larger than the width of the trail immediately before the bifurcation. Our data contradict the “trail addition hypothesis” and support the “border effect hypothesis” to explain this pattern. First, the width of the widest branch was smaller than the width of the trail upstream of the bifurcation, an unexpected result if one branch is merely the continuation of the basal trail. Second, ants collided with obstacles more often in the margin than in the central portion of the trail, relocated ants from central to margin trail sectors reduced their speed, and ant flow was higher in the central sections of the trail. Since the delaying effect of trail margins increases as the trail width decreases, ants may build branches wider than expected to reduce the border effect. The delaying effect of trail margins should be included in the analysis of costs and benefits to fully understand the adaptive value of the design of ant trail networks.     

Recruitment and Other Communication Behavior in the Ponerine Ant Ectatomma ruidum

The ponerine ant Ectatomma ruidum, though previously reported to possess only rudimentary recruitment ability, was found to lay chemical trails for mass recruitment to rich or difficult food sources. The pheromone originates from the Dufour's gland, a new source of trail pheromones in the primitive ant subfamily Ponerinae. During nest emigrations, E. ruidum practices stereotyped social carrying in the myrmicine mode. The discovery of this form of social carrying and of a recruitment pheromone in the Dufour's gland secretions support the hypothesis that the subfamily Myrmicinae is derived from an ectatommine ancestor. Other communication behaviors exhibited by E. ruidum include exchange of liquid food carried between the mandibles, chemical alarm communication, nest entrance marking, and an additional social carrying posture previously unknown in ants.     

Impact of seed abundance on seed processing and dispersal by the red ant Myrmica rubra

1. Myrmecochory sensu stricto is an ant–plant mutualism in which non‐granivorous ants disperse plant diaspores after feeding on their nutrient‐rich seed appendage, the elaiosome. Phenological traits associated with the diaspore can influence the behaviour of ants and thus their ultimate efficiency as seed dispersers. 2. This study investigated how a contrasting availability of seeds (20 vs. 200 seeds) from the diplochorous Chelidonium majus (Papaveraceae, Linnaeus) plant species influences the behaviour of Myrmica rubra (Formicidae, Linnaeus) ants, from the retrieval of seeds until their dispersal outside the ant nest. 3. Regardless of seed abundance, the ants collected the first diaspores at similar rates. Then, seed retrieval sped up over time for large seed sources until satiation took place with only one‐third of the tested colonies wholly depleting abundant seed sources. 4. No active recruitment by trail‐laying ants was triggered, even to an abundant seed source 5. In both conditions of seed abundance, the majority of the diaspores retrieved inside the nest were discarded with the elaiosome removed and were dispersed at similar distances from the nest. 6. The paper concludes with a discussion of how the quantity of seeds released by a plant with a dual mode of dispersal can potentially influence the behaviour of ant dispersers and hence the dispersal efficiency derived from myrmecochory.     

Pheromone trail following in the ant Lasius niger: high accuracy and variability but no effect of task state

The use of pheromone trails in ant colony organization is an important model for understanding collective decision‐making and complex adaptive systems. The ant Lasius niger L. (Hymenoptera: Fomicidae) is one of the main model organisms used for such studies. Key to understanding pheromone trail use by ants is knowing how well trails are followed. The results of a previous study suggest that L. niger trail following is poor, with between 60% and 70% accuracy at a T bifurcation. It is hypothesized that the true trail following accuracy is higher, and that the low accuracy reported previously is the result of a methodological error. Specifically, it is hypothesized that ‘task state’ (i.e. what the ants ‘thought they were doing’) affected pheromone following accuracy. In the present study, the task state of the ants is set experimentally to one of three states: scouting (completely naive), recruited (having information that food has been found, but not where it is) and shuttling (having a strong memory of the location of a food source). Trail following accuracy is tested for each group. Trail following is found to be more accurate than previously reported: 83%, 82% and 74% correct decisions for scouts, recruits and shuttlers, respectively. However, the difference between the three groups is not significant. Importantly, very high inter‐trial variation is reported both in the present study and in experiments from other research groups. This variation is unexplainable by trail strengths or colony‐level differences, and is highlighted as an important factor when experimentally measuring trail following.     

Colony Budding and its Effects on Food Allocation in the Highly Polygynous Ant, Monomorium pharaonis

To advance our understanding of the causes and the consequences of budding (colony multiplication by fragmentation of main nests), we investigated nest movement in the facultatively polydomous Pharaoh ant, Monomorium pharaonis. Demographic data revealed that Pharaoh ants are highly polygynous and have a relatively low worker to queen ratio of 12.86. Budding experiments demonstrated that the number of available bud nests has a significant effect on colony fragmentation and increasing the number of bud nests resulted in smaller colony fragments. The overall distribution among bud nests was uneven, even though there was no evidence that the different life stages and castes partitioned unevenly among the bud nests and the analysis of individual colonies revealed no evidence of an uneven split in any of the colonies. This demonstrates that Pharaoh ants have the ability to exert social control over colony size and caste proportions during budding, which may contribute to their success as an invasive ant. The intensity of nest disturbance had a significant effect on whether or not the ants migrated into bud nests. Major disturbance resulted in the ants abandoning the source nest and migrating to bud nests and minor disturbance did not stimulate the ants to abandon the source nest. The results of the successive budding experiment which allowed the ants the opportunity to bud into progressively smaller nest fragments demonstrate that Pharaoh ants maintain a preferred minimum group size of 469 ± 28 individuals. Food allocation experiments utilizing protein marking revealed that nest fragmentation in Pharaoh ants has no negative impact on intracolony food distribution. Overall, our results suggest that nest units in the Pharaoh ant behave like cooperative, rather than competitive, entities. Such cooperation is most likely facilitated by the fact that individuals in all bud nests are genetically related, remain in close proximity to each other, and may continue to exchange individuals after budding.     

Effect of Short-Term GSM Radiation at Representative Levels in Society on a Biological Model: The Ant Myrmica sabuleti

Well-controlled electromagnetic exposure conditions were set up at a representative societal GSM radiation intensity level, 1.5 V/m, which is the legally allowed level in Brussels. Two nests of the ant species Myrmica sabuleti were repeatedly irradiated during 10 min. before their behavior was observed, based on the analysis of the ant trajectories. Under these exposure conditions, behavioral effects were detected. The ants’ locomotion slightly changed. The ants’ orientation towards their attractive alarm pheromone statistically became of lower quality. The ants still presented their trail following behavior but less efficiently. In this controversial issue, ants could be considered as possible bioindicators.     

An agent-based model to investigate the roles of attractive and repellent pheromones in ant decision making during foraging

Pharaoh's ants organise their foraging system using three types of trail pheromone. All previous foraging models based on specific ant foraging systems have assumed that only a single attractive pheromone is used. Here we present an agent-based model based on trail choice at a trail bifurcation within the foraging trail network of a Pharaoh's ant colony which includes both attractive (positive) and repellent (negative) trail pheromones. Experiments have previously shown that Pharaoh's ants use both types of pheromone. We investigate how the repellent pheromone affects trail choice and foraging success in our simulated foraging system. We find that both the repellent and attractive pheromones have a role in trail choice, and that the repellent pheromone prevents random fluctuations which could otherwise lead to a positive feedback loop causing the colony to concentrate its foraging on the unrewarding trail. An emergent feature of the model is a high level of variability in the level of repellent pheromone on the unrewarding branch. This is caused by the repellent pheromone exerting negative feedback on its own deposition. We also investigate the dynamic situation where the location of the food is changed after foraging trails are established. We find that the repellent pheromone has a key role in enabling the colony to refocus the foraging effort to the new location. Our results show that having a repellent pheromone is adaptive, as it increases the robustness and flexibility of the colony's overall foraging response.     

Spatial and temporal variation in pheromone composition of ant foraging trails

Many social insects use pheromones to communicate and coordinatetheir activities. Investigation of intraspecific differencesin pheromone use is a new area of social insect research. Forexample, interindividual variation in alarm pheromone contenthas been found in physical castes of polymorphic ants. Manyant species use multiple trail pheromones. Here we present novelresearch into trail pheromone variations between behavioralsubcastes of pharaoh ants, Monomorium pharaonis. Monomoriumpharaonis is attracted to trail pheromones found in its poisonglands (monomorines) and Dufour's glands (faranal). We showthat the most abundant monomorines, I (M1) and III (M3), canbe readily detected in pheromone trails. A behaviorally distinctsubcaste known as "pathfinder" foragers can relocate long-livedpheromone trails. Chemical analysis showed that pathfinder foragershad low M3:M1 ratios (mean 3.09 ± 1.53, range 1.03–7.10).Nonpathfinder foragers had significantly greater M3:M1 ratios(38.3 ± 60.0, range 3.54–289). We found that M3:M1ratio did not differ between foragers of different age but wascorrelated with behavioral subcaste at all ages. The relativeabundance of M3:M1 on foraging trails ranged from 3.03–41.3over time during pheromone trail build-up. M3:M1 ratio alsovaried spatially throughout trail networks, being lowest ontrail sections closest to a food source (M3:M1 = 1.9–3.61)and highest near the nest (M3:M1 = 67–267). Our resultsindicate a functional role for differences in pheromone trailcomposition, whereby pathfinder foragers might preferentiallymark sections of pheromone trail networks for future exploration.     

Trail‐sharing among tropical ants: interspecific use of trail pheromones?

1. Trail‐sharing between different ant species is rare and restricted to a small number of species pairs. Its underlying mechanisms are largely unknown. For trail‐sharing to occur, two factors are required: (i) one or both species must recognise the other species or its pheromone trails and (ii) both species must tolerate each other to a certain extent to allow joint use of the trail. A species that follows another's trails can efficiently exploit the other's information on food sources contained in the pheromone trails. Hence, food competition and thus aggressive interactions between a species following another's trail and the species being followed, seem likely. 2. In the present study, we investigated interspecific trail following and interspecific aggression in trail sharing associations (i) among Polyrhachis ypsilon, Camponotus saundersi, and Dolichoderus cuspidatus, and (ii) among Camponotus rufifemur and Crematogaster modiglianii. We tested whether trail‐sharing species follow each other's pheromone trails, and whether the ants tolerated or attacked their trail‐sharing partners. In both associations, we confronted workers with pheromone trails of their associated species, and, for the former association, measured interspecific aggression among the trail‐sharing species. 3. In our assays, D. cuspidatus and C. rufifemur regularly followed heterospecific pheromone trails of P. ypsilon and C. modiglianii, respectively. However, only few workers of the remaining species followed heterospecific pheromone trails. Thus, shared trails of P. ypsilon and C. saundersi cannot be explained by interspecific trail‐following. 4. Interspecific aggression among P. ypsilon, C. saundersi, and D. cuspidatus was strongly asymmetric, C. saundersi being submissive to the other two. All three species differentiated between heterospecific workers from the same or another site, suggesting habituation to the respective trail‐sharing partners. We therefore hypothesise that differential tolerance by dominant ant species may be mediated by selective habituation towards submissive species and this way determines the assembly of trail‐sharing associations.