Elva Robinson | University of York (original) (raw)

Papers by Elva Robinson

Research paper thumbnail of The relationship between canopy cover and colony size of the wood ant Formica lugubris--implications for the thermal effects on a keystone ant species

PloS one, 2014

Climate change may affect ecosystems and biodiversity through the impacts of rising temperature o... more Climate change may affect ecosystems and biodiversity through the impacts of rising temperature on species' body size. In terms of physiology and genetics, the colony is the unit of selection for ants so colony size can be considered the body size of a colony. For polydomous ant species, a colony is spread across several nests. This study aims to clarify how climate change may influence an ecologically significant ant species group by investigating thermal effects on wood ant colony size. The strong link between canopy cover and the local temperatures of wood ant's nesting location provides a feasible approach for our study. Our results showed that nests were larger in shadier areas where the thermal environment was colder and more stable compared to open areas. Colonies (sum of nests in a polydomous colony) also tended to be larger in shadier areas than in open areas. In addition to temperature, our results supported that food resource availability may be an additional fact...

Research paper thumbnail of How collective comparisons emerge without individual comparisons of the options

Proceedings. Biological sciences / The Royal Society, Jan 22, 2014

Collective decisions in animal groups emerge from the actions of individuals who are unlikely to ... more Collective decisions in animal groups emerge from the actions of individuals who are unlikely to have global information. Comparative assessment of options can be valuable in decision-making. Ant colonies are excellent collective decision-makers, for example when selecting a new nest-site. Here, we test the dependency of this cooperative process on comparisons conducted by individual ants. We presented ant colonies with a choice between new nests: one good and one poor. Using individually radio-tagged ants and an automated system of doors, we manipulated individual-level access to information: ants visiting the good nest were barred from visiting the poor one and vice versa. Thus, no ant could individually compare the available options. Despite this, colonies still emigrated quickly and accurately when comparisons were prevented. Individual-level rules facilitated this behavioural robustness: ants allowed to experience only the poor nest subsequently searched more. Intriguingly, som...

Research paper thumbnail of Resource redistribution in polydomous ant nest networks: local or global?

Behavioral ecology : official journal of the International Society for Behavioral Ecology, Sep 1, 2014

An important problem facing organisms in a heterogeneous environment is how to redistribute resou... more An important problem facing organisms in a heterogeneous environment is how to redistribute resources to where they are required. This is particularly complex in social insect societies as resources have to be moved both from the environment into the nest and between individuals within the nest. Polydomous ant colonies are split between multiple spatially separated, but socially connected, nests. Whether, and how, resources are redistributed between nests in polydomous colonies is unknown. We analyzed the nest networks of the facultatively polydomous wood ant Formica lugubris. Our results indicate that resource redistribution in polydomous F. lugubris colonies is organized at the local level between neighboring nests and not at the colony level. We found that internest trails connecting nests that differed more in their amount of foraging were stronger than trails between nests with more equal foraging activity. This indicates that resources are being exchanged directly from nests w...

Research paper thumbnail of NURSE HOME VISITS TO HELP FAMILIES MANAGE ASTHMATIC-CHILDREN-A COST-BENEFIT ANALYSIS

Research paper thumbnail of A bibliophile is born

Research paper thumbnail of Foraging in invasive species: evading the limitations of polydomy

Research paper thumbnail of What is the relationship between altitude and ant colony size?

Research paper thumbnail of How temporally stable are polydomous wood ant networks?

Research paper thumbnail of How do polydomous ant colonies correct nutritional imbalances between nests?

Research paper thumbnail of Family stress and childhood asthma

Eighty-six families, each with an asthmatic child, were studied in their normal environment. The ... more Eighty-six families, each with an asthmatic child, were studied in their normal environment. The relationships between the physical severity of asthma and the families' knowledge and feelings about the asthma were measured. The mean peak expiratory flow rate recorded prospectively for one week correlated (R = 0.33) with the symptom score. In the 73 families who knew their child had asthma, there was a significant relationship (R = +0.28) between the knowledge of physical signs of attack severity and knowledge of the drugs available for emergency therapy. The mean peak expiratory flow rate was inversely related to the practical effectiveness of the inhaler technique (R = 0.62) only when the 12 families in whom the child's mean peak expiratory flow rate was less than 90 per cent of predicted were examined. The parents' feelings of worry and fear, measured independently, related to the family's knowledge of physical indications of the severity of asthma attacks. Although the families' overall knowledge scores were poor, they were apparently adjusting appropriately to asthma as a psychological stress as it appeared that they sought more information if they were worried. Their comprehension should improve if such knowledge was correctly delivered. There is a need to explore more effective techniques for the education of such families.

Research paper thumbnail of Social networking in the world of ants

Trends in Ecology & Evolution, 2011

Research paper thumbnail of Experience, corpulence and decision making in ant foraging

Journal of Experimental Biology, 2012

Social groups are structured by the decisions of their members. Social insects typically divide l... more Social groups are structured by the decisions of their members. Social insects typically divide labour: some decide to stay in the nest while others forage for the colony. Two sources of information individuals may use when deciding whether to forage are their own experience of recent task performance and their own physiology, e.g. fat reserves (corpulence). The former is primarily personal information; the latter may give an indication of the food reserves of the whole colony. These factors are hard to separate because typically leaner individuals are also more experienced foragers. We designed an experiment to determine whether foraging specialisation is physiological or experience based (or both). We invented a system of automatic doors controlled by radio-tag information to manipulate task access and decouple these two sources of information. Our results show that when information from corpulence and recent experience conflict, ants behave only in accordance with their corpulence. However, among ants physiologically inclined to forage (less corpulent ants), recent experience of success positively influenced their propensity to forage again. Hence, foraging is organised via long-term physiological differences among individuals resulting in a relatively stable response threshold distribution, with fine-tuning provided by short-term learning processes. Through these simple rules, colonies can organise their foraging effort both robustly and flexibly.

Research paper thumbnail of Preemptive Defensive Self‐Sacrifice by Ant Workers

The American Naturalist, 2008

Worker insects altruistically sacrifice their own reproduction to rear nondescendant kin. This sa... more Worker insects altruistically sacrifice their own reproduction to rear nondescendant kin. This sacrifice reaches its most spectacular level in suicidal colony defense. Suicidal defense, such as when the sting of a honeybee worker embeds in a predator and then breaks off, is normally a facultative response. Here we describe the first example of preemptive self-sacrifice in nest defense. In the Brazilian ant Forelius pusillus, the nest entrance is closed at sunset. One to eight workers finish the job from the outside and, in doing so, sacrifice their lives.

Research paper thumbnail of Do ants make direct comparisons?

Proceedings of the Royal Society B: Biological Sciences, 2009

Many individual decisions are informed by direct comparison of the alternatives. In collective de... more Many individual decisions are informed by direct comparison of the alternatives. In collective decisions, however, only certain group members may have the opportunity to compare options. Emigrating ant colonies (Temnothorax albipennis) show sophisticated nest-site choice, selecting superior sites even when they are nine times further away than the alternative. How do they do this? We used radio-frequency identification-tagged ants to monitor individual behaviour. Here we show for the first time that switching between nests during the decision process can influence nest choice without requiring direct comparison of nests. Ants finding the poor nest were likely to switch and find the good nest, whereas ants finding the good nest were more likely to stay committed to that nest. When ants switched quickly between the two nests, colonies chose the good nest. Switching by ants that had the opportunity to compare nests had little effect on nest choice. We suggest a new mechanism of collective nest choice: individuals respond to nest quality by the decision either to commit or to seek alternatives. Previously proposed mechanisms, recruitment latency and nest comparison, can be explained as side effects of this simple rule. Colony-level comparison and choice can emerge, without direct comparison by individuals.

Research paper thumbnail of Flexible task allocation and the organization of work in ants

Proceedings of the Royal Society B: Biological Sciences, 2009

Research paper thumbnail of Crystal structure of the SOS cell division inhibitor SulA and in complex with FtsZ

Proceedings of the National Academy of Sciences, 2003

SulA halts cell division in Escherichia coli by binding to the major component of the division ma... more SulA halts cell division in Escherichia coli by binding to the major component of the division machinery FtsZ. We have solved the crystal structure of SulA alone and in complex with FtsZ from Pseudomonas aeruginosa. SulA is expressed when the SOS response is induced. This is a mechanism to inhibit cell division and repair DNA in the event of DNA damage. FtsZ is a tubulin-like protein that forms polymers, with the active-site GTPase split across two monomers. One monomer provides the GTP-binding site and the other, through its T7 loop nucleotide hydrolysis. Our structures show that SulA is a dimer, and that SulA inhibits cell division neither by binding the nucleotide-binding site nor by inducing conformational changes in FtsZ. Instead, SulA binds the T7 loop surface of FtsZ, opposite the nucleotide-binding site, blocking polymer formation. These findings explain why GTP hydrolysis and polymer turnover are required for SulA inhibition.

Research paper thumbnail of A Simple Threshold Rule Is Sufficient to Explain Sophisticated Collective Decision-Making

PLoS ONE, 2011

Decision-making animals can use slow-but-accurate strategies, such as making multiple comparisons... more Decision-making animals can use slow-but-accurate strategies, such as making multiple comparisons, or opt for simpler, faster strategies to find a 'good enough' option. Social animals make collective decisions about many group behaviours including foraging and migration. The key to the collective choice lies with individual behaviour. We present a case study of a collective decision-making process (house-hunting ants, Temnothorax albipennis), in which a previously proposed decision strategy involved both quality-dependent hesitancy and direct comparisons of nests by scouts. An alternative possible decision strategy is that scouting ants use a very simple quality-dependent threshold rule to decide whether to recruit nestmates to a new site or search for alternatives. We use analytical and simulation modelling to demonstrate that this simple rule is sufficient to explain empirical patterns from three studies of collective decision-making in ants, and can account parsimoniously for apparent comparison by individuals and apparent hesitancy (recruitment latency) effects, when available nests differ strongly in quality. This highlights the need to carefully design experiments to detect individual comparison. We present empirical data strongly suggesting that best-of-n comparison is not used by individual ants, although individual sequential comparisons are not ruled out. However, by using a simple threshold rule, decision-making groups are able to effectively compare options, without relying on any form of direct comparison of alternatives by individuals. This parsimonious mechanism could promote collective rationality in group decision-making.

Research paper thumbnail of Record Dynamics in Ants

PLoS ONE, 2010

The success of social animals (including ourselves) can be attributed to efficiencies that arise ... more The success of social animals (including ourselves) can be attributed to efficiencies that arise from a division of labour. Many animal societies have a communal nest which certain individuals must leave to perform external tasks, for example foraging or patrolling. Staying at home to care for young or leaving to find food is one of the most fundamental divisions of labour. It is also often a choice between safety and danger. Here we explore the regulation of departures from ant nests. We consider the extreme situation in which no one returns and show experimentally that exiting decisions seem to be governed by fluctuating record signals and ant-ant interactions. A record signal is a new 'high water mark' in the history of a system. An ant exiting the nest only when the record signal reaches a level it has never perceived before could be a very effective mechanism to postpone, until the last possible moment, a potentially fatal decision. We also show that record dynamics may be involved in first exits by individually tagged ants even when their nest mates are allowed to re-enter the nest. So record dynamics may play a role in allocating individuals to tasks, both in emergencies and in everyday life. The dynamics of several complex but purely physical systems are also based on record signals but this is the first time they have been experimentally shown in a biological system.

Research paper thumbnail of Insect communication: ‘No entry’ signal in ant foraging

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

Journal of Theoretical Biology, 2008

Pharaoh's ants organise their foraging system using three types of trail pheromone. All previous ... more 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.

Research paper thumbnail of The relationship between canopy cover and colony size of the wood ant Formica lugubris--implications for the thermal effects on a keystone ant species

PloS one, 2014

Climate change may affect ecosystems and biodiversity through the impacts of rising temperature o... more Climate change may affect ecosystems and biodiversity through the impacts of rising temperature on species' body size. In terms of physiology and genetics, the colony is the unit of selection for ants so colony size can be considered the body size of a colony. For polydomous ant species, a colony is spread across several nests. This study aims to clarify how climate change may influence an ecologically significant ant species group by investigating thermal effects on wood ant colony size. The strong link between canopy cover and the local temperatures of wood ant's nesting location provides a feasible approach for our study. Our results showed that nests were larger in shadier areas where the thermal environment was colder and more stable compared to open areas. Colonies (sum of nests in a polydomous colony) also tended to be larger in shadier areas than in open areas. In addition to temperature, our results supported that food resource availability may be an additional fact...

Research paper thumbnail of How collective comparisons emerge without individual comparisons of the options

Proceedings. Biological sciences / The Royal Society, Jan 22, 2014

Collective decisions in animal groups emerge from the actions of individuals who are unlikely to ... more Collective decisions in animal groups emerge from the actions of individuals who are unlikely to have global information. Comparative assessment of options can be valuable in decision-making. Ant colonies are excellent collective decision-makers, for example when selecting a new nest-site. Here, we test the dependency of this cooperative process on comparisons conducted by individual ants. We presented ant colonies with a choice between new nests: one good and one poor. Using individually radio-tagged ants and an automated system of doors, we manipulated individual-level access to information: ants visiting the good nest were barred from visiting the poor one and vice versa. Thus, no ant could individually compare the available options. Despite this, colonies still emigrated quickly and accurately when comparisons were prevented. Individual-level rules facilitated this behavioural robustness: ants allowed to experience only the poor nest subsequently searched more. Intriguingly, som...

Research paper thumbnail of Resource redistribution in polydomous ant nest networks: local or global?

Behavioral ecology : official journal of the International Society for Behavioral Ecology, Sep 1, 2014

An important problem facing organisms in a heterogeneous environment is how to redistribute resou... more An important problem facing organisms in a heterogeneous environment is how to redistribute resources to where they are required. This is particularly complex in social insect societies as resources have to be moved both from the environment into the nest and between individuals within the nest. Polydomous ant colonies are split between multiple spatially separated, but socially connected, nests. Whether, and how, resources are redistributed between nests in polydomous colonies is unknown. We analyzed the nest networks of the facultatively polydomous wood ant Formica lugubris. Our results indicate that resource redistribution in polydomous F. lugubris colonies is organized at the local level between neighboring nests and not at the colony level. We found that internest trails connecting nests that differed more in their amount of foraging were stronger than trails between nests with more equal foraging activity. This indicates that resources are being exchanged directly from nests w...

Research paper thumbnail of NURSE HOME VISITS TO HELP FAMILIES MANAGE ASTHMATIC-CHILDREN-A COST-BENEFIT ANALYSIS

Research paper thumbnail of A bibliophile is born

Research paper thumbnail of Foraging in invasive species: evading the limitations of polydomy

Research paper thumbnail of What is the relationship between altitude and ant colony size?

Research paper thumbnail of How temporally stable are polydomous wood ant networks?

Research paper thumbnail of How do polydomous ant colonies correct nutritional imbalances between nests?

Research paper thumbnail of Family stress and childhood asthma

Eighty-six families, each with an asthmatic child, were studied in their normal environment. The ... more Eighty-six families, each with an asthmatic child, were studied in their normal environment. The relationships between the physical severity of asthma and the families' knowledge and feelings about the asthma were measured. The mean peak expiratory flow rate recorded prospectively for one week correlated (R = 0.33) with the symptom score. In the 73 families who knew their child had asthma, there was a significant relationship (R = +0.28) between the knowledge of physical signs of attack severity and knowledge of the drugs available for emergency therapy. The mean peak expiratory flow rate was inversely related to the practical effectiveness of the inhaler technique (R = 0.62) only when the 12 families in whom the child's mean peak expiratory flow rate was less than 90 per cent of predicted were examined. The parents' feelings of worry and fear, measured independently, related to the family's knowledge of physical indications of the severity of asthma attacks. Although the families' overall knowledge scores were poor, they were apparently adjusting appropriately to asthma as a psychological stress as it appeared that they sought more information if they were worried. Their comprehension should improve if such knowledge was correctly delivered. There is a need to explore more effective techniques for the education of such families.

Research paper thumbnail of Social networking in the world of ants

Trends in Ecology & Evolution, 2011

Research paper thumbnail of Experience, corpulence and decision making in ant foraging

Journal of Experimental Biology, 2012

Social groups are structured by the decisions of their members. Social insects typically divide l... more Social groups are structured by the decisions of their members. Social insects typically divide labour: some decide to stay in the nest while others forage for the colony. Two sources of information individuals may use when deciding whether to forage are their own experience of recent task performance and their own physiology, e.g. fat reserves (corpulence). The former is primarily personal information; the latter may give an indication of the food reserves of the whole colony. These factors are hard to separate because typically leaner individuals are also more experienced foragers. We designed an experiment to determine whether foraging specialisation is physiological or experience based (or both). We invented a system of automatic doors controlled by radio-tag information to manipulate task access and decouple these two sources of information. Our results show that when information from corpulence and recent experience conflict, ants behave only in accordance with their corpulence. However, among ants physiologically inclined to forage (less corpulent ants), recent experience of success positively influenced their propensity to forage again. Hence, foraging is organised via long-term physiological differences among individuals resulting in a relatively stable response threshold distribution, with fine-tuning provided by short-term learning processes. Through these simple rules, colonies can organise their foraging effort both robustly and flexibly.

Research paper thumbnail of Preemptive Defensive Self‐Sacrifice by Ant Workers

The American Naturalist, 2008

Worker insects altruistically sacrifice their own reproduction to rear nondescendant kin. This sa... more Worker insects altruistically sacrifice their own reproduction to rear nondescendant kin. This sacrifice reaches its most spectacular level in suicidal colony defense. Suicidal defense, such as when the sting of a honeybee worker embeds in a predator and then breaks off, is normally a facultative response. Here we describe the first example of preemptive self-sacrifice in nest defense. In the Brazilian ant Forelius pusillus, the nest entrance is closed at sunset. One to eight workers finish the job from the outside and, in doing so, sacrifice their lives.

Research paper thumbnail of Do ants make direct comparisons?

Proceedings of the Royal Society B: Biological Sciences, 2009

Many individual decisions are informed by direct comparison of the alternatives. In collective de... more Many individual decisions are informed by direct comparison of the alternatives. In collective decisions, however, only certain group members may have the opportunity to compare options. Emigrating ant colonies (Temnothorax albipennis) show sophisticated nest-site choice, selecting superior sites even when they are nine times further away than the alternative. How do they do this? We used radio-frequency identification-tagged ants to monitor individual behaviour. Here we show for the first time that switching between nests during the decision process can influence nest choice without requiring direct comparison of nests. Ants finding the poor nest were likely to switch and find the good nest, whereas ants finding the good nest were more likely to stay committed to that nest. When ants switched quickly between the two nests, colonies chose the good nest. Switching by ants that had the opportunity to compare nests had little effect on nest choice. We suggest a new mechanism of collective nest choice: individuals respond to nest quality by the decision either to commit or to seek alternatives. Previously proposed mechanisms, recruitment latency and nest comparison, can be explained as side effects of this simple rule. Colony-level comparison and choice can emerge, without direct comparison by individuals.

Research paper thumbnail of Flexible task allocation and the organization of work in ants

Proceedings of the Royal Society B: Biological Sciences, 2009

Research paper thumbnail of Crystal structure of the SOS cell division inhibitor SulA and in complex with FtsZ

Proceedings of the National Academy of Sciences, 2003

SulA halts cell division in Escherichia coli by binding to the major component of the division ma... more SulA halts cell division in Escherichia coli by binding to the major component of the division machinery FtsZ. We have solved the crystal structure of SulA alone and in complex with FtsZ from Pseudomonas aeruginosa. SulA is expressed when the SOS response is induced. This is a mechanism to inhibit cell division and repair DNA in the event of DNA damage. FtsZ is a tubulin-like protein that forms polymers, with the active-site GTPase split across two monomers. One monomer provides the GTP-binding site and the other, through its T7 loop nucleotide hydrolysis. Our structures show that SulA is a dimer, and that SulA inhibits cell division neither by binding the nucleotide-binding site nor by inducing conformational changes in FtsZ. Instead, SulA binds the T7 loop surface of FtsZ, opposite the nucleotide-binding site, blocking polymer formation. These findings explain why GTP hydrolysis and polymer turnover are required for SulA inhibition.

Research paper thumbnail of A Simple Threshold Rule Is Sufficient to Explain Sophisticated Collective Decision-Making

PLoS ONE, 2011

Decision-making animals can use slow-but-accurate strategies, such as making multiple comparisons... more Decision-making animals can use slow-but-accurate strategies, such as making multiple comparisons, or opt for simpler, faster strategies to find a 'good enough' option. Social animals make collective decisions about many group behaviours including foraging and migration. The key to the collective choice lies with individual behaviour. We present a case study of a collective decision-making process (house-hunting ants, Temnothorax albipennis), in which a previously proposed decision strategy involved both quality-dependent hesitancy and direct comparisons of nests by scouts. An alternative possible decision strategy is that scouting ants use a very simple quality-dependent threshold rule to decide whether to recruit nestmates to a new site or search for alternatives. We use analytical and simulation modelling to demonstrate that this simple rule is sufficient to explain empirical patterns from three studies of collective decision-making in ants, and can account parsimoniously for apparent comparison by individuals and apparent hesitancy (recruitment latency) effects, when available nests differ strongly in quality. This highlights the need to carefully design experiments to detect individual comparison. We present empirical data strongly suggesting that best-of-n comparison is not used by individual ants, although individual sequential comparisons are not ruled out. However, by using a simple threshold rule, decision-making groups are able to effectively compare options, without relying on any form of direct comparison of alternatives by individuals. This parsimonious mechanism could promote collective rationality in group decision-making.

Research paper thumbnail of Record Dynamics in Ants

PLoS ONE, 2010

The success of social animals (including ourselves) can be attributed to efficiencies that arise ... more The success of social animals (including ourselves) can be attributed to efficiencies that arise from a division of labour. Many animal societies have a communal nest which certain individuals must leave to perform external tasks, for example foraging or patrolling. Staying at home to care for young or leaving to find food is one of the most fundamental divisions of labour. It is also often a choice between safety and danger. Here we explore the regulation of departures from ant nests. We consider the extreme situation in which no one returns and show experimentally that exiting decisions seem to be governed by fluctuating record signals and ant-ant interactions. A record signal is a new 'high water mark' in the history of a system. An ant exiting the nest only when the record signal reaches a level it has never perceived before could be a very effective mechanism to postpone, until the last possible moment, a potentially fatal decision. We also show that record dynamics may be involved in first exits by individually tagged ants even when their nest mates are allowed to re-enter the nest. So record dynamics may play a role in allocating individuals to tasks, both in emergencies and in everyday life. The dynamics of several complex but purely physical systems are also based on record signals but this is the first time they have been experimentally shown in a biological system.

Research paper thumbnail of Insect communication: ‘No entry’ signal in ant foraging

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

Journal of Theoretical Biology, 2008

Pharaoh's ants organise their foraging system using three types of trail pheromone. All previous ... more 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.