Istvan Karsai

Food Quantity Affects Traits of Offspring in the Paper Wasp Polistes metricus (Hymenoptera: Vespidae)

Abstract The effects of food quantity on the morphology and development of the paper wasp Polistes metricus Say are studied, and experimental results are compared with predictions of the parental manipulation hypothesis. Food deprivation led to smaller female offspring. By hand feeding larvae we used a technique that counteracts the queen’s hypothesized ability to restrict food provisioning. Hand feeding larvae did not result in larger offspring, but their abdomen was wider and heavier and the hand-fed wasps survived longer in a cold test. We infer that hand-fed colonies produced more gynes and fewer workers than did control colonies. Results of a restricted nourishment treatment do not support the differential feeding hypothesis, because in fasting colonies the emergence of all larvae was delayed by a month, and we did not detect discriminatory feeding of particular larvae for faster emergence. Although fasting colonies produced fewer offspring, the sex ratio did not show significant differences from the other groups. These data suggest that Polistes metricus colonies are partly able to respond to different nutritional conditions by allocating excess food to increase the number of gynes at the expense of workers.

Organization and Regulation of Nest Construction Behavior in Metapolybia Wasps

Field observations and experiments revealed that construction behavior of Metapolybia wasps is based on parallel processing and distributed decision making. Sixteen behaviors were used to separate five behavioral groupings: specialized water forager, flexible pulpforager, active builder, active generalist, and idle. The idle category proved to be the source and the sink of the other task groups, although specialist foragers tend to retain their duties or take over other active roles. Nest construction is partitioned into three tasks. Pulp foragers transfer wood-pulp to the nest where other wasps (builders) distribute and process it further. The builders incorporate this material into the nest structure on the basis of individual decisions. Water foragers provide the extra water necessary for both building and pulp collecting. Material exchange takes place on the nest between pairs or in small groups. The duration and frequency of different behaviors, the number of wasps belonging to different behavioral groups, and the different scale of specialization in different groups suggest that the colony-level performance and speed are governed by the activity of the pulp foragers, who receive information about both the water saturation level of the colony and the activity of the builders through local interactions. Several predictions of this hypothesis were supported by disturbing the normal construction behavior through removing or decreasing the number of individuals belonging to different behavioral groups or supplying additional building material.

Decentralized control of construction behavior in paper wasps: an overview of the stigmergy approach

Grass [26] coined the term stigmergy (previous work directs and triggers new building actions) to describe a mechanism of decentralized pathway of information flow in social insects. In general, all kinds of multi-agent groups require coordination for their effort and it seems that stigmergy is a very powerful means to coordinate activity over great spans of time and space in a wide variety of systems. In a situation in which many individuals contribute to a collective effort, such as building a nest, stimuli provided by the emerging structure itself can provide a rich source of information for the working insects. The current article provides a detailed review of this stigmergic paradigm in the building behavior of paper wasps to show how stigmergy influenced the understanding of mechanisms and evolution of a particular biological system. The most important feature to understand is how local stimuli are organized in space and time to ensure the emergence of a coherent adaptive structure and to explain how workers could act independently yet respond to stimuli provided through the common medium of the environment of the colony.

Dynamics of colony development in Polistes dominulus: a modeling approach

Abstract A simple model based on feedback mechanisms is developed to describe the dynamics of brood production and colony development of primitively eusocial paper wasps. The presence of pupae and empty cells stimulate egg laying, which varies between a basic rate and a physiological maximum. Newly hatched larvae are fed eggs, causing fluctuations in brood demography and forming cohorts of offspring. The basic feedback mechanisms produce emergent colony-level properties such as synchronized development of the brood and mature nest size. Results suggest that it is incorrect to imply colony decline from lack of nest growth, and that production of waves of offspring can be interpreted as the inevitable result of these simple feedback mechanisms rather than the solution to ultimate optimality criteria. Simulations using the parameters estimated in Polistes dominulus Christ are compared to studies of live wasps to test the validity of the model. Comparing simulated results with a perturbation experiment in nature suggests that feedback relationships establish a system that is robust and resilient against severe disturbance.

Nest shapes in paper wasps: can the variability of forms be deduced from the same construction algorithm?

We constructed a simple model to show that the variability of nest structures in the social wasp genus Polistes could result from a simple algorithm modulated by a single parameter. The interplay between the simple algorithm and the geometry of the nest structure resulted in isotropic growth. New construction on the sides of the comb was initiated according to the size of the sides. Due to the geometry of the comb a new rule emerged: if one side of the structure is extended, it decreases the chance that that side will be further extended in the next step (because the number of possible initiation positions decreases), but at the same time construction on one side increases the probability of the extension of that sides neighbours (because their possible initiation positions increase). This emergent isotropic growth keeps the nest almost circular, regardless of nest size, and it ensures well–centred nests. Eccentric nests reflect differences in initiation preference towards particular sides of the previously built structure. Interplay between structural constraints and the simple behavioural algorithm, which is governed by both structural and ‘external’ information, resulted in nests of different shapes. The model was a good predictor of the range of Polistes nest forms found in nature.

The Crossroads between Biology and Mathematics: The Scientific Method as the Basics of Scientific Literacy

Biology is changing and becoming more quantitative. Research is creating new challenges that need to be addressed in education as well. New educational initiatives focus on combining laboratory procedures with mathematical skills, yet it seems that most curricula center on a single relationship between scientific knowledge and scientific method: that of the validity of knowledge claims, judged in terms of their consistency with data. Collecting data and obtaining results (however quantitative) are commonly part of science, but are not science itself. We envision that the operative use of the complete scientific method will play a critical role in providing the necessary underpinning for the integration of math and biology at various professional levels.

Time Delay Implies Cost on Task Switching: A Model to Investigate the Efficiency of Task Partitioning

Task allocation, and task switching have an important effect on the efficiency of distributed, locally controlled systems such as social insect colonies. Both efficiency and workload distribution are global features of the system which are not directly accessible to workers and can only be sampled locally by an individual in a distributed system. To investigate how the cost of task switching affects global performance we use social wasp societies as a metaphor to construct a simple model system with four interconnected tasks. Our goal is not the accurate description of the behavior of a given species, but to seek general conclusions on the effect of noise and time delay on a behavior that is partitioned into subtasks. In our model a nest structure needs to be constructed by the cooperation of individuals that carry out different tasks: builders, pulp and water foragers, and individuals storing water. We report a simulation study based on a model using delay-differential equations to analyze the trade-off between task switching costs and keeping a high degree of adaptivity in a dynamic, noisy environment. Combining the methods of time-delayed equations and stochastic processes we are able to represent the influence of swarm size and task switching sensitivity. We find that the system is stable for reasonable choices of parameters but shows oscillations for extreme choices of parameters and we find that the system is resilient to perturbations. We identify a trade-off between reaching equilibria of high performance and having short transients.

Sting, Carry and Stock: How Corpse Availability Can Regulate De-Centralized Task Allocation in a Ponerine Ant Colony

We develop a model to produce plausible patterns of task partitioning in the ponerine ant Ectatomma ruidum based on the availability of living prey and prey corpses. The model is based on the organizational capabilities of a “common stomach” through which the colony utilizes the availability of a natural (food) substance as a major communication channel to regulate the income and expenditure of the very same substance. This communication channel has also a central role in regulating task partitioning of collective hunting behavior in a supply&demand-driven manner. Our model shows that task partitioning of the collective hunting behavior in E. ruidum can be explained by regulation due to a common stomach system. The saturation of the common stomach provides accessible information to individual ants so that they can adjust their hunting behavior accordingly by engaging in or by abandoning from stinging or transporting tasks. The common stomach is able to establish and to keep stabilized an effective mix of workforce to exploit the prey population and to transport food into the nest. This system is also able to react to external perturbations in a de-centralized homeostatic way, such as to changes in the prey density or to accumulation of food in the nest. In case of stable conditions the system develops towards an equilibrium concerning colony size and prey density. Our model shows that organization of work through a common stomach system can allow Ectatomma ruidum to collectively forage for food in a robust, reactive and reliable way. The model is compared to previously published models that followed a different modeling approach. Based on our model analysis we also suggest a series of experiments for which our model gives plausible predictions. These predictions are used to formulate a set of testable hypotheses that should be investigated empirically in future experimentation.

Mentoring Interdisciplinary Undergraduate Students via a Team Effort

We describe how a team approach that we developed as a mentoring strategy can be used to recruit, advance, and guide students to be more interested in the interdisciplinary field of mathematical biology, and lead to success in undergraduate research in this field. Students are introduced to research in their first semester via lab rotations. Their participation in the research of four faculty members—two from biology and two from mathematics—gives them a first-hand overview of research in quantitative biology and also some initial experience in research itself. However, one of the primary goals of the lab rotation experience is that of developing teams of students and faculty that combine mathematics and statistics with biology and the life sciences, teams that subsequently mentor undergraduate research in genuine interdisciplinary environments. Thus, the team concept serves not only as a means of establishing interdisciplinary research, but also as a means of incorporating new students into existing research efforts that will then track those students into meaningful research of their own. We report how the team concept is used to support undergraduate research in mathematical biology and what types of team-building strategies have worked for us.

The ‘common stomach’ as information source for the regulation of construction behaviour of the swarm

The construction of nests in insect societies requires building materials, pulp and water foragers, builders and also an organized workforce for effective construction. The central hypothesis of this study is that wasp societies developed a social crop, or common stomach, which stores water and provides a mechanism for worker connectivity, which in turn regulates construction behaviour. Inspired by the construction behaviour of social wasps, an agent-based model is presented to show that via the usage of the common stomach, larger colonies enjoy the benefit of having highly effective foragers, while most of the swarm stays on the nest and only a few engage in highly risky foraging trips. We also demonstrate how colony efficiency changes as a function of colony size and the constitution of the labour distribution, as well as how indirect interactions can increase efficiency of labour in wasp societies.