Eusocial insect colonies have often been imagined as “superorganisms” exhibiting tight homeostasis at the colony level. However, colonies lack the tight spatial and organizational integration that many multicellular, unitary organisms exhibit. Precise regulation requires rapid feedback, which is often not possible when nestmates are distributed across space, making decisions asynchronously. Thus, one should expect poorer regulation in superorganisms than unitary organisms.Here, I investigate aspects of regulation in collective foraging behaviors that involve both slow and rapid feedback processes. In Chapter 2, I examine a tightly coupled system with near-instantaneous signaling: teams of weaver ants cooperating to transport massive prey items back to their nest. I discover that over an extreme range of scenarios—even up vertical surfaces—the efficiency per transporter remains constant. My results suggest that weaver ant colonies are maximizing their total intake rate by regulating the allocation of transporters among loads. This is an exception that “proves the rule;” the ant teams are recapitulating the physical integration of unitary organisms. Next, I focus on a process with greater informational constraints, with loose temporal and spatial integration. In Chapter 3, I measure the ability of solitarily foraging Ectatomma ruidum colonies to balance their collection of protein and carbohydrates given different nutritional environments. Previous research has found that ant species can precisely collect a near-constant ratio between these two macronutrients, but I discover these studies were using flawed statistical approaches. By developing a quantitative measure of regulatory effect size, I show that colonies of E. ruidum are relatively insensitive to small differences in food source nutritional content, contrary to previously published claims. In Chapter 4, I design an automated, micro-RFID ant tracking system to investigate how the foraging behavior of individuals integrates into colony-level nutrient collection. I discover that spatial fidelity to food resources, not individual specialization on particular nutrient types, best predicts individual forager behavior. These findings contradict previously published experiments that did not use rigorous quantitative measures of specialization and confounded the effects of task type and resource location.

Chemical Reaction Networks (CRNs) provide a useful framework for modeling andcontrolling large numbers of agents that undergo stochastic transitions between a set of states in a manner similar to chemical compounds. By utilizing CRN models to design agent control policies, some of the computational challenges in the coordination of multi-agent systems can be overcome. In this thesis, a CRN model is developed that defines agent control policies for a multi-agent construction task. The use of surface CRNs to overcome the tradeoff between speed and accuracy of task performance is explained. The computational difficulties involved in coordinating multiple agents to complete collective construction tasks is then discussed. A method for stochastic task and motion planning (TAMP) is proposed to explain how a TAMP solver can be applied with CRNs to coordinate multiple agents. This work defines a collective construction scenario in which a group of noncommunicating agents must rearrange blocks on a discrete domain with obstacles into a predefined target distribution. Four different construction tasks are considered with 10, 20, 30, or 40 blocks, and a simulation of each scenario with 2, 4, 6, or 8 agents is performed. As the number of blocks increases, the construction problem becomes more complex, and a given population of agents requires more time to complete the task. Populations of fewer than 8 agents are unable to solve the 30-block and 40-block problems in the allotted simulation time, suggesting an inflection point for computational feasibility, implying that beyond that point the solution times for fewer than 8 agents would be expected to increase significantly. For a group of 8 agents, the time to complete the task generally increases as the number of blocks increases, except for the 30-block problem, which has specifications that make the task slightly easier for the agents to complete compared to the 20-block problem. For the 10-block and 20- block problems, the time to complete the task decreases as the number of agents increases; however, the marginal effect of each additional two agents on this time decreases. This can be explained through the pigeonhole principle: since there area finite number of states, when the number of agents is greater than the number of available spaces, deadlocks start to occur and the expectation is that the overall solution time to tend to infinity.

Variation in living systems and how it cascades across organizational levels is central to biology. To understand the constraints and amplifications of variation in collective systems, I mathematically study how group-level differences emerge from individual variation in eusocial-insect colonies, which are inherently diverse and easily observable individually and collectively. Considering collective processes in three species where increasing degrees of heterogeneity are relevant, I address how individual variation scales to colony-level variation and to what degree it is adaptive. In Chapter 2, I introduce a Markov-chain decision model for stochastic individual quorum-based recruitment decisions of rock-ant workers during house hunting, and how they determine collective speed--accuracy balance. Differences in the average threshold-dependent response characteristics of workers between colonies cause collective differences in decision-making. Moreover, noisy behavior may prevent drastic collective cascading into poor nests. In Chapter 3, I develop an ordinary differential equation (ODE) model to study how cognitive diversity among honey-bee foragers influences collective attention allocation between novel and familiar resources. Results provide a mechanistic basis for changes in foraging activity and preference with group composition. Moreover, sensitivity analysis reveals that the main individual driver for foraging allocation shifts from recruitment (communication) to persistence (independent effort) as colony composition changes. This might favor specific degrees of heterogeneity that best amplify communication in wild colonies. Lastly, in Chapter 4, I consider diversity in size, age, and task for nest defense in stingless bees. To better understand how these dimensions of diversity interact to balance defensive demands with other colony needs, I study their effect on colony size and task allocation through a demographic Filippov ODE model. Along each dimension, variation is beneficial in a certain range, outside of which colony adaptation and survival are compromised. This work elucidates how variation in collective properties emerges from nonlinear interactions between varying components in eusocial insects, but it can be generalized to other biological systems with similar fundamental characteristics but less empirical tractability. Moreover, it has the potential of inspiring algorithms that capitalize on heterogeneity in engineered systems where simple components with limited information and no central control must solve complex tasks.

The flexibility and robustness of social insect colonies, when they cope with challenges as integrated units, raise many questions, such as how hundreds and thousands of individual local responses are coordinated without a central controlling process. Answering such questions requires: 1. Quantifiable collective responses of colonies under specific scenarios; 2. Decomposability of the collective colony-level response into individual responses; and 3. Mechanisms to integrate the colony- and individual-level responses. In the first part of my dissertation, I explore coordinated collective responses of colonies in during the alarm response to an alarmed nestmate (chapter 2&3). I develop a machine-learning approach to quantitatively estimate the collective and individual alarm response (chapter 2). Using this methodology, I demonstrate that colony alarm responses to the introduction of alarmed nestmates can be decomposed into immediately cascading, followed by variable dampening processes. Each of those processes are found to be modulated by variation in individual alarm responsiveness, as measured by alarm response threshold and persistence of alarm behavior. This variation is modulated in turn by environmental context, in particular with task-related social context (chapter 3). In the second part of my dissertation, I examine the mechanisms responsible for colonial changes in metabolic rate during ontogeny. Prior studies have found that larger ant colonies (as for larger organisms) have lower mass-specific metabolic rates, but the mechanisms remain unclear. In a 3.5-year study on 25 colonies, metabolic rates of colonies and colony components were measured during ontogeny (chapter 4). The scaling of metabolic rate during ontogeny was fit better by segmented regression or quadratic regression models than simple linear regression models, showing that colonies do not follow a universal power-law of metabolism during the ontogenetic development. Furthermore, I showed that the scaling of colonial metabolic rates can be primarily explained by changes in the ratio of brood to adult workers, which nonlinearly affects colonial metabolic rates. At high ratios of brood to workers, colony metabolic rates are low because the metabolic rate of larvae and pupae are much lower than adult workers. However, the high colony metabolic rates were observed in colonies with moderate brood: adult ratios, because higher ratios cause adult workers to be more active and have higher metabolic rates, presumably due to the extra work required to feed more brood.

Astrobiology is premised on the idea that life beyond Earth can exist. Yet, everything known about life is derivative from life on Earth. To understand life beyond Earth, then, requires a definition of life that is abstracted beyond a particular geophysical context. To do this requires a formal understanding of the physical mechanisms by which matter is animated into life. At current, such descriptions are completely lacking for the emergence of life, but do exist for the emergence of consciousness. Namely, contemporary neuroscience offers definitions for universal physical processes that are in one-to-one correspondence with conscious experience. Since consciousness is a sufficient condition for life, these universal definitions of consciousness offer an interesting way forward in terms of the search for life in the cosmos. In this work, I systematically examine Integrated Information Theory (IIT), a well-established theory of consciousness, with the aim of applying it in both biological and astrobiological settings. Surprisingly, I discover major problems with Integrated Information Theory on two fronts: mathematical and epistemological. On the mathematical side, I show how degeneracies buried deep within the theory render it mathematically ill-defined, while on the epistemological side, I prove that the postulates of IIT are scientifically unfalsifiable and inherently metaphysical. Given that IIT is the preeminent theory of consciousness in modern neuroscience, these results have far-reaching implications in this field. In addition, I show that the epistemic issues of falsifiability that hamstring IIT apply quite generally to all contemporary theories of consciousness, which suggests a major reframing of the problem is necessary. The problems that I reveal in regard to defining consciousness offer an important parallel in regard to defining life, as both fields seek to define their topic of study in absence of an existing theoretical framework. To avoid metaphysical problems related to falsifiability, universal theories of both life and consciousness must be framed with respect to independent empirical observations that can be used to benchmark predictions from the theory. In this regard, I argue that the epistemic debate over scientific theories of consciousness should be used to inform the discussion regarding theoretical definitions of life.

In a multi-robot system, locating a team robot is an important issue. If robots

can refer to the location of team robots based on information through passive action

recognition without explicit communication, various advantages (e.g. improving security

for military purposes) can be obtained. Specifically, when team robots follow

the same motion rule based on information about adjacent robots, associations can

be found between robot actions. If the association can be analyzed, this can be a clue

to the remote robot. Using these clues, it is possible to infer remote robots which are

outside of the sensor range.

In this paper, a multi-robot system is constructed using a combination of Thymio

II robotic platforms and Raspberry pi controllers. Robots moving in chain-formation

take action using motion rules based on information obtained through passive action

recognition. To find associations between robots, a regression model is created using

Deep Neural Network (DNN) and Long Short-Term Memory (LSTM), one of state-of-art technologies.

The input data of the regression model is divided into historical data, which

are consecutive positions of the robot, and observed data, which is information about the

observed robot. Historical data is sequence data that is analyzed through the LSTM

layer. The accuracy of the regression model designed using DNN can vary depending

on the quantity and quality of the input. In this thesis, three different input situations

are assumed for comparison. First, the amount of observed data is different, second, the

type of observed data is different, and third, the history length is different. Comparative

models are constructed for each case, and prediction accuracy is compared to analyze

the effect of input data on the regression model. This exploration validates that these

methods from deep learning can reduce the communication demands in coordinated

motion of multi-robot systems

What makes living systems different than non-living ones? Unfortunately this question is impossible to answer, at least currently. Instead, we must face computationally tangible questions based on our current understanding of physics, computation, information, and biology. Yet we have few insights into how living systems might quantifiably differ from their non-living counterparts, as in a mathematical foundation to explain away our observations of biological evolution, emergence, innovation, and organization. The development of a theory of living systems, if at all possible, demands a mathematical understanding of how data generated by complex biological systems changes over time. In addition, this theory ought to be broad enough as to not be constrained to an Earth-based biochemistry. In this dissertation, the philosophy of studying living systems from the perspective of traditional physics is first explored as a motivating discussion for subsequent research. Traditionally, we have often thought of the physical world from a bottom-up approach: things happening on a smaller scale aggregate into things happening on a larger scale. In addition, the laws of physics are generally considered static over time. Research suggests that biological evolution may follow dynamic laws that (at least in part) change as a function of the state of the system. Of the three featured research projects, cellular automata (CA) are used as a model to study certain aspects of living systems in two of them. These aspects include self-reference, open-ended evolution, local physical universality, subjectivity, and information processing. Open-ended evolution and local physical universality are attributed to the vast amount of innovation observed throughout biological evolution. Biological systems may distinguish themselves in terms of information processing and storage, not outside the theory of computation. The final research project concretely explores real-world phenomenon by means of mapping dominance hierarchies in the evolution of video game strategies. Though the main question of how life differs from non-life remains unanswered, the mechanisms behind open-ended evolution and physical universality are revealed.