Inferring COVID-19 testing and vaccination behavior from New Jersey testing data.

Characterizing the relationship between disease testing behaviors and infectious disease dynamics is of great importance for public health. Tests for both current and past infection can influence disease-related behaviors at the individual level, while population-level knowledge of an epidemic's course may feed back to affect one's likelihood of taking a test. The COVID-19 pandemic has generated testing data on an unprecedented scale for tests detecting both current infection (PCR, antigen) and past infection (serology); this opens the way to characterizing the complex relationship between testing behavior and infection dynamics. Leveraging a rich database of individualized COVID-19 testing histories in New Jersey, we analyze the behavioral relationships between PCR and serology tests, infection, and vaccination. We quantify interactions between individuals' test-taking tendencies and their past testing and infection histories, finding that PCR tests were disproportionately taken by people currently infected, and serology tests were disproportionately taken by people with past infection or vaccination. The effects of previous positive test results on testing behavior are less consistent, as individuals with past PCR positives were more likely to take subsequent PCR and serology tests at some periods of the epidemic time course and less likely at others. Lastly, we fit a model to the titer values collected from serology tests to infer vaccination trends, finding a marked decrease in vaccination rates among individuals who had previously received a positive PCR test. These results exemplify the utility of individualized testing histories in uncovering hidden behavioral variables affecting testing and vaccination.

Individual costs and societal benefits of interventions during the COVID-19 pandemic.

Individual and societal reactions to an ongoing pandemic can lead to social dilemmas: In some cases, each individual is tempted to not follow an intervention, but for the whole society, it would be best if they did. Now that in most countries, the extent of regulations to reduce SARS-CoV-2 transmission is very small, interventions are driven by individual decision-making. Assuming that individuals act in their best own interest, we propose a framework in which this situation can be quantified, depending on the protection the intervention provides to a user and to others, the risk of getting infected, and the costs of the intervention. We discuss when a tension between individual and societal benefits arises and which parameter comparisons are important to distinguish between different regimes of intervention use.

Non-equilibrium early-warning signals for critical transitions in ecological systems.

Complex systems can exhibit sudden transitions or regime shifts from one stable state to another, typically referred to as critical transitions. It becomes a great challenge to identify a robust warning sufficiently early that action can be taken to avert a regime shift. We employ landscape-flux theory from nonequilibrium statistical mechanics as a general framework to quantify the global stability of ecological systems and provide warning signals for critical transitions. We quantify the average flux as the nonequilibrium driving force and the dynamical origin of the nonequilibrium transition while the entropy production rate as the nonequilibrium thermodynamic cost and thermodynamic origin of the nonequilibrium transition. Average flux, entropy production, nonequilibrium free energy, and time irreversibility quantified by the difference in cross-correlation functions forward and backward in time can serve as early warning signals for critical transitions much earlier than other conventional predictors. We utilize a classical shallow lake model as an exemplar for our early warning prediction. Our proposed method is general and can be readily applied to assess the resilience of many other ecological systems. The early warning signals proposed here can potentially predict critical transitions earlier than established methods and perhaps even sufficiently early to avert catastrophic shifts.

Evolutionary dynamics within and among competing groups.

Biological and social systems are structured at multiple scales, and the incentives of individuals who interact in a group may diverge from the collective incentive of the group as a whole. Mechanisms to resolve this tension are responsible for profound transitions in evolutionary history, including the origin of cellular life, multicellular life, and even societies. Here, we synthesize a growing literature that extends evolutionary game theory to describe multilevel evolutionary dynamics, using nested birth-death processes and partial differential equations to model natural selection acting on competition within and among groups of individuals. We analyze how mechanisms known to promote cooperation within a single group-including assortment, reciprocity, and population structure-alter evolutionary outcomes in the presence of competition among groups. We find that population structures most conducive to cooperation in multiscale systems can differ from those most conducive within a single group. Likewise, for competitive interactions with a continuous range of strategies we find that among-group selection may fail to produce socially optimal outcomes, but it can nonetheless produce second-best solutions that balance individual incentives to defect with the collective incentives for cooperation. We conclude by describing the broad applicability of multiscale evolutionary models to problems ranging from the production of diffusible metabolites in microbes to the management of common-pool resources in human societies.

Spreading processes with mutations over multilayer networks.

A key scientific challenge during the outbreak of novel infectious diseases is to predict how the course of the epidemic changes under countermeasures that limit interaction in the population. Most epidemiological models do not consider the role of mutations and heterogeneity in the type of contact events. However, pathogens have the capacity to mutate in response to changing environments, especially caused by the increase in population immunity to existing strains, and the emergence of new pathogen strains poses a continued threat to public health. Further, in the light of differing transmission risks in different congregate settings (e.g., schools and offices), different mitigation strategies may need to be adopted to control the spread of infection. We analyze a multilayer multistrain model by simultaneously accounting for i) pathways for mutations in the pathogen leading to the emergence of new pathogen strains, and ii) differing transmission risks in different settings, modeled as network layers. Assuming complete cross-immunity among strains, namely, recovery from any infection prevents infection with any other (an assumption that will need to be relaxed to deal with COVID-19 or influenza), we derive the key epidemiological parameters for the multilayer multistrain framework. We demonstrate that reductions to existing models that discount heterogeneity in either the strain or the network layers may lead to incorrect predictions. Our results highlight that the impact of imposing/lifting mitigation measures concerning different contact network layers (e.g., school closures or work-from-home policies) should be evaluated in connection with their effect on the likelihood of the emergence of new strains.

Coupled models of genomic surveillance and evolving pandemics with applications for timely public health interventions.

Disease surveillance systems provide early warnings of disease outbreaks before they become public health emergencies. However, pandemics containment would be challenging due to the complex immunity landscape created by multiple variants. Genomic surveillance is critical for detecting novel variants with diverse characteristics and importation/emergence times. Yet, a systematic study incorporating genomic monitoring, situation assessment, and intervention strategies is lacking in the literature. We formulate an integrated computational modeling framework to study a realistic course of action based on sequencing, analysis, and response. We study the effects of the second variant's importation time, its infectiousness advantage and, its cross-infection on the novel variant's detection time, and the resulting intervention scenarios to contain epidemics driven by two-variants dynamics. Our results illustrate the limitation in the intervention's effectiveness due to the variants' competing dynamics and provide the following insights: i) There is a set of importation times that yields the worst detection time for the second variant, which depends on the first variant's basic reproductive number; ii) When the second variant is imported relatively early with respect to the first variant, the cross-infection level does not impact the detection time of the second variant. We found that depending on the target metric, the best outcomes are attained under different interventions' regimes. Our results emphasize the importance of sustained enforcement of Non-Pharmaceutical Interventions on preventing epidemic resurgence due to importation/emergence of novel variants. We also discuss how our methods can be used to study when a novel variant emerges within a population.

Coexistence in diverse communities with higher-order interactions.

A central assumption in most ecological models is that the interactions in a community operate only between pairs of species. However, two species may interactively affect the growth of a focal species. Although interactions among three or more species, called higher-order interactions, have the potential to modify our theoretical understanding of coexistence, ecologists lack clear expectations for how these interactions shape community structure. Here we analytically predict and numerically confirm how the variability and strength of higher-order interactions affect species coexistence. We found that as higher-order interaction strengths became more variable across species, fewer species could coexist, echoing the behavior of pairwise models. If interspecific higher-order interactions became too harmful relative to self-regulation, coexistence in diverse communities was destabilized, but coexistence was also lost when these interactions were too weak and mutualistic higher-order effects became prevalent. This behavior depended on the functional form of the interactions as the destabilizing effects of the mutualistic higher-order interactions were ameliorated when their strength saturated with species' densities. Last, we showed that more species-rich communities structured by higher-order interactions lose species more readily than their species-poor counterparts, generalizing classic results for community stability. Our work provides needed theoretical expectations for how higher-order interactions impact species coexistence in diverse communities.

Sharp thresholds limit the benefit of defector avoidance in cooperation on networks.

Consider a cooperation game on a spatial network of habitat patches, where players can relocate between patches if they judge the local conditions to be unfavorable. In time, the relocation events may lead to a homogeneous state where all patches harbor the same relative densities of cooperators and defectors, or they may lead to self-organized patterns, where some patches become safe havens that maintain an elevated cooperator density. Here we analyze the transition between these states mathematically. We show that safe havens form once a certain threshold in connectivity is crossed. This threshold can be analytically linked to the structure of the patch network and specifically to certain network motifs. Surprisingly, a forgiving defector avoidance strategy may be most favorable for cooperators. Our results demonstrate that the analysis of cooperation games in ecological metacommunity models is mathematically tractable and has the potential to link topics such as macroecological patterns, behavioral evolution, and network topology.

Fundamental limitations on efficiently forecasting certain epidemic measures in network models.

The ongoing COVID-19 pandemic underscores the importance of developing reliable forecasts that would allow decision makers to devise appropriate response strategies. Despite much recent research on the topic, epidemic forecasting remains poorly understood. Researchers have attributed the difficulty of forecasting contagion dynamics to a multitude of factors, including complex behavioral responses, uncertainty in data, the stochastic nature of the underlying process, and the high sensitivity of the disease parameters to changes in the environment. We offer a rigorous explanation of the difficulty of short-term forecasting on networked populations using ideas from computational complexity. Specifically, we show that several forecasting problems (e.g., the probability that at least a given number of people will get infected at a given time and the probability that the number of infections will reach a peak at a given time) are computationally intractable. For instance, efficient solvability of such problems would imply that the number of satisfying assignments of an arbitrary Boolean formula in conjunctive normal form can be computed efficiently, violating a widely believed hypothesis in computational complexity. This intractability result holds even under the ideal situation, where all the disease parameters are known and are assumed to be insensitive to changes in the environment. From a computational complexity viewpoint, our results, which show that contagion dynamics become unpredictable for both macroscopic and individual properties, bring out some fundamental difficulties of predicting disease parameters. On the positive side, we develop efficient algorithms or approximation algorithms for restricted versions of forecasting problems.

Sociocultural determinants of global mask-wearing behavior.

Behavioral responses influence the trajectories of epidemics. During the COVID-19 pandemic, nonpharmaceutical interventions (NPIs) reduced pathogen transmission and mortality worldwide. However, despite the global pandemic threat, there was substantial cross-country variation in the adoption of protective behaviors that is not explained by disease prevalence alone. In particular, many countries show a pattern of slow initial mask adoption followed by sharp transitions to high acceptance rates. These patterns are characteristic of behaviors that depend on social norms or peer influence. We develop a game-theoretic model of mask wearing where the utility of wearing a mask depends on the perceived risk of infection, social norms, and mandates from formal institutions. In this model, increasing pathogen transmission or policy stringency can trigger social tipping points in collective mask wearing. We show that complex social dynamics can emerge from simple individual interactions and that sociocultural variables and local policies are important for recovering cross-country variation in the speed and breadth of mask adoption. These results have implications for public health policy and data collection.

Understanding the coevolution of mask wearing and epidemics: A network perspective.

Nonpharmaceutical interventions (NPIs) such as mask wearing can be effective in mitigating the spread of infectious diseases. Therefore, understanding the behavioral dynamics of NPIs is critical for characterizing the dynamics of disease spread. Nevertheless, standard infection models tend to focus only on disease states, overlooking the dynamics of "beneficial contagions," e.g., compliance with NPIs. In this work, we investigate the concurrent spread of disease and mask-wearing behavior over multiplex networks. Our proposed framework captures both the competing and complementary relationships between the dueling contagion processes. Further, the model accounts for various behavioral mechanisms that influence mask wearing, such as peer pressure and fear of infection. Our results reveal that under the coupled disease-behavior dynamics, the attack rate of a disease-as a function of transition probability-exhibits a critical transition. Specifically, as the transmission probability exceeds a critical threshold, the attack rate decreases abruptly due to sustained mask-wearing responses. We empirically explore the causes of the critical transition and demonstrate the robustness of the observed phenomena. Our results highlight that without proper enforcement of NPIs, reductions in the disease transmission probability via other interventions may not be sufficient to reduce the final epidemic size.

Robots as models of evolving systems.

Experimental robobiological physics can bring insights into biological evolution. We present a development of hybrid analog/digital autonomous robots with mutable diploid dominant/recessive 6-byte genomes. The robots are capable of death, rebirth, and breeding. We map the quasi-steady-state surviving local density of the robots onto a multidimensional abstract "survival landscape." We show that robot death in complex, self-adaptive stress landscapes proceeds by a general lowering of the robotic genetic diversity, and that stochastically changing landscapes are the most difficult to survive.

When can higher-order interactions produce stable coexistence?

Most ecological models are based on the assumption that species interact in pairs. Diverse communities, however, can have higher-order interactions, in which two or more species jointly impact the growth of a third species. A pitfall of the common pairwise approach is that it misses the higher-order interactions potentially responsible for maintaining natural diversity. Here, we explore the stability properties of systems where higher-order interactions guarantee that a specified set of abundances is a feasible equilibrium of the dynamics. Even these higher-order interactions which lead to equilibria do not necessarily produce stable coexistence. Instead, these systems are more likely to be stable when the pairwise interactions are weak or facilitative. Correlations between the pairwise and higher-order interactions, however, do permit robust coexistence even in diverse systems. Our work not only reveals the challenges in generating stable coexistence through higher-order interactions but also uncovers interaction patterns that can enable diversity.

On the limits to invasion prediction using coexistence outcomes.

The dynamics of ecological communities in nature are typically characterized by probabilistic processes involving invasion dynamics. Because of technical challenges, however, the majority of theoretical and experimental studies have focused on coexistence dynamics. Therefore, it has become central to understand the extent to which coexistence outcomes can be used to predict analogous invasion outcomes relevant to systems in nature. Here, we study the limits to this predictability under a geometric and probabilistic Lotka-Volterra framework. We show that while individual survival probability in coexistence dynamics can be fairly closely translated into invader colonization probability in invasion dynamics, the translation is less precise between community persistence and community augmentation, and worse between exclusion probability and replacement probability. These results provide a guiding and testable theoretical framework regarding the translatability of outcomes between coexistence and invasion outcomes when communities are represented by Lotka-Volterra dynamics under environmental uncertainty.

Host heterogeneity and epistasis explain punctuated evolution of SARS-CoV-2.

Identifying drivers of viral diversity is key to understanding the evolutionary as well as epidemiological dynamics of the COVID-19 pandemic. Using rich viral genomic data sets, we show that periods of steadily rising diversity have been punctuated by sudden, enormous increases followed by similarly abrupt collapses of diversity. We introduce a mechanistic model of saltational evolution with epistasis and demonstrate that these features parsimoniously account for the observed temporal dynamics of inter-genomic diversity. Our results provide support for recent proposals that saltational evolution may be a signature feature of SARS-CoV-2, allowing the pathogen to more readily evolve highly transmissible variants. These findings lend theoretical support to a heightened awareness of biological contexts where increased diversification may occur. They also underline the power of pathogen genomics and other surveillance streams in clarifying the phylodynamics of emerging and endemic infections. In public health terms, our results further underline the importance of equitable distribution of up-to-date vaccines.

Sunsetting as an adaptive strategy.

Major financial legislation is invariably enacted in the wake of a financial crisis. However, legislating following a crisis is hazardous because information is scarce regarding causes of the crisis, let alone what would be an appropriate response. Compounding the lack of information, crisis-driven legislation is sticky, but financial markets are dynamically innovative, which can undermine the efficacy of regulation. As a result, it is foreseeable that such legislation will contain at least some provisions that are inapt or inadequate or, more often, have consequences that are not well understood or even knowable. This article advocates the use of sunsetting as a mechanism for mitigating the potentially adverse consequences of crisis-driven financial legislation. With sunsetting, after a fixed time span, legislation and its implementing regulation must be reenacted to remain in force. This approach has parallels in evolutionary biology, in which a central issue is the ability to adapt to changing environments. Sunsetting does not mean simply discarding (or reenacting) existing regulations, but revisiting them and improving them, much as mutation and recombination do in the evolutionary process.

Link recommendation algorithms and dynamics of polarization in online social networks.

The level of antagonism between political groups has risen in the past years. Supporters of a given party increasingly dislike members of the opposing group and avoid intergroup interactions, leading to homophilic social networks. While new connections offline are driven largely by human decisions, new connections on online social platforms are intermediated by link recommendation algorithms, e.g., "People you may know" or "Whom to follow" suggestions. The long-term impacts of link recommendation in polarization are unclear, particularly as exposure to opposing viewpoints has a dual effect: Connections with out-group members can lead to opinion convergence and prevent group polarization or further separate opinions. Here, we provide a complex adaptive-systems perspective on the effects of link recommendation algorithms. While several models justify polarization through rewiring based on opinion similarity, here we explain it through rewiring grounded in structural similarity-defined as similarity based on network properties. We observe that preferentially establishing links with structurally similar nodes (i.e., sharing many neighbors) results in network topologies that are amenable to opinion polarization. Hence, polarization occurs not because of a desire to shield oneself from disagreeable attitudes but, instead, due to the creation of inadvertent echo chambers. When networks are composed of nodes that react differently to out-group contacts, either converging or polarizing, we find that connecting structurally dissimilar nodes moderates opinions. Overall, our study sheds light on the impacts of social-network algorithms and unveils avenues to steer dynamics of radicalization and polarization in online social networks.

Unifying deterministic and stochastic ecological dynamics via a landscape-flux approach.

The frequency distributions can characterize the population-potential landscape related to the stability of ecological states. We illustrate the practical utility of this approach by analyzing a forest-savanna model. Savanna and forest states coexist under certain conditions, consistent with past theoretical work and empirical observations. However, a grassland state, unseen in the corresponding deterministic model, emerges as an alternative quasi-stable state under fluctuations, providing a theoretical basis for the appearance of widespread grasslands in some empirical analyses. The ecological dynamics are determined by both the population-potential landscape gradient and the steady-state probability flux. The flux quantifies the net input/output to the ecological system and therefore the degree of nonequilibriumness. Landscape and flux together determine the transitions between stable states characterized by dominant paths and switching rates. The intrinsic potential landscape admits a Lyapunov function, which provides a quantitative measure of global stability. We find that the average flux, entropy production rate, and free energy have significant changes near bifurcations under both finite and zero fluctuation. These may provide both dynamical and thermodynamic origins of the bifurcations. We identified the variances in observed frequency time traces, fluctuations, and time irreversibility as kinematic measures for bifurcations. This framework opens the way to characterize ecological systems globally, to uncover how they change among states, and to quantify the emergence of quasi-stable states under stochastic fluctuations.

Interindividual cooperation mediated by partisanship complicates Madison's cure for "mischiefs of faction".

Political theorists have long argued that enlarging the political sphere to include a greater diversity of interests would cure the ills of factions in a pluralistic society. While the scope of politics has expanded dramatically over the past 75 y, polarization is markedly worse. Motivated by this paradox, we take a bottom-up approach to explore how partisan individual-level dynamics in a diverse (multidimensional) issue space can shape collective-level factionalization via an emergent dimensionality reduction. We extend a model of cultural evolution grounded in evolutionary game theory, in which individuals accumulate benefits through pairwise interactions and imitate (or learn) the strategies of successful others. The degree of partisanship determines the likelihood of learning from individuals of the opposite party. This approach captures the coupling between individual behavior, partisan-mediated opinion dynamics, and an interaction network that changes endogenously according to the evolving interests of individuals. We find that while expanding the diversity of interests can indeed improve both individual and collective outcomes, increasingly high partisan bias promotes a reduction in issue dimensionality via party-based assortment that leads to increasing polarization. When party bias becomes extreme, it also boosts interindividual cooperation, thereby further entrenching extreme polarization and creating a tug-of-war between individual cooperation and societal cohesion. These dangers of extreme partisanship are highest when individuals' interests and opinions are heavily shaped by peers and there is little independent exploration. Overall, our findings highlight the urgency to study polarization in a coupled, multilevel context.

Segregation and clustering of preferences erode socially beneficial coordination.

Polarization on various issues has increased in many Western democracies over the last decades, leading to divergent beliefs, preferences, and behaviors within societies. We develop a model to investigate the effects of polarization on the likelihood that a society will coordinate on a welfare-improving action in a context in which collective benefits are acquired only if enough individuals take that action. We examine the impacts of different manifestations of polarization: heterogeneity of preferences, segregation of the social network, and the interaction between the two. In this context, heterogeneity captures differential perceived benefits from coordinating, which can lead to different intentions and sensitivity regarding the intentions of others. Segregation of the social network can create a bottleneck in information flows about others' preferences, as individuals may base their decisions only on their close neighbors. Additionally, heterogeneous preferences can be evenly distributed in the population or clustered in the local network, respectively reflecting or systematically departing from the views of the broader society. The model predicts that heterogeneity of preferences alone is innocuous and it can even be beneficial, while segregation can hamper coordination, mainly when local networks distort the distribution of valuations. We base these results on a multimethod approach including an online group experiment with 750 individuals. We randomize the range of valuations associated with different choice options and the information respondents have about others. The experimental results reinforce the idea that, even in a situation in which all could stand to gain from coordination, polarization can impede social progress.