by Javad Mohamadichamgavi
As I began my research as an Early Stage Researcher in the EvoGamePlus project, a question consistently emerged, first in my mind and later from audiences at my presentations: Why study delays in evolutionary games? This blog post will delve into the rationale behind investigating the impact of delays on evolutionary game theory (EGT), drawing on real-world examples of delays we encounter in biological and social systems. First, some examples of delays observed in real-world phenomena are presented. Following this, we provide a brief literature review on the incorporation of delay in evolutionary game models.
Time delays in biological systems
Time delays are an important feature in many biological systems and are widely used in mathematical biology, with applications ranging from population dynamics, gene expression, virus dynamics, and cancer growth. In ecological and biological systems, interactions among population agents play a crucial role in the growth and persistence of populations. These agents, whether they are cells within an organism or animals in nature, can interact and exchange information, and based on this information, some action occurs. The effects of these interactions are not always immediate, and often, there is a time lag between the initial interaction and its observable impact on the population. Therefore, delay is a common phenomenon across different scales of biological organization. For instance, when a gene is triggered to produce a protein, this doesn’t happen immediately. Instead, the cell undergoes stages such as transcription (DNA to RNA copying) and translation (using RNA for protein synthesis). Each of these stages requires time, leading to a lag between the gene’s activation and the appearance of the protein product. These delays can also be observed in communication and evolutionary changes in populations. It has been observed that these time lags significantly influence the behavior of systems. They can induce changes like fluctuations in protein levels, enhance stability, or alter existing patterns. In cancer studies, researchers leverage time delays to gain insights into tumor growth mechanisms and immune responses against them. These delays reflect the duration needed for processes to unfold, such as cancer cell proliferation or immune cell identification and attack on tumors. For instance, there exists a delay from when an immune cell recognizes a cancer cell to when it becomes prepared to combat it.
These are just a few examples of delays observed in biological systems. Looking at past research, you can find many other biological processes where delays are important.
Time delays in social systems
Time delays are essential in social science research as they provide insights into social interactions and decision-making processes. In social science, we study groups of people with different traits who interact with each other. These interactions can be simple, like following someone on Facebook, or more complex, like how people interact in a city or university. Therefore, it is possible to observe a delay between when people interact and when they take certain actions. For example, in phenomena like the spread of innovations and social trends, the gap between exposure and adoption introduces characteristics that are influenced by factors such as reinforcement and information accumulation. This temporal element poses challenges to modeling methods while enriching our comprehension of how ideas and behaviors propagate within societies. In the realm of economics, time delays are observable in the adoption of technologies. Companies often delay implementing innovations due to cost considerations, uncertainties regarding profitability, or expectations of advancements. This deliberate delay allows for information gathering and risk management, underscoring the significance of timing in business decision-making. Researchers also utilize concepts of time delays to explore reciprocity and strategic behavior in controlled experiments. The spread of viruses among people is also a common example of delay in both biological and social systems together. In a pandemic, we look at how a virus moves from infected to healthy (susceptible) people. The spread can vary depending on the type of virus. Sometimes, a person may catch a virus after interaction with an infected individual but not show symptoms right away. This delay can greatly affect how the epidemic spreads. The length of the delay and the severity of the virus can lead to social problems, and therefore, scientists always try to consider it in their epidemic models.
Time delays in evolutionary game theory
Until now, we have presented several examples of real-world phenomena exhibiting time delays. However, these examples represent only a fraction of the many situations in which time delay can be observed. The occurrence of time delays is widespread across various fields and systems. Now is the time to answer why we are interested in investigating time delays in EGT. In general, EGT offers a framework for modeling various social and biological processes as populations of interacting individuals. These populations can range from animals sharing food to people spreading news on social networks, or even cells in our bodies. A crucial aspect of EGT is the interaction between individuals within these systems. Agents in a population interact and, based on these interactions and other factors, receive points that influence their ability to persist, spread, compete, or potentially become extinct. Traditionally, EGT models assume that the interactions between individuals and the resulting responses are immediate. However, to create more realistic models, we should consider the possibility of time delays between actions and their consequences. Of course, the scale of these delays can vary depending on the type of population.
The replicator dynamic is usually the first and most famous model that new researchers learn about in EGT. It is a mathematical equation that shows how the fraction of different strategies in a population changes over time. Therefore, it is obvious that replicator dynamics are considered the earliest work of studying time delay in evolutionary games. Alboszta and Miękisz, in their research on discrete systems, identified two distinct categories of time delay in replicator dynamics: social delay and biological delay. Social delay exclusively impacts payoffs, with individuals reproducing based on past payoffs at each time step. In contrast, biological delay affects both payoffs and frequencies, representing scenarios where individuals born in the past reproduce according to historical payoffs. Their findings demonstrated that these different types of time delays exert varying influences on the stability of mixed evolutionarily stable strategies. They demonstrated that for small time delays in the social type of delay, the stable stationary state of replicator dynamics remains unchanged. However, as the delay increases beyond a critical threshold, a Hopf bifurcation occurs, leading to oscillatory behavior. This finding aligns with previous research on payoff delays, which Alboszta and Miękisz later termed “social type of delay.” In contrast, biological delays do not affect the stability of the stationary state, with one stable state persisting even under significant time delays. Interestingly, further studies revealed that when biological delay is strategy-dependent, it can shift the stationary state’s value, such that increasing the delay for a particular strategy disadvantages that strategy.
Subsequent researchers incorporated time delay into game theory models, exploring both fixed delays and those following distributions. They investigated delay in conjunction with other factors such as mutation, migration, punishment, and penalties, discovering that in certain scenarios, delay could significantly alter previously established results. More recently, studies have examined the impact of time delay in other frameworks, including stochastic processes affecting fixation and cooperative behavior. These investigations have yielded novel insights into the role of time delay factors in evolutionary game models.
Conclusion
In this short blog post, I try to answer this question: why do we study time delay in evolutionary game theory? Real-world phenomena often exhibit a lag between interaction and action, necessitating the inclusion of time delay for more accurate representations. By examining various examples where such delays occur naturally, we underscore the importance of this factor in creating realistic models of evolutionary games. While numerous researchers have investigated the impact of time delay in evolutionary game models, many questions remain unanswered.
References
Feng, J., Sevier, S. A., Huang, B., Jia, D., & Levine, H. (2016). Modeling delayed processes in biological systems. Physical Review E, 94(3), 032408.
Orosz, G., Moehlis, J., & Murray, R. M. (2010). Controlling biological networks by time-delayed signals. Philosophical Transactions of the Royal Society A: Mathematical, Physical and Engineering Sciences, 368(1911), 439-454.
Sargood, A., Gaffney, E. A., & Krause, A. L. (2022). Fixed and distributed gene expression time delays in reaction–diffusion systems. Bulletin of Mathematical Biology, 84(9), 98.
Dehingia, K., Sarmah, H. K., Alharbi, Y., & Hosseini, K. (2021). Mathematical analysis of a cancer model with time-delay in tumor-immune interaction and stimulation processes. Advances in Difference Equations, 2021, 1-27.
Rahman, B. (2020). Time-delay systems: An overview. Nonlinear Phenomena in Complex Systems, 23(07).
Richard, J. P. (2003). Time-delay systems: an overview of some recent advances and open problems. Automatica, 39(10), 1667-1694.
Fanelli, V., & Maddalena, L. (2012). A time delay model for the diffusion of a new technology. Nonlinear Analysis: Real World Applications, 13(2), 643-649.
Neo, W. S., Yu, M., Weber, R. A., & Gonzalez, C. (2013). The effects of time delay in reciprocity games. Journal of Economic Psychology, 34, 20-35.
Wang, W., Stanley, H. E., & Braunstein, L. A. (2018). Effects of time-delays in the dynamics of social contagions. New Journal of Physics, 20(1), 013034.
Zhang, Z., Kundu, S., Tripathi, J. P., & Bugalia, S. (2020). Stability and Hopf bifurcation analysis of an SVEIR epidemic model with vaccination and multiple time delays. Chaos, Solitons & Fractals, 131, 109483.
Alboszta, J., & Mie, J. (2004). Stability of evolutionarily stable strategies in discrete replicator dynamics with time delay. Journal of theoretical biology, 231(2), 175-179.
Miękisz, J., & Bodnar, M. (2021). Evolution of populations with strategy-dependent time delays. Physical Review E, 103(1), 012414.
Yi, T., & Zuwang, W. (1997). Effect of time delay and evolutionarily stable strategy. Journal of theoretical biology, 187(1), 111-116.