{"title":"Modeling how hunting strategies and pack size shape each other","authors":"Denys Dutykh , Ramón Escobedo , Lee Spector","doi":"10.1016/j.jtbi.2026.112390","DOIUrl":"10.1016/j.jtbi.2026.112390","url":null,"abstract":"<div><div>Social foraging exhibits unexpected features, such as the existence of a group size threshold above which hunting success is not improved, mainly because, above this threshold, additional individuals are free-riders that withhold effort. These observations have been supported by computational models of group hunting, which reveal a mechanism that causes hunting success to peak at small group sizes. In the model, hunters follow two simple rules: approach the prey until a safe distance is reached, and when closer to the prey than a critical avoidance distance, move away from other hunters. The mechanism is that the spatial configuration that the hunters adopt during the hunt is disrupted by the excessive number of participants. Direct observations of wolves (<em>Canis lupus</em>) in Yellowstone Park have shown that the group size threshold when hunting bison (<em>Bison bison</em>), their most formidable prey, is nearly three times greater than when hunting elk (<em>Cervus elaphus</em>). However, the relationship between prey type and the threshold pack size is complex and non-linear, driven by a feedback loop: hunting strategies are adjusted based on prey size and behaviour, which in turn affects the formation and effectiveness of the pack.</div><div>This study explores how prey size influences the optimal pack size and vice versa. Our analysis confirms the non-linearity of this relationship. As the size and danger of the prey change, the optimal pack size does not follow a simple linear pattern. Instead, it reflects a more complex interaction, where both prey characteristics and wolf hunting strategies determine the most effective group size. This complexity arises from the need to balance the pressure exerted on the prey with the spatial arrangement of the pack. The feedback loop between hunting success and pack size illustrates how adaptations in hunting strategies lead to changes in pack organization, which then impact hunting success. This dynamic interaction underlines the need for models that account for these complex interactions to better understand and predict the behavior of wolf packs in different prey scenarios.</div></div>","PeriodicalId":54763,"journal":{"name":"Journal of Theoretical Biology","volume":"623 ","pages":"Article 112390"},"PeriodicalIF":2.0,"publicationDate":"2026-05-21","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"146088222","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"数学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Natalia G. Lavalle , Jerónimo Miranda-Rodríguez , Emanuel Cura Costa , Augusto Borges , Oriol Viader-Llargués , Hernán López-Schier , Osvaldo Chara
{"title":"Local control of cellular proliferation underlies neuromast regeneration in zebrafish","authors":"Natalia G. Lavalle , Jerónimo Miranda-Rodríguez , Emanuel Cura Costa , Augusto Borges , Oriol Viader-Llargués , Hernán López-Schier , Osvaldo Chara","doi":"10.1016/j.jtbi.2026.112389","DOIUrl":"10.1016/j.jtbi.2026.112389","url":null,"abstract":"<div><div>Biological systems are never in equilibrium, yet they maintain stability in the face of continuous external disturbances. A prime example of this is organ regeneration, during which organs are reliably rebuilt through controlled cellular proliferation. In this study, we employ a cell-based computational modelling approach to investigate the proliferative response of an organ after injury. We developed a minimal two-dimensional Cellular Potts Model (CPM) using empirical data from regenerating neuromasts in larval zebrafish. Remarkably, the CPM both qualitatively and quantitatively recapitulates the regenerative response of neuromasts following laser-mediated cell ablation. Assuming that cell proliferation is locally regulated by a delayed switch, we discovered that mitotic activity ceases once the type-dependent number of neighbouring cells exceeds a deterministic critical threshold. An intriguing corollary of our findings is that a local negative feedback loop among identical cells may represent a general mechanism underlying organ-level proportional homeostasis.</div></div>","PeriodicalId":54763,"journal":{"name":"Journal of Theoretical Biology","volume":"623 ","pages":"Article 112389"},"PeriodicalIF":2.0,"publicationDate":"2026-05-21","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"146094835","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"数学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"Equilibrium red blood shape configurations to Canham-Hilfrich energy: Analytical study","authors":"Houda Fahim , Mohammed Guedda , Nour Eddine Alaa","doi":"10.1016/j.jtbi.2026.112397","DOIUrl":"10.1016/j.jtbi.2026.112397","url":null,"abstract":"<div><div>In this study, we revisit the Canham-Helfrich energy to analytically determine the equilibrium configurations of the red blood cell (RBC). By extending the work of Au and Wan (2003), we establish a sufficient analytical condition ensuring the existence of the biconcave equilibrium shape. This result provides a rigorous complement to previous asymptotic arguments, clarifying the mechanical balance underlying RBC morphology. Moreover, the analytical derivation of the equilibrium shape function from the Helfrich energy naturally reproduces the empirical contour equation proposed by Evans and Fung (1972). This demonstrates that their experimental results can be obtained via the energy-minimization framework, thereby linking experimental observations with a consistent theoretical foundation. The present results provide both a mathematical completion and a physical unification of previous models, confirming that the biconcave RBC geometry arises as an equilibrium configuration governed by simple curvature constraints.</div></div>","PeriodicalId":54763,"journal":{"name":"Journal of Theoretical Biology","volume":"624 ","pages":"Article 112397"},"PeriodicalIF":2.0,"publicationDate":"2026-05-07","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"146144299","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"数学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Verdiana Mustaro , Vincenzo Casolaro , Antonio Di Crescenzo
{"title":"On the dynamics of antigen receptors on the B-cell membrane through a two-dimensional stochastic process","authors":"Verdiana Mustaro , Vincenzo Casolaro , Antonio Di Crescenzo","doi":"10.1016/j.jtbi.2026.112376","DOIUrl":"10.1016/j.jtbi.2026.112376","url":null,"abstract":"<div><div>B cells are important components of the adaptive immune system, responsible for antibody production and working as antigen-presenting cells. B cells display protein receptors on their membrane, which bind with foreign antigens and process them before presenting them to T cells. In this work, we present a stochastic process modeling the dynamics of such receptors on the B cell. The model consists of a two-dimensional birth-death process <span><math><mstyle><mrow><mo>{</mo><mo>(</mo><mi>X</mi><mo>(</mo><mi>t</mi><mo>)</mo><mo>,</mo><mi>Y</mi><mo>(</mo><mi>t</mi><mo>)</mo><mo>)</mo><mo>,</mo><mspace></mspace><mi>t</mi><mo>≥</mo><mn>0</mn><mo>}</mo></mrow></mstyle></math></span> having linear transition rates, where <em>X</em>(<em>t</em>) and <em>Y</em>(<em>t</em>) represent the number of free and occupied receptors, respectively. After determining the partial differential equation for the probability generating function of the process, we compute the main moments of the process, including the covariance. The transient and asymptotic behavior of the means of <em>X</em>(<em>t</em>) and <em>Y</em>(<em>t</em>) is also studied. Throughout the paper, we provide insights into the biological significance of each parameter on the system’s dynamics. In addition, we conduct a sensitivity analysis to assess how variations in the model parameters affect the first-order moments. Such analysis shows that minimal variations of the parameters representing the binding frequency of antigens and B-cell receptors, when happening in the initial instants of the process, result in noticeable alterations of the number of occupied receptors.</div></div>","PeriodicalId":54763,"journal":{"name":"Journal of Theoretical Biology","volume":"622 ","pages":"Article 112376"},"PeriodicalIF":2.0,"publicationDate":"2026-04-07","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"145994734","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"数学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Isobel R. Abell , Thao P. Le , Jennifer A. Flegg , Christopher M. Baker
{"title":"Modelling the spread and management of Varroa destructor in naive european honeybee populations","authors":"Isobel R. Abell , Thao P. Le , Jennifer A. Flegg , Christopher M. Baker","doi":"10.1016/j.jtbi.2025.112329","DOIUrl":"10.1016/j.jtbi.2025.112329","url":null,"abstract":"<div><div><em>Varroa destructor</em> is a significant European honeybee pest, impacting agricultural industries globally. Since arriving in 2022, Australia faces the possibility that <em>Varroa</em> will become established in European honeybee colonies nationally. Australia initially pursued a strategy of testing and subsequently eliminating hives infested with <em>Varroa</em>. These management efforts raise interesting questions about the interplay between hive testing and elimination, and the spread of <em>Varroa</em> between hives. This study uses mathematical modelling to investigate how combined hive testing and elimination strategies impact the spread of <em>Varroa</em> through a network of European honeybee hives. We develop a model of both within-hive reproduction of <em>Varroa</em> and hive testing, and between-hive movement of <em>Varroa</em> on a network of hives. This model is used to assess the impact of various testing and hive elimination strategies on the total number of hives eliminated on the network of hives. Each model simulation starts with a single infested hive, and from this we observed one of two dynamics: either the infestation is caught before spreading, or <em>Varroa</em> spreads extensively through the network before being caught by testing. Within our model we implement two common hive testing methods – sugar shake and alcohol testing. A shared limitation of these testing methods is that they can only detect mites in a specific stage of their lifecycle. As such, testing is not only dependent on how many <em>Varroa</em> mites are in a hive, but also on what lifecycle stage the mites are in at the time of testing. By varying testing and movement parameters, we see that this testing limitation greatly impacts the number of hives eliminated in various scenarios. Furthermore, testing earlier, or testing more frequently, does not guarantee a smaller invasion. Our model results suggest irregular testing schedules, e.g. testing multiple times in close succession rather than just once in a given timeframe, may help overcome the limitations of common hive testing strategies.</div></div>","PeriodicalId":54763,"journal":{"name":"Journal of Theoretical Biology","volume":"622 ","pages":"Article 112329"},"PeriodicalIF":2.0,"publicationDate":"2026-04-07","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"145688186","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"数学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"Modeling tumor progression in heterogeneous microenvironments: A cellular automata approach","authors":"Yue Deng , Mingjing Li , Jinzhi Lei","doi":"10.1016/j.jtbi.2026.112388","DOIUrl":"10.1016/j.jtbi.2026.112388","url":null,"abstract":"<div><div>Understanding how microenvironmental heterogeneity influences tumor progression is essential for advancing both cancer biology and therapeutic strategies. In this study, we develop a cellular automata (CA) model to simulate tumor growth under varying microenvironmental conditions and genetic mutation rates, addressing a gap in existing studies that rarely integrate these two factors to explain tumor dynamics. The model explicitly incorporates the cellular heterogeneity of stem and non-stem cells, dynamic cell-cell interactions, and tumor-microenvironment crosstalk. Using computational simulations, we examine the synergistic effects of gene mutation rate, initial tumor burden, and microenvironmental state on tumor progression. Our results demonstrate that lowering the mutation rate significantly mitigates tumor expansion and preserves microenvironmental integrity. Interestingly, the initial tumor burden has a limited impact, whereas the initial condition of the microenvironment critically shapes tumor dynamics. A supportive microenvironment promotes proliferation and spatial invasion, while inhibitory conditions suppress tumor growth. These findings highlight the key role of microenvironmental modulation in tumor evolution and provide computational insights that may inform more effective cancer therapies.</div></div>","PeriodicalId":54763,"journal":{"name":"Journal of Theoretical Biology","volume":"622 ","pages":"Article 112388"},"PeriodicalIF":2.0,"publicationDate":"2026-04-07","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"146047329","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"数学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Edwin van Leeuwen , Frank G. Sandmann , Rosalind M. Eggo , Peter J. White
{"title":"Social mixing and time use throughout the year: Potential changes in disease transmission and age distribution of cases","authors":"Edwin van Leeuwen , Frank G. Sandmann , Rosalind M. Eggo , Peter J. White","doi":"10.1016/j.jtbi.2025.112349","DOIUrl":"10.1016/j.jtbi.2025.112349","url":null,"abstract":"<div><div>Susceptibility in children is a key driver of heterogeneity in the transmission of different respiratory viruses. For example, SARS-CoV-2 is associated with low susceptibility in children, while for the influenza and respiratory syncytial viruses it is thought that children have higher susceptibility, because adults will have built up natural immunity. We modelled seasonal changes in time use and social mixing based on age-stratified contact rates using historical nationally-representative surveys. We explored changes in the reproduction number and the age distribution of infections of respiratory diseases during the early phase of an epidemic, for different assumptions of susceptibility in children aged 0-15. Across ages, the estimated <em>R</em><sub>0</sub> and the age distribution of incidence fluctuated due to changes in time use. For the scenarios where adults have acquired natural immunity through past infection <em>R</em><sub>0</sub> fell and relative incidence decreased in children aged 0-15 but increased in other ages during holiday periods. If children were less susceptible than adults these changes were less pronounced. Our modelling findings suggest that changing contacts during the holiday periods may shift the age distribution of cases from children towards adults. Given that severity and deaths rise with age for many diseases, more intergenerational mixing risks the disease moving into the more vulnerable following the holidays even if the absolute number of infections did not increase.</div></div>","PeriodicalId":54763,"journal":{"name":"Journal of Theoretical Biology","volume":"622 ","pages":"Article 112349"},"PeriodicalIF":2.0,"publicationDate":"2026-04-07","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"145936505","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"数学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Stéphane Urcun , Yasmin El Mahi , Raluca Eftimie , Zélie Dirand , Gwenaël Rolin , Stéphane P.A. Bordas
{"title":"Simple cellular potts model of scratch assays on healthy and keloid fibroblasts driven by contact inhibition of locomotion","authors":"Stéphane Urcun , Yasmin El Mahi , Raluca Eftimie , Zélie Dirand , Gwenaël Rolin , Stéphane P.A. Bordas","doi":"10.1016/j.jtbi.2025.112365","DOIUrl":"10.1016/j.jtbi.2025.112365","url":null,"abstract":"<div><div>In this study, we propose and validate a simple agent-based model to study cell-cell interactions and cell migration during in vitro scratch assays in the context of cutaneous fibrosis (keloid). For model parametrization, we collect data from in vitro experiments performed with healthy or keloid fibroblasts treated (or not) with type 1 or 2 macrophages secretome to mimic specific in vivo environments. All experiments were performed with mitomycin to inhibit cell proliferation, and subsequently isolate the sole contribution of migration to wound filling over time. The scratch assays are modeled within the cellular Potts model framework. The calibration process, via Levenberg-Maquart algorithm, gives a mean error of 4.53 ± 0.77% across the four modalities (healthy, control, M1 and M2 secretum) and the evaluation dataset gives a mean error of 10.55 ± 0.77%. With the help of this model, we test whether the hypothesis of contact inhibition of locomotion (CIL) can explain the movement of keloid fibroblasts. The simulation results and their comparison with the experimental data suggest that CIL might not characterize the movement of keloid fibroblasts, which is in contrast to the importance of CIL for the movement of healthy fibroblasts.</div></div>","PeriodicalId":54763,"journal":{"name":"Journal of Theoretical Biology","volume":"622 ","pages":"Article 112365"},"PeriodicalIF":2.0,"publicationDate":"2026-04-07","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"146020818","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"数学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"Mathematical modeling of tumor-immune interactions in breast cancer must model tumor-immune interactions in breast cancer","authors":"Heiko Enderling","doi":"10.1016/j.jtbi.2026.112375","DOIUrl":"10.1016/j.jtbi.2026.112375","url":null,"abstract":"","PeriodicalId":54763,"journal":{"name":"Journal of Theoretical Biology","volume":"622 ","pages":"Article 112375"},"PeriodicalIF":2.0,"publicationDate":"2026-04-07","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"146004805","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"数学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
{"title":"An individual-based simulation framework exploring the ecology and mechanistic underpinnings of larval crowding in laboratory populations of Drosophila","authors":"Srikant Venkitachalam , Amitabh Joshi","doi":"10.1016/j.jtbi.2026.112378","DOIUrl":"10.1016/j.jtbi.2026.112378","url":null,"abstract":"<div><div>The study of larval competition in laboratory populations of <em>Drosophila</em>, implemented via the crowding of larval cultures, has contributed greatly to the understanding of the ecology of competition, the evolution of larval competitive ability, and formed the basis of rigorous testing of the theory of density-dependent selection. Earlier studies led to the view that the outcomes of larval competition, and resulting evolutionary consequences of crowding-adaptation, could largely be understood by varying the starting density of individuals in a crowded culture. However, recent studies have shown that the results of adaptation to larval crowding may not be well predicted by the overall larval density (i.e., total starting individuals/total volume of food). Cultures raised at the same overall density but at different egg number and food volume combinations were shown to have different underlying density-specific fitness functions, and crowding-adaptation in each of these cultures was attained through different evolutionary trajectories as well. A recent study showed that cultures with not just the same density, but the same egg and food volume combination, achieved through food columns of differing diameter and height, could also differ greatly in fitness-related trait outcomes. In that study, the density of larvae in the feeding band (volume of food close to the surface in contact with air, to which larval feeding is largely restricted) was a very important factor in predicting the outcomes of larval competition. Given these recent findings, it is important to understand the overall role of feeding band density, and how it influences density-specific fitness functions in different kinds of crowded cultures. As the older models of larval competition are now insufficient to capture current empirical data, we constructed an individual-based simulation framework informed in part by these more recent findings, in order to better understand the evolutionary ecology and mechanistic underpinnings of larval competition, and predict robust experiments for expanding our understanding of the process of larval competition in <em>Drosophila</em>.</div></div>","PeriodicalId":54763,"journal":{"name":"Journal of Theoretical Biology","volume":"622 ","pages":"Article 112378"},"PeriodicalIF":2.0,"publicationDate":"2026-04-07","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"146004789","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"数学","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}