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Group-based phylogenetic models on 3-sunlet networks. 基于群的3-sunlet网络系统发育模型。
IF 2.2 4区 数学 Q2 BIOLOGY Pub Date : 2025-08-18 DOI: 10.1007/s11538-025-01506-1
Shelby Cox, Elizabeth Gross, Samuel Martin

Phylogenetic networks describe the evolution of a set of taxa for which reticulate events have occurred at some point in their evolutionary history. Of particular interest is when the evolutionary history between a set of just three taxa has a reticulate event. In molecular phylogenetics, substitution models can model the process of evolution at the genetic level, and the case of three taxa with a reticulate event can be modeled using a substitution model on a semi-directed graph called a 3-sunlet. We investigate a class of substitution models called group-based phylogenetic models on 3-sunlet networks. In particular, we investigate the discrete geometry of the parameter space and how this relates to the dimension of the phylogenetic variety associated to the model. This enables us to give a dimension formula for this variety for general group-based models when the order of the group is odd.

系统发育网络描述了一组类群的进化,对于这些类群,网状事件在它们的进化史上的某个时刻发生过。特别令人感兴趣的是,当一组仅三个分类群之间的进化史发生网状事件时。在分子系统发育学中,替代模型可以在遗传水平上模拟进化过程,具有网状事件的三个类群的情况可以在称为3-sunlet的半有向图上使用替代模型进行建模。我们研究了一类基于群的3-太阳网络系统发育模型。特别是,我们研究了参数空间的离散几何,以及这与与模型相关的系统发育多样性的维度之间的关系。这使我们能够在群的顺序为奇数时,为一般基于群的模型给出这种变化的维数公式。
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引用次数: 0
Evolutionary Dispersal of Species with Starvation-Driven Diffusion Incorporating Perceptual Constraints in Competition Models in Heterogeneous Habitats. 异质生境竞争模型中包含感知约束的饥饿驱动扩散的物种进化扩散。
IF 2.2 4区 数学 Q2 BIOLOGY Pub Date : 2025-08-15 DOI: 10.1007/s11538-025-01513-2
Youngseok Chang, Wonhyung Choi, Inkyung Ahn

This study examines a competition model featuring nonuniform dispersal, referred to as starvation-driven-type diffusion (SDTD). This model incorporates the motility of species that adhere to a starvation-driven diffusion (SDTD). paradigm while also factoring in perceptual constraints within a spatially heterogeneous region. In the proposed model, our study aims to understand the impact of SDTD on system dynamics in a spatially heterogeneous environment. To achieve this, we consider a Lotka-Volterra-type competition model exhibiting identical population dynamics under no-flux boundary conditions. To illuminate the evolutionary implications of SDTD, this study contrasts two different models. The first model involves two species with distinct, uniform diffusion rates. In comparison, the second model features one species that adheres to a constant diffusion rate and another that operates under the principles of SDTD. This study focuses on analyzing the fitness variation of competing species based on their respective diffusion dynamics: (i) The study examines how the fitness of a species following SDTD changes compared to another species that diffuses at a constant rate. (ii) We investigate the dynamics of fitness alteration within a competitive two-species model wherein one species exhibits a constant diffusion rate while the other species alternates between constant-rate diffusion and SDTD. In addition, we examine how the shifting diffusion strategies of one species affect its fitness relative to a species with a fixed, constant diffusion rate. Our conclusions suggest that a species adhering to SDTD may enhance its fitness, consistent with the model incorporating SDD. Nonetheless, we show that certain circumstances may exist where SDTD does not result in increased fitness for a species, mainly due to the perceptual limitations of that species.

本研究考察了一个具有非均匀扩散特征的竞争模型,即饥饿驱动型扩散(SDTD)。该模型结合了坚持饥饿驱动扩散(SDTD)的物种的运动性。范式,同时也考虑到空间异质区域内的感知约束。在提出的模型中,我们的研究旨在了解SDTD对空间异构环境中系统动力学的影响。为了实现这一点,我们考虑了lotka - voltera型竞争模型,该模型在无通量边界条件下具有相同的种群动态。为了阐明SDTD的进化意义,本研究对比了两种不同的模型。第一个模型涉及两个具有不同的均匀扩散速率的物种。相比之下,第二个模型的特征是一个物种遵循恒定的扩散速率,另一个物种遵循SDTD原则。本研究的重点是分析竞争物种在各自扩散动力学基础上的适应度变化:(i)研究了一个物种在SDTD后的适应度变化与另一个物种在恒定速率扩散时的适应度变化。(ii)研究了两物种竞争模型中适合度变化的动态,其中一个物种表现出恒定的扩散速率,而另一个物种在恒定速率扩散和SDTD之间交替。此外,我们还研究了一个物种的移动扩散策略如何影响其相对于具有固定、恒定扩散速率的物种的适合度。我们的结论表明,加入SDTD的物种可能会提高其适应度,这与加入SDD的模型一致。尽管如此,我们表明,在某些情况下,SDTD可能不会导致一个物种的适应度增加,这主要是由于该物种的感知限制。
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引用次数: 0
Insights on Influenza Control Through Vaccination for the 2023-2024 Season in the USA: Mathematical Modeling, Optimal Control, and Sensitivity Analysis. 美国2023-2024年流感季节疫苗接种控制的见解:数学建模、最优控制和敏感性分析
IF 2.2 4区 数学 Q2 BIOLOGY Pub Date : 2025-08-14 DOI: 10.1007/s11538-025-01494-2
Ana-Maria Croicu

This study examines influenza transmission dynamics through mathematical modeling, vaccination strategies using optimal control, and the impact of key model parameters to the optimal control for the 2023-2024 influenza season in the United States. Using data from the CDC's FluView Interactive and FluVaxView databases, key parameters were estimated, revealing significant variability across states. Tailored vaccination strategies, designed to align with state-specific data and conditions, effectively reduced infections and reproduction numbers in all states. Sensitivity analysis of the optimal vaccination strategy and reproduction number revealed complex interactions among model parameters. These findings emphasize the need for carefully designed public health strategies to manage influenza effectively, supported by key recommendations presented in this paper.

本研究通过数学建模、采用最优控制的疫苗接种策略,以及关键模型参数对美国2023-2024年流感季节最优控制的影响,研究了流感传播动力学。使用来自CDC的FluView Interactive和FluVaxView数据库的数据,估计了关键参数,揭示了各州之间的显著差异。量身定制的疫苗接种战略旨在与各州的具体数据和条件保持一致,有效地减少了所有州的感染和繁殖数量。最优接种策略和繁殖数的敏感性分析揭示了模型参数之间复杂的相互作用。这些发现强调需要精心设计公共卫生战略,以有效管理流感,并得到本文提出的关键建议的支持。
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引用次数: 0
Mathematical Modelling and Uncertainty Quantification for Analysis of Biphasic Coral Reef Recovery Patterns. 双相珊瑚礁恢复模式分析的数学模型和不确定性量化。
IF 2.2 4区 数学 Q2 BIOLOGY Pub Date : 2025-08-12 DOI: 10.1007/s11538-025-01512-3
David J Warne, Kerryn Crossman, Grace E M Heron, Jesse A Sharp, Wang Jin, Paul Pao-Yen Wu, Matthew J Simpson, Kerrie Mengersen, Juan-C Ortiz

Coral reefs are increasingly subjected to major disturbances threatening the health of marine ecosystems. Substantial research is underway to develop intervention strategies that assist reefs in recovery from, and resistance to, inevitable future climate and weather extremes. To assess potential benefits of interventions, mechanistic understanding of coral reef recovery and resistance patterns is essential. Recent evidence suggests that more than half of the reefs surveyed across the Great Barrier Reef (GBR) exhibit deviations from standard recovery modelling assumptions when the initial coral cover is low ( 10 %). New modelling is necessary to account for these observed patterns to better inform management strategies. We consider a new model for reef recovery at the coral cover scale that accounts for biphasic recovery patterns. The model is based on a multispecies Richards' growth model that includes a change point in the recovery patterns. Bayesian inference is applied for uncertainty quantification of key parameters for assessing reef health and recovery patterns. This analysis is applied to benthic survey data from the Australian Institute of Marine Science (AIMS). We demonstrate agreement between model predictions and data across every recorded recovery trajectory with at least two years of observations following disturbance events occurring between 1992-2020. This new approach will enable new insights into the biological, ecological and environmental factors that contribute to the duration and severity of biphasic coral recovery patterns across the GBR. These new insights will help to inform managements and monitoring practice to mitigate the impacts of climate change on coral reefs.

珊瑚礁日益受到威胁海洋生态系统健康的重大干扰。大量的研究正在进行中,以制定干预策略,帮助珊瑚礁从不可避免的未来气候和极端天气中恢复并抵抗。为了评估干预措施的潜在效益,对珊瑚礁恢复和抵抗模式的机制理解是必不可少的。最近的证据表明,在大堡礁(GBR)的调查中,超过一半的珊瑚礁在初始珊瑚覆盖率较低(≤10%)时表现出与标准恢复模型假设的偏差。有必要建立新的模型来解释这些观察到的模式,以便更好地为管理策略提供信息。我们考虑了一种新的珊瑚覆盖范围内的珊瑚礁恢复模型,该模型考虑了双相恢复模式。该模型基于多物种理查兹的增长模型,其中包括恢复模式的变化点。贝叶斯推理应用于评估珊瑚礁健康和恢复模式的关键参数的不确定性量化。该分析应用于澳大利亚海洋科学研究所(AIMS)的底栖生物调查数据。我们证明了在1992-2020年发生扰动事件后至少两年的观测中,模型预测和每一个记录的恢复轨迹的数据之间的一致性。这种新方法将使人们对生物、生态和环境因素有了新的认识,这些因素导致了大堡礁双相珊瑚恢复模式的持续时间和严重程度。这些新的见解将有助于为管理和监测实践提供信息,以减轻气候变化对珊瑚礁的影响。
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引用次数: 0
Modeling Combination Therapies and T Cell Exhaustion Dynamics in the Tumor Under Immune Checkpoint Blockade. 免疫检查点阻断下肿瘤中联合治疗和T细胞衰竭动力学的建模。
IF 2.2 4区 数学 Q2 BIOLOGY Pub Date : 2025-08-11 DOI: 10.1007/s11538-025-01507-0
Xiulan Lai, Teng Yu

Chronic antigen exposure in the tumor microenvironment drives CD 8 + T cell exhaustion, marked by increased inhibitory receptors and diminished effector functions. Immune checkpoint blockade seeks to prevent or reverse exhaustion, but its success relies on the pre-existing state of tumor-infiltrating T cells. To investigate this, we developed a mathematical model examining: (1) how T cell exhaustion disrupts tumor-immune equilibrium, (2) anti-PD-L1 efficacy across exhaustion states, and (3) efficacy of next-generation therapies (e.g., IFN α -anti-PD-L1, PD1-IL2v). Stability analysis and simulations reveal that tumor PD-L1 expression critically influences immune dynamics, particularly the bistability of tumor-free and tumorous states. High PD-1 expression and exhaustion rates correlate with growth of tumor and impaired expansion of less-exhausted CD 8 + T cells. While anti-PD-L1 efficacy depends on baseline exhaustion, severe exhaustion enables immune escape. Next-generation therapies enhancing cytotoxicity and sustaining less-exhausted T cell populations show improved tumor control, suggesting combination strategies may overcome resistance.

肿瘤微环境中的慢性抗原暴露驱动cd8 + T细胞衰竭,其特征是抑制受体增加和效应功能减弱。免疫检查点阻断旨在防止或逆转衰竭,但其成功依赖于肿瘤浸润T细胞的预先存在状态。为了研究这一点,我们建立了一个数学模型来检验:(1)T细胞衰竭如何破坏肿瘤免疫平衡,(2)在衰竭状态下抗pd - l1的功效,以及(3)下一代疗法(例如,IFN α -抗pd - l1, PD1-IL2v)的功效。稳定性分析和模拟表明,肿瘤PD-L1表达对免疫动力学具有重要影响,特别是无瘤和肿瘤状态的双稳定性。高PD-1表达和耗竭率与肿瘤生长和较少耗竭的cd8 + T细胞的扩张受损相关。虽然抗pd - l1的有效性取决于基线耗竭,但严重的耗竭会使免疫逃逸。增强细胞毒性和维持较少耗竭的T细胞群的新一代疗法显示出更好的肿瘤控制,表明联合策略可能克服耐药性。
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引用次数: 0
Optimal Control of Immune Checkpoint Inhibitor Therapy in a Heart-Tumour Model. 免疫检查点抑制剂治疗在心脏肿瘤模型中的最优控制。
IF 2.2 4区 数学 Q2 BIOLOGY Pub Date : 2025-08-11 DOI: 10.1007/s11538-025-01468-4
Solveig A van der Vegt, Ruth E Baker, Sarah L Waters

Autoimmune myocarditis, or cardiac muscle inflammation, is a rare but frequently fatal side-effect of immune checkpoint inhibitors (ICIs), a class of cancer therapies. Despite the dangers that side-effects such as these pose to patients, they are rarely, if ever, included explicitly when mechanistic mathematical modelling of cancer therapy is used for optimization of treatment. In this paper, we develop a two-compartment mathematical model which incorporates the impact of ICIs on both the heart and the tumour. Such a model can be used to inform the conditions under which autoimmune myocarditis may develop as a consequence of treatment. We use this model in an optimal control framework to design optimized dosing schedules for three types of ICI therapy that balance the positive and negative effects of treatment. We show that including the negative side-effects of ICI treatment explicitly within the mathematical framework significantly impacts the predictions for the optimized dosing schedule, thus stressing the importance of a holistic approach to optimizing cancer therapy regimens.

自身免疫性心肌炎或心肌炎症是免疫检查点抑制剂(ICIs)的一种罕见但经常致命的副作用,这是一类癌症治疗方法。尽管诸如此类的副作用会给患者带来危险,但当癌症治疗的机械数学模型用于优化治疗时,它们很少(如果有的话)被明确地包括在内。在本文中,我们开发了一个双室数学模型,其中包含了ICIs对心脏和肿瘤的影响。这样的模型可以用来告知自身免疫性心肌炎作为治疗结果可能发展的条件。我们在最优控制框架中使用该模型来设计三种ICI治疗的优化给药方案,以平衡治疗的积极和消极影响。我们的研究表明,在数学框架中明确包括ICI治疗的负面副作用会显著影响优化给药方案的预测,从而强调了整体方法优化癌症治疗方案的重要性。
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引用次数: 0
Switch-like Behavior in the Heme Receptor for Vibrio Vulnificus. 创伤弧菌血红素受体的开关行为。
IF 2.2 4区 数学 Q2 BIOLOGY Pub Date : 2025-08-09 DOI: 10.1007/s11538-025-01505-2
Kathryn S Lynch, James P Keener

Switch-like behavior and bistability are important features in gene regulatory networks, allowing cells to distinguish between changing environments and express certain genes only under the appropriate conditions. Vibrio vulnificus, an opportunistic Gram-negative marine pathogen, has iron as a limiting growth factor. When inside a human host, this bacteria utilizes heme as a source of iron, necessitating the ability to turn this heme acquisition system off and on in response to environmental pressures. As establishment of infection depends on V. vulnificus's ability to change from a marine to human environment, the ability to switch on the heme-intake system is an important part of establishment of initial infection. In particular, the protein HupA is a key part of the bacteria's heme importation complex, and is regulated primarily by a divergently transcribed protein, HupR. The dynamics of this regulation result in a genetic switch, allowing the bacteria to differentiate between high iron or high heme environments, determining which source of iron should be used. Bifurcation analysis of this network uncovers a saddle-node bifurcation, which encodes this switch-like behavior into the regulation of the heme transport system and allows different levels of expression for HupA depending on external concentrations of heme and iron. The influences of other parameters in this system are also investigated; in particular, promoter leakage is found to be required to enable this bistability, indicating the importance of imperfect regulation in a cell's ability to respond to the environment.

开关样行为和双稳定性是基因调控网络的重要特征,使细胞能够区分不断变化的环境,并仅在适当的条件下表达某些基因。创伤弧菌是一种机会性革兰氏阴性海洋病原体,它的限制性生长因子是铁。当在人类宿主体内时,这种细菌利用血红素作为铁的来源,需要有能力关闭和打开这个血红素获取系统来应对环境压力。由于感染的建立取决于创伤弧菌从海洋环境转变为人类环境的能力,因此开启血红素摄取系统的能力是建立初始感染的重要组成部分。特别是,蛋白质HupA是细菌血红素输入复合物的关键部分,主要由发散转录的蛋白质HupR调节。这种调控的动态导致了一种基因开关,允许细菌区分高铁或高血红素环境,决定应该使用哪种铁来源。对该网络的分岔分析揭示了一个鞍节点分岔,该分岔将这种开关样行为编码为血红素运输系统的调节,并允许HupA根据外部血红素和铁浓度的不同水平表达。研究了其他参数对系统性能的影响;特别是,启动子泄漏被发现是实现这种双稳定性所必需的,这表明了细胞对环境反应能力的不完美调节的重要性。
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引用次数: 0
Pattern Formation Driven by Nonlocal Perception in a Delayed Pine Wilt Disease Model with Top-Hat Kernel. 具有顶帽核的延迟松材萎蔫病模型的非局部感知驱动模式形成。
IF 2.2 4区 数学 Q2 BIOLOGY Pub Date : 2025-08-09 DOI: 10.1007/s11538-025-01504-3
Jia Li, Yuting Ding, Yongli Song

Nonlocal perception plays a crucial role in studying animal cognitive movement modeling. In this paper, the impact of nonlocal perception on pattern formation is analyzed, and it is applied to study the control of pine wilt disease. It turns out that perceptual movement can provide a theoretical scientific basis for the multi-point outbreaks and spatiotemporal aggregation of pine wilt disease. For the top-hat kernel, we concentrate on the joint effect of perception scale and delay on the stability, and find that Turing-Hopf bifurcation occurs due to their interaction. Besides, the patterns near the bifurcation points are simulated in detail by adopting parameters with actual biological meaning, which are selected by analyzing real data, and diverse complicated spatiotemporal patterns are obtained, such as peak alternating periodic patterns and spatiotemporal aggregation patterns. Finally, we demonstrate that the artificial release of the parasitic natural enemy of the pest can drive the populations to reach stability in the form of the steady state or periodic solutions. The obtained results not only well explain the transmission mechanism of pine wilt disease, but also contribute to the study of biological phenomena such as the formations of flocks and swarms.

非局部知觉在动物认知运动建模研究中起着至关重要的作用。本文分析了非局部感知对图案形成的影响,并将其应用于松材萎蔫病的防治研究。结果表明,感知运动可以为松材萎蔫病的多点爆发和时空聚集提供理论科学依据。对于顶帽核,我们重点研究了感知尺度和延迟对稳定性的共同影响,并发现由于它们的相互作用而产生图灵-霍普夫分岔。此外,通过对实际数据的分析,选取具有实际生物学意义的参数,对分岔点附近的模式进行了详细模拟,得到了峰值交替周期模式和时空聚集模式等多种复杂的时空模式。最后,我们证明了人工释放害虫的寄生天敌可以驱动种群以稳态或周期解的形式达到稳定。研究结果不仅很好地解释了松材枯萎病的传播机制,而且有助于研究松材枯萎病的群、群形成等生物学现象。
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引用次数: 0
Modelling the Impact of Phenotypic Heterogeneity on Cell Migration: A Continuum Framework Derived from Individual-Based Principles. 模拟表型异质性对细胞迁移的影响:基于个体原则的连续框架。
IF 2.2 4区 数学 Q2 BIOLOGY Pub Date : 2025-08-08 DOI: 10.1007/s11538-025-01502-5
Rebecca M Crossley, Philip K Maini, Ruth E Baker

Collective cell migration plays a crucial role in numerous biological processes, including tumour growth, wound healing, and the immune response. Often, the migrating population consists of cells with various different phenotypes. This study derives a general mathematical framework for modelling cell migration in the local environment, which is coarse-grained from an underlying individual-based model that captures the dynamics of cell migration that are influenced by the phenotype of the cell, such as random movement, proliferation, phenotypic transitions, and interactions with the local environment. The resulting, flexible, and general model provides a continuum, macroscopic description of cell invasion, which represents the phenotype of the cell as a continuous variable and is much more amenable to simulation and analysis than its individual-based counterpart when considering a large number of phenotypes. We showcase the utility of the generalised framework in three biological scenarios: range expansion; cell invasion into the extracellular matrix; and T cell exhaustion. The results highlight how phenotypic structuring impacts the spatial and temporal dynamics of cell populations, demonstrating that different environmental pressures and phenotypic transition mechanisms significantly influence migration patterns, a phenomenon that would be computationally very expensive to explore using an individual-based model alone. This framework provides a versatile and robust tool for understanding the role of phenotypic heterogeneity in collective cell migration, with potential applications in optimising therapeutic strategies for diseases involving cell migration.

集体细胞迁移在许多生物过程中起着至关重要的作用,包括肿瘤生长、伤口愈合和免疫反应。通常,迁移群体由各种不同表型的细胞组成。本研究导出了一个用于模拟局部环境中细胞迁移的一般数学框架,该框架是基于潜在的基于个体的模型的粗粒度模型,该模型捕获了受细胞表型影响的细胞迁移动力学,如随机运动、增殖、表型转变以及与局部环境的相互作用。由此产生的、灵活的、通用的模型提供了一个连续的、宏观的细胞入侵描述,它将细胞的表型表示为一个连续的变量,在考虑大量表型时,它比基于个体的模型更易于模拟和分析。我们展示了广义框架在三种生物情景中的效用:范围扩展;细胞侵入细胞外基质;和T细胞衰竭。研究结果强调了表型结构如何影响细胞群体的时空动态,表明不同的环境压力和表型过渡机制显著影响迁移模式,这一现象如果仅使用基于个体的模型进行探索,在计算上是非常昂贵的。该框架为理解表型异质性在集体细胞迁移中的作用提供了一个通用且强大的工具,在优化涉及细胞迁移的疾病的治疗策略方面具有潜在的应用。
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引用次数: 0
Estimating the Size of the Aedes Mosquitoes' Population Involved in Outbreaks of Dengue and Chikungunya Using a Mathematical Model. 利用数学模型估计与登革热和基孔肯雅热暴发有关的伊蚊种群规模。
IF 2.2 4区 数学 Q2 BIOLOGY Pub Date : 2025-08-08 DOI: 10.1007/s11538-025-01489-z
Francisco Antônio Bezerra Coutinho, Marcos Amaku, Fernanda Castro Boulos, José Alfredo de Sousa Moreira, Eliana Nogueira Castro de Barros, Esper Georges Kallas, Eduardo Massad

Aedes aegypti continues to cause many cases of dengue, chikungunya and Zika fever in affected areas of the tropical world. After being eradicated from Brazil in the decades of 1940 and 1950, Aedes aegypti returned with full force in the early 1970s. Knowing the total number of mosquitoes transmitting Aedes-borne infections is crucial for quantifying the intensity of transmission of these infections. In this paper, we propose a model to estimate the distribution of the number of Aedes mosquitoes' populations during an outbreak of either dengue or chikungunya. The model assumes that the mosquitoes' distribution follows a Gaussian Mesa Function (GMF), which has 5 parameters and allows for variable asymmetry. These 5 parameters are adjusted so that they fit indirectly, from a modified Ross‒Macdonald model, the incidence of dengue or chikungunya infections (see main text). Therefore, the observed incidence becomes a function of the parameters of the GMF. We illustrate the model with dengue and chikungunya data from 5 cities in the state of Minas Gerais in the southeastern region of Brazil for the 2023-2024 transmission season. The model shows that it is possible to estimate the size of the mosquitoes' population from incidence data, circumventing the logistic hurdles involved in the actual counting of mosquitoes. This is the most important practical contribution of this paper. The paper also contains several theoretical innovations, such as a modification of the Ross‒Macdonald model, which is usually presented for a constant mosquitoes' population, which, of course, is very unrealistic.

在热带世界的受影响地区,埃及伊蚊继续引起许多登革热、基孔肯雅热和寨卡热病例。埃及伊蚊于1940年和1950年在巴西被消灭后,于20世纪70年代初全面卷土重来。了解传播伊蚊传播感染的蚊子总数对于量化这些感染的传播强度至关重要。在本文中,我们提出了一个模型来估计伊蚊种群在登革热或基孔肯雅热暴发期间的分布。该模型假设蚊子的分布遵循高斯台面函数(GMF),该函数有5个参数,并允许可变不对称。对这5个参数进行了调整,以便从一个改进的Ross-Macdonald模型间接拟合登革热或基孔肯雅热感染的发病率(见正文)。因此,观测到的入射成为GMF参数的函数。我们使用巴西东南部米纳斯吉拉斯州5个城市在2023-2024年传播季节的登革热和基孔肯雅热数据来说明该模型。该模型表明,根据发病率数据估计蚊子种群的规模是可能的,从而绕过了实际计数蚊子所涉及的后勤障碍。这是本文最重要的实践贡献。这篇论文还包含了几个理论创新,比如对Ross-Macdonald模型的修改,该模型通常是针对恒定的蚊子数量提出的,当然,这是非常不现实的。
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