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Waiting times in a branching process model of colorectal cancer initiation 结直肠癌起始分支过程模型中的等待时间
IF 1.4 4区 生物学 Q2 Agricultural and Biological Sciences Pub Date : 2023-06-01 DOI: 10.1016/j.tpb.2023.04.001
Ruibo Zhang, Obinna A. Ukogu, Ivana Bozic

We study a multi-stage model for the development of colorectal cancer from initially healthy tissue. The model incorporates a complex sequence of driver gene alterations, some of which result in immediate growth advantage, while others have initially neutral effects. We derive analytic estimates for the sizes of premalignant subpopulations, and use these results to compute the waiting times to premalignant and malignant genotypes. This work contributes to the quantitative understanding of colorectal tumor evolution and the lifetime risk of colorectal cancer.

我们从最初的健康组织中研究了结直肠癌癌症发展的多阶段模型。该模型包含了一系列复杂的驱动基因改变,其中一些改变会立即产生生长优势,而另一些则最初具有中性影响。我们推导了癌前亚群大小的分析估计,并使用这些结果来计算癌前和恶性基因型的等待时间。这项工作有助于定量了解结直肠癌的演变和结直肠癌癌症的终生风险。
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引用次数: 0
Copuling population dynamics and diel migration patterns 交配种群动态与死亡迁移模式
IF 1.4 4区 生物学 Q2 Agricultural and Biological Sciences Pub Date : 2023-06-01 DOI: 10.1016/j.tpb.2023.03.004
Emil F. Frølich

The diel vertical migration is one of the main drivers of population dynamics in the ocean. Population dynamical models of the ocean typically do not incorporate the behavioral aspects of the migration. We demonstrate a model with coupled population dynamics and behavior with the diel vertical migration emerging. We study the population dynamics and behavioral dynamics of a predator–prey system. We impose a cost of motion for both consumers and prey, and model each individual as following an Itô stochastic differential equation. We study the fixed-points of the ecosystem. Our modeling shows that as we increase the basal resource load, the strength of the diel vertical migration increases, as well as maximal velocity. In addition, a bimodal pattern emerges both for predators and consumers. The increase in the magnitude of the diel vertical migration causes a change in the allocation of copepod resources.

昼夜垂直迁移是海洋种群动态的主要驱动因素之一。海洋的种群动力学模型通常不包括迁徙的行为方面。我们展示了一个具有耦合种群动态和行为的模型,其中出现了昼夜垂直迁移。我们研究了捕食者-猎物系统的种群动力学和行为动力学。我们对消费者和猎物都施加了运动成本,并按照随机微分方程对每个个体进行建模。我们研究生态系统的固定点。我们的模型表明,随着基础资源负荷的增加,昼夜垂直迁移的强度和最大速度都会增加。此外,捕食者和消费者都出现了双峰模式。昼夜垂直迁移幅度的增加导致桡足类资源分配的变化。
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引用次数: 1
Evolution with recombination as Gibbs sampling 用吉布斯采样法进行重组进化
IF 1.4 4区 生物学 Q2 Agricultural and Biological Sciences Pub Date : 2023-06-01 DOI: 10.1016/j.tpb.2023.03.005
Jenny M. Poulton , Lee Altenberg , Chris Watkins

This work presents a population genetic model of evolution, which includes haploid selection, mutation, recombination, and drift. The mutation-selection equilibrium can be expressed exactly in closed form for arbitrary fitness functions without resorting to diffusion approximations. Tractability is achieved by generating new offspring using n-parent rather than 2-parent recombination. While this enforces linkage equilibrium among offspring, it allows analysis of the whole population under linkage disequilibrium. We derive a general and exact relationship between fitness fluctuations and response to selection. Our assumptions allow analytical calculation of the stationary distribution of the model for a variety of non-trivial fitness functions. These results allow us to speak to genetic architecture, i.e., what stationary distributions result from different fitness functions. This paper presents methods for exactly deriving stationary states for finite and infinite populations. This method can be applied to many fitness functions, and we give exact calculations for four of these. These results allow us to investigate metastability, tradeoffs between fitness functions, and even consider error-correcting codes.

这项工作提出了一个群体遗传进化模型,包括单倍体选择、突变、重组和漂移。对于任意适应度函数,突变选择平衡可以精确地以闭合形式表示,而不需要采用扩散近似。可牵引性是通过使用n-亲本而不是2-亲本重组产生新的后代来实现的。虽然这加强了后代之间的连锁平衡,但它允许在连锁不平衡下分析整个种群。我们推导了适应度波动和对选择的反应之间的一般而精确的关系。我们的假设允许对各种非平凡适应度函数的模型的平稳分布进行分析计算。这些结果使我们能够谈论遗传结构,即不同适应度函数产生的平稳分布。本文给出了精确推导有限和无限总体平稳状态的方法。这种方法可以应用于许多适应度函数,我们给出了其中四个函数的精确计算。这些结果使我们能够研究亚稳态、适应度函数之间的权衡,甚至考虑纠错码。
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引用次数: 0
A population genetics theory for piRNA-regulated transposable elements pirna调控转座因子的群体遗传学理论
IF 1.4 4区 生物学 Q2 Agricultural and Biological Sciences Pub Date : 2023-04-01 DOI: 10.1016/j.tpb.2023.02.001
Siddharth S. Tomar, Aurélie Hua-Van, Arnaud Le Rouzic

Transposable elements (TEs) are self-reproducing selfish DNA sequences that can invade the genome of virtually all living species. Population genetics models have shown that TE copy numbers generally reach a limit, either because the transposition rate decreases with the number of copies (transposition regulation) or because TE copies are deleterious, and thus purged by natural selection. Yet, recent empirical discoveries suggest that TE regulation may mostly rely on piRNAs, which require a specific mutational event (the insertion of a TE copy in a piRNA cluster) to be activated — the so-called TE regulation “trap model”. We derived new population genetics models accounting for this trap mechanism, and showed that the resulting equilibria differ substantially from previous expectations based on a transposition–selection equilibrium. We proposed three sub-models, depending on whether or not genomic TE copies and piRNA cluster TE copies are selectively neutral or deleterious, and we provide analytical expressions for maximum and equilibrium copy numbers, as well as cluster frequencies for all of them. In the full neutral model, the equilibrium is achieved when transposition is completely silenced, and this equilibrium does not depend on the transposition rate. When genomic TE copies are deleterious but not cluster TE copies, no long-term equilibrium is possible, and active TEs are eventually eliminated after an active incomplete invasion stage. When all TE copies are deleterious, a transposition–selection equilibrium exists, but the invasion dynamics is not monotonic, and the copy number peaks before decreasing. Mathematical predictions were in good agreement with numerical simulations, except when genetic drift and/or linkage disequilibrium dominates. Overall, the trap-model dynamics appeared to be substantially more stochastic and less repeatable than traditional regulation models.

可转座元件(TE)是一种自我繁殖的自私DNA序列,可以入侵几乎所有生物的基因组。群体遗传学模型表明,TE拷贝数通常达到极限,要么是因为转座率随着拷贝数的增加而降低(转座调节),要么是由于TE拷贝是有害的,因此被自然选择清除。然而,最近的经验发现表明,TE调节可能主要依赖于piRNA,这需要激活特定的突变事件(在piRNA簇中插入TE拷贝),即所谓的TE调节“陷阱模型”。我们推导了解释这种陷阱机制的新的群体遗传学模型,并表明由此产生的平衡与以前基于换位-选择平衡的预期有很大不同。我们提出了三个子模型,这取决于基因组TE拷贝和piRNA簇TE拷贝是否是选择性中性或有害的,我们提供了最大拷贝数和平衡拷贝数的分析表达式,以及所有这些的簇频率。在完全中性模型中,当转置完全静音时,达到平衡,并且这种平衡不取决于转置速率。当基因组TE拷贝是有害的,但不是簇TE拷贝时,不可能实现长期平衡,并且活性TE在活性不完全入侵阶段后最终被消除。当所有TE拷贝都是有害的时,存在换位-选择平衡,但入侵动力学不是单调的,拷贝数在减少之前达到峰值。除了遗传漂移和/或连锁不平衡占主导地位外,数学预测与数值模拟非常一致。总体而言,陷阱模型动力学似乎比传统的监管模型更具随机性和可重复性。
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引用次数: 0
Stability of Rosenzweig–MacArthur models with non-diffusive dispersal on non-regular networks 非正则网络上具有非扩散扩散扩散的Rosenzweig–MacArthur模型的稳定性
IF 1.4 4区 生物学 Q2 Agricultural and Biological Sciences Pub Date : 2023-04-01 DOI: 10.1016/j.tpb.2023.02.002
Ryusuke Kon , Dinesh Kumar

This paper examines the stability of the Rosenzweig–MacArthur model distributed to identical discrete habitat patches. Migration between patches is assumed to follow the non-diffusive rule that individuals have a fixed rate of leaving their local habitat patch and migrating to another. Under this non-diffusive migration rule, we found that population dispersal on a non-regular and connected habitat network can both stabilize and destabilize the Rosenzweig–MacArthur model. It is also shown that our non-diffusive migration rule apparently becomes diffusive if the habitat network is regular.

本文研究了Rosenzweig–MacArthur模型分布在相同离散生境斑块上的稳定性。斑块之间的迁移被认为遵循非扩散规则,即个体离开当地栖息地斑块并迁移到另一个栖息地斑块的速率是固定的。在这种非扩散迁移规则下,我们发现种群在非规则和连通的栖息地网络上的扩散既可以稳定Rosenzweig–MacArthur模型,也可以破坏其稳定。研究还表明,如果栖息地网络是规则的,我们的非扩散迁移规则显然会变得具有扩散性。
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引用次数: 0
Continuous irregular dynamics with multiple neutral trajectories permit species coexistence in competitive communities 具有多个中性轨迹的连续不规则动力学允许物种在竞争群落中共存
IF 1.4 4区 生物学 Q2 Agricultural and Biological Sciences Pub Date : 2023-02-01 DOI: 10.1016/j.tpb.2022.12.003
Atsushi Yamauchi , Koichi Ito , Shota Shibasaki , Toshiyuki Namba

The colonization model formulates competition among propagules for habitable sites to colonize, which serves as a mechanism enabling coexistence of multiple species. This model traditionally assumes that encounters between propagules and sites occur as mass action events, under which species distribution can eventually reach an equilibrium state with multiple species in a constant environment. To investigate the effects of encounter mode on species diversity, we analyzed community dynamics in the colonization model by varying encounter processes. The analysis indicated that equilibrium is approximately neutrally-stable under perfect ratio-dependent encounter, resulting in temporally continuous variation of species’ frequencies with irregular trajectories even under a constant environment. Although the trajectories significantly depend on initial conditions, they are considered to be “strange nonchaotic attractors” (SNAs) rather than chaos from the asymptotic growth rates of displacement. In addition, trajectories with different initial conditions remain different through time, indicating that the system involves an infinite number of SNAs. This analysis presents a novel mechanism for transient dynamics under competition.

定殖模型规定了繁殖体之间对可居住地点的竞争,这是一种使多个物种共存的机制。该模型传统上假设繁殖体和地点之间的相遇是作为群体作用事件发生的,在这种情况下,物种分布最终可以在恒定的环境中与多个物种达到平衡状态。为了研究相遇模式对物种多样性的影响,我们通过不同的相遇过程分析了定殖模型中的群落动力学。分析表明,在完全比例相关的遭遇下,平衡是近似中性稳定的,即使在恒定的环境下,物种的频率也会以不规则的轨迹在时间上连续变化。尽管轨迹在很大程度上取决于初始条件,但从位移的渐进增长率来看,它们被认为是“奇怪的非混沌吸引子”(SNA),而不是混沌。此外,具有不同初始条件的轨迹随着时间的推移保持不同,这表明该系统涉及无限数量的SNA。该分析为竞争条件下的瞬态动力学提供了一种新的机制。
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引用次数: 1
Improving the realism of neutral ecological models by incorporating transient dynamics with temporal changes in community size 通过将瞬态动力学与群落规模的时间变化相结合,提高中性生态模型的真实性
IF 1.4 4区 生物学 Q2 Agricultural and Biological Sciences Pub Date : 2023-02-01 DOI: 10.1016/j.tpb.2022.12.001
Tak Fung, Ryan A. Chisholm

Neutral models in ecology assume that all species are demographically equivalent, such that their abundances differ ultimately because of demographic stochasticity rather than selection. In spite of their simplicity, neutral models have been found to accurately reproduce static patterns of biodiversity for diverse communities. However, the same neutral models have been found to exhibit species abundance dynamics that are far too slow compared to reality, resulting in poor fits to temporally dynamic patterns of biodiversity. Here, we show that one of the root causes of these slow dynamics is the additional assumption that a community has reached an equilibrium with a fixed community size, with species that have a net growth rate close to zero. We removed this additional assumption by constructing and analyzing a neutral model with an expected community size that can change over time and is not necessarily at equilibrium, which thus allows the historical formation of a community to be represented explicitly. Our analysis demonstrated that for the general scenario where a small community rapidly grows in size to a carrying capacity, representing recovery from ecological disturbance or assembly of a new community, the model produced much larger changes in species abundances and much shorter species ages than a neutral model at an equilibrium with fixed community size. In addition, the species abundance distribution was biphasic with a subset of abundant species arising from a founder effect. We confirmed these new results in applications of the new model to the specific scenario of recovery of the Amazon tree community after the end-Cretaceous bolide impact, which involved periods of increasing and decreasing community size. We conclude that incorporating transient dynamics in neutral models improves realism by allowing explicit consideration of how a community is formed over realistic time-scales.

生态学中的中立模型假设所有物种在人口统计学上是相等的,因此它们的丰度最终是由于人口统计学的随机性而不是选择而不同的。尽管它们很简单,但已经发现中性模型可以准确地再现不同社区的生物多样性的静态模式。然而,人们发现,同样的中性模型显示出的物种丰度动态与现实相比太慢,导致与生物多样性的时间动态模式不匹配。在这里,我们表明,这些缓慢动态的根本原因之一是额外的假设,即一个群落已经在固定的群落规模下达到平衡,物种的净增长率接近零。我们通过构建和分析一个中性模型来消除这一额外的假设,该模型的预期社区规模可以随着时间的推移而变化,不一定处于平衡状态,因此可以明确地表示社区的历史形成。我们的分析表明,对于一个小群落的规模迅速增长到承载能力的一般情况,代表着从生态干扰或新群落的组装中恢复,该模型在物种丰度方面产生了更大的变化,物种年龄也比在固定群落规模的平衡下的中性模型短得多。此外,物种丰度分布是双相的,丰富物种的子集由奠基者效应引起。我们在新模型的应用中证实了这些新结果,该模型适用于白垩纪末的玻利维亚影响后亚马逊树木群落恢复的特定场景,该场景涉及群落规模的增加和减少时期。我们得出的结论是,在中性模型中结合瞬态动力学可以通过明确考虑社区是如何在现实时间尺度上形成的来提高现实性。
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引用次数: 0
Analytical Bayesian approach for the design of surveillance and control programs to assess pest-eradication success 用分析贝叶斯方法设计监测和控制计划,以评估消灭虫害的成功
IF 1.4 4区 生物学 Q2 Agricultural and Biological Sciences Pub Date : 2023-02-01 DOI: 10.1016/j.tpb.2022.11.003
B. Barnes , M. Parsa , F. Giannini , D. Ramsey

Large invasive species eradication programs are undertaken to protect native biodiversity and agriculture. Programs are typically followed by a series of surveys to assess the likelihood of eradication success and, when residual pests are detected, small-scale control or ‘mop-ups’ are implemented to eliminate these infestations. Further surveys follow to confirm absence with ‘freedom’ declared when a target probability of absence is reached. Such biosecurity programs comprise many interacting processes — stochastic biological processes including growth, and response and control interventions — and are an important component of post-border biosecurity. Statistical frameworks formulated to contribute to the design and efficiency of these surveillance and control programs are few and, those available, rely on the simulation of the component processes. In this paper we formulate an analytical Bayesian framework for a general biosecurity program with multiple components to assess pest-eradication success. Our model incorporates stochastic growth and detection processes, and several pest control mechanisms. Survey results and economic considerations are also taken into account to support a range of biosecurity management decisions. Using a case study we demonstrate that solutions match published simulation results and extend the available analysis. Principally, we show how analytical solutions can offer a powerful tool to support the design of effective and cost-efficient biosecurity systems, and we establish some general principles that guide and contribute to robust design.

为保护当地生物多样性和农业,实施了大型入侵物种根除计划。项目之后通常会进行一系列调查,以评估根除成功的可能性,当发现残留害虫时,会实施小规模控制或“扫荡”以消除这些虫害。随后进行了进一步的调查,以确认缺席,并在达到缺席的目标概率时宣布“自由”。此类生物安全计划包括许多相互作用的过程——包括生长、反应和控制干预在内的随机生物过程——是边境后生物安全的重要组成部分。为有助于这些监督和控制程序的设计和效率而制定的统计框架很少,而且这些可用的统计框架依赖于对组成过程的模拟。在本文中,我们为具有多个组成部分的通用生物安全计划制定了一个分析贝叶斯框架,以评估害虫根除的成功率。我们的模型结合了随机生长和检测过程,以及几种害虫控制机制。还考虑了调查结果和经济因素,以支持一系列生物安全管理决策。通过案例研究,我们证明了解决方案与已发布的模拟结果相匹配,并扩展了可用的分析。主要是,我们展示了分析解决方案如何提供强大的工具来支持有效和成本效益高的生物安全系统的设计,并制定了一些指导和促进稳健设计的一般原则。
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引用次数: 0
Distributions of cherries and pitchforks for the Ford model 福特车型的樱桃和干草叉分布
IF 1.4 4区 生物学 Q2 Agricultural and Biological Sciences Pub Date : 2023-02-01 DOI: 10.1016/j.tpb.2022.12.002
Gursharn Kaur , Kwok Pui Choi , Taoyang Wu

Distributional properties of tree shape statistics under random phylogenetic tree models play an important role in investigating the evolutionary forces underlying the observed phylogenies. In this paper, we study two subtree counting statistics, the number of cherries and that of pitchforks for the Ford model, the alpha model introduced by Daniel Ford. It is a one-parameter family of random phylogenetic tree models which includes the proportional to distinguishable arrangement (PDA) and the Yule models, two tree models commonly used in phylogenetics. Based on a non-uniform version of the extended Pólya urn models in which negative entries are permitted for their replacement matrices, we obtain the strong law of large numbers and the central limit theorem for the joint distribution of these two statistics for the Ford model. Furthermore, we derive a recursive formula for computing the exact joint distribution of these two statistics. This leads to exact formulas for their means and higher order asymptotic expansions of their second moments, which allows us to identify a critical parameter value for the correlation between these two statistics. That is, when the number of tree leaves is sufficiently large, they are negatively correlated for 0α1/2 and positively correlated for 1/2<α<1.

随机系统发育树模型下树形统计的分布特性在研究所观察到的系统发育背后的进化力方面发挥着重要作用。在本文中,我们研究了Daniel Ford引入的alpha模型Ford模型的两个子树计数统计数据,即樱桃数量和干草叉数量。它是一个单参数的随机系统发育树模型家族,包括系统发育学中常用的比例可区分排列(PDA)和Yule模型。基于扩展的Pólya-urn模型的非均匀版本,其中它们的替换矩阵允许负项,我们得到了Ford模型的强数定律和这两个统计量联合分布的中心极限定理。此外,我们还导出了计算这两个统计量的精确联合分布的递归公式。这导致了它们的平均值的精确公式和二阶矩的高阶渐近展开式,这使我们能够确定这两个统计量之间相关性的关键参数值。也就是说,当树叶数量足够大时,它们在0≤α≤1/2时呈负相关,在1/2<;α<;1.
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引用次数: 1
Modeling temporal dynamics of genetic diversity in stage-structured plant populations with reference to demographic genetic structure 基于种群遗传结构的阶段结构植物群体遗传多样性时间动态建模
IF 1.4 4区 生物学 Q2 Agricultural and Biological Sciences Pub Date : 2022-12-01 DOI: 10.1016/j.tpb.2022.11.001
Yoichi Tsuzuki, Takenori Takada, Masashi Ohara

Predicting temporal dynamics of genetic diversity is important for assessing long-term population persistence. In stage-structured populations, especially in perennial plant species, genetic diversity is often compared among life history stages, such as seedlings, juveniles, and flowerings, using neutral genetic markers. The comparison among stages is sometimes referred to as demographic genetic structure, which has been regarded as a proxy of potential genetic changes because individuals in mature stages will die and be replaced by those in more immature stages over the course of time. However, due to the lack of theoretical examination, the basic property of the stage-wise genetic diversity remained unclear. We developed a matrix model which was made up of difference equations of the probability of non-identical-by-descent of each life history stage at a neutral locus to describe the dynamics and the inter-stage differences of genetic diversity in stage-structured plant populations. Based on the model, we formulated demographic genetic structure as well as the annual change rate of the probability of non-identical-by-descent (denoted as η). We checked if theoretical expectations on demographic genetic structure and η obtained from our model agreed with computational results of stochastic simulation using randomly generated 3,000 life histories. We then examined the relationships of demographic genetic structure with effective population size Ne, which is the determinants of diversity loss per generation time. Theoretical expectations on η and demographic genetic structure fitted well to the results of stochastic simulation, supporting the validity of our model. Demographic genetic structure varied independently of Ne and η, while having a strong correlation with stable stage distribution: genetic diversity was lower in stages with fewer individuals. Our results indicate that demographic genetic structure strongly reflects stable stage distribution, rather than temporal genetic dynamics, and that inferring future genetic diversity solely from demographic genetic structure would be misleading. Instead of demographic genetic structure, we propose η as an useful tool to predict genetic diversity at the same time scale as population dynamics (i.e., per year), facilitating evaluation on population viability from a genetic point of view.

预测遗传多样性的时间动态对于评估种群的长期持久性具有重要意义。在阶段结构种群中,特别是在多年生植物物种中,遗传多样性经常使用中性遗传标记在生活史阶段(如幼苗、幼体和花期)进行比较。阶段之间的比较有时被称为人口遗传结构,它被认为是潜在遗传变化的代表,因为成熟阶段的个体会死亡,并随着时间的推移被更不成熟阶段的个体所取代。然而,由于缺乏理论检验,分阶段遗传多样性的基本性质尚不清楚。本文建立了一个由中性位点上各生活史阶段遗传变异概率的差分方程组成的矩阵模型,用以描述阶段结构植物群体遗传多样性的动态和阶段间差异。在此基础上,我们给出了人口统计遗传结构以及非同血统概率的年变化率(记为η)。我们用随机生成的3000个生活史进行了随机模拟,验证了从我们的模型中得到的人口统计学遗传结构和η的理论预期是否与随机模拟的计算结果一致。然后,我们研究了人口统计学遗传结构与有效种群大小Ne的关系,这是每代时间多样性损失的决定因素。对η和人口统计学遗传结构的理论预测与随机模拟结果吻合较好,支持了模型的有效性。种群遗传结构的变化与Ne和η无关,但与稳定分期分布有较强的相关性,个体较少的分期遗传多样性较低。我们的研究结果表明,人口遗传结构强烈地反映了稳定的阶段分布,而不是时间遗传动态,仅从人口遗传结构推断未来的遗传多样性将是误导性的。代替人口遗传结构,我们提出η作为一种有用的工具来预测与种群动态(即每年)相同时间尺度的遗传多样性,有助于从遗传角度评估种群生存能力。
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引用次数: 0
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Theoretical Population Biology
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