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Nonequilibrium Membrane Dynamics Induced by Active Protein Interactions and Chemical Reactions: A Review 活性蛋白质相互作用和化学反应诱导的非平衡膜动力学:综述
Pub Date : 2024-07-22 DOI: arxiv-2407.15371
Hiroshi Noguchi
Biomembranes wrapping cells and organelles are not only the partitions thatseparate the insides but also dynamic fields for biological functionsaccompanied by membrane shape changes. In this review, we discuss thespatiotemporal patterns and fluctuations of membranes under nonequilibriumconditions. In particular, we focus on theoretical analyses and simulations.Protein active forces enhance or suppress the membrane fluctuations; themembrane height spectra are deviated from the thermal spectra. Protein bindingor unbinding to the membrane is activated or inhibited by other proteins andchemical reactions, such as ATP hydrolysis. Such active binding processes caninduce traveling waves, Turing patterns, and membrane morphological changes.They can be represented by the continuum reaction-diffusion equations anddiscrete lattice/particle models with state flips. The effects of structuralchanges in amphiphilic molecules on the molecular-assembly structures are alsodiscussed.
包裹细胞和细胞器的生物膜不仅是分隔内部的隔板,也是伴随膜形状变化的生物功能动态场。在这篇综述中,我们将讨论非平衡条件下膜的时空模式和波动。蛋白质活性力增强或抑制膜波动;膜高度谱偏离热谱。蛋白质与膜的结合或解除结合会被其他蛋白质和化学反应(如 ATP 水解)激活或抑制。这种活跃的结合过程可引发行波、图灵模式和膜形态变化,可用连续反应-扩散方程和具有状态翻转的离散晶格/粒子模型来表示。此外,还讨论了两亲分子结构变化对分子组装结构的影响。
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引用次数: 0
Binding and dimerization control phase separation in a compartment 结合和二聚化控制隔室中的相分离
Pub Date : 2024-07-21 DOI: arxiv-2407.15179
Riccardo Rossetto, Gerrit Wellecke, David Zwicker
Biological cells exhibit a hierarchical spatial organization, where variouscompartments harbor condensates that form by phase separation. Cells cancontrol the emergence of these condensates by affecting compartment size, theamount of the involved molecules, and their physical interactions. Whilephysical interactions directly affect compartment binding and phase separation,they can also cause oligomerization, which has been suggested as a controlmechanism. Analyzing an equilibrium model, we illustrate that oligomerizationamplifies compartment binding and phase separation, which reinforce each other.This nonlinear interplay can also induce multistability, which providesadditional potential for control. Our work forms the basis for derivingthermodynamically consistent kinetic models to understand how biological cellscan regulate phase separation in their compartments.
生物细胞呈现出层次分明的空间组织结构,不同的细胞间隙蕴藏着通过相分离形成的凝聚体。细胞可以通过影响隔室的大小、参与分子的数量以及它们之间的物理相互作用来控制这些凝聚体的出现。虽然物理相互作用会直接影响细胞间隙的结合和相分离,但它们也会导致寡聚化,这也被认为是一种控制机制。通过分析一个平衡模型,我们说明了低聚作用会放大区室结合和相分离,从而相互加强。我们的工作为推导热力学上一致的动力学模型奠定了基础,从而了解生物细胞如何调节其区室中的相分离。
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引用次数: 0
Generalized Langevin equation for a tagged monomer in a Gaussian semiflexible polymer 高斯半柔性聚合物中标记单体的广义朗温方程
Pub Date : 2024-07-20 DOI: arxiv-2407.14886
Xavier Durang, Jae-Hyung Jeon
In this study, we present a comprehensive analysis of the motion of a taggedmonomer within a Gaussian semiflexible polymer model. We carefully derived thegeneralized Langevin Equation (GLE) that governs the motion of a tagged centralmonomer. This derivation involves integrating out all the other degrees offreedom within the polymer chain, thereby yielding an effective description ofthe viscoelastic motion of the tagged monomer. A critical component of ouranalysis is the memory kernel that appears in the GLE. By examining thiskernel, we characterized the impact of bending rigidity on the non-Markoviandiffusion dynamics of the tagged monomer. Furthermore, we calculated themean-squared displacement of the tagged monomer using the derived GLE. Ourresults not only show remarkable agreement with previously known results incertain limiting cases but also provide dynamic features over the entiretimescale.
在本研究中,我们对高斯半柔性聚合物模型中的标记单体运动进行了全面分析。我们仔细推导了支配标记中心单体运动的广义朗文方程(GLE)。这一推导涉及对聚合物链中的所有其他自由度进行积分,从而得出对标记单体粘弹性运动的有效描述。我们分析的一个关键部分是出现在 GLE 中的记忆核。通过研究该记忆核,我们确定了弯曲刚度对标记单体非马尔可夫扩散动力学的影响。此外,我们还利用推导出的 GLE 计算了标记单体的主题位移平方。我们的结果不仅与之前已知的某些极限情况下的结果显著一致,而且还提供了整个时间尺度上的动态特征。
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引用次数: 0
Growing tissue sheets on substrates: buds, buckles, and pores 在基质上生长组织片:芽、扣和孔
Pub Date : 2024-07-18 DOI: arxiv-2407.13187
Hiroshi Noguchi, Jens Elgeti
Many tissues take the form of thin sheets, being only a single cell thick,but millions of cells wide. These tissue sheets can bend and buckle in thethird dimension. In this work, we investigated the growth of suspended andsupported tissue sheets using particle-based simulations. We combineparticle-based tissue growth and meshless membrane models to simulate thegrowth of tissue sheets with mechanical feedback. Free suspended growingtissues exhibit wrinkling when growth is sufficiently fast. Conversely, tissueson a substrate form buds when the adhesion to the substrate is weak and/or whenthe friction with the substrate is strong. These buds undergo amembrane-mediated attraction and subsequently fuse. The complete detachment oftissues from the substrate and straight buckled bump formation are alsoobtained at very weak adhesion and/or fast growth rates. Tissue pores grow viaOstwald ripening and coalescence. The reported dynamics can also be applied inresearch on the detachment dynamics of different tissues with weakenedadhesion.
许多组织呈薄片状,只有一个细胞厚,但却有数百万个细胞宽。这些组织薄片可以在三维空间弯曲和折叠。在这项工作中,我们利用基于粒子的模拟研究了悬浮和支撑组织薄片的生长。我们将基于粒子的组织生长和无网格膜模型相结合,模拟了具有机械反馈的组织片的生长。当生长速度足够快时,自由悬浮生长的组织会出现皱褶。相反,当与基底的粘附力较弱和/或与基底的摩擦力较强时,基底上的组织会形成芽。这些芽经过膜介导的吸引,随后融合。在粘附力很弱和/或生长速度很快的情况下,组织也会完全脱离基底并形成直扣凸起。组织孔隙通过奥斯特瓦尔德成熟和凝聚而生长。所报告的动力学也可用于研究粘附力减弱的不同组织的脱离动力学。
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引用次数: 0
Dissipation at limited resolutions: Power law and detection of hidden dissipative scales 有限分辨率下的耗散:幂律和隐藏耗散尺度的检测
Pub Date : 2024-07-18 DOI: arxiv-2407.13707
Qiwei Yu, Pedro E. Harunari
Nonequilibrium systems, in particular living organisms, are maintained byirreversible transformations of energy that drive diverse functions.Quantifying their irreversibility, as measured by energy dissipation, isessential for understanding the underlying mechanisms. However, existingtechniques usually overlook experimental limitations, either by assuming fullinformation or by employing a coarse-graining method that requires knowledge ofthe structure behind hidden degrees of freedom. Here, we study the inference ofdissipation from finite-resolution measurements by employing a recentlydeveloped model-free estimator that considers both the sequence ofcoarse-grained transitions and the waiting time distributions:$sigma_2=sigma_2^ell + sigma_2^t$. The dominant term $sigma_2^ell$originates from the sequence of observed transitions; we find that it scaleswith resolution following a power law. Comparing the scaling exponent with aprevious estimator highlights the importance of accounting for fluxcorrelations at lower resolutions. $sigma_2^t$ comes from asymmetries inwaiting time distributions, with its peak revealing characteristic scales ofthe underlying dissipative process. Alternatively, the characteristic scale canbe detected in a crossover of the scaling of $sigma_2^ell$. This provides anovel perspective for extracting otherwise hidden characteristic dissipativescales directly from dissipation measurements. We illustrate these results inbiochemical models as well as complex networks. Overall, this study highlightsthe significance of resolution considerations in nonequilibrium systems,providing insights into the interplay between experimental resolution, entropyproduction, and underlying complexity.
非平衡系统,特别是生物体,是通过能量的不可逆转化来维持的,而能量的不可逆转化驱动着各种功能。然而,现有的技术通常忽略了实验的局限性,要么假定信息充分,要么采用需要了解隐藏自由度背后结构的粗粒度方法。在这里,我们研究了从有限分辨率测量中推断耗散的方法,采用了最近开发的无模型估计器,该估计器同时考虑了粗粒度转换序列和等待时间分布:$sigma_2=sigma_2^ell + sigma_2^t$。主要项$sigma_2^ell$源自观测到的转换序列;我们发现它与分辨率呈幂律关系。将缩放指数与以前的估计值进行比较,突出了在较低分辨率下考虑通量相关性的重要性。$sigma_2^t$ 来自等待时间分布的不对称性,其峰值揭示了底层耗散过程的特征尺度。或者,特征尺度可以从 $sigma_2^ell$ 的比例交叉中检测到。这为直接从耗散测量中提取隐藏的特征耗散尺度提供了一个新的视角。我们用生化模型和复杂网络来说明这些结果。总之,这项研究强调了在非平衡系统中考虑分辨率的重要性,提供了对实验分辨率、熵产生和潜在复杂性之间相互作用的见解。
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引用次数: 0
Encoding spatiotemporal asymmetry in artificial cilia with a ctenophore-inspired soft-robotic platform 用栉水母启发的软机器人平台编码人工纤毛的时空不对称性
Pub Date : 2024-07-18 DOI: arxiv-2407.13894
David J. Peterman, Margaret L. Byron
A remarkable variety of organisms use metachronal coordination (i.e.,numerous neighboring appendages beating sequentially with a fixed phase lag) toswim or pump fluid. This coordination strategy is used by microorganisms tobreak symmetry at small scales where viscous effects dominate and flow istime-reversible. Some larger organisms use this swimming strategy atintermediate scales, where viscosity and inertia both play important roles.However, the role of individual propulsor kinematics - especially acrosshydrodynamic scales - is not well-understood, though the details of propulsormotion can be crucial for the efficient generation of flow. To investigate thisbehavior, we developed a new soft robotic platform using magnetoactive siliconeelastomers to mimic the metachronally coordinated propulsors found in swimmingorganisms. Furthermore, we present a method to passively encode spatiallyasymmetric beating patterns in our artificial propulsors. We investigated thekinematics and hydrodynamics of three propulsor types, with varying degrees ofasymmetry, using Particle Image Velocimetry and high-speed videography. We findthat asymmetric beating patterns can move considerably more fluid relative tosymmetric beating at the same frequency and phase lag, and that asymmetry canbe passively encoded into propulsors via the interplay between elastic andmagnetic torques. Our results demonstrate that nuanced differences in propulsorkinematics can substantially impact fluid pumping performance. Our soft roboticplatform also provides an avenue to explore metachronal coordination at themeso-scale, which in turn can inform the design of future bioinspired pumpingdevices and swimming robots.
多种多样的生物利用元周期协调(即众多相邻附肢以固定的相位滞后顺序跳动)来游泳或泵送流体。在粘性效应占主导地位、流动具有时间可逆性的小尺度上,微生物利用这种协调策略来打破对称性。然而,单个推进器运动学的作用--尤其是跨流体动力尺度的作用--尚未得到很好的理解,尽管推进器运动的细节对于高效产生流动可能至关重要。为了研究这种行为,我们开发了一种新的软机器人平台,使用磁活性硅胶弹性体来模拟游泳生物中的元协调推进器。此外,我们还提出了一种在人造推进器中被动编码空间不对称跳动模式的方法。我们利用粒子图像测速仪和高速摄像技术研究了三种不同对称程度的推进器的运动学和流体力学。我们发现,相对于相同频率和相位滞后的对称跳动,不对称跳动模式可以移动更多的流体,而且不对称可以通过弹性和磁力矩之间的相互作用被动地编码到推进器中。我们的研究结果表明,推进器运动学的细微差别会对流体泵送性能产生重大影响。我们的软机器人平台还为探索主题尺度的元协调提供了一条途径,这反过来又能为未来生物启发泵设备和游泳机器人的设计提供信息。
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引用次数: 0
A statistical mechanics investigation of Unfolded Protein Response across organisms 跨生物体的折叠蛋白反应统计力学研究
Pub Date : 2024-07-17 DOI: arxiv-2407.12464
Nicole Luchetti, Keith M. Smith, Margherita A. G. Matarrese, Alessandro Loppini, Simonetta Filippi, Letizia Chiodo
Living systems rely on coordinated molecular interactions, especially thoserelated to gene expression and protein activity. The Unfolded Protein Responseis a crucial mechanism in eukaryotic cells, activated when unfolded proteinsexceed a critical threshold. It maintains cell homeostasis by enhancing proteinfolding, initiating quality control, and activating degradation pathways whendamage is irreversible. This response functions as a dynamic signaling network,with proteins as nodes and their interactions as edges. We analyze theseprotein-protein networks across different organisms to understand theirintricate intra-cellular interactions and behaviors. In this work, analyzingtwelve organisms, we assess how fundamental measures in network theory canindividuate seed-proteins and specific pathways across organisms. We employnetwork robustness to evaluate and compare the strength of the investigated PPInetworks, and the structural controllability of complex networks to find andcompare the sets of driver nodes necessary to control the overall networks. Wefind that network measures are related to phylogenetics, and advanced networkmethods can identify main pathways of significance in the complete UnfoldedProtein Response mechanism.
生命系统依赖于协调的分子相互作用,尤其是与基因表达和蛋白质活性有关的相互作用。折叠蛋白反应是真核细胞中的一种重要机制,当折叠蛋白超过临界阈值时就会被激活。它通过加强蛋白质折叠、启动质量控制以及在损伤不可逆转时激活降解途径来维持细胞的平衡。这种反应就像一个动态信号网络,蛋白质是节点,它们之间的相互作用是边。我们分析了不同生物体的这些蛋白质-蛋白质网络,以了解它们错综复杂的细胞内相互作用和行为。在这项工作中,我们分析了十二种生物,评估了网络理论中的基本测量方法如何在不同生物间划分种子蛋白和特定通路。我们利用网络鲁棒性来评估和比较所研究的 PPInetworks 的强度,并利用复杂网络的结构可控性来寻找和比较控制整个网络所需的驱动节点集。我们发现,网络度量与系统发生学有关,先进的网络方法可以识别完整的蛋白折叠反应机制中重要的主要通路。
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引用次数: 0
Metabolic activity controls the emergence of coherent flows in microbial suspensions 代谢活动控制着微生物悬浮液中相干流的出现
Pub Date : 2024-07-13 DOI: arxiv-2407.09884
Alexandros A. Fragkopoulos, Florian Böhme, Nicole Drewes, Oliver Bäumchen
Photosynthetic microbes have evolved and successfully adapted to theever-changing environmental conditions in complex microhabitats throughoutalmost all ecosystems on Earth. In the absence of light, they can sustain theirbiological functionalities through aerobic respiration, and even in anoxicconditions through anaerobic metabolic activity. For a suspension ofphotosynthetic microbes in an anaerobic environment, individual cellularmotility is directly controlled by its photosynthetic activity, i.e. theintensity of the incident light absorbed by chlorophyll. The effects of themetabolic activity on the collective motility on the population level, however,remain elusive so far. Here, we demonstrate that at high light intensities, asuspension of photosynthetically active microbes exhibits a stable reversesedimentation profile of the cell density due to the microbes' natural bias tomove against gravity. With decreasing photosynthetic activity, and thereforesuppressed individual motility, the living suspension becomes unstable givingrise to coherent bioconvective flows. The collective motility is fullyreversible and manifests as regular, three-dimensional plume structures, inwhich flow rates and cell distributions are directly controlled via the lightintensity. The coherent flows emerge in the highly unfavourable condition oflacking both light and oxygen and, thus, might help the microbial collective toexpand the exploration of their natural habitat in search for better survivalconditions.
在地球上几乎所有的生态系统中,光合微生物都在复杂的微生境中进化并成功地适应了不断变化的环境条件。在无光条件下,它们可以通过有氧呼吸维持其生物功能,甚至在缺氧条件下通过厌氧代谢活动维持其生物功能。对于处于厌氧环境中的光合微生物悬浮液来说,单个细胞的运动能力直接受其光合作用活动(即叶绿素吸收入射光的强度)的控制。然而,迄今为止,光合作用对种群集体运动性的影响仍然难以捉摸。在这里,我们证明了在高光强度下,光合作用活跃的微生物悬浮液中的细胞密度会呈现稳定的反向沉降曲线,这是由于微生物逆重力运动的自然偏好所致。随着光合作用活性的降低,个体运动受到抑制,生物悬浮液变得不稳定,从而产生相干的生物对流。集体运动是完全可逆的,表现为规则的三维羽流结构,其中的流速和细胞分布直接受光照强度控制。相干流出现在缺乏光和氧气的非常不利的条件下,因此可能有助于微生物集体扩大对其自然栖息地的探索,以寻求更好的生存条件。
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引用次数: 0
Modeling Ion-Specific Effects in Polyelectrolyte Brushes: A Modified Poisson-Nernst-Planck Model 聚电解质刷中的离子特异性效应建模:修正的泊松-纳斯特-普朗克模型
Pub Date : 2024-07-12 DOI: arxiv-2407.09633
William J Ceely, Marina Chugunova, Ali Nadim, James D Sterling
Polyelectrolyte brushes consist of a set of charged linear macromoleculeseach tethered at one end to a surface. An example is the glycocalyx whichrefers to hair-like negatively charged sugar molecules that coat the outsidemembrane of all cells. We consider the transport and equilibrium distributionof ions, and the resulting electrical potential, when such a brush is immersedin a salt buffer containing monovalent cations (sodium and/or potassium). TheGouy-Chapman model for ion screening at a charged surface captures the effectsof the Coulombic force that drives ion electrophoresis and diffusion, butneglects non-Coulombic forces and ion pairing. By including the distinctbinding affinities of these counter-ions with the brush, and their so-calledBorn radii, which account for Born forces acting on them when the permittivityis non-uniform, we propose modified Poisson-Nernst-Planck continuum models thatshow the distinct profiles that may result depending on those ion-specificproperties.
聚电解质刷由一组带电的线性大分子组成,每个大分子的一端都拴在表面。例如,糖萼膜(glyocalyx)指的是包裹在所有细胞膜外的毛发状带负电的糖分子。我们考虑的是当这种刷子浸入含有一价阳离子(钠和/或钾)的盐缓冲液中时,离子的迁移和平衡分布以及由此产生的电动势。古伊-查普曼(Gouy-Chapman)电荷表面离子筛选模型捕捉到了驱动离子电泳和扩散的库仑力的影响,但忽略了非库仑力和离子配对。通过将这些反离子与电刷的不同结合亲和力及其所谓的博恩半径(当介电常数不均匀时作用于它们的博恩力)包括在内,我们提出了修正的泊松-奈恩斯特-普朗克连续模型,该模型显示了根据这些离子的特定性质可能产生的不同剖面。
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引用次数: 0
Directed Motion and Spatial Coherence in the Cell Nucleus 细胞核中的定向运动和空间一致性
Pub Date : 2024-07-12 DOI: arxiv-2407.08899
M. Hidalgo-Soria, Y. Haddad, E. Barkai, Y. Garini, S. Burov
Investigating the dynamics of chromatin and the factors that are affectingit, has provided valuable insights into the organization and functionality ofthe genome in the cell nucleus. We control the expression of Lamin-A, animportant organizer protein of the chromatin and nucleus structure. Bysimultaneously tracking tens of chromosomal loci (telomeres) in each nucleus,we find that the motion of chromosomal loci in Lamin-A depleted cells is bothfaster and more directed on a scale of a few micrometers, which coincides withthe size of chromosome territories. Moreover, in the absence of Lamin-A wereveal the existence of correlations among neighboring telomeres. We show howthese pairwise correlations are linked with the intermittent and persistentcharacter of telomere trajectories, underscoring the importance of Lamin-Aprotein in chromosomal organization.
研究染色质的动态和影响染色质的因素为了解细胞核中基因组的组织和功能提供了宝贵的信息。我们控制染色质和细胞核结构的重要组织蛋白 Lamin-A 的表达。通过同时追踪每个细胞核中的数十个染色体位点(端粒),我们发现在缺乏Lamin-A的细胞中,染色体位点的运动速度更快,而且在几微米的范围内更有方向性,这与染色体区域的大小相吻合。此外,在缺乏Lamin-A的情况下,相邻端粒之间存在相关性。我们展示了这些成对相关性是如何与端粒轨迹的间歇性和持续性特征联系在一起的,从而强调了Lamin-A蛋白在染色体组织中的重要性。
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引用次数: 0
期刊
arXiv - PHYS - Biological Physics
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