Pub Date : 2024-01-11DOI: 10.1140/epje/s10189-023-00398-0
Akira Onuki
We investigate electric-field effects in dilute electrolytes with nonlinear polarization. As a first example of such systems, we add a dipolar component with a relatively large dipole moment (mu _0) to an aqueous electrolyte. As a second example, the solvent itself exhibits nonlinear polarization near charged objects. For such systems, we present a Ginzburg-Landau free energy and introduce field-dependent chemical potentials, entropy density, and stress tensor, which satisfy general thermodynamic relations. In the first example, the dipoles accumulate in high-field regions, as predicted by Abrashikin et al.[Phys.Rev.Lett. 99, 077801 (2007)]. Finally, we consider the case, where Bjerrum ion pairs form a dipolar component with nonlinear polarization. The Bjerrum dipoles accumulate in high-field regions, while field-induced dissociation was predicted by Onsager [J. Chem. Phys.2, 599 (1934)]. We present an expression for the field-dependent association constant K(E), which depends on the field strength nonmonotonically.
{"title":"Ions and dipoles in electric field: nonlinear polarization and field-dependent chemical reaction","authors":"Akira Onuki","doi":"10.1140/epje/s10189-023-00398-0","DOIUrl":"10.1140/epje/s10189-023-00398-0","url":null,"abstract":"<p>We investigate electric-field effects in dilute electrolytes with nonlinear polarization. As a first example of such systems, we add a dipolar component with a relatively large dipole moment <span>(mu _0)</span> to an aqueous electrolyte. As a second example, the solvent itself exhibits nonlinear polarization near charged objects. For such systems, we present a Ginzburg-Landau free energy and introduce field-dependent chemical potentials, entropy density, and stress tensor, which satisfy general thermodynamic relations. In the first example, the dipoles accumulate in high-field regions, as predicted by Abrashikin <i>et al</i>.[Phys.Rev.Lett. <b>99</b>, 077801 (2007)]. Finally, we consider the case, where Bjerrum ion pairs form a dipolar component with nonlinear polarization. The Bjerrum dipoles accumulate in high-field regions, while field-induced dissociation was predicted by Onsager [J. Chem. Phys.<b>2</b>, 599 (1934)]. We present an expression for the field-dependent association constant <i>K</i>(<i>E</i>), which depends on the field strength nonmonotonically.</p>","PeriodicalId":790,"journal":{"name":"The European Physical Journal E","volume":"47 1","pages":""},"PeriodicalIF":1.8,"publicationDate":"2024-01-11","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"139416060","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"物理与天体物理","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Pub Date : 2024-01-11DOI: 10.1140/epje/s10189-023-00391-7
Dima Bolmatov, C. Patrick Collier, John Katsaras, Maxim O. Lavrentovich
Electrical signals may propagate along neuronal membranes in the brain, thus enabling communication between nerve cells. In doing so, lipid bilayers, fundamental scaffolds of all cell membranes, deform and restructure in response to such electrical activity. These changes impact the electromechanical properties of the membrane, which then physically store biological memory. This memory can exist either over a short or long period of time. Traditionally, biological memory is defined by the strengthening or weakening of transmissions between individual neurons. Here, we show that electrical stimulation may also alter the properties of the lipid membrane, thus pointing toward a novel mechanism for memory storage. Furthermore, based on the analysis of existing electrophysiological data, we study molecular mechanisms underlying the long-term potentiation in phospholipid membranes. Finally, we examine possible relationships between the memory capacitive properties of lipid membranes, neuronal learning, and memory.
{"title":"Physical insights into biological memory using phospholipid membranes","authors":"Dima Bolmatov, C. Patrick Collier, John Katsaras, Maxim O. Lavrentovich","doi":"10.1140/epje/s10189-023-00391-7","DOIUrl":"10.1140/epje/s10189-023-00391-7","url":null,"abstract":"<p>Electrical signals may propagate along neuronal membranes in the brain, thus enabling communication between nerve cells. In doing so, lipid bilayers, fundamental scaffolds of all cell membranes, deform and restructure in response to such electrical activity. These changes impact the electromechanical properties of the membrane, which then physically store biological memory. This memory can exist either over a short or long period of time. Traditionally, biological memory is defined by the strengthening or weakening of transmissions between individual neurons. Here, we show that electrical stimulation may also alter the properties of the lipid membrane, thus pointing toward a novel mechanism for memory storage. Furthermore, based on the analysis of existing electrophysiological data, we study molecular mechanisms underlying the long-term potentiation in phospholipid membranes. Finally, we examine possible relationships between the memory capacitive properties of lipid membranes, neuronal learning, and memory.\u0000</p>","PeriodicalId":790,"journal":{"name":"The European Physical Journal E","volume":"47 1","pages":""},"PeriodicalIF":1.8,"publicationDate":"2024-01-11","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"139416062","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"物理与天体物理","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Pub Date : 2024-01-11DOI: 10.1140/epje/s10189-023-00399-z
Reinhard Lipowsky
Phase separation of biomembranes into two fluid phases, a and b, leads to the formation of vesicles with intramembrane a- and b-domains. These vesicles can attain multispherical shapes consisting of several spheres connected by closed membrane necks. Here, we study the morphological complexity of these multispheres using the theory of curvature elasticity. Vesicles with two domains form two-sphere shapes, consisting of one a- and one b-sphere, connected by a closed ab-neck. The necks’ effective mean curvature is used to distinguish positive from negative necks. Two-sphere shapes of two-domain vesicles can attain four different morphologies that are governed by two different stability conditions. The closed ab-necks are compressed by constriction forces which induce neck fission and vesicle division for large line tensions and/or large spontaneous curvatures. Multispherical shapes with one ab-neck and additional aa- and bb-necks involve several stability conditions, which act to reduce the stability regimes of the multispheres. Furthermore, vesicles with more than two domains form multispheres with more than one ab-neck. The multispherical shapes described here represent generalized constant-mean-curvature surfaces with up to four constant mean curvatures. These shapes are accessible to experimental studies using available methods for giant vesicles prepared from ternary lipid mixtures.
将生物膜相分离成 a 和 b 两种流体相会形成具有膜内 a 和 b 域的囊泡。这些囊泡可呈多球形,由封闭的膜颈连接的多个球体组成。在此,我们利用曲率弹性理论研究了这些多球的形态复杂性。具有两个领域的囊泡形成双球体形状,由一个 a 球体和一个 b 球体组成,并由一个封闭的 ab 形颈连接。颈部的有效平均曲率用于区分正颈和负颈。双域囊泡的双球形状可以达到四种不同的形态,它们受两种不同稳定性条件的制约。当线张力较大和/或自发曲率较大时,封闭的ab-necks会受到收缩力的压缩,从而导致颈部裂开和囊泡分裂。具有一个ab-颈和额外的aa-和bb-颈的多球体形状涉及多个稳定性条件,这些条件降低了多球体的稳定性。此外,具有两个以上结构域的囊泡会形成具有一个以上ab-颈的多球体。这里描述的多球体形状代表了具有多达四个恒定平均曲率的广义恒定平均曲率表面。使用现有方法对三元脂质混合物制备的巨型囊泡进行实验研究,可以获得这些形状。
{"title":"Multispherical shapes of vesicles with intramembrane domains","authors":"Reinhard Lipowsky","doi":"10.1140/epje/s10189-023-00399-z","DOIUrl":"10.1140/epje/s10189-023-00399-z","url":null,"abstract":"<p>Phase separation of biomembranes into two fluid phases, <i>a</i> and <i>b</i>, leads to the formation of vesicles with intramembrane <i>a</i>- and <i>b</i>-domains. These vesicles can attain multispherical shapes consisting of several spheres connected by closed membrane necks. Here, we study the morphological complexity of these multispheres using the theory of curvature elasticity. Vesicles with two domains form two-sphere shapes, consisting of one <i>a</i>- and one <i>b</i>-sphere, connected by a closed <i>ab</i>-neck. The necks’ effective mean curvature is used to distinguish positive from negative necks. Two-sphere shapes of two-domain vesicles can attain four different morphologies that are governed by two different stability conditions. The closed <i>ab</i>-necks are compressed by constriction forces which induce neck fission and vesicle division for large line tensions and/or large spontaneous curvatures. Multispherical shapes with one <i>ab</i>-neck and additional <i>aa</i>- and <i>bb</i>-necks involve several stability conditions, which act to reduce the stability regimes of the multispheres. Furthermore, vesicles with more than two domains form multispheres with more than one <i>ab</i>-neck. The multispherical shapes described here represent generalized constant-mean-curvature surfaces with up to four constant mean curvatures. These shapes are accessible to experimental studies using available methods for giant vesicles prepared from ternary lipid mixtures.</p>","PeriodicalId":790,"journal":{"name":"The European Physical Journal E","volume":"47 1","pages":""},"PeriodicalIF":1.8,"publicationDate":"2024-01-11","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"https://link.springer.com/content/pdf/10.1140/epje/s10189-023-00399-z.pdf","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"139416061","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"物理与天体物理","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"OA","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Pub Date : 2024-01-06DOI: 10.1140/epje/s10189-023-00396-2
Euan T. Smithers, Jingxi Luo, Rosemary J. Dyson
Plant cell growth is regulated through manipulation of the cell wall network, which consists of oriented cellulose microfibrils embedded within a ground matrix incorporating pectin and hemicellulose components. There remain many unknowns as to how this manipulation occurs. Experiments have shown that cellulose reorients in cell walls as the cell expands, while recent data suggest that growth is controlled by distinct collections of hemicellulose called biomechanical hotspots, which join the cellulose molecule together. The enzymes expansin and Cel12A have both been shown to induce growth of the cell wall; however, while Cel12A’s wall-loosening action leads to a reduction in the cell wall strength, expansin’s has been shown to increase the strength of the cell wall. In contrast, members of the XTH enzyme family hydrolyse hemicellulose but do not appear to cause wall creep. This experimentally observed behaviour still awaits a full explanation. We derive and analyse a mathematical model for the effective mechanical properties of the evolving cell wall network, incorporating cellulose microfibrils, which reorient with cell growth and are linked via biomechanical hotspots made up of regions of crosslinking hemicellulose. Assuming a visco-elastic response for the cell wall and using a continuum approach, we calculate the total stress resultant of the cell wall for a given overall growth rate. By changing appropriate parameters affecting breakage rate and viscous properties, we provide evidence for the biomechanical hotspot hypothesis and develop mechanistic understanding of the growth-inducing enzymes.
{"title":"A continuum mechanics model of the plant cell wall reveals interplay between enzyme action and cell wall structure","authors":"Euan T. Smithers, Jingxi Luo, Rosemary J. Dyson","doi":"10.1140/epje/s10189-023-00396-2","DOIUrl":"10.1140/epje/s10189-023-00396-2","url":null,"abstract":"<p>Plant cell growth is regulated through manipulation of the cell wall network, which consists of oriented cellulose microfibrils embedded within a ground matrix incorporating pectin and hemicellulose components. There remain many unknowns as to how this manipulation occurs. Experiments have shown that cellulose reorients in cell walls as the cell expands, while recent data suggest that growth is controlled by distinct collections of hemicellulose called biomechanical hotspots, which join the cellulose molecule together. The enzymes expansin and Cel12A have both been shown to induce growth of the cell wall; however, while Cel12A’s wall-loosening action leads to a reduction in the cell wall strength, expansin’s has been shown to increase the strength of the cell wall. In contrast, members of the XTH enzyme family hydrolyse hemicellulose but do not appear to cause wall creep. This experimentally observed behaviour still awaits a full explanation. We derive and analyse a mathematical model for the effective mechanical properties of the evolving cell wall network, incorporating cellulose microfibrils, which reorient with cell growth and are linked via biomechanical hotspots made up of regions of crosslinking hemicellulose. Assuming a visco-elastic response for the cell wall and using a continuum approach, we calculate the total stress resultant of the cell wall for a given overall growth rate. By changing appropriate parameters affecting breakage rate and viscous properties, we provide evidence for the biomechanical hotspot hypothesis and develop mechanistic understanding of the growth-inducing enzymes.</p>","PeriodicalId":790,"journal":{"name":"The European Physical Journal E","volume":"47 1","pages":""},"PeriodicalIF":1.8,"publicationDate":"2024-01-06","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"https://link.springer.com/content/pdf/10.1140/epje/s10189-023-00396-2.pdf","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"139110591","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"物理与天体物理","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"OA","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Pub Date : 2023-12-26DOI: 10.1140/epje/s10189-023-00395-3
Alexander Farutin, Suhail M. Rizvi, Wei-Fan Hu, Te-Sheng Lin, Salima Rafai, Chaouqi Misbah
Autonomous locomotion is a ubiquitous phenomenon in biology and in physics of active systems at microscopic scale. This includes prokaryotic, eukaryotic cells (crawling and swimming) and artificial swimmers. An outstanding feature is the ability of these entities to follow complex trajectories, ranging from straight, curved (circular, helical...), to random-like ones. The non-straight nature of these trajectories is often explained as a consequence of the asymmetry of the particle or the medium in which it moves, or due to the presence of bounding walls, etc... Here, we show that for a particle driven by a concentration field of an active species, straight, circular and helical trajectories emerge naturally in the absence of asymmetry of the particle or that of suspending medium. Our proof is based on general considerations, without referring to an explicit form of a model. We show that these three trajectories correspond to self-congruent solutions. Self-congruency means that the states of the system at different moments of time can be made identical by an appropriate combination of rotation and translation of the coordinate space. We show that these solutions are exhibited by spherically symmetric particles as a result of a series of pitchfork bifurcations, leading to spontaneous symmetry breaking in the concentration field driving the particle motility. Self-congruent dynamics in one and two dimensions are analyzed as well. Finally, we present a simple explicit nonlinear exactly solvable model of fully isotropic phoretic particle that shows the transitions from a non-motile state to straight motion to circular motion to helical motion as a series of spontaneous symmetry-breaking bifurcations. Whether a system exhibits or not a given trajectory only depends on the numerical values of parameters entering the model, while asymmetry of swimmer shape, or anisotropy of the suspending medium, or influence of bounding walls are not necessary.
{"title":"Motility and swimming: universal description and generic trajectories","authors":"Alexander Farutin, Suhail M. Rizvi, Wei-Fan Hu, Te-Sheng Lin, Salima Rafai, Chaouqi Misbah","doi":"10.1140/epje/s10189-023-00395-3","DOIUrl":"10.1140/epje/s10189-023-00395-3","url":null,"abstract":"<p>Autonomous locomotion is a ubiquitous phenomenon in biology and in physics of active systems at microscopic scale. This includes prokaryotic, eukaryotic cells (crawling and swimming) and artificial swimmers. An outstanding feature is the ability of these entities to follow complex trajectories, ranging from straight, curved (circular, helical...), to random-like ones. The non-straight nature of these trajectories is often explained as a consequence of the asymmetry of the particle or the medium in which it moves, or due to the presence of bounding walls, etc... Here, we show that for a particle driven by a concentration field of an active species, straight, circular and helical trajectories emerge naturally in the absence of asymmetry of the particle or that of suspending medium. Our proof is based on general considerations, without referring to an explicit form of a model. We show that these three trajectories correspond to self-congruent solutions. Self-congruency means that the states of the system at different moments of time can be made identical by an appropriate combination of rotation and translation of the coordinate space. We show that these solutions are exhibited by spherically symmetric particles as a result of a series of pitchfork bifurcations, leading to spontaneous symmetry breaking in the concentration field driving the particle motility. Self-congruent dynamics in one and two dimensions are analyzed as well. Finally, we present a simple explicit nonlinear exactly solvable model of fully isotropic phoretic particle that shows the transitions from a non-motile state to straight motion to circular motion to helical motion as a series of spontaneous symmetry-breaking bifurcations. Whether a system exhibits or not a given trajectory only depends on the numerical values of parameters entering the model, while asymmetry of swimmer shape, or anisotropy of the suspending medium, or influence of bounding walls are not necessary.</p>","PeriodicalId":790,"journal":{"name":"The European Physical Journal E","volume":"46 12","pages":""},"PeriodicalIF":1.8,"publicationDate":"2023-12-26","publicationTypes":"Journal Article","fieldsOfStudy":null,"isOpenAccess":false,"openAccessPdf":"","citationCount":null,"resultStr":null,"platform":"Semanticscholar","paperid":"139037280","PeriodicalName":null,"FirstCategoryId":null,"ListUrlMain":null,"RegionNum":4,"RegionCategory":"物理与天体物理","ArticlePicture":[],"TitleCN":null,"AbstractTextCN":null,"PMCID":"","EPubDate":null,"PubModel":null,"JCR":null,"JCRName":null,"Score":null,"Total":0}
Pub Date : 2023-12-21DOI: 10.1140/epje/s10189-023-00394-4
Joelle A. Labastide, David A. Quint, Reilly K. Cullen, Bryan Maelfeyt, Jennifer L. Ross, Ajay Gopinathan
Active, motor-based cargo transport is important for many cellular functions and cellular development. However, the cell interior is complex and crowded and could have many weak, non-specific interactions with the cargo being transported. To understand how cargo-environment interactions will affect single motor cargo transport and multi-motor cargo transport, we use an artificial quantum dot cargo bound with few (~ 1) to many (~ 5–10) motors allowed to move in a dense microtubule network. We find that kinesin-driven quantum dot cargo is slower than single kinesin-1 motors. Excitingly, there is some recovery of the speed when multiple motors are attached to the cargo. To determine the possible mechanisms of both the slow down and recovery of speed, we have developed a computational model that explicitly incorporates multi-motor cargos interacting non-specifically with nearby microtubules, including, and predominantly with the microtubule on which the cargo is being transported. Our model has recovered the experimentally measured average cargo speed distribution for cargo-motor configurations with few and many motors, implying that numerous, weak, non-specific interactions can slow down cargo transport and multiple motors can reduce these interactions thereby increasing velocity.