B. Žekš’s research while affiliated with University of Ljubljana and other places

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Publications (76)


Nonlocal membrane bending: A reflection, the facts and its relevance
  • Literature Review

June 2014

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47 Reads

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18 Citations

Advances in Colloid and Interface Science

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B. Žekš

About forty years ago it was realized that phospholipid membranes, because they are composed of two layers, exhibit particular, specific mechanical properties , and . This led to the concept of nonlocal membrane bending, often called area difference elasticity. We present a short history of the development of the concept, followed by arguments for a proper definition of the corresponding elastic constant. The effects of the nonlocal bending energy on vesicle shape are explained. It is demonstrated that lipid vesicles, cells and cellular aggregates exhibit phenomena that can only be described in a complete manner by considering nonlocal bending.


FERROELECTRIC LIQUID CRYSTALS: FROM THE PLANE WAVE TO THE MULTISOLITON LIMIT

May 2012

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15 Reads

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8 Citations

International Journal of Modern Physics B

In the presence of external fields or in restricted geometries, the originally continuous helical symmetry of the Sm C* phase is broken by the appearence of field- or geometry-induced soliton-like domain walls. As a result of this symmetry breaking, a crossover between the plane-wave-like and soliton-like regime occurs in both static and dynamic properties which is responsible for some remarkable phenomena such as field-induced optical biaxiality or a field-induced band structure of collective excitations. Whereas we find in the plane-wave-like regime a degenerate soft mode which splits below the Sm A→Sm C* transition into a symmetry recovering Goldstone-phason-mode and an amplitudon mode, we find in the soliton regime a splitting of the phason mode into acoustic and optic-like branches separated by a band gap. Within the same framework we also discuss other remarkable and extraordinary properties such as reentrant phases, Lifshitz points, one dimensional photonic band gaps and thickness dependent phase diagrams.


Proton order-disorder in KH2PO4-type ferroelectrics: Slater theory and ising model in a transverse tunneling field

February 2011

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50 Reads

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37 Citations

Ferroelectrics

Recent developments in the theory of phase transitions in KH2PO4-type crystals are reviewed together with some relevant experiments. The nature of the isotope effect in the static and dynamic properties of these systems on replacing hydrogen by deuterium is discussed. Expressions for the static and dynamic properties derived by four-cluster analysis are compared with those found from a mean field treatment of the Ising model.


FIGURE 1 Nucleosomal core particle: Biochemical view. Histone proteins and DNA are shown using ribbon presentation. Side view of the particle has a typical V-like shape which can be clearly distinguished in the electron microscopy experiments. 
FIGURE 2 Nucleosomal core particle: Biophysical view. DNA is wrapped around the cylindrical core formed by the octamer of four histone proteins. Superhelix DNA twisting around the histone core makes the particle chiral. The entry-exit point of the DNA free ends marks a polar dyadic axis P . a – Top view. 1 – histone core; 2 – 1.7 superhelical turns of the DNA; 3 – histone tails. b – Side view. c – Schematic notation of the NCP and its polar dyadic axis. 
FIGURE 3 Structure of the lamellar phase [17]. a – Tubules observed by optical microscopy. b – d – Lamellar structure of the NCP columns revealed by cryoelectron microscopy. b, c – Images perpendicular to the columns axes. d – Image parallel to the columns axes. The orientations of the dyadic axes of individual NCP are shown in d. The orientations of the average polar axes directions are shown in c. 
FIGURE 4 Dyadic axes orientation. a – Red point marks the entry-exit point of the DNA superhelix; the arrow indicates the orientation of the axis. b – dyadic axes orientations disordered along the column in the hexagonal phase. c – dyadic axes are correlated along the column in the lamellar phase; each column carries a non-zero polarization vector P . 
FIGURE 5 Hexagonal and lamellar structures. a – Hexagonal parent structure: columns noted as 1, 2, 3, and 4 indicate the nodes of a unit cell in the 2D hexagonal structure; k 1 , k 2 , and k 3 are the wave vectors of the periodic polar vector field which drives the transition; X and Y are the hexagonal 

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Polarity and Chirality in NCP Mesophases and Chromatin Fibers
  • Article
  • Full-text available

December 2007

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203 Reads

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2 Citations

Molecular Crystals and Liquid Crystals

We propose a unified mechanism of phase transitions in the high density solutions of Nucleosomal Core Particles. We demonstrate its relation with the high shape and charge anisotropy of the NCP due to the DNA wrapping around the histone protein core. Hexagonal-to-lamellar transition in the solution of NCP columnar aggregates is shown to be driven by the condensation of the periodic antiparallel polar vector field. The transition results in the polar dyadic axes correlations. Fine structure of the NCP organization is also explained, namely the correlated tilting of NCP with respect to the column axis.

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Short Pitch Structures in Smectics with Interactions over More Than Two Layers

September 2006

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27 Reads

Molecular Crystals and Liquid Crystals Science and Technology Section A Molecular Crystals and Liquid Crystals

Phenomenological model of chiral polar smectics is introduced with interactions up to the fourth neighboring layers. The minimization of the free energy gives three stable structures: the ferroelectric Sm C* phase, the antiferroelectric Sm CA * phase and the hellicoidally modulated structure of the Sm CH * phase. The Sm CH * phase can be recognized as the Sm Cα* phase below Sm A phase and as the Sm CFI1* phase or as the Sm CFI1 phase when appearing between the Sm C* phase and the Sm CA* phase. Stability of these phases is analyzed and the phase diagram in the space of model parameters is presented.



Figure 1: Crystalline lamellar phase (c) as a common subphase phase of the 2D hexagonal columnar (a) and the decorrelated smectic lamellar phase (b). Effect of chirality on the S-axes of the NCPs (d). The red circles and black squares denote the front (F) and the back (B) direction of the NCP dyadic axis [6] (lower right). 
Figure 2: Displacement fields for the transition from the hexagonal to lamellar ( η, 0 , 0) (columns forming a single lamella are enclosed within a gray rect- angle), orthorhombic ( η, η, 0) and inverted hexagonal ( η, η, η ) phases. Below, the phase digram stemming from Eqs. 5, 6. 
Correlated and Decorrelated Positional and Orientational Order in the Nucleosomal Core Particle Mesophases

February 2004

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77 Reads

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9 Citations

We investigate the orientational order of transverse polarization vectors of long columns of nucleosomal core particles and their coupling to positional order in high density mesophases discovered recently. Inhomogeneous polar ordering of these columns precipitates crystalization of the 2D sections with different orientations of the transverse polarization vector on each column in the unit cell. We propose possible scenarios for going from the 2D hexagonal phase into distorted lamellar and related phases observed experimentally.


Ferro- and Antiferroelectric Liquid Crystals

January 2003

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420 Reads

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67 Citations

B. Žekš

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S. A. Pikin

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In this section a historical overview will be presented of the development of phenomenological theoretical models of ferroelectric and antiferroelectric liquid crystals. In Sect. 5.1.2 the two order parameters of ferroelectric smectic C* (Sm C*) liquid crystals will be introduced, which are the tilt of long axes of molecules from the normal to the smectic layers and the in-plane electric polarization, that corresponds to the ordering of molecules transverse to their long axes. The phenomenological models of ferroelectric liquid crystals (FLC) will be presented, that correspond to the Landau expansion of the free-energy density in the tilt and the polarization order parameters. An extension of the phenomenological model of FLC will also be demonstrated, that includes as the additional order parameter the transverse quadrupolar ordering and that describes some anomalies of FLC’s, that cannot be accounted by the model with only two order parameters. In these models the layered structure of smectic liquid crystals is not explicitly taken into account and the same is true for the phenomenological models of antiferroelectric liquid crystals (AFLC), that are presented and described in Sect. 5.1.3. Such models cannot describe all existing phases in AFLC and therefore discrete models of AFLC’s were introduced, which take explicitly smectic layers into account and assume some inter-layer interactions, which must have longer range then only between nearest neighboring layers. Discrete models are described in Sect. 5.1.4, while in Sect. 5.1.5 the nature and the structure of the inter-layer interactions is presented together with possible phases predicted by such models.


Theoretical and experimental study of the intermediate Sm CFI 2∗ and the Sm CFI 1∗ phases in antiferroelectric liquid crystals

July 2002

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17 Reads

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65 Citations

Within the new phenomenological model of antiferroelectric liquid crystals, structures of the two phases, the Sm CFI2∗ and the Sm CFI1∗ that appear in the temperature range between the Sm C∗ and the Sm CA∗ phase, are found. The structure of the Sm CFI2∗ and the Sm CFI1∗ phases is consistent with extensive optical, electro-optic, and dielectric measurements performed on the liquid crystal R-10OTBBB1M7, for which resonant x-ray scattering studies have previously proven that these two phases have the three-layer (Sm CFI1∗) and the four-layer (Sm CFI2∗) periodic structure, respectively. The biaxiality of the four-layer structure of the Sm CFI2∗ phase is directly observed experimentally for the first time at the temperature where the optical pitch is unwound. The theoretical model allows for experimentally consistent phase sequence, the order of transitions, and the temperature dependence of optical properties including the optical helicoidal modulation inversion within the temperature range of the Sm CFI2∗ phase. © 2002 American Institute of Physics.


Mechanical properties of closed lamellar membranes and cellular processes

August 2001

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7 Reads

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6 Citations

Nonlinear Analysis

This paper considers cellular processes that involve changes in membrane conformation. A general overview of the relationship between the mechanical properties of lamellar membranes and the shapes of vesicles and cells is given. Variational procedures are described for calculating the equilibrium shapes of simple vesicular objects, based on the assumption that these shapes correspond to the minimum of the sum of the membrane local and nonlocal bending energies. The two properties of closed lamellar membranes which could form a physical basis for biological processes axe proposed to be the symmetry breaking in vesicle shape behavior and vesicle budding. The cellular processes are discussed on the basis of the hypothesis that the shape behavior of simple vesicular objects may serve as a framework for their analysis. It is proposed that the description of certain cellular processes requires the inclusion of energy contributions additional to the membrane bending energy, and/or the involvement of cytoplasmic structures which exert forces on the membrane. Several specific examples are used to demonstrate that such generalizations may help reveal the functional role of certain cell constituents.


Citations (44)


... As it will be shown, this macroscopic alignment of the SmC* phase in the presence of an external magnetic field results in an apparent decrease in the orientational order measured through NMR by decreasing the temperature [57][58][59][60]. The effect of the magnetic field on the supramolecular structure of the SmC* phase was first investigated by applying the magnetic field perpendicular to the helical axis, similarly to what was done to study the effect of the electric field [73]. However, in 1998, Zalar et al. [74] reported for the first time the phenomenon of helix unwinding and helix distortion by performing 2 H NMR experiments at variable angles between the helical axes of the ferroelectric phases and the magnetic field. ...

Reference:

Supramolecular Arrangement and Conformational and Dynamic Properties of Chiral Smectic Liquid Crystals Obtained Through Nuclear Magnetic Resonance: A Brief Review
The Physics of Ferroelectric and Antiferroelectric Liquid Crystals
  • Citing Book
  • July 2000

... It has been demonstrated that an increased in surface anchoring energy tends to increase the devices working voltages and the grayscale dynamic range is deteriorated. Certainly, surface anchoring is essential for FLCs when the cell thickness is in the range of the surface coupling, and [21,22]. In contrast, conventional polyimides produce strong anchoring layers, where the average anchoring energies, W, are in the order of W≈10 −4 J/m 2 , whereas alignment layers like Nylon, photoalignment layers or layers with soft rubbing conditions, produce weak anchoring that is one or two orders of magnitude smaller [23,24]. ...

Ferro- and Antiferroelectric Liquid Crystals
  • Citing Chapter
  • January 2003

... Regarding the morphometric changes in hBMSC shape, the effect of pH on the morphology of hBMSCs could be associated with a change in the difference between the outer and the inner monolayer areas of the cell membrane lipid bilayer, as described previously [13,14]. The cell membrane consists of a phospholipid bilayer and the underlying membrane skeleton, which is a complex network of interlinking filaments and tubules that extend from the nucleus throughout the cytoplasm, towards the plasma membrane. ...

A role of membrane skeleton in discontinuous red blood cell shape transformations
  • Citing Article
  • January 1996

Cellular & Molecular Biology Letters

... It is important to mention that others have used a quite different approach to understand the basis for the red blood cell's biconcave shape. These types of studies have used solutions of theoretical models that are mainly based on the red blood cell's known physical parameters, such as the membrane's bending energy and elastic properties (e.g., [5][6][7][8][9][10][11] in addition to lateral inhomogeneities in the M/CS (12). These analyses have remarkably generated not only the cell's biconcave shape but also many other types of shapes that the cells are known to assume. ...

Cytoskeleton and red cell shape
  • Citing Article
  • January 1996

Cellular & Molecular Biology Letters

... At higher concentrations of CPP more CPP induced pores are formed, so that osmotic pressure is changed rapidly and cannot be compensated by the opening of transient pores. Consequently, GUVs burst and disintegrate [32]. ...

Lipid-polypeptide interactions studied by optical observation of a single giant lipid vesicle
  • Citing Article
  • January 2006

... Nonlocal bending models such as these have indeed been proposed in the past [26][27][28][29][30][31], but they have typically been viewed as alternatives to the local Helfrich energy, not as an addition. Taking the second view, the total bilayer energy is of the form ...

Lipid bilayer elasticity and the bilayer couple interpretation of red cell shape transformations and lysis
  • Citing Article
  • January 1985

... [4][5][6][7] The structures of these compounds comprise infinite sheets of corner-shared MCl 6 octahedra, 8 with the interlayer space occupied by alkylammonium ions oriented perpendicular or nearly perpendicular to the above sheets, and charged NH 3 groups hydrogen-bonded to the chloride atoms of MCl 6 units. 9,10 The compound with n ¼ 1 and M ¼ Cd, (CH 3 NH 3 ) 2 CdCl 4 , undergoes three structural phase transitions at 484, 283, and 163 K, [11][12][13] with the respective phases denoted as I (tetragonal high-temperature), II (orthorhombic room-temperature), III (tetragonal low-temperature), and IV (monoclinic low-temperature). It is known that the tetragonal (I) to orthorhombic (II) transition is second-order and the orthorhombic (II) to tetragonal (III) is first-order, and tetragonal (III) to monoclinic (IV) is firstorder. ...

Model of structural phase transitions in (CH 3 NH 3 ) 2 CdCl 4 -type compounds
  • Citing Article
  • April 1978

Ferroelectrics

... It cannot be excluded, especially in view of the results on FAPI, that also the sharp frequency independent stiffening just below T βγ is related to the loss of reorientational modes. In fact, one can construct symmetrized coordinates from the linear combinations of the populations of the possible orientations of MA, as it has been done for hybrid layered perovskites [38][39][40]. Some of these coordinates act as OP of the phase transitions and the others are associated with relaxations such as that in γ − MAPI. ...

Dynamics of structural phase transitions in (CH_{3}NH_{3})_{2}CdCl_{4}-type compounds
  • Citing Article
  • April 1979

Physical review. B, Condensed matter

... This phenomenology is widely observed in the whole family of H-bonded ferroelectric compounds [6][7][8][9][10][11][12][13] including organic ferroelectrics that were recently discovered which attract much attention because of their potential for ecofriendly technological applications [14,15]. Ab initio (AI) calculations have shown that tunneling and geometrical effects are involved in a self-consistent phenomenon that greatly amplifies the effect of their correlation leading to the huge isotope effect observed [7,16]. ...

Proton order-disorder in KH2PO4-type ferroelectrics: Slater theory and ising model in a transverse tunneling field
  • Citing Article
  • February 2011

Ferroelectrics

... According to the differential elasticity model, differences in the areas of the inner and outer bilayer of vesicle membranes dictate vesicle morphologies. [47,48] We hypothesize that accumulation of FFM in the inner leaflet creates an asymmetry that causes inward bulging and drives vesicle division. As shown in Movie S1, daughter vesicles are propelled by internal currents generated by changes in interfacial tension due to peptide deposition. ...

Nonlocal membrane bending: A reflection, the facts and its relevance
  • Citing Article
  • June 2014

Advances in Colloid and Interface Science