Cellular Tip Research Articles

Cellular tip splitting instability during transient growth

In this work, cellular growth under transient conditions is investigated by using a developed quantitative phase-field model in which both the pulling speed and thermal gradient are time-dependent variables. Cellular tip shapes during transient growth are first characterized using the three-dimensional Saffman-Taylor viscous finger shape equation. Simulation results show that cellular tip and shoulders under non-steady-state conditions can be described by this simple mathematical model even at large Peclet number. The problem of cellular pattern evolution during directional solidification in the laser molten pool is possibly that of viscous finger in fluid mechanics. Previous tip splitting criterion is not applicable in our work. The mechanism of cellular tip splitting and later tip crack deepening under transient conditions is first studied, this interesting phenomenon may result from the coupling effect between morphological instabilities which can be measured by the shape factor and cellular spacing, and non-equilibrium of tip solute concentration which provides the driving force for deepening of the bifurcation.

Tip-splitting instability in directional solidification based on bias field method**Project supported by the National Basic Research Program of China (Grant No. 2011CB610401), the National Natural Science Foundation of China (Grant No. 51371151), and the Free Research Fund of State Key Laboratory of Solidification Processing, China (Grant No. 100-QP-2014).

Tip splitting instability of cellular interface morphology in directional solidification is analyzed based on the bias field method proposed recently by Glicksman. The physical mechanism of tip instability is explained by analyzing the interface potential, the tangential energy flux, and the normal energy flux. A rigorous criterion for tip-splitting instability is established analytically, i.e., the ratio of the cellular tip radius to the cellular width , which is in good agreement with simulation results. This study also reveals that the cellular tip splitting instability is attributable to weak Gibbs–Thomson energy acting on the interface.

Mercury-sensitive water channels as possible sensors of water potentials in pollen

The growing pollen tube is central to plant reproduction and is a long-standing model for cellular tip growth in biology. Rapid osmotically driven growth is maintained under variable conditions, which requires osmosensing and regulation. This study explores the mechanism of water entry and the potential role of osmosensory regulation in maintaining pollen growth. The osmotic permeability of the plasmalemma of Lilium pollen tubes was measured from plasmolysis rates to be 1.32±0.31×10–3 cm s–1. Mercuric ions reduce this permeability by 65%. Simulations using an osmotic model of pollen tube growth predict that an osmosensor at the cell membrane controls pectin deposition at the cell tip; inhibiting the sensor is predicted to cause tip bursting due to cell wall thinning. It was found that adding mercury to growing pollen tubes caused such a bursting of the tips. The model indicates that lowering the osmotic permeability per se does not lead to bursting but rather to thickening of the tip. The time course of induced bursting showed no time lag and was independent of mercury concentration, compatible with a surface site of action. The submaximal bursting response to intermediate mercuric ion concentration was independent of the concentration of calcium ions, showing that bursting is not due to a competitive inhibition of calcium binding or entry. Bursting with the same time course was also shown by cells growing on potassium-free media, indicating that potassium channels (implicated in mechanosensing) are not involved in the bursting response. The possible involvement of mercury-sensitive water channels as osmosensors and current knowledge of these in pollen cells are discussed.

INVESTIGATION OF EFFECT OF INTERFACE ENERGY ANISOTROPY ON DENDRITIC GROWTH IN UNIDIRECTIONAL SOLIDIFICATION BY FRONT TRACKING SIMULATION

The dendritic growth with the different solid/liquid (S/L) interface energy anisotropies in the unidirectional solidification has been investigated using the self-consistent front tracking model. It is found that, for a given solidification condition, there were two kind of interface shape solutions with the different spacing Peclect number ranges. The interface shape with the small spacing Peclect number range was similar with cellular tip, and that with the large spacing Peclect number range referred to dendritic tip. The higher S/L interface energy anisotropy was in favor of the widening of the dendritic growth solution range. There was a certain power exponential relationship between the dendritic tip marginal stability parameter σ ∗ and the S/L interface energy anisotropic parameter E4. A modified Fisher dendritic tip solution, which considered the effect of S/L interface energy anisotropy, was obtained as follows: RIMS =2 .5646( ΓD L Vk 0ΔT0 ) 0.5 E −0.1905 4 , ΔT0 = mC0(k0 − 1)/k0. The undercooling in front of the S/L interface decreased with increasing the anisotropic parameter. The primary dendritic spacing mainly depended on the interaction of solute diffusion field between the adjacent dendrite, and the S/L interface energy had little influence on

Influence of liquid flow on cellular and dendritic spacings

The cellular and dendritic formations are two kinds of typical morphology in the solidification, and there are many theoretical models and experimental researches on them. Most models and researches are based on purely diffusive transport mechanism. However, convection effects are of importance in the evolution of cellular and dendritic growth. Since the metal materials are not transparency and the researches on microstructure only after quenching, it is difficult to observe the dynamic microstructure evolution in real time. In this paper, the effect of liquid flow on the cellular and dendritic growth was investigated by the in-situ observation of SCN-1.8 wt% Ace transparent alloy during the directional solidification under the liquid flow. The cellular tip splitting is found in the presence of liquid flow and the cellular microstructure is smaller after the cellular tip splitting. The cellular spacing decreases as the flow rate becomes larger, but the spacing will become steady ultimately. At high growth rate the dendritic spacing increases with the increase of the flow rate, because the upstream side branches, which are accelerated by liquid flow, will suppress adjacent branches. But, at low growth rate the dendritic spacing decreases with the increase of the flow rate, because the lateral branches will exceed the tip of dendrite to form new dendrite by liquid flow.

An osmotic model of the growing pollen tube.

Pollen tube growth is central to the sexual reproduction of plants and is a longstanding model for cellular tip growth. For rapid tip growth, cell wall deposition and hardening must balance the rate of osmotic water uptake, and this involves the control of turgor pressure. Pressure contributes directly to both the driving force for water entry and tip expansion causing thinning of wall material. Understanding tip growth requires an analysis of the coordination of these processes and their regulation. Here we develop a quantitative physiological model which includes water entry by osmosis, the incorporation of cell wall material and the spreading of that material as a film at the tip. Parameters of the model have been determined from the literature and from measurements, by light, confocal and electron microscopy, together with results from experiments made on dye entry and plasmolysis in Lilium longiflorum. The model yields values of variables such as osmotic and turgor pressure, growth rates and wall thickness. The model and its predictive capacity were tested by comparing programmed simulations with experimental observations following perturbations of the growth medium. The model explains the role of turgor pressure and its observed constancy during oscillations; the stability of wall thickness under different conditions, without which the cell would burst; and some surprising properties such as the need for restricting osmotic permeability to a constant area near the tip, which was experimentally confirmed. To achieve both constancy of pressure and wall thickness under the range of conditions observed in steady-state growth the model reveals the need for a sensor that detects the driving potential for water entry and controls the deposition rate of wall material at the tip.

Effect of surface tension anisotropy on the growth patterns of cellulars in directional solidification

The growth patterns of cellulars in directional solidification are investigated numerically using the cellular automata (CA) model in two dimensions. A criterion which determine whether the cellulars reach stable state is derived from the analysis of simulated results. The simulated results also show that it is easy for tip splitting to appear for cellulars when the surface tention anisotropy is very small. So it is hard to obtain stable cellular arrays. However, if the amplitude of surface tention anisotropy is strong enough, it is easy to obtain stable cellular arrays. And the intensity of surface energy anisotropy can considerably influence the stable cellular patterns. The stronger the surface energy anisotropy, the smaller the stable cellular spacing and the cellular tip radius are, and the smaller the ratio between tip radius and cellular spacing, the smaller the tip concentration and the tip undercooling are.

Quantitative investigation of cellular growth in directional solidification by phase-field simulation

Using a quantitative phase-field model, a systematic investigation of cellular growth in directional solidification is carried out with emphasis on the selection of cellular tip undercooling, tip radius, and cellular spacing. Previous analytical models of cellular growth are evaluated according to the phase-field simulation results. The results show that cellular tip undercooling and tip radius not only depend on the pulling velocity and thermal gradient, but also depend on the cellular interaction related to the cellular spacing. The cellular interaction results in a finite stable range of cellular spacing. The lower limit is determined by the submerging mechanism while the upper limit comes from the tip splitting instability corresponding to the absence of the cellular growth solution, both of which can be obtained from phase-field simulation. Further discussions on the phase-field results also present an analytical method to predict the lower limit. Phase-field simulations on cell elimination between cells with equal spacing validate the finite range of cellular spacing and give deep insight into the cellular doublon and oscillatory instability between cell elimination and tip splitting.

Spatial control of Cdc42 activation determines cell width in fission yeast

The fission yeast Schizosaccharomyces pombe is a rod-shaped cell that grows by linear extension at the cell tips, with a nearly constant width throughout the cell cycle. This simple geometry makes it an ideal system for studying the control of cellular dimensions. In this study, we carried out a near-genome-wide screen for mutants wider than wild-type cells. We found 11 deletion mutants that were wider; seven of the deleted genes are implicated in the control of the small GTPase Cdc42, including the Cdc42 guanine nucleotide exchange factor (GEF) Scd1 and the Cdc42 GTPase-activating protein (GAP) Rga4. Deletions of rga4 and scd1 had additive effects on cell width, and the proteins localized independently of one another, with Rga4 located at the cell sides and Scd1 at the cell tips. Activated Cdc42 localization is altered in rga4Δ, scd1Δ, and scd2Δ mutants. Delocalization and ectopic retargeting experiments showed that the localizations of Rga4 and Scd1 are crucial for their roles in determining cell width. We propose that the GAP Rga4 and the GEF Scd1 establish a gradient of activated Cdc42 within the cellular tip plasma membrane, and it is this gradient that determines cell growth-zone size and normal cell width.

Histological heterogeneity of glomerular segmental lesions in focal segmental glomerulosclerosis

Focal segmental glomerulosclerosis (FSGS) involves considerable histological heterogeneity in terms of location and quality of the glomerular segmental lesions. The present study investigated the heterogeneity of segmental lesions in each variant of FSGS, determined by the Columbia classification, and its clinical relevance. All glomerular segmental lesions of 80 cases of primary FSGS were evaluated histologically based on location [tip (TIP), perihilar (PH), or not otherwise specified (NOS)], and quality (cellular or fibrous). Among the 1,299 glomeruli of the 80 biopsy specimens, 210 glomeruli (16.2%) had segmental lesions, comprising 57 (27%) cellular TIP, 4 (2%) fibrous TIP, 42 (20%) cellular NOS, 86 (41%) fibrous NOS, and 21 (10%) fibrous PH lesions. Each case was also classified into one of the five histological variants of the Columbia classification: collapsing (COL), TIP, cellular (CEL), PH, or NOS. Overlap of segmental lesions in different location categories was seen in the COL, TIP, and PH variants, and heterogeneity of quality was apparent in the COL and CEL variants. Histological findings of the CEL variant (endocapillary hypercellularity) were observed in nine of the 13 COL variants. Both location and quality correlated with disease duration, degree of proteinuria, and histological severity of global glomerular sclerosis and tubulo-interstitial lesions. These results demonstrated the histological heterogeneity of glomerular segmental lesions in all variants of the Columbia classification, except NOS. However, the fidelity of location and dominance of histological features were generally conserved in the TIP and PH variants. The COL and CEL variants warrant further investigation because of their overlapping histological findings and apparent histological heterogeneity in the glomerular segmental lesions.Electronic supplementary materialThe online version of this article (doi:10.1007/s11255-011-9932-y) contains supplementary material, which is available to authorized users.

Phase field investigation on cellular tip splitting during directional solidification

Cellular growth during directional solidification with different cellular spacings is investigated by using the quantitative phase field model. Simulation results reveal that the cellular tip splitting is not noise-induced but deterministic. The non-monotonic curvature distribution from cellular tip to grooves reveals the tip splitting instability. The monotonicity of curvature distribution is determined by the ratio of the cellular tip radius to the cellular width.

The knock‐out of ARP3a gene affects F‐actin cytoskeleton organization altering cellular tip growth, morphology and development in moss Physcomitrella patens

The seven subunit Arp2/3 complex is a highly conserved nucleation factor of actin microfilaments. We have isolated the genomic sequence encoding a putative Arp3a protein of the moss Physcomitrella patens. The disruption of this ARP3A gene by allele replacement has generated loss-of-function mutants displaying a complex developmental phenotype. The loss-of function of ARP3A gene results in shortened, almost cubic chloronemal cells displaying affected tip growth and lacking differentiation to caulonemal cells. In moss arp3a mutants, buds differentiate directly from chloronemata to form stunted leafy shoots having differentiated leaves similar to wild type. Yet, rhizoids never differentiate from stem epidermal cells. To characterize the F-actin organization in the arp3a-mutated cells, we disrupted ARP3A gene in the previously described HGT1 strain expressing conditionally the GFP-talin marker. In vivo observation of the F-actin cytoskeleton during P. patens development demonstrated that loss-of-function of Arp3a is associated with the disappearance of specific F-actin cortical structures associated with the establishment of localized cellular growth domains. Finally, we show that constitutive expression of the P. patens Arp3a and its Arabidopsis thaliana orthologs efficiently complement the mutated phenotype indicating a high degree of evolutionary conservation of the Arp3 function in land plants.

Effects of electric current pulse on stability of solid/liquid interface of Al–4.5 wt.% Cu alloy during directional solidification

This paper investigates the effects of the electric current pulse (ECP) on the interface stability of Al–4.5% Cu alloy during the directional solidification. Experimental results show both the cellular spacing and the mushy zone depth decrease with increasing current density, moreover, the solid–liquid interface morphology transforms from dendritic to cellular or even planar interface. The secondary dendrites are suppressed due to the homogeneous distribution of the solute and the increase of temperature gradient originated from ECP. The decrease of the cellular spacing results from branching of the cellular tip caused by Joule heat of the current exerting on cellular tip.

The expression of an extensin-like protein correlates with cellular tip growth in tomato.

Extensins are abundant proteins presumed to determine physical characteristics of the plant cell wall. We have cloned a cDNA encoding LeExt1 from a tomato (Lycopersicon esculentum Mill.) root hair cDNA library. The deduced sequence of the LeExt1 polypeptide defined a novel type of extensin-like proteins in tomato. Patterns of mRNA distribution indicated that expression of the LeExt1 gene was initiated in the root hair differentiation zone of the tomato rhizodermis. Cloning of the corresponding promoter and fusion to the -glucuronidase (GUS) reporter gene allowed detailed examination of LeExt1 expression in transgenic tomato plants. Evidence is presented for a direct correlation between LeExt1 expression and cellular tip growth. LeExt1/GUS expression was detectable in trichoblasts (=root hair-bearing cells), but not in atrichoblasts of the tomato rhizodermis. Both hair formation and LeExt1 expression was inducible by the plant hormone ethylene. Comparative analysis of the LeExt1/GUS expression was performed in transgenic tomato, potato (Solanum tuberosum), tobacco (Nicotiana tabacum), and Arabidopsis plants. In the apical/basal dimension, GUS staining was absent from the root cap and undifferentiated cells at the root tip in all species investigated. It was induced at the distal end of the differentiation zone and remained high proximally to the root/hypocotyl boundary. In the radial dimension, GUS expression was root hair specific in the solanaceous species. Whereas LeExt1 mRNA was exclusively detectable in the rhizodermis, root hair-specific expression correlated with GUS expression in germinating pollen tubes. This is correlative evidence for a role of LeExt1 in root hair tip growth [corrected].

Tip60 Inhibits Activation of CREB Protein by Protein Kinase A

We have previously identified a cDNA encoding a cellular protein, Tip60 (Tat interactive protein, 60 kDa), that specifically interacts with the Tat (transactivating transcriptional regulator) protein of the human immunodeficiency virus-1 (HIV-1). In this report, we have characterized cellular Tip and find that it is a 60 kDa nuclear protein expressed in a wide variety of differentiated cell lines from insects to man. To identify cellular functions of Tip, we have assayed the effects of Tip on cellular pathways that Tat has been reported to affect. Overexpression of Tip results in an almost complete block in activation of a Gal4-CREB (cAMP response element binding protein) fusion protein by cyclic AMP dependent protein kinase A (PKA). This inhibition appears to be mediated through direct interaction of Tip and CREB, since Tip directly binds to CREB protein in vitro. We show that amino acid substitutions of two conserved amino acids found in the putative acetyl coenzyme A binding motif of Tip completely abolishes the histone acetyltransferase (HAT) activity of recombinant Tip. Inhibition of CREB activation by Tip is not diminished in a HAT negative Tip mutant, indicating that Tip can negatively regulate gene expression independent of HAT activity. Recently, Tip has also been shown to be a transcriptional coactivator of nuclear hormone receptors; therefore, Tip can both activate transcription factors of one signaling pathway (nuclear hormone receptors) and bind to a different transcription factor (CREB) and inhibit activation of another signaling pathway.

In situ observations of stress-induced defect formation at the solid—liquid interface

Directional solidification experiments have been carried out to examine in situ the stress generated at the interface which lead to the formation of defects. A transparent system, which is also birefringence, is selected for the study to observe simultaneously both the morphology of the interface and the stress fringes. For a cusp orientation, the instability of the interface is shown to give rise to stress generation, and when sufficient stress accumulates, a grain boundary is formed to relieve the stress. A stress-induced parallel grain boundary formation is shown to be the mechanism for planar-to-cellular interface transition. The same mechanism is shown to cause cellular tip splitting to adjust the cellular spacing. A simple model of stress generation due to concentration gradients near the perturbed interface is developed to show the possible mechanism of stress generation.

Numerical simulation of cellular tip growth.

Numerical simulations of a model for cellular tip growth are performed, using a Green's function formalism. In this model, cellular growth is coupled to a morphogen concentration outside the cell. Typical shapes are obtained, and compare satisfactorily to experimental results. \textcopyright{} 1996 The American Physical Society.

Improved analysis of tip undercooling in cellular solidification of a binary alloy

Classical expressions for the undercooling, Δ T, at the tip of a cell during arrayed-directional solidification of a binary alloy, are compared with the experimental data on succinonitrile-acetone alloys. The range of validity of existing cellular models on cell tip undercooling is then determined. It is shown that cellular growth is an intricate situation in which the cell shape and the solute partitioning coefficient need to be properly included in the analysis. An improved analysis of cellular tip undercooling is developed, and the results presented in terms of an intrinsic solutal undercooling, Δ T s, and a part, Δ T SMI, that is associated with the simultaneous existence of a shape (S) and some morphological instability (MI).