Cell division orientation and planar cell polarity pathways

The orientation of cell division has a crucial role in early embryo body plan specification, axis determination and cell fate diversity generation, as well as in the morphogenesis of tissues and organs. In many instances, cell division orientation is regulated by the planar cell polarity (PCP) pathways: the Wnt/Frizzled non-canonical pathway or the Fat/Dachsous/Four-jointed pathway. Firstly, using asymmetric cell division in both Drosophila and C. elegans, we describe the central role of the Wnt/Frizzled pathway in the regulation of asymmetric cell division orientation, focusing on its cooperation with either the Src kinase pathway or the heterotrimeric G protein pathway. Secondly, we describe our present understanding of the mechanisms by which the planar cell polarity pathways drive tissue morphogenesis by regulating the orientation of symmetric cell division within a field of cells. Finally, we will discuss the important avenues that need to be explored in the future to better understand how planar cell polarity pathways control embryo body plan determination, cell fate specification or tissue morphogenesis by mitotic spindle orientation.     

Cell division in leaves ofNicotiana

Summary Young leaves ofNicotiana tabacum were fixed in glutaraldehyde-formaldehyde followed by osmium tetroxide. The fine structure of dividing cells was studied. Before prophase a band of microtubules was observed between the nucleus and the cell wall at a position judged as the future plane of division. The microtubules in the band are 4–6 units deep and relatively closely packed, giving sections of the band a characteristic appearance. Micro-tubules of the mitotic spindle, the phragmoplast, and the preprophase band are morphologically similar. Some of the microtubules of the mitotic spindle and the phragmoplast have an undulate appearance. It is suggested that the undulate microtubules may have been fixed at a time when microwaves were traveling along them. The cell plate is formed by a fusion of small smooth surfaced vesicles and small coated vesicles. Fusion of small vesicles results first in larger vesicles and then in a meshwork of new cell-wall material surrounded by new regions of plasma membrane. Most of the vesicles are derived from dictyosomes and may be produced before and during prophase as well as during later stages of division. The ER may also contribute some vesicles to the cell plate.     

有丝分裂方向的影响因素

有丝分裂方向不仅关系到细胞增殖过程中子细胞的排列方式,而且对细胞分化、组织器官形态结构具有决定性作用。从细胞内环境和所处的微环境2个方面,讨论有丝分裂方向的影响因素。     

Cell division in baeocyte producing cyanobacteria

Summary An ultrastructural examination of cell division in two baeocyte producing cyanobacteria,Pleurocapsa minor andDermocarpa violaceae, reveals two distinct patterns of binary (transverse) fission. Septate binary fission, inPleurocapsa minor, involves centripetal synthesis and deposition of the mucopolymer cell wall layer (L 2). The ingrowth of the cytoplasmic membrane and L 1 cell wall layer, along with the synthesis of the L 2 cell wall layer, results in the formation of a prominent septum. Partitioning of the cell occurs by the constriction of the outer cell wall layers (L 3 and L 4) through the septum. InDermocarpa violaceae, constrictive binary fission occurs by the simultaneous ingrowth or constriction of the cytoplasmic membrane and all cell wall layers (L1, L2, L3, L4). Septate and constrictive binary fission may proceed symmetrically (medially) or asymmetrically (nonmedially). Multiple fission occurs regularly inDermocarpa violaceae and provides for a rapid means of reproduction when compared to binary fission. Successive radial and tangential divisions of the protoplast result in formation of many small daughter cells (baeocytes). The process of multiple fission is similar to septate binary fission with reduced septa being formed. However, constriction of the outer cell wall layers, through the septa, proceeds concurrently with septum formation.     

Photocontrol of the orientation of cell division in Adiantum

Summary Two-dimensional prothallia of Adiantum capillus-veneris always expanded in a plane which was at a right angle to any given direction of irradiation with continuous white light. The expansion began with a longitudinal division of the apical cell, in the filamentous protonema, and the orientation of the mitotic cell plate of this first longitudinal division as well as the subsequent divisions was always parallel to the direction of the incident light. When three irradiations with white light, interrupted by periods of darkness, were given, two transverse and one subsequent longitudinal division were induced. When the last two irradiations were given from the same direction, the cell plate of the first longitudinal division in most protonemata was oriented parallel to the direction of light. However, when the direction of light during the third irradiation was at right angle to that during the second, the frequency of the longitudinal division greatly decreased but that of the third transverse division increased. Thus, the orientation of the first longitudinal division appeared to be controlled in some way not only by the irradiation which actually induced the third division but also by that inducing the preceding transverse division, while the direction of light for the first transverse division had little effect on the orientation of the third division.     

Cell polarity from cell division

Apical-basal polarity of epithelial cells is critical for their symmetric versus asymmetric division and commonly thought to be established in interphase. In a novel type of cell division termed "mirror-symmetric", apical cell constituents accumulate during M-phase at the cleavage furrow, resulting in epithelial daughter cells with opposite apical-basal polarity.     

Cell division: mid-level management

When a fission yeast cell divides, the anillin-like protein mid1p helps to position the contractile ring in the cell middle. Recent experiments from two groups have shown how the cell-polarity factor pom1p negatively regulates the distribution of mid1p.     

Cell division by swelling stresses

It is demonstrated that, if the variations of viscosity throughout a cell are considered, swelling stresses may produce elongation and division. To do this it is necessary to generalize Betti's theorem to cover systems containing viscosity gradients and such a generalization is presented. On the basis of two special assumptions it is shown that most of the results of the diffusion drag theory of cell division may be duplicated by the present theory.     

Photocontrol of the orientation of cell division inAdiantum

The rate of transition from one- to two-dimensional growth of fernAdiantum gametophytes under white light depends on the age of gametophyte cultured under red light. When gametophytes were cultured for longer period under red light, the rate of transition decreased and the number of abnormal gametophytes increased. Although the first step of the transition was the first longitudinal cell division following the two transverse ones (Wada and Furuya, 1970), the time-lapse-video study revealed that the apical cell of protonemata became flattened in the plane perpendicular to the incident ray of white light before the first longitudinal cell division. Analytical study of growing part of the apical cell with grains of activated charcoal as markers revealed that the apical cell flattening occurred evenly throughout the equatorial circumference of the cell even in the shaded side of the protonemata as well as in the side irradiated with white light.