Unique Characteristics of Cell Division in Vascular Plants


In eukaryotes, both microtubules (MTs) and microfilaments (MFs) are cytoskeletal polymers involved in meristematic cell proliferation. While animal cells build their MT arrays from structured organelles, such as centrosomes, and while they depolymerise their MFs and become round during mitosis, vascular plant cells lack centrosomes, maintain a filamentous actin cage around the spindle and are surrounded by a cell wall, preventing cellular mobility. During the cell cycle, plants activate specific dispersed MT nucleating sites, revealing successive plant‐specific cytoskeletal arrays. The MF meshwork surrounding the spindle eventually drives the centrifugal growth of MTs, which leads Golgi‐derived vesicles to fuse and separate each daughter cell during cytokinesis. The main orientation of actin fibres is parallel to that of spindle MTs, while a perpendicular constriction ring ensures animal daughter cell separation. However, despite the differences in their cytoskeleton behaviour and dynamics, both cell types succeed in controlled chromosome segregation.

Key Concepts

  • Mitosis is a short cell cycle period during which the cytoskeleton ensures balanced segregation of duplicated sister chromatids into two daughter cells.
  • Plant cells have to deal with their pecto‐cellulosic cell wall, constraining the orientation of the division axis in order to control morphogenesis.
  • In plants, primary stem cells are located in shoot and root apical meristems (SAM and RAM) and secondary growth is ensured by cambium activation.
  • Besides the conservation of microtubules (MTs) and actin filaments (MFs), required for successful karyokinesis and cytokinesis, respectively, in all eukaryotes, plant mitotic cells present an original cytoskeleton organisation.
  • All the somatic cells of vascular plants lack centrosomes, and microtubule organising centres are spread all over the nuclear envelope, at the cell cortex and along pre‐existing microtubules.
  • Preceding the mitotic step per se, maintenance of centromere/kinetochore cohesion and integrity is required for the building of a bipolar spindle and accurate chromosome segregation.
  • During mitosis, the activity of MT nucleating complexes (TuRCs) and MT‐associated proteins (MAPs) leads to a succession of MT arrays that reorganise in the cytoplasm: cortical MTs/perinuclear MTs/pre‐prophase band MTs/spindle MTs/phragmoplast MTs. The spindle apparatus, consisting of kinetochore fibres and interpolar MTs, is barrel‐shaped and lacks polar asters. Some of the MAPs also connect MFs.
  • Gamma‐tubulin containing complexes and augmin complexes participate in the nucleation of new highly dynamic MTs that ensure spindle robustness.
  • Actin filaments intermingle with MTs and do not depolymerise during mitosis, except in the PPB, which becomes an actin‐deplete zone (ADZ).
  • The cell plate expands centrifugally to the cell edges thanks to actin filaments, preceding MTs on which Golgi‐derived vesicles move towards the equator and fuse together.

Keywords: plants; cell division; mitosis; cytoskeleton; microtubules; actin microfilaments; preprophase band; spindle; phragmoplast; centromere/kinetochore; centromere cohesion

Figure 1. Establishment of division polarity. Immunolabelling of MTs (red) and DAPI staining of chromatin (blue). (a, b) During G2 phase of the cell cycle, MTs are nucleated at the nuclear surface from dispersed γTuRC complexes. Then, they reorganise into small asters (arrows), which further converge to poles (arrow heads). They progressively form a bipolar prospindle. Simultaneously in the cortex, while polar cortical MTs depolymerise, a preprophase band formed by dense MT bundles (bracket) forecasts the location of the future cell plate. (c) z‐Stack of six views separated in the z axis by 0.3 µm. (d) Details of an equatorial focal plane in the same prophase cell. Bars 2 µm.
Figure 2. MT dynamics throughout the cell cycle. (a) Immunolabelling of a root tip with anti‐tubulin antibodies, revealed using secondary antibodies marked by Alexa 568 fluorochrome. Chromatin staining using DAPI (blue). (b) Interphase cell with MTs radiating from the nuclear surface towards the cell cortex. (c) Early PPB and (d) mature PPB (brackets) in the cortex around the G2 nucleus. (e) Prophase cell with two polar caps and MT arrays pushing towards the nucleus. (f) Prometaphase spindle starting to spread at the poles. (g) Metaphase spindle with barrel‐shaped spindle poles. (h–j) Anaphase spindles refocalising the MT at poles through the activity of MAPs (e.g. ATK1). (k) Early telophase with phragmoplast MTs between daughter nuclei. (l) Late telophase with a wreath of short phragmoplast MTs growing centrifugically towards the cortex. Bars = 2 µm.
Figure 3. GIP1‐GFP expression under the control of its native promoter in gip1gip2 mutant background. Chromosomes are labelled blue, using DAPI staining. GIP1 is a γTuRC component required for spindle robustness and genome maintenance. It localises on the spindle in metaphase (a), especially kinetochore fibres in anaphase (b) and relocates in telophase on the phragmoplast (c). Bars = 2 µm.
Figure 4. Phragmoplast MT immunolabelling in late telophase. (a–f) Six focal planes separated by 1 µm in the z axis show the wreath‐like organisation of phragmoplast MTs. (a, c) Cell surface. Two sets of MTs, parallel to the spindle axis but with opposite polarity intermingle at the centre of the interzone. Their (+) end is located at the cell plate. (d–f) In a more central part of the cell where daughter nuclei become visible after DAPI staining (blue), the growing cell plate (dotted line in f) forms by Golgi‐derived vesicle fusion. MTs depolymerise in the central part of the cell and new MT assembly takes place at the periphery, allowing centrifugal extension and further fusion of the cell plate with the mother plasma membrane. Bars = 2 µm.
Figure 5. Schematic drawing of MT and MF dynamics throughout the cell cycle of plant cells. Proteins involved in each peculiar cytoskeletal array are mentioned on the sides.


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Further Reading

Batzenschlager M, Lermontova I, Schubert V, et al. (2015) Arabidopsis MZT1 homologs GIP1 and GIP2 are essential for centromere architecture. Proceedings of the National Academy of Sciences of the United States of America 112 (28): 8656–8660.

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Li S, Sun T, Ren H, et al. (2015) The functions of the cytoskeleton and associated proteins during mitosis and cytokinesis in plant cells. Frontiers in Plant Science 6: 282.

Lipka E, Herrmann A and Mueller S (2015) Mechanisms of plant cell division. WIREs Developmental Biology 4: 391–405. DOI: 10.1002/wdev.186.

Rasmussen CG, Wright AJ and Müller S (2013) The role of the cytoskeleton and associated proteins in determination of the plant cell division plane. Plant Journal 75 (2): 258–269.

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Chabouté, Marie‐Edith, and Schmit, Anne‐Catherine(Jul 2016) Unique Characteristics of Cell Division in Vascular Plants. In: eLS. John Wiley & Sons Ltd, Chichester. http://www.els.net [doi: 10.1002/9780470015902.a0001686.pub3]