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. Author manuscript; available in PMC: 2021 Oct 1.
Published in final edited form as: Mol Cell. 2020 Aug 28;80(1):9–20. doi: 10.1016/j.molcel.2020.08.007

Table 1.

Summary of Phase-separated Condensates in Cell Division

Structure Key components Properties Biological Function System
Spindle poles Centrosome
SPD-5 (C. elegans), Plk4 (X. laevis extracts and cultured human cells)

  • SPD-5 condensates promoted by Plk1, incorporated centrosome factors, nucleated MTs, hardened over time

  • Concentrates tubulin and generates microtubule asters; promotes MTOC formation

 C. elegans embryos (SPD-5) (Woodruff et al., 2017)

  • Plk4 condensates promoted by autophosphorylation, hardened over time, promoted centrosome maturation and stability

  • Centrosome maturation

 Recombinant Plk4 and X. laevis extracts (Gouveia et al., 2019), cultured human cells (Park et al., 2019)
Acentrosomal
Centrosome and centrosome-associated proteins

  • LISD promoted by Aurora A kinase activity, TACC3 binding to CHC17 and to microtubules

  • Promotes spindle assembly and formation of MTOC

 Recombinant proteins and mammalian oocytes (So et al., 2019)
Spindle body Tubulin, microtubule motors and crosslinkers

  • Tubulin polymers form tactoids that can fuse with other tactoids

  • Diffusion of microtubule-associated proteins

  • Two bipolar spindles will fuse into one bipolar spindle in a dynein- dependent manner

  • Local range liquid-like state preserves microtubule spindle assembly and homeostasis

  • Regulates microtubule orientation and density

 X. laevis extracts (Brugues and Needleman, 2014; Gatlin et al., 2009), purified tubulin and recombinant proteins (Edozie et al., 2019)
Tau

  • Concentrates tubulin, promotes nucleation, elongation, and bundling of MTs

  • Unknown role in spindle assembly

 Recombinant proteins (Hernandez-Vega et al., 2017)
TPX2

  • Concentrates tubulin, promotes nucleation of MTs

  • Promotes mitotic spindle density and assembly

 Recombinant proteins, X. laevis extracts (King and Petry, 2020)
BuGZ

  • Concentrates tubulin, promotes nucleation and bundling of MTs

  • Incorporates and promotes activity of Aurora A kinase

  • Promotes mitotic spindle density and assembly

 X. laevis extracts, recombinant proteins, and cultured human cells (Huang et al., 2018; Jiang et al., 2015)
Centromere Chromosomal passenger complex (INCENP, Borealin, Survivin)

  • Borealin condensates promoted by Cdk1/Cyclin B kinase activity, microtubules, a-satellite DNA, and nucleosomes

  • Nucleates microtubules; hardens in vitro and in cells

  • Promotes localization of CPC to inner centromere and midzone, spindle assembly checkpoint, and mitotic timing

 Cultured human cells (Trivedi et al., 2019)
Chromosomes Mitotic
Ki-67

  • Ki-67 protein surfactant properties mediated via steric hindrance and/or electrostatic interactions

  • Ki-67 promotes chromosome dispersal and promotes MT spindle formation

 Cultured human cells (Cuylen et al., 2016)
Meiotic
RNA binding proteins and lncRNA

  • Meiotic lncRNA droplets fuse with droplets of same RNA species

  • lncRNA droplets promote pairing of homologous chromosomes

 S. pombe (Ding et al., 2019)
Other Mostly stress granules, RNA splicing speckles, centrosome/pericentri olar matrix

  • DYRK3 promotes condensate dissolution at NEBD to release mitotic factors

  • Promotes spindle formation and mitotic timing

 Cultured human cells (Rai et al., 2018)

Abbreviations: MT, microtubule; MTOC, microtubule organizing center; NEBD, nuclear envelope breakdown