Abstract
The emergence of eukaryotes around two billion years ago provided new challenges for the chromosome segregation machineries: the physical separation of multiple large and linear chromosomes from the microtubule-organizing centres by the nuclear envelope. In this review, we set out the diverse solutions that eukaryotic cells use to solve this problem, and show how stepping away from ‘mainstream’ mitosis can teach us much about the mechanisms and mechanics that can drive chromosome segregation. We discuss the evidence for a close functional and physical relationship between membranes, nuclear pores and kinetochores in generating the forces necessary for chromosome segregation during mitosis.
Highlights
The accurate segregation of replicated genomes into daughter cells during cell division is a prerequisite for life in all three domains of life: Archaea, bacteria, Eukaryota
All mechanisms known so far can be reduced to two basic components: (i) directional force-generating mechanisms that consume chemical energy provided by hydrolysis of nucleotide triphosphates (NTP), and (ii) an adaptor that physically links this machinery to the carriers of genetic information
The most obvious route, and the one taken by higher eukaryotes, is to disassemble the nucleus at the onset of mitosis. This process, called nuclear envelope (NE) breakdown (NEBD), begins with dispersal of nucleoporins, followed by the physical deformation and tearing of the NE by forces generated by dynein motors and microtubules, and the disassembly of the nuclear lamina, which is required for the structural integrity of the nucleus [29]
Summary
The emergence of eukaryotes around two billion years ago provided new challenges for the chromosome segregation machineries: the physical separation of multiple large and linear chromosomes from the microtubule-organizing centres by the nuclear envelope. We set out the diverse solutions that eukaryotic cells use to solve this problem, and show how stepping away from ‘mainstream’ mitosis can teach us much about the mechanisms and mechanics that can drive chromosome segregation. We discuss the evidence for a close functional and physical relationship between membranes, nuclear pores and kinetochores in generating the forces necessary for chromosome segregation during mitosis
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