Centromeres are essential for proper chromosome segregation to the daughter cells during mitosis and meiosis. Chromosomes of most eukaryotes studied so far have regional centromeres that form primary constrictions on metaphase chromosomes. These monocentric chromosomes vary from point centromeres to so-called "meta-polycentromeres", with multiple centromere domains in an extended primary constriction, as identified in Pisum and Lathyrus species. However, in various animal and plant lineages centromeres are distributed along almost the entire chromosome length. Therefore, they are called holocentromeres. In holocentric plants, centromere-specific proteins, at which spindle fibers usually attach, are arranged contiguously (line-like), in clusters along the chromosomes or in bands. Here, we summarize findings of ultrastructural investigations using immunolabeling with centromere-specific antibodies and super-resolution microscopy to demonstrate the structural diversity of plant centromeres. A classification of the different centromere types has been suggested based on the distribution of spindle attachment sites. Based on these findings we discuss the possible evolution and advantages of holocentricity, and potential strategies to segregate holocentric chromosomes correctly.

Biology Centre Czech Academy of Sciences 37005 České Budějovice Czech Republic

Department of Biological Sciences Chungnam National University Daejeon 34134 Korea

Department of Botany Federal University of Pernambuco Recife 50670 901 Pernambuco Brazil

Department of Chromosome Biology Max Planck Institute for Plant Breeding Research 50829 Cologne Germany

Leibniz Institute of Plant Genetics and Crop Plant Research Gatersleben 06466 Seeland Germany

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