Chromatin Diminution

Chromatin diminution is defined as chromosomal fragmentation, followed by the elimination of part of the chromosome during mitosis. The process was first observed in early cleavage divisions of the Parascaris equorum embryo by Boveri in 1887, and later on was also found to occur in some other parasitic nematodes, copepods and ciliates.

Keywords: chromatin diminution; DNA elimination; programmed genome rearrangement; germline-soma differentiation; nematode development

Figure 1. Chromatin diminution in Parascaris univalens. (a) Anaphase of the first cleavage division. (b) Anaphase of the second cleavage division. Chromatin diminution occurs in the upper S1 cell but not in the lower P1 cell. (c) Four-cell stage after completion of the second cleavage division. The cells S1a, S1b and S2 give rise to the somatic cells, while the P2 cell represents the germline. P0, zygote; C, centromere; E, eliminated chromatin; N, nucleus; P0-P2, germline; S1, S1a, S1b, S2, presomatic cells. From Müller et al. (1996).
Figure 2. Cell lineage of the early embryo of Parascaris univalens. Chromatin diminution is indicated by a broken bar. The germline lineage is drawn in thick red lines. Presomatic cell lineages before elimination are represented in thin red lines and somatic cell lineages in broken red lines. From Müller et al. (1996).
Figure 3. Process of chromatin diminution (in chromosomal breakage region ( CBR)-1; blue box) and of the organization of the DNA in the germline, somatic and eliminated chromatin in Ascaris suum. Chromosomal breakage may occur at many different sites within the CBR-1. This process is followed by the addition of telomeric (TTAGGC)n repeats to all broken chromosomal ends (hatched portions), and by the degradation of the eliminated DNA in the cytoplasm. The gene localized to the left of CBR-1 (red box) is retained in the somatic genome, but becomes located close to the telomere upon chromatin diminution, whereas the gene to the right of CBR-1 (purple box) will be eliminated from all somatic cells. The different diameter of the white arrows indicates the relative amounts of retained somatic and eliminated DNA upon chromatin diminution. The small arrow on the red box denotes the direction of transcription. Modified from Müller et al. (1996).
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 References
    Aeby P, Spicher A, de Chastonay Y, Müller F and Tobler H (1986) Structure and genomic organization of proretrovirus-like elements partially eliminated from the somatic genome of Ascaris lumbricoides. EMBO Journal 5: 3353–3360.
    Bennett KL and Ward S (1986) Neither a germline-specific nor several somatically expressed genes are lost or rearranged during embryonic chromatin diminution in the nematode Ascaris lumbricoides var. suum. Developmental Biology 118: 141–147.
    book Boveri T (1910) "Die Potenzen der Ascaris-Blastomeren bei abgeänderter Furchung. Zugleich ein Beitrag zur Frage qualitativ-ungleicher Chromosomen-Teilung" In: Festschrift für R. Hertwig. Jena: Fischer.
    Esteban MR, Giovinazzo G and Goday C (1995) Chromatin diminution is strictly correlated to somatic behavior in early development of the nematode Parascaris univalens. Journal of Cell Science 108: 2393–2404.
    Etter A, Aboutanos M, Tobler H and Müller F (1991) Eliminated chromatin of Ascaris contains a gene that encodes a putative ribosomal protein. Proceedings of the National Academy of Sciences of the USA 88: 1593–1596.
    Etter A, Bernard V, Kenzelmann M, Tobler H and Müller F (1994) Ribosomal heterogeneity from chromatin diminution in Ascaris lumbricoides. Science 265: 954–956.
    Goday C and Pimpinelli S (1986) Cytological analysis of chromosomes in the two species Parascaris univalens and P. equorum. Chromosoma 94: 1–10.
    Goday C, Gonzales-Garcia JM, Esteban MR, Giovinazzo G and Pimpinelli S (1992) Kinetochores and chromatin diminution in early embryos of Parascaris univalens. Journal of Cell Biology 118: 23–32.
    Huang Y-J, Stoffel R, Tobler H and Müller F (1996) A newly formed telomere in Ascaris suum does not exert a telomere position effect on a nearby gene. Molecular and Cellular Biology 16: 130–134.
    Magnenat L, Tobler H and Müller F (1999) Developmentally regulated telomerase activity is correlated with chromosomal healing during chromatin diminution in Ascaris suum. Molecular and Cellular Biology 19: 3457–3465.
    Moritz KB (1967) Die Blastomerendifferenzierung für Soma und Keimbahn bei Parascaris equorum. II. Untersuchungen mittels UV-Bestrahlung und Zentrifugierung. Roux' Archiv für Entwicklungsmechanik 159: 203–266.
    Moritz KB and Roth GE (1976) Complexity of germline and somatic DNA in Ascaris. Nature 259: 55–57.
    Müller F, Walker P, Aeby P et al. (1982) Nucleotide sequence of satellite DNA contained in the eliminated genome of Ascaris lumbricoides. Nucleic Acids Research 10: 7493–7510.
    Müller F, Wicky C, Spicher A and Tobler H (1991) New telomere formation after developmentally regulated chromosomal breakage during the process of chromatin diminution in Ascaris lumbricoides. Cell 67: 815–822.
    Müller F, Bernard V and Tobler H (1996) Chromatin diminution in nematodes. BioEssays 18: 133–138.
    Spicher A, Etter A, Bernard V, Tobler H and Müller F (1994) Extremely stable transcripts may compensate for the elimination of the gene fert-1 from all Ascaris lumbricoides somatic cells. Developmental Biology 164: 72–86.
    book Tobler H (1986) "The differentiation of germ and somatic cell lines in nematodes". In: Hennig W (ed.) Germline-Soma Differentiation; Results and Problems in Cell Differentiation, pp. 1–69. Berlin: Springer
 Further Reading
    Goday C and Pimpinelli S (1993) The occurrence, role and evolution of chromatin diminution in nematodes. Parasitology Today 9: 319–322.
    Pimpinelli S and Goday C (1989) Unusual kinetochores and chromatin diminution in Parascaris. Trends in Genetics 5: 310–315.
    Tobler H, Etter A and Müller F (1992) Chromatin diminution in nematode development. Trends in Genetics 8: 427–432.
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Tobler, Heinz, and Müller, Fritz(Apr 2001) Chromatin Diminution. In: eLS. John Wiley & Sons Ltd, Chichester. http://www.els.net [doi: 10.1038/npg.els.0001181]