Abstract

The retention of supernumerary chromosomes in the germ-line of Sciara coprophila is part of a highly-intricate pattern of chromosome behaviours that have fascinated cytogeneticists for over 80 years. Germ-line limited (termed L or “limited”) chromosomes are cytologically heterochromatic and late-replicating, with more recent studies confirming they possess epigenetic hallmarks characteristic of constitutive heterochromatin. Little is known about their genetic constitution although they have been found to undergo cycles of condensation and de-condensation at different stages of development. Unlike most supernumeraries, the L chromosomes in S. coprophila are thought to be indispensable, although in two closely related species Sciara ocellaris and Sciara reynoldsi the L chromosomes, have been lost during evolution. Here, we review what we know about L chromosomes in Sciara coprophila. We end by discussing how study of the L chromosome condensation cycle has provided insight into the site and timing of both the erasure of parental “imprints” and also the placement of a putative “imprint” that might be carried by the sperm into the egg.

Highlights

  • Sciara coprophila possesses a complicated, sometimes bizarre, pattern of chromosomal behaviours that involve both the regular chromosomes and the supernumerary germ-line limited or L chromosomes [1,2,3]

  • L chromosome behaviour is best described as part of the whole

  • Studies on the sciarid species, Trichosia pubescens, have shown that in spermatocytes, L chromosomes become de-condensed and indistinguishable from ordinary chromosomes during the interphase between the meiotic divisions [12]. Based on this observation it was suggested that the de-condensation represents gene activity likely required for proper spermatogenesis, a notion supported by the unpublished results of Crouse who observed that spermatocytes having no L chromosomes were smaller, develop more slowly and do not undergo typical meiosis

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Summary

Introduction

Sciara coprophila possesses a complicated, sometimes bizarre, pattern of chromosomal behaviours that involve both the regular chromosomes and the supernumerary germ-line limited or L chromosomes [1,2,3]. The cycles of both kinds of chromosome are intimately associated. L chromosome behaviour is best described as part of the whole. Looking at the chromosomal complements provided by each parent, the female pro-nucleus provides 5 chromosomes

L Chromosome Elimination in the Soma
L Chromosome Elimination in the Germ-Line
L Chromosome Behaviour During Male Meiosis
The Epigenotype of L Chromosomes
L Chromosomes and Chromosomal Imprinting of the Regular Chromosomes
L Chromosomes and Imprinting of the Controlling Element
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