The
B chromosome system of the grasshopper Eyprepocnemis plorans has
provided support for our dynamic model on the evolution of parasite
B chromosomes, by which Bs are neutralized by the evolution of drive
suppressor genes in the A genome, so that they are condemned to extinction
through stochastic loss (and the low fitness of individuals carrying
four or more Bs) lasting hundreds or thousands of generations (depending
on population size), unless a new selfish B variant may emerge and regenerate
the polymorphism. With the present project, we propose to reach three
general objectives:
i)
to determine the molecular composition of B chromosomes in this species,
ii)
to reveal the possible expression of the DNA contained in the Bs, or possible
regulatory changes in genes located on the A chromosomes caused by B presence,
and
iii)
to disentangle the coevolution mechanisms underlying the neutralization-regeneration
process of these parasite chromosomes.
In
relation with the first objective, we will continue the analysis of the
B2 and B24 DNA libraries built in the former project, we will characterize
cytogenetically and molecularly the B chromosomes that have recently been
reported in natural populations from the Caucasus, we will analyse the
presence of retroelements in the B chromosomes that might explain their
high mutability, and will compare the potential capability to capture
centromere proteins among B chromosomes differing in transmission ratio.
In relation with the second objective, we plan to build a cDNA library
from individuals possessing B chromosomes, and to analyse the distribution
of 5-metil-cytosin on the A chromosomes in the presence of different
B chromosome types. With regard to the third objective, we will continue
analysing the evolution of B frequency in the population where Bs were
introduced in 1993, and will perform a population survey of the Torrox
region, an interesting locality where B chromosome substitutions are presently
taking place.
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