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There may also be selection on modifiers of recombination that influence the stage of meiosis at which distorters gain a transmission advantage (Haig 2010; Brandvain and Coop 2012).
Under this model, genetic modifiers of recombination rate are themselves plastic, and the rate of recombination caused by a modifier depends on the condition of the organism in which it is found.
However, he also found evidence for the existence of genetic, rather than structural, modifiers of recombination and was unable to determine the relative contributions of these two phenomena with respect to the suppression of recombination in this region.
It is hypothesized that a gradual reduction of crossover frequencies is due to the spread of genetic modifiers of recombination rates or to chromosome rearrangements such as inversions [ 2].
Of course, modifiers of recombination are well known to Drosophila (Baker et al. 1976; Cattani et al. 2012), and the theory of conditions that favor increased (or decreased) recombination is rich (Barton 1995, 2010; Feldman et al. 1996; Otto and Michalakis 1998; Lenormand and Otto 2000; Otto and Barton 2001; Otto and Lenormand 2002; Martin et al. 2006).
In the early stages of sex-chromosome differentiation, it is suggested that interactions between sex-determining genes and sexually antagonistic genes can drive selection for reduced recombination via gradual reduction of crossover frequencies, due to the spread of genetic modifiers of recombination rates and/or chromosome rearrangements such as inversions [ 2].
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Testing this prediction requires modifier-of-recombination theory (e.g., Otto and Barton 1997; Martin et al. 2006; Roze and Barton 2006; Kermany and Lessard 2012).
We suggest that future theoretical studies (i) obtain analogous approximations for bi- rather than unidirectional gene flow, (ii) account for epistasis and dominance, (iii) incorporate the distribution of fitness effects of beneficial mutations, and (iv) employ a stochastic modifier-of-recombination model to assess the importance of nonzero optimal recombination rates.
The frequency change of the modifier, averaged over a large number of samples, is then used as a proxy for the selection on the modifier of recombination.
Another potential modifier of recombination, originally implicated in empirical work (McGaugh et al. 2012), was confirmed here by observing genome-wide elevated recombination in historical recombination rates of D. miranda despite a much lower effective population size.
The concept that gene expression may act as a "plastic" and heritable modifier of recombination, directly or epigenetically, is particularly relevant to evolutionary models on the maintenance of recombination.
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