scholarly journals EXPECTED LINKAGE DISEQUILIBRIUM FOR A NEUTRAL LOCUS LINKED TO A CHROMOSOMAL ARRANGEMENT

Genetics ◽  
1983 ◽  
Vol 103 (3) ◽  
pp. 545-555
Author(s):  
Curtis Strobeck

ABSTRACT The expected value of the squared linkage disequilibrium is derived for a neutral locus associated with a chromosomal arrangement that is maintained in the population by strong balancing selection. For a given value of recombination, the expected squared linkage disequilibrium is shown to decrease as the intensity of selection maintaining the arrangement increases. The transient behavior of the expected square linkage disequilibrium is also derived. This theory applies to loci that are closely linked to inversions in Drosophila species and to loci closely linked to the differential segments of the translocation complexes in ring-forming species of Oenothera. In both cases the strong linkage disequilibria that have been observed in natural populations can be explained by random drift.

Genetics ◽  
1979 ◽  
Vol 93 (2) ◽  
pp. 497-523
Author(s):  
M Loukas ◽  
C B Krimbas ◽  
Y Vergini

ABSTRACT Gametic frequencies were obtained in four natural populations of D. subobscura by extracting wild chromosomes and subsequently analyzing them for inversions and allozymes. The genes Lap and Pept-1, both located within the same inversions of chromosome 0, were found in striking nonrandom associations with them of the same kind and degree in all populations studied. On the contrary, the gene Acph, also located within the previously mentioned inversions, was found in linkage disequilibrium with them only in two populations and of opposite directions. This is also the case for the genes Est-9 and Hk, both located within chromosome E inversions. While the gene Est-9 was in strong linkage disequilibrium with the inversions, of the same kind and degree in all populations studied, Hk was found to be in linkage equilibrium. Allele frequencies for the 29 genes studied do not show geographical variation except for the genes Lap, Pept-1 and Est-9, the ones found in linkage disequilibria with the geographically varying gene arrangements. Although mechanical or historical explanations for these equilibria cannot be ruled out, these data cannot be explained satisfactorily by the "middle gene explanation," which states that loci displaying such linkage disequilibria are the ones located near the break points of inversions, while the ones displaying linkage equilibria with them are located in the middle of them. There is no evidence for consistent linkage disequilibria between pairs of loci, except for the closely linked genes of the complex locus, Est-9. This would imply, if it is not a peculiarity of the Est-9 complex, that the linkage disequilibria aye found only between very closely linked loci or that, far less closely linked genes, the associations are too weak to be detected by the usual samples sizes.


1970 ◽  
Vol 16 (2) ◽  
pp. 165-177 ◽  
Author(s):  
Tomoko Ohta ◽  
Motoo Kimura

SUMMARYAssociative overdominance arises at an intrinsically neutral locus through its non-random association with overdominant loci. In finite populations, even if fitness is additive between loci, non-random association will be created by random genetic drift.The magnitude of such associative overdominance is roughly proportional to the sum of between the neutral and the surrounding over-dominant loci, where is the squared standard linkage deviation, defined between any two loci by the relationin which p and 1 – p are frequencies of alleles A1 and A2 in the first locus, q and 1 – q are frequencies of alleles B1 and B2 in the second locus, and D is the coefficient of linkage disequilibrium. A theory was developed based on diffusion models which enables us to obtain formulae for under various conditions, and Monte Carlo experiments were performed to check the validity of those formulae.It was shown that if A1 and A2 are strongly overdominant while B1 and B2 are selectively neutral, we have approximatelyprovided that 4Nec ≫ 1, where Ne is the effective population size and c is the recombination fraction between the two loci. This approximation formula is also valid between two strongly overdominant as well as weakly overdominant loci, if 4Nec ≫ 1.The significance of associative overdominance for the maintenance of genetic variability in natural populations was discussed, and it was shown that Nes′, that is, the product between effective population size and the coefficient of associative overdominance, remanis constant with varying Ne, if the total segregational (overdominant) load is kept constant.The amount of linkage disequilibrium expected due to random drift in experimental populations was also discussed, and it was shown that in the first generation, if it is produced by extracting n chromosomes from a large parental population in which D = 0.


Genetics ◽  
1979 ◽  
Vol 92 (4) ◽  
pp. 1295-1314
Author(s):  
C C Laurie-Ahlberg ◽  
B S Weir

ABSTRACT Nine laboratory populations of D. melanogaster were surveyed by starch gel electrophoresis for variation at 17 enzyme loci. A single-fly extract could be assayed for all 17 enzymes, so that the data consist of 17-locus genotypes.— Pairwise linkage disequilibria were estimated from the multilocus genotypic frequencies, using both BURROWS' and HILL'S methods. Large amounts of link-age disequilibrium were found, in contrast to the results reported for natural populations.—Knowledge of the approximate sizes of these populations was used to compare the observed heterozygosities and linkage disequilibria with predictions of the neutral allele hypothesis. The relatively large amount of linkage disequilibrium is consistent with the small sizes of the populations. However, the levels of heterozygosity in at least some populations suggest that some mechanism has been operating to retard the rate of decay by random drift. Several examples of significant deviation from Hardy-Weinberg frequencies and the large amount of linkage disequilibnim present in these populations indicate that a likely mechanism is selective effects associated with neutral alleles because of linkage disequilibrium with selected loci (e.g., "associative overdominance"). The results are therefore consistent with both neutralist, and selectionist hypotheses, but suggest the importance of considering linkage disequilibrium between neutral and selected loci when attempting to explain the dynamics of enzyme polymorphisms.


Genetics ◽  
2001 ◽  
Vol 157 (2) ◽  
pp. 899-909
Author(s):  
Rongling Wu ◽  
Zhao-Bang Zeng

Abstract A new strategy for studying the genome structure and organization of natural populations is proposed on the basis of a combined analysis of linkage and linkage disequilibrium using known polymorphic markers. This strategy exploits a random sample drawn from a panmictic natural population and the open-pollinated progeny of the sample. It is established on the principle of gene transmission from the parental to progeny generation during which the linkage between different markers is broken down due to meiotic recombination. The strategy has power to simultaneously capture the information about the linkage of the markers (as measured by recombination fraction) and the degree of their linkage disequilibrium created at a historic time. Simulation studies indicate that the statistical method implemented by the Fisher-scoring algorithm can provide accurate and precise estimates for the allele frequencies, recombination fractions, and linkage disequilibria between different markers. The strategy has great implications for constructing a dense linkage disequilibrium map that can facilitate the identification and positional cloning of the genes underlying both simple and complex traits.


1983 ◽  
Vol 25 (2) ◽  
pp. 139-145 ◽  
Author(s):  
C. Strobeck ◽  
G. B. Golding

The variance of three-locus linkage disequilibria for an equilibrium infinite alleles model is solved numerically on a computer, using identity coefficients. It is shown that the variance of three-locus linkage disequilibrium created by random drift, although smaller than the variance of two-locus linkage disequilibrium, is of the same order of magnitude. Hence third-order disequilibria are not necessarily good indications of selection. The formula for the variance of linkage disequilibrium is given when there is no recombination between the genes. This model can also be interpreted as intragenic recombination between three sites within a gene.


1987 ◽  
Vol 50 (3) ◽  
pp. 187-193
Author(s):  
Catherine Montchamp-Moreau ◽  
Mariano Katz

SummaryLinkage disequilibrium between five polymorphic enzymic loci of the third chromosome (Esterase-6, Phosphoglucomutase, Esterase-C, Aldehyde Oxidase and Acid Phosphatase) was studied in experimental populations of Drosophila simulans. Gametic data were obtained by mating sampled males with homozygous females at the five loci. Four cage populations were initiated with flies caught from natural populations. Extensive linkage disequilibrium was detected after 25 or 34 generations. The effective size of these populations was estimated about 400. Monte-Carlo simulations were performed in order to determine whether the observed disequilibria could be due to genetic drift. The observed probability distribution of the experimental values of r (the gametic correlation coefficient) was consistent with the distribution expected under random genetic drift. Our results are thus in accordance with the neutralist hypothesis.


Genetics ◽  
1977 ◽  
Vol 85 (3) ◽  
pp. 543-556
Author(s):  
E Zouros ◽  
G B Golding ◽  
Trudy F C MacKay

ABSTRACT When alleles are combined into few detectable classes, linkage correlations are underestimated most of the time. The probability that the linkage correlation will be underestimated is a function of the actual degree of correlation and the evenness of the allelic distribution, but is mainly determined by the distribution of alleles into distinguishable classes. With only two alleles per class this probability will usually be higher than 0.7. Also, the consistency in the sign of the linkage disequilibrium over many populations may escape detection. An increase of sample size by one order of magnitude or more may be required to compensate for the loss in detection power. It follows that the available electrophoretic studies of linkage correlations, although negative in their majority, do not suggest that epistatic interactions and linkage disequilibria are rare in natural populations.


PeerJ ◽  
2018 ◽  
Vol 6 ◽  
pp. e5110 ◽  
Author(s):  
Jacob A. Tennessen

The fates of genetic polymorphisms maintained by balancing selection depend on evolutionary dynamics at linked sites. While coevolution across linked, epigenetically-interacting loci has been extensively explored, such supergenes may be relatively rare. However, genes harboring adaptive variation can occur in close physical proximity while generating independent effects on fitness. Here, I present a model in which two linked loci without epistasis are both under balancing selection for unrelated reasons. Using forward-time simulations, I show that recombination rate strongly influences the retention of adaptive polymorphism, especially for intermediate selection coefficients. A locus is more likely to retain adaptive variation if it is closely linked to another locus under balancing selection, even if the two loci have no interaction. Thus, two linked polymorphisms can both be retained indefinitely even when they would both be lost to drift if unlinked. While these results may be intuitive, they have important implications for genetic architecture: clusters of mutually reinforcing genes may underlie phenotypic variation in natural populations, and such genes cannot be assumed to be functionally associated. Future studies that measure selection coefficients and recombination rates among closely linked genes will be fruitful for characterizing the extent of this phenomenon.


2017 ◽  
Author(s):  
Jacob A Tennessen

The fates of genetic polymorphisms maintained by balancing selection depend on evolutionary dynamics at linked sites. While coevolution across linked, epigenetically-interacting loci has been extensively explored, such supergenes may be relatively rare. However, genes harboring adaptive variation can occur in close physical proximity while generating independent effects on fitness. Here, I present a model in which two linked loci without epistasis are both under balancing selection for unrelated reasons. Using forward-time simulations, I show that recombination rate strongly influences the retention of adaptive polymorphism, especially for intermediate selection coefficients. A locus is more likely to retain adaptive variation if it is closely linked to another locus under balancing selection, even if the two loci have no interaction. Thus, two linked polymorphisms can both be retained indefinitely even when they would both be lost to drift if unlinked. Such clusters of mutually reinforcing genes may underlie phenotypic variation in natural populations. Future studies that measure selection coefficients and recombination rates among closely linked genes will be fruitful for characterizing the extent of this phenomenon.


1978 ◽  
Vol 32 (3) ◽  
pp. 215-229 ◽  
Author(s):  
Charles H. Langley ◽  
Diana B. Smith ◽  
F. M. Johnson

SUMMARYLinkage disequilibria between pairs of 8 polymorphic enzyme loci (αGpdh, Mdh, Adh, Est-6, Pgm, Odh, Est-C and Acph) in some 100 natural population samples of Drosophila melanogaster were examined. The estimates of linkage disequilibrium were made from zygotic frequencies. The magnitude of linkage disequilibria are small and similar to those in previous reports. Variation in linkage disequilibrium among related subpopulations was analysed by analysis of variance of the correlation coefficients. Despite the small absolute value of linkage disequilibrium there is a suggestion of a correlation among related subpopulations. The magnitude of linkage disequilibrium was observed to be positively correlated with linkage. Two cage populations were observed to demonstrate large amounts of linkage disequilibrium between closely linked loci in contrast to the situation in natural populations. This is attributable to the finite sizes of these cage populations.


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