arXiv subjects

Kavita Jain

Publications and source records attributed to Kavita Jain.

At least 37 records · Page 2Linked to original sources

arXiv2016

Response of polygenic traits under stabilising selection and mutation when loci have unequal effects

We consider an infinitely large population under stabilising selection and mutation in which the allelic effects determining a polygenic trait vary between loci. We obtain analytical expressions for the stationary genetic variance as a function of the distribution of effects, mutation rate and selection coefficient. We also study the dynamics of the allele frequencies, focussing on short-term evolution of the phenotypic mean as it approaches the optimum after an environmental change. We find that when most effects are small, the genetic variance does not change appreciably during adaptation, and the time until the phenotypic mean reaches the optimum is short if the number of loci is large. However, when most effects are large, the change of the variance during the adaptive process cannot be neglected. In this case, the short-term dynamics may be described by that of a single locus of large effect. Our results may be used to understand polygenic selection driving rapid adaptation.

q-bio.PE↗

arXiv2015

Fixation probability of rare nonmutator and evolution of mutation rates

Although mutations drive the evolutionary process, the rates at which the mutations occur are themselves subject to evolutionary forces. Our purpose here is to understand the role of selection and random genetic drift in the evolution of mutation rates, and we address this question in asexual populations at mutation-selection equilibrium neglecting selective sweeps. Using a multitype branching process, we calculate the fixation probability of a rare nonmutator in a large asexual population of mutators, and find that a nonmutator is more likely to fix when the deleterious mutation rate of the mutator population is high. Compensatory mutations in the mutator population are found to decrease the fixation probability of a nonmutator when the selection coefficient is large. But, surprisingly, the fixation probability changes nonmonotonically with increasing compensatory mutation rate when the selection is mild. Using these results for the fixation probability and a drift-barrier argument, we find a novel relationship between the mutation rates and the population size. We also discuss the time to fix the nonmutator in an adapted population of asexual mutators, and compare our results with experiments.

q-bio.PE↗

arXiv2015

Real eigenvalues of non-Gaussian random matrices and their products

We study the properties of the eigenvalues of real random matrices and their products. It is known that when the matrix elements are Gaussian-distributed independent random variables, the fraction of real eigenvalues tends to unity as the number of matrices in the product increases. Here we present numerical evidence that this phenomenon is robust with respect to the probability distribution of matrix elements, and is therefore a general property that merits detailed investigation. Since the elements of the product matrix are no longer distributed as those of the single matrix nor they remain independent random variables, we study the role of these two factors in detail. We study numerically the properties of the Hadamard (or Schur) product of matrices and also the product of matrices whose entries are independent but have the same marginal distribution as that of normal products of matrices, and find that under repeated multiplication, the probability of all eigenvalues to be real increases in both cases, but saturates to a constant below unity showing that the correlations amongst the matrix elements are responsible for the approach to one. To investigate the role of the non-normal nature of the probability distributions, we present a thorough analytical treatment of the $2 \times 2$ single matrix for several standard distributions. Within the class of smooth distributions with zero mean and finite variance, our results indicate that the Gaussian distribution has the maximum probability of real eigenvalues, but the Cauchy distribution characterised by infinite variance is found to have a larger probability of real eigenvalues than the normal. We also find that for the two-dimensional single matrices, the probability of real eigenvalues lies in the range [5/8,7/8].

math-ph↗

arXiv2015

Effect of drift, selection and recombination on the equilibrium frequency of deleterious mutations

We study the stationary state of a population evolving under the action of random genetic drift, selection and recombination in which both deleterious and reverse beneficial mutations can occur. We find that the equilibrium fraction of deleterious mutations decreases as the population size is increased. We calculate exactly the steady state frequency in a nonrecombining population when population size is infinite and for a neutral finite population, and obtain bounds on the fraction of deleterious mutations. We also find that for small and very large populations, the number of deleterious mutations depends weakly on recombination, but for moderately large populations, recombination alleviates the effect of deleterious mutations. An analytical argument shows that recombination decreases disadvantageous mutations appreciably when beneficial mutations are rare as is the case in adapting microbial populations, whereas it has a moderate effect on codon bias where the mutation rates between the preferred and unpreferred codons are comparable.

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arXiv2014

Two-point correlation function of an exclusion process with hole-dependent rates

We consider an exclusion process on a ring in which a particle hops to an empty neighbouring site with a rate that depends on the number of vacancies $n$ in front of it. In the steady state, using the well known mapping of this model to the zero range process, we write down an exact formula for the partition function and the particle-particle correlation function in the canonical ensemble. In the thermodynamic limit, we find a simple analytical expression for the generating function of the correlation function. This result is applied to the hop rate $u(n)=1+(b/n)$ for which a phase transition between high-density laminar phase and low-density jammed phase occurs for $b > 2$. For these rates, we find that at the critical density, the correlation function decays algebraically with a continuously varying exponent $b-2$. We also calculate the two-point correlation function above the critical density, and find that the correlation length diverges with a critical exponent $ν=1/(b-2)$ for $b < 3$ and $1$ for $b > 3$. These results are compared with those obtained using an exact series expansion for finite systems.

cond-mat.stat-mech↗

arXiv2014

Length of adaptive walk on uncorrelated and correlated fitness landscapes

We consider the adaptation dynamics of an asexual population that walks uphill on a rugged fitness landscape which is endowed with large number of local fitness peaks. We work in a parameter regime where only those mutants that are single mutation away are accessible, as a result of which the population eventually gets trapped at a local fitness maximum and the adaptive walk terminates. We study how the number of adaptive steps taken by the population before reaching a local fitness peak depends on the initial fitness of the population, the extreme value distribution of the beneficial mutations and correlations amongst the fitnesses. Assuming that the relative fitness difference between successive steps is small, we analytically calculate the average walk length for both uncorrelated and correlated fitnesses in all extreme value domains for a given initial fitness. We present numerical results for the model where the fitness differences can be large, and find that the walk length behavior differs from that in the former model in the Fréchet domain of extreme value theory. We also discuss the relevance of our results to microbial experiments.

q-bio.PE↗

arXiv2014

Purifying selection, drift and reversible mutation with arbitrarily high mutation rates

Some species exhibit very high levels of DNA sequence variability; there is also evidence for the existence of heritable epigenetic variants that experience state changes at a much higher rate than sequence variants. In both cases, the resulting high diversity levels within a population (hyperdiversity) mean that standard population genetics methods are not trustworthy. We analyze a population genetics model that incorporates purifying selection, reversible mutations and genetic drift, assuming a stationary population size. We derive analytical results for both population parameters and sample statistics, and discuss their implications for studies of natural genetic and epigenetic variation. In particular, we find that (1) many more intermediate frequency variants are expected than under standard models, even with moderately strong purifying selection (2) rates of evolution under purifying selection may be close to, or even exceed, neutral rates. These findings are related to empirical studies of sequence and epigenetic variation.

q-bio.PE↗

arXiv2013

Adaptive walks and distribution of beneficial fitness effects

We study the adaptation dynamics of a maladapted asexual population on rugged fitness landscapes with many local fitness peaks. The distribution of beneficial fitness effects is assumed to belong to one of the three extreme value domains, viz. Weibull, Gumbel and Fr{é}chet. We work in the strong selection-weak mutation regime in which beneficial mutations fix sequentially, and the population performs an uphill walk on the fitness landscape until a local fitness peak is reached. A striking prediction of our analysis is that the fitness difference between successive steps follows a pattern of diminishing returns in the Weibull domain and accelerating returns in the Fr{é}chet domain, as the initial fitness of the population is increased. These trends are found to be robust with respect to fitness correlations. We believe that this result can be exploited in experiments to determine the extreme value domain of the distribution of beneficial fitness effects. Our work here differs significantly from the previous ones that assume the selection coefficient to be small. On taking large effect mutations into account, we find that the length of the walk shows different qualitative trends from those derived using small selection coefficient approximation.

q-bio.PE↗

arXiv2012

Fixation of mutators in asexual populations: the role of genetic drift and epistasis

We study the evolutionary dynamics of an asexual population of nonmutators and mutators on a class of epistatic fitness landscapes. We consider the situation in which all mutations are deleterious and mutators are produced from nonmutators continually at a constant rate. We find that in an infinitely large population, a minimum nonmutator-to-mutator conversion rate is required to fix the mutators but an arbitrarily small conversion rate results in the fixation of mutators in a finite population. We calculate analytical expressions for the mutator fraction at mutation-selection balance and fixation time for mutators in a finite population when mutational effects are weaker (regime I) and stronger (regime II) than the selective effects. Our main result is that in regime I, the mutator fraction and the fixation time are independent of epistasis but in regime II, mutators are rarer and take longer to fix when the decrease in fitness with the number of deleterious mutations occurs at an accelerating rate (synergistic epistasis) than at a diminishing rate (antagonistic epistasis). Our analytical results are compared with numerics and their implications are discussed.

q-bio.PE↗

arXiv2011

Number of adaptive steps to a local fitness peak

We consider a population of genotype sequences evolving on a rugged fitness landscape with many local fitness peaks. The population walks uphill until it encounters a local fitness maximum. We find that the statistical properties of the walk length depend on whether the underlying fitness distribution has a finite mean. If the mean is finite, all the walk length cumulants grow with the sequence length but approach a constant otherwise. Experimental implications of our analytical results are also discussed.

q-bio.PE↗

arXiv2011

Multiple adaptive substitutions during evolution in novel environments

We consider an asexual population under strong selection-weak mutation conditions evolving on rugged fitness landscapes with many local fitness peaks. Unlike the previous studies in which the initial fitness of the population is assumed to be high, here we start the adaptation process with a low fitness corresponding to a population in a stressful novel environment. For generic fitness distributions, using an analytic argument we find that the average number of steps to a local optimum varies logarithmically with the genotype sequence length and increases as the correlations amongst genotypic fitnesses increase. When the fitnesses are exponentially or uniformly distributed, using an evolution equation for the distribution of population fitness, we analytically calculate the fitness distribution of fixed beneficial mutations and the walk length distribution.

q-bio.PE↗

arXiv2011

Extreme value distributions for weakly correlated fitnesses in block model

We study the limit distribution of the largest fitness for two models of weakly correlated and identically distributed random fitnesses. The correlated fitness is given by a linear combination of a fixed number of independent random variables drawn from a common parent distribution. We find that for certain class of parent distributions, the extreme value distribution for correlated random variables can be related either to one of the known limit laws for independent variables or the parent distribution itself. For other cases, new limiting distributions appear. The conditions under which these results hold are identified.

cond-mat.stat-mech↗

arXiv2011

Nonlinear deterministic equations in biological evolution

We review models of biological evolution in which the population frequency changes deterministically with time. If the population is self-replicating, although the equations for simple prototypes can be linearised, nonlinear equations arise in many complex situations. For sexual populations, even in the simplest setting, the equations are necessarily nonlinear due to the mixing of the parental genetic material. The solutions of such nonlinear equations display interesting features such as multiple equilibria and phase transitions. We mainly discuss those models for which an analytical understanding of such nonlinear equations is available.

q-bio.PE↗

arXiv2011

Evolutionary advantage of small populations on complex fitness landscapes

Background: Recent experimental and theoretical studies have shown that small asexual populations evolving on complex fitness landscapes may achieve a higher fitness than large ones due to the increased heterogeneity of adaptive trajectories. Here we introduce a class of haploid three-locus fitness landscapes that allows to investigate this scenario in a precise and quantitative way. Results: Our main result derived analytically shows how the probability of choosing the path of largest initial fitness increase grows with the population size. This makes large populations more likely to get trapped at local fitness peaks and implies an advantage of small populations at intermediate time scales. The range of population sizes where this effect is operative coincides with the onset of clonal interference. Additional studies using ensembles of random fitness landscapes show that the results achieved for a particular choice of three-locus landscape parameters are robust and also persist as the number of loci increases. Conclusions: Our study indicates that an advantage for small populations is likely whenever the fitness landscape contains local maxima. The advantage appears at intermediate time scales, which are long enough for trapping at local fitness maxima to have occurred but too short for peak escape by the creation of multiple mutants.

q-bio.PE↗

arXiv2010

Evolutionary dynamics on strongly correlated fitness landscapes

We study the evolutionary dynamics of a maladapted population of self-replicating sequences on strongly correlated fitness landscapes. Each sequence is assumed to be composed of blocks of equal length and its fitness is given by a linear combination of four independent block fitnesses. A mutation affects the fitness contribution of a single block leaving the other blocks unchanged and hence inducing correlations between the parent and mutant fitness. On such strongly correlated fitness landscapes, we calculate the dynamical properties like the number of jumps in the most populated sequence and the temporal distribution of the last jump which is shown to exhibit a inverse square dependence as in evolution on uncorrelated fitness landscapes. We also obtain exact results for the distribution of records and extremes for correlated random variables.

q-bio.PE↗

arXiv2009

Time to fixation in the presence of recombination

We study the evolutionary dynamics of a haploid population of infinite size recombining with a probability $r$ in a two locus model. Starting from a low fitness locus, the population is evolved under mutation, selection and recombination until a finite fraction of the population reaches the fittest locus. An analytical method is developed to calculate the fixation time $T$ to the fittest locus for various choices of epistasis. We find that (1) for negative epistasis, $T$ decreases slowly for small $r$ but decays fast at larger $r$ (2) for positive epistasis, $T$ increases linearly for small $r$ and mildly for large $r$ (3) for compensatory mutation, $T$ diverges as a power law with logarithmic corrections as the recombination fraction approaches a critical value. Our calculations are seen to be in good agreement with the exact numerical results.

q-bio.PE↗

arXiv2009

Exact and limit distributions of the largest fitness on correlated fitness landscapes

We study the distribution of the maximum of a set of random fitnesses with fixed number of mutations in a model of biological evolution. The fitness variables are not independent and the correlations can be varied via a parameter $\ell=1,...,L$. We present analytical calculations for the following three solvable cases: (i) one-step mutants with arbitrary $\ell$ (ii) weakly correlated fitnesses with $\ell=L/2$ (iii) strongly correlated fitnesses with $\ell=2$. In all these cases, we find that the limit distribution for the maximum fitness is not of the standard Gumbel form.

cond-mat.stat-mech↗

arXiv2009

Exact phase diagram of quasispecies model with mutation rate modifier

We consider an infinite asexual population with a mutator allele which can elevate mutation rates. With probability $f$, a transition from nonmutator to mutator state occurs but the reverse transition is forbidden. We find that at $f=0$, the population is in the state with minimum mutation rate and at $f=f_c$, a phase transition occurs between a mixed phase with both nonmutators and mutators and a pure mutator phase. We calculate the critical probability $f_c$ and the total mutator fraction $Q$ in the mixed phase exactly. Our predictions for $Q$ are in agreement with those seen in microbial populations in static environments.

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