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Measuring inbreeding Measuring inbreeding Connection to population - - PowerPoint PPT Presentation

Causes of inbreeding depression Read Chapter 11 (pp 376-399) Reduced heterozygosity Alter, S. E., Rynes, E. and Palumbi, S. R. (2007) Increased exposure of recessive deleterious alleles in DNA evidence for historic population size and


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Alter, S. E., Rynes, E. and Palumbi, S. R. (2007) DNA evidence for historic population size and past ecosystem impacts of gray whales. PNAS 104: 15162–15167

Causes of inbreeding depression

Reduced heterozygosity Increased exposure of recessive deleterious alleles in homozygotes Recessive deleterious alleles are common in large

  • populations. These alleles are at low frequencies and

typically occur mainly in heterozygotes and are therefore not purged from the populations because there is no associated penalty for the heterozygotes. In small populations, just by chance, they might get fixed What is inbreeding? What is inbreeding depression? Why are small populations a problem? What is Mutation - Drift balance?

Read Chapter 11 (pp 376-399)

Measuring inbreeding

F = HExpected − Hobserved HExpected

for population we use the heterozygosity as a proxy

HExpected = 2pq using Hardy-Weinberg proportions and two alleles

F measures the deviation from a random mating population

Measuring inbreeding

F = He − Ho He

if Hois zero, F is maximal and 1 if Ho is equal to He, F is 0

Connection to population size

F = 1 − (1 − 1 2Ne )t

t = number of generations Probability that variability is maintained

  • ver t generations

Homozygosity Inbreeding coefficient

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Connection to population size

F = 1 − (1 − 1 2Ne )t Ne F t F F Time t Ne = 100 Ne = 1000 Ne = 10, 000 Ne = 100, 000

Connection to population size

F = 1 − (1 − 1 2Ne )t Ne F t F

Extinction vortex

The frequency of matings between close relative rises INBREEDING Heterozygosity is reduced in

  • ffspring, reducing the ability to

respond to environmental change Semilethal recessive alleles are expressed in homozygous conditions As a result of this expression, fecundity is reduced and mortality is increased (inbreeding depression) Population becomes smaller

“the worse it gets, the worse it gets.”

Caughley 1994 EXTINCTION

Predicting inbreeding depression

Scenario

What is the chance of suffering “Inbreeding depression” when mating with close relatives

Genetic load Large population High

(large number of rare semilethal alleles)

Low Recently small population Intermediate

(purged some detrimentals of large effect)

Intermediate

(fixed some detrimentals with small effect)

Long-term small population Low

(purged detrimentals of medium or large effect)

High

(fixed many detrimentals with small effect) Genetic load = Reduction of the mean fitness resulting from detrimental variation for a population compared to a population without lowered fitness.

Hedrick, P. W. 2001. Conservation Genetics: where are we now? Trends in Ecology and Evolution 16:629-636

Clarkia – a naturally self-pollinating plant that does not show inbreeding depression

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Heredity (1996) 76, 83–91; doi:10.1038/hdy.1996.11

Variation in inbreeding depression among families and populations of Clarkia tembloriensis (Onagraceae)

Timothy P Holtsford1

1Division of Biological Sciences, University of Missouri, Columbia, MO 65203, USA

Received 9 June 1995. Top

  • f page

Abstract The relationship between the self-fertilization rate of a population and the severity of inbreeding depression is difficult to predict because the underlying genetics may be complex. However, it is important to understand this relationship, and the degree to which inbreeding depression varies among families within populations, if we are to understand the evolution of plant mating systems. Inbreeding depression was studied in glasshouse trials using seed derived from two populations of Clarkia tembloriensis, Cantua Creek and Idria. These had very different rates of self-fertilization (s) and inbreeding coefficients (F) (s = 0.74, F = 0.77 in the CC-1 population; 5 = 0.16, F = 0.10 in the I-1 population). Outcrossing between these populations was equivalent to outcrossing within them; there was no evidence for outbreeding depression or increased heterosis from interpopulation

  • crosses. The more self-fertilizing CC-1 population had fewer recessive lethal genes than the
  • utbreeding I-1 population. However, cumulative inbreeding depression at the end of flowering was

not significantly different between these populations. Inbreeding has seemingly purged the lethal genes from the CC-1 population but overall mutational load, especially in characters that are manifest late in the life history, is still substantial despite a history of inbreeding. Variation in inbreeding depression among families within these two populations was surprisingly large. The range

  • f variation among families was five to seven times larger than the difference in inbreeding

depression between populations. Variation in inbreeding depression among families should make these populations more susceptible to the invasion of genes which increase the rate of self- fertilization.

Keywords: Clarkia, inbreeding depression, mating system, self-fertilization

Colonial naked mole rats do not show much inbreeding depression

VIRAL EPIZOOTIC REVEALS INBREEDING DEPRESSION IN A HABITUALLY INBREEDING MAMMAL

Adin Ross-Gillespie 1,2,3 , M. Justin O'Riain 1 , and Lukas F. Keller 4 1 Department of Zoology, University of Cape Town, Rondebosch 7700, South Africa 2 E-mail: A.Ross-Gillespie@ed.ac.uk 4 Zoologisches Museum, Universität Zürich, Winterthurerstrasse 190, CH-8057 Zürich, Switzerland Correspondence to 3 Present address: Institute of Evolutionary Biology, University of Edinburgh, West Mains Road, Edinburgh EH9 3JT, United Kingdom. Associate Editor: H. Kokko Correspondence to 3 Present address: Institute of Evolutionary Biology, University of Edinburgh, West Mains Road, Edinburgh EH9 3JT, United Kingdom. KEYWORDS Coronavirus • disease susceptibility • Heterocephalus glaber • inbreeding depression • purging ABSTRACT Inbreeding is typically detrimental to fitness. However, some animal populations are reported to inbreed without incurring inbreeding depression, ostensibly due to past "purging" of deleterious alleles. Challenging this is the position that purging can, at best, only adapt a population to a particular environment; novel selective regimes will always uncover additional inbreeding load. We consider this in a prominent test case: the eusocial naked mole-rat (Heterocephalus glaber), one of the most inbred of all free-living mammals. We investigated factors affecting mortality in a population of naked mole-rats struck by a spontaneous, lethal coronavirus outbreak. In a multivariate model, inbreeding coefficient strongly predicted mortality, with closely inbred mole-rats (F 0.25) over 300% more likely to die than their outbred counterparts. We demonstrate that, contrary to common assertions, strong inbreeding depression is evident in this

  • species. Our results suggest that loss of genetic diversity through inbreeding may render populations vulnerable to local extinction from

emerging infectious diseases even when other inbreeding depression symptoms are absent. Received January 29, 2007 Accepted May 8, 2007 DIGITAL OBJECT IDENTIFIER (DOI) 10.1111/j.1558-5646.2007.00177.x

Predicting inbreeding depression

Scenario

What is the chance of suffering “Inbreeding depression” when mating with close relatives

Genetic load Large population High

(large number of rare semilethal alleles)

Low Recently small population Intermediate

(purged some detrimentals of large effect)

Intermediate

(fixed some detrimentals with small effect)

Long-term small population Low

(purged detrimentals of medium or large effect)

High

(fixed many detrimentals with small effect) Genetic load = Reduction of the mean fitness resulting from detrimental variation for a population compared to a population without lowered fitness.

Hedrick, P. W. 2001. Conservation Genetics: where are we now? Trends in Ecology and Evolution 16:629-636

Inbreeding avoidance

Plants Self-incompatiblity Male and female parts flower at different times Heterostyly (females and male parts are far from each

  • ther)

Male and females are different plants Animals Mate choice (mice smell whether they belong to the same MHC type or not) Migration behavior (Young lions get driven from their pack)

Larkspur Delphinium nuttallianum

0.08 a 30 0.65 b 10 0.42 b 3 0.30 a 1 Overall fitness Distance between parents (m)

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Outbreeding depression

(=lower fitness due to breeding with unrelated individuals)

Local adaptation: differences in alleles frequencies due to

different selection pressures in different places Coadapted gene complexes: group of traits that have high fitness when they occur together, but low fitness when occurring with other

  • traits. Populations that are separated a long time were selected for

different combinations, bringing them together might be problematic: Largemouth bass Increased Infectious Disease Susceptibility Resulting from Outbreeding Depression

Authors: GOLDBERG, TONY L.; GRANT, EMILY C.1; INENDINO, KATE R.1; KASSLER, TODD W.

2; CLAUSSEN, JULIE E.3; PHILIPP, DAVID P.

Source: Conservation Biology, Volume 19, Number 2, April 2005 , pp. 455-462(8) Publisher: Blackwell Publishing Abstract: The mechanisms by which outbreeding depression leads to reduced fitness are poorly

  • understood. We considered the hypothesis that
  • utbreeding can depress fitness by increasing the

susceptibility of hybrid individuals and populations to infectious disease. Competitive breeding trials in experimental ponds indicated that outbred largemouth bass (Micropterus salmoides) crossed from two geographically and genetically distinct populations suffered a reduction in fitness of approximately 14% relative to parental stocks. We measured the comparative susceptibility of these same outbred stocks to a novel viral pathogen, largemouth bass virus. Following experimental inoculation, F2 generation hybrids suffered mortality at a rate 3.6 times higher than either F1 generation hybrids or wild-type parental fish. Analysis of viral loads indicated that viral replication was more rapid in F2 fish than in F1 hybrids or wild-type parental fish. We attribute these results to the disruption of coadapted gene complexes in the immune systems of outbred fish in the F2 generation. Increased susceptibility to infectious disease may be an important but underappreciated mechanism by which outbreeding reduces the fitness of individuals and populations and by which novel infectious diseases emerge in populations of hybrid organisms. Keywords: hybrids; largemouth bass virus; reduced fitness; híbridos; reducción de adaptabilidad; virus de la lobina Micropterus salmoides

Hybrids and Admixture

Two separated populations come together Individual of two different species have

  • ffspring

(sometimes fertile) Wahlund-effect: Heterozygote deficiency

Hybrid Zone

Bombina variegata Bombina bombina

Hybrids

Foto from Website of Beate Nürnberger, Edinburgh, now Munich (toads are anesthetized)

Hybridization, Admixture

Florida panther has a cowlick and a kinked tail because

  • f inbreeding

Hedrick, P. W. 2001. Conservation Genetics: where are we now? Trends in Ecology and Evolution 16:629-636

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Structured populations

Mainland Island Stepping Stone model Metapopulation

Subpopulation

Continuous populations

High density of individuals Some density of individuals Low density of individuals Low density of individuals

Isolation by distance: the genetic distance between individuals is correlated with geographic distance

Population (Genetics)

A population is a group of individuals that interbreed and that is somewhat isolated from other populations.

Species Population Population Population

Population versus Deme

A deme is a group of individuals that interbreed and that is somewhat isolated from other demes.

Species Deme Deme Deme Population