Uneven distribution of progeny Uneven distribution of progeny Red - - PDF document

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Uneven distribution of progeny Uneven distribution of progeny Red - - PDF document

Uneven distribution of progeny Uneven distribution of progeny Red Drum (redfish) N e = 4 N c 2 2 N c N e 2 + 2 0 100 199 Family Sciaenidae, DRUMS Sciaenops ocellatus 1 100 133 2 variance of family size Description: chin


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SLIDE 1

100 199 1 100 133 2 100 100 5 100 57 10 100 33

Uneven distribution of progeny

σ2

variance of family size

Nc

census size

σ2 Nc Ne

Ne = 4Nc − 2 σ2 + 2

σ2

random mating = 2(1 − 1/Nc)

Uneven distribution of progeny

Red Drum (redfish)

Family Sciaenidae, DRUMS Sciaenops ocellatus

Description: chin without barbels; copper bronze body, lighter shade in clear waters; one to many spots at base of tail (rarely no spots); mouth horizontal and openng downward; scales large. Similar Fish: black drum, Pogonias cromis. Where found: juveniles are an INSHORE fish, migrating out of the estuaries at about 30 inches (4 years) and joining the spawning population OFFSHORE. Size: one of 27 inches weighs about 8 pounds. *Florida Record: 51 lbs., 8 ozs. Remarks: red drum are an INSHORE species until they attain roughly 30 inches (4 years), then they migrate to join the NEARSHORE population; spawning occurs from August to November in NEARSHORE waters; sudden cold snaps may kill red drum in shallow, INSHORE waters; feeds on crustaceans, fish and mollusks; longevity to 20 years

  • r more.

Turner, T. F., J.P. Wares, and J.R. Gold. 2002. Genetic Effective Size is Three Orders of Magnitude lower than Adult Census Size in an Abundant, Estuarine Dependent Marine Fish (Sciaenops ocellatus). Genetics

Ne = 1854 Nc = 3, 400, 000

Genetic drift and mutation balance

With mutations a random mating population of diploids has the chance to acquire 2N new alleles every generation. A population looses variability at a rate of

1 2Ne ∆H = µ − 1 2Ne mutation rate loss rate ∆H = µ − 1 2Ne

Small population loose alleles faster than they arrive in by mutation Small population are not at mutation-drift equilibrium. Heterozygosity H will decrease over time

Small populations

Relationship between Heterozygosity and Population size

Red-cockaded woodpecker

Founder effect

Locality Heterozygosity Expected Heterozygosity 8 Selçuk 0.114 0.132 13 Samos 0.097 0.119 14 Ikaria 0.042 0.050

13

When only few individuals from a large population colonize an island (or other isolated habitat), only a small number of alleles will be present in the new population.

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SLIDE 2

Reduction of population size

Genetic drift (loss of alleles is larger than gain of alleles) Founder effects Bottlenecks Inbreeding

Inbreeding

Breeding with close relatives

Inbreeding depression

Breeding with close relatives

Reduced heterozygosity and increased mortality of offspring caused by mating of close relatives

Disease susceptibility in California sea lion

Figure 2 Internal relatedness in sea lions and the incidence of different disease classes. Carcinoma, n = 13; helminth infection, n = 72; nonspecific, n = 51; bacterial infection, n = 98; algal toxin, n = 101; trauma (control), n = 36. Values are means s.e. The mean internal relatedness value of trauma animals (-0.004) is indistinguishable from zero, as would be expected for individuals born to randomly mated parents5.

Experiment showing inbreeding in a cricket

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SLIDE 3

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

Measuring inbreeding

AB CD

Dad Mom Daughter

Measuring inbreeding

Dad Mom Daughter

AB CD

Son

What is the probability that the son got two copies of the same allele from his dad? = inbreeding coefficient F

Measuring inbreeding

AB CD p(A)=0.5 p(A)=0.5 p(A)=0.5 p(AA) = 0.5 * 0.5 * 0.5 = 0.125

Measuring inbreeding

AB CD p(A)=0.5 p(A)=0.5 p(A)=0.5 p(AA) = 0.5 * 0.5 * 0.5 = 0.125 p(BB) = 0.5 * 0.5 * 0.5 = 0.125 F = p(AA) + p(BB) = 0.125 + 0.125 = 0.25 p(B)=0.5 p(B)=0.5 p(B)=0.5

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

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SLIDE 4

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

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

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SLIDE 5

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

Colonial naked mole rats do not show much inbreeding

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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SLIDE 6

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

Hybrids and Admixture

Two separated populations come together Individual of two different species have

  • ffspring

(sometimes fertile)

Hybrid Zone

Bombina variegata Bombina bombina

Hybrids

Foto from Website of Beate Nürnberger, Edinburgh

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