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Inbreeding and Heterosis

This document discusses inbreeding, heterosis, and their effects on genetic variation. 1. Inbreeding occurs when related individuals mate, often resulting in changes to trait means due to increased homozygosity. It can be intentionally practiced in breeding or occur unintentionally in small populations. 2. Crossing inbred lines (heterosis) leads to increased mean values in the offspring (F1) compared to the parental lines, known as hybrid vigor. This effect declines in subsequent generations as heterozygosity decreases. 3. Inbreeding increases variation between populations but decreases variation within populations. The magnitude of changes in trait means under inbreeding depends on gene frequencies and dominance effects.

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114 views

Inbreeding and Heterosis

This document discusses inbreeding, heterosis, and their effects on genetic variation. 1. Inbreeding occurs when related individuals mate, often resulting in changes to trait means due to increased homozygosity. It can be intentionally practiced in breeding or occur unintentionally in small populations. 2. Crossing inbred lines (heterosis) leads to increased mean values in the offspring (F1) compared to the parental lines, known as hybrid vigor. This effect declines in subsequent generations as heterozygosity decreases. 3. Inbreeding increases variation between populations but decreases variation within populations. The magnitude of changes in trait means under inbreeding depends on gene frequencies and dominance effects.

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© © All Rights Reserved
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Lecture 6:

Inbreeding and Heterosis


Inbreeding
• Inbreeding = mating of related individuals
• Often results in a change in the mean of a trait
• Inbreeding is intentionally practiced to:
– create genetic uniformity of laboratory stocks
– produce stocks for crossing (animal and plant
breeding)
• Inbreeding is unintentionally generated:
– by keeping small populations (such as is found at
zoos)
– during selection
Genotype frequencies under inbreeding

• The inbreeding coefficient, F


• F = Prob(the two alleles within an individual
are IBD) -- identical by descent
• Hence, with probability F both alleles in an
individual are identical, and hence a
homozygote
• With probability 1-F, the alleles are
combined at random
A1 A1
p A1 A1
F
q
A1 1-F A1 A2
p
Random
Alleles IBDmating
p A2 A1
q A2 1-F
q
F A2 A2
A2 A2

Genotype Alleles IBD Alleles not IBD frequency

A1A1 Fp (1-F)p2 p2 + Fpq


A2A1 0 (1-F)2pq (1-F)2pq

A 2A 2 Fq (1-F)q2 q2 + Fpq
Changes in the mean under inbreeding
Genotypes A1A1 A1A2 A2A2
0 a+d 2a

freq(A1) = p, freq(A2) = q

Using the genotypic frequencies under inbreeding, the


population mean mF under a level of inbreeding F is
related to the mean m0 under random mating by

mF = m0 - 2Fpqd
For k loci, the change in mean is
Xk
š F = š 0 ° 2F pi qi di = š 0 ° B F
i= 1

Here B is the reduction in mean under X


complete inbreeding (F=1) , where B= 2 pi qi di

• There will be a change of mean value dominance is present (d not zero)

• For a single locus, if d > 0, inbreeding will decrease the mean value
of the trait. If d < 0, inbreeding will increase the mean

• For multiple loci, a decrease (inbreeding depression) requires


directional dominance --- dominance effects di tending to be positive.

• The magnitude of the change of mean on inbreeding depends on gene


frequency, and is greatest when p = q = 0.5
Inbreeding Depression and Fitness
traits

Inbred Outbred
Define ID = 1-mF/m0 = 1-(m0-B)/m0 = B/m0
Drosophila Trait Lab-measured ID = B/m0
Viability 0.442 (0.66, 0.57, 0.48, 0.44, 0.06)
Female fertility 0.417 (0.81, 0.35, 0.18)
Female reproductive rate 0.603 (0.96, 0.57, 0.56, 0.32)
Male mating ability 0.773 (0.92, 0.76, 0.52)
Competitive ability 0.905 (0.97, 0.84)
Male fertility 0.11 (0.22, 0)
Male longevity 0.18
Male weight 0.085 (0.1, 0.07)
Female weight -0.10
Abdominal bristles 0.077 (0.06, 0.05, 0)
Sternopleural bristles -.005 (-0.001, 0)
Wing length 0.02 (0.03, 0.01)
Thorax length 0.02
Why do traits associated with fitness
show inbreeding depression?
• Two competing hypotheses:
– Overdominance Hypothesis: Genetic variance for fitness is
caused by loci at which heterozygotes are more fit than both
homozygotes. Inbreeding decreases the frequency of
heterozygotes, increases the frequency of homozygotes, so
fitness is reduced.
– Dominance Hypothesis: Genetic variance for fitness is
caused by rare deleterious alleles that are recessive or partly
recessive; such alleles persist in populations because of
recurrent mutation. Most copies of deleterious alleles in the
base population are in heterozygotes. Inbreeding increases
the frequency of homozygotes for deleterious alleles, so
fitness is reduced.
Estimating B
In many cases, lines cannot be completely inbred due to
either time constraints and/or because in many species
lines near complete inbreeding are nonviable

In such cases, estimate B from the regression of mF on F,


mF = m0 - BF
m0
mF
m0 - B

0 F 1

If epistasis is present, this regression is non-linear,


with CkFk for k-th order epistasis
mF

F
Minimizing the Rate of Inbreeding
• Avoid mating of relatives
• Maximum effective population size Ne
• Ne maximized with equal representation
– Contribution (number of sibs) from each parent as
equal as possible
– Sex ratio as close to 1:1 as possible
– When sex ratio skewed (r dams/sires ), every male
should contribute (exactly) one son and r daughters,
while every female should leave one daughter and
also with probability 1/r contribute a son
•
Line Crosses: Heterosis
When inbred lines are crossed, the progeny show an increase in mean
for characters that previously suffered a reduction from inbreeding.

This increase in the mean


P1over
x the
P2 average value of the
parents is called hybrid vigor or heterosis
F1
š P1 + š P 2
H F1 = š F 1 F°2
2
A cross is said to show heterosis if H > 0, so that the
F1 mean is average than the average of both parents.
Expected levels of heterosis
If pi denotes the frequency of Qi in line 1, let pi + di denote
the frequency of Qi in line 2.

The expected amount of heterosis becomes


Xn
H F1 = (±pi ) 2 d i
i= 1

• Heterosis depends on dominance: d = 0 = no inbreeding depression and no.


heterosis as with inbreeding depression, directional dominance is required for heterosis.

•H is proportional to the square of the difference in gene frequency


Between populations. H is greatest when alleles are fixed in one population and
lost in the other (so that | di| = 1). H = 0 if d = 0.

• H is specific to each particular cross. H must be determined empirically,


since we do not know the relevant loci nor their gene frequencies.
Heterosis declines in the F2
In the F1, all offspring are heterozygotes. In the F2,
Random mating has occurred, reducing the frequency
of heterozygotes.

As a result, there is a reduction of the amount of


heterosis in the F2 relative to the F1,

š P 1 + š P2 (±p) 2 d HF 1
HF 2 = š F 2 ° = =
2 2 2
Since random mating occurs in the F2 and subsequent
generations, the level of heterosis stays at the F2 level.
Agricultural importance of heterosis
Crosses often show high-parent heterosis, wherein the
F1 not only beats the average of the two parents
(mid-parent heterosis), it exceeds the best parent.

Crop % planted % yield Annual Annual Annual land


as hybrids advantage added added savings
yield: % yield: tons
Maize 65 15 10 55 x 106 13 x 106 ha

Sorghum 48 40 19 13 x 106 9 x 106 ha

Sunflower 60 50 30 7 x 106 6 x 106 ha

Rice 12 30 4 15 x 106 6 x 106 ha


Variance Changes Under Inbreeding
reduces variation
Inbreeding increases withinbetween
the variation each population
populations
(i.e., variation in the means of the populations)

F = 1/4
0
3/4
1
Variance Changes Under Inbreeding

General F=1 F=0

Between lines 2FVA 2VA 0


Within Lines (1-F) VA 0 VA
Total (1+F) VA 2VA VA
Mutation and Inbreeding
• Eventually these
As lines lose two forces
genetic balance,
variation leading
from drift, to
mutation
an equilibrium
introduces newlevel of genetic variance reflecting
variation
the balance between loss from drift, gain from mutation
VM = new mutation variation each generation, typically
VM = 10-3 VE
Assuming:
Strictly neutral mutations
Strictly additive mutations
Symmetrical distribution of mutational effects

VA = VG = 2N e VM
Between-line Divergence
The between-line variance in the mean (VB) in generation
t is

° t =2 N e
VB = 2VM [ t ° 2N e (1 ° e )]
For large t, the asymptotic rate is 2VMt

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