mating system evolution...evolution of selfing: • selfing poplns thought to be evolved from...
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Mating System Evolution
Inbreeding and Outbreeding
Lecture 8: Spring 2013
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Mating system = who you mate with
• Outcrossing rate = t
• Ranges from 0 to 1.0 where 0 is total
obligate selfing and 1.0 is obligate
outcrossing.
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Mating system & breeding system of
plants & animals
Animals:
Mobility
Most dioecious, but some hermaphroditic (still require
mate), fission, and parthenogenetic
Plants:
Lack mobility
Many hermaphroditic
selfing -----> outcrossing -----> dioecy
Breeding system = Attributes of the flowers within an individual that may influence gamete transfer among conspecifics
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Evolution of Mating Systems
Breeding System Variation in Plants
Selfing -------------------- Outcrossing
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Evolution of selfing:
• Selfing poplns thought to be evolved from outcrossing
poplns
• Selfing has evolved repeatedly in plant kingdom
• Darwin’s early observation in 1876
“Nature abhors self-fertilization”
• A lot of theory but not a lot of empirical data on many of the
hypotheses for the evolution of selfing
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Inbreeding depression = relative performance of progeny
resulting from matings between relatives versus
between two unrelated individuals
Fitness = the quantity and quality of progeny produced
by an individual.
Selfing or self mating = a decline in heterozygosity by 50%
each generation selfing occurs.
The most extreme form of inbreeding.
Terminology
Outcrossing or outcrossed mating= mating between
unrelated individuals
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Issues in Conservation and Restoration Biology
Anthropogenic:
• Land development
• Over exploitation
• Species Translocations
• Pollution
Ecological:
• Environmental fluctuations
• Environmental catastrophes
• Fragmentation
• Meta-population dynamics
• Allele and edge effects
• Small effective number (Ne) (Lande 1997)
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Issues in Conservation and Restoration Biology
Genetic:
• Maladaptive hybrids
• Selective breeding and harvesting
• Small population size
• Decreased genetic variation
• Inbreeding depression
• Fixation of new mutations
(Lande 1997)
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Mating System Variation:
Animals and Plants
0.4
0.1
0.3
0
0.2
0-0.2 >0.2-0.4 >0.4-0.6 >0.6-0.8 >0.8-1
Selfing ------- Mixed-mating------ Outcrossing
Animal Plant
(Redrawn from Jarne & Auld 2006)
Pro
po
rtio
n o
f S
pecie
s
9
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Evolution of Mating Systems
Ecological Advantages to Selfing:
Pollination/reproductive assurance (Darwin 1876)
If pollinators become rare
Survival at low population densities
Promotion of local adaptation
Lower costs of reproduction
Colonizing ability: long distance colonization (Baker 1955)
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Reproductive Assurance
(1) Reproductive assurance (RA)
– 1st proposed by Darwin 1876
– Initially viewed as a rare event, favored by natural
selection under special environmental conditions
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Reproductive Assurance or
enhancement of gamete transmission?
(Lloyd 1979)
• Prior selfing - ~ to complete selfing
• Competing selfing –
– self and outcross pollen compete for the same ovules
– Pollen equally effective
• Delayed selfing – increase opportunity for
outcrossing first, then selfing allowed if all ovules
not fertilized
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RA Hypotheses
(2) Lack of pollinators
– Darlington 1939 – autogamy arises under environmental
conditions occur that prevent effective pollination by
normal vectors
– Hagerup 1951 – suggested that extra moisture in Northern
oceanic climates limit good weather days for flying by
insects
– Barrett 1980s – tristyly found in continental Brazil but a
selfing morph found on Jamiaca at edge of range of
Eichhornia paniculata
https://encrypted-tbn0.gstatic.com/images?q=tbn:ANd9GcQxLT64_NAFGVd3iXrsxgkFDOQmFN-
sNjQzqT4tpNBWyJDIk1iN
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RA Hypotheses
(3) Long distance colonization (Baker’s rule 1955)
– Single propagule following long distance dispersal will
succeed only if individual is autogamous thus allowing a
new population to establish
– Autogamy is a characteristic of a successful colonizing
species (Baker and Stebbins 1965; Stebbins 1970)
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RA Hypotheses
(4) Competition for limited # of pollinator vectors
among plant species
– Grant and Grant 1965
– May be more intense for SI species (Levin 1972)
– May expect a divergence in flowering phenology
between two competing species for pollinator
visits
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RA Hypotheses
(5) Local adaptation
– Selection for autogamy arises when environmental
conditions allow only a narrow range of genotypes
to survive or reproduce (i.e., following a
bottleneck) Mather 1953;1973
– The gain in immediate fitness via an increase in
local adaptation might overcompensate for the
decrease in long-term fitness through a loss of
genetic flexibility (Jain 1976)
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RA Hypotheses
(6) Restrict gene flow
– Autogamy arises as a mechanism to restrict gene flow
between closely related sympatric species
– Restricted gene flow allows for population differentiation at
a local level between adjacent populations. (Antonovics,
J. 1968. Evolution in closely adjacent plant populations. V. Evolution
of self-fertility. Heredity 23: 219-238. )
– Genetic neighborhood small (Fenster 1991 a, b; Evolution)
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RA Hypothesis
(7) Lower cost of reproduction – shunt resources for
larger flowers to other traits that might improve
survival.
https://encrypted-tbn0.gstatic.com/images?q=tbn:ANd9GcQxLT64_NAFGVd3iXrsxgkFDOQmFN-
sNjQzqT4tpNBWyJDIk1iN
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Selfing Mimulus micranthus
Mixed-mating Mimulus guttatus
However, a recent review has noted that many plant species
with specialize pollination systems also exhibit
Reproductive Assurance, since there is often more
variation in pollinator service
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Fenster & Marten-Rodriguez 2007 (former BEES student) Int. J. Plant Sci. 168: 215-228.
Reproductive assurance of specialized flowers:
delayed selfing
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RA Hypotheses
(8) Automatic selection hypothesis
– Fisher 1941
– Gene for autogamy spreads throughout an
outcrossing population without significant inbreeding
depression (>50%) and no pollen discounting.
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Gene Contribution to the Next Generation
Automatic selection hypothesis (Fisher 1941)
50% transmission advantage of selfing
Genetic Advantage to selfing
Outcross Self
Maternal seed 0.5 1.0
Sired seed 0.5 0.5
Total 1.0 1.5
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Evolution of Mating Systems
Ecological Advantages to Selfing:
Reproductive Assurance Hypothesis (Darwin 1876)
Genetic Advantage of Selfing:
50% Genetic Transmission Advantage (Fisher 1941)
https://encryptedtbn1.gstatic.com/images?q=tbn:ANd9GcQWSQ
cStDSNqC_0mtCOJRbFzORyjoT9SzauCNSR2fSdN9UFR_1k
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Why aren’t all populations
selfing or inbreeding?
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Why aren’t all populations
selfing or inbreeding?
Inbreeding Depression
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Gene Contribution to the Next Generation
Automatic selection hypothesis (Fisher 1941)
50% transmission advantage of selfing
Genetic Advantage to selfing
Outcross Self
Maternal seed 0.5 1.0
Sired seed 0.5 0.5
Total 1.0 1.5
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Thus:
• If only outcross, one contributes 50% of genes
in both their pollen and ovules compared to
another individual who can both self and
outcross, having a 1.5 advantage.
• If Inbreeding depression > 50% then
outcrossing maintained
• If inbreeding depression < 50% then selfing
may evolve
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Evolution of Mating Systems
Ecological Advantages to Selfing:
Reproductive Assurance Hypothesis (Darwin 1876)
Genetic Advantage of Selfing:
50% Genetic Transmission Advantage (Fisher 1941)
Genetic Disadvantage of Selfing
Inbreeding Depression (Darwin 1876)
https://encryptedtbn1.gstatic.com/images?q=tbn:ANd9GcQWSQ
cStDSNqC_0mtCOJRbFzORyjoT9SzauCNSR2fSdN9UFR_1k
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Issues with respect to expression of
inbreeding depression
• Population level
• Individual level
• Species level
• Role of environment on detection of inbreeding
depression?
• Abiotic and biotic factors?
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• Genetic basis of inbreeding depression?
Partial Dominance (Dominance) vs. Overdominance
AA Aa aa vs. AA Aa aa
• Mating system?
Outcrossing Species vs. Selfing Species
M. guttatus M. micranthus
What factors determine whether a population
can purge its genetic load?
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Evolution of Mating Systems
Genetic Basis
• Dominance
• Overdominance
Purging of Genetic Load
• Selfer >> Outcrosser
• Selfer = Outcrosser
The relationship between mating system
and the genetics of inbreeding depression
Carr, D.E., and M.R. Dudash. 2003. Recent approaches into the genetic basis of inbreeding
depression in plants. Philosophical Transactions of the Royal Society of London Series B. 358:
1071-1084.
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Glasshouse Inbreeding Depression
Studies: Tests of Purging
1) Comparison of inbreeding depression in selfing M.
micranthus and mixed-mating M. guttatus.
2) Consequences of 5 generations of serial inbreeding
in M. guttatus across the entire lifespan.
3) Quantifying the level of dominance responsible for
the expression of inbreeding depression in M.
guttatus and M. micranthus.
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0
0.2
0.4
0.6
0.8
1
1.2
1.4
M. guttatus M. micranthus
Self Outcross
Bio
mass
*
*
Carr and Dudash (AJB 1996)
1st Approach:
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0
200
400
600
800
1000
1200
1400
1600
M. guttatus M. micranthus
Self
Outcross*
Carr and Dudash (AJB 1996)
0
50
100
150
200
250
300
M. guttatus M. micranthus
Self
Outcross
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Relative
Performance
In Flower Production
Dudash et al. Evolution 1997
2nd Approach:
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(Dudash & Carr Nature1998)
Genetic basis of inbreeding depression:
Can genetic load be purged?
M. guttatus M. micranthus
3rd Approach:
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• Genetic basis of inbreeding depression?
Partial Dominance (Dominance) : AA Aa aa
• Mating system?
Selfing Species has Lower Inbreeding Depression
than Outcrossing Species
M. guttatus M. micranthus
What factors determine whether a population
can purge its genetic load?
(Carr & Dudash 1996 AJB, Carr & Dudash 1997 Evolution, Dudash et al.1997 Evolution, Carr et al. 1997 Evolution,
Dudash & Carr Nature1998)
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Factors that influence ability to purge
• Genetic basis of inbreeding depression (dominance vs.
overdominance)
• Natural selection reduces frequency of deleterious alleles
• Effects of prior inbreeding in reducing effects of new
inbreeding
• Impact of new mutations
• Role of environment in expression of inbreeding depression
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Mating System Variation: Animals and Plants
0.4
0.1
0.3
0
0.2
0-0.2 >0.2-0.4 >0.4-0.6 >0.6-0.8 >0.8-1
Animal Plant
(Lande & Schemske 1985, Redrawn from Jarne & Auld 2006)
Pro
po
rtio
n o
f S
pecie
s Why mixed mating?
Selfing ------- Mixed-mating------ Outcrossing
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How can mixed-mating strategies
be maintained in nature?
• Balance of immediate & long-term fitness (Mather 1943)
• Ecological conditions (Stebbins 1950)
• Loss of local adaptation in a novel habitat (Ronce et al. 2009)
• Environment dependent inbreeding depression (Dudash
1990, Goodwillie et al 2005, Jarne & Auld 2006, Cheptou & Donohue 2011, Winn et al. 2011)
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Selfing and subsequent inbreeding
depression in Animals
~ 1/3 of all animals capable of some form of selfing ~
hermaphrodites
Inbreeding avoidance mechanisms?
Prezygotic -
Postzygotic -
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Animals with Hermaphroditic phyla (Jarne & Auld 2006)
• Molluscs
• Trematodes
• Cnidarians
Hydras, anemones ,jelly fish, corals
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THE INBREEDING COEFFICIENT (F) The inbreeding coefficient of an individual refers to how closely related its parents are.
When parents are unrelated, offspring F = 0, for completely inbred individuals F = 1.
Levels of inbreeding in offspring for different kinds of relationships among parents are:
______________________________________________________________
Parents Offspring F
______________________________________________________________
Unrelated 0
Brother-sister, mother-son, or father-daughter 0.25
Half brother-half-sister (half sibs) 0.125
First-cousins 0.0625
Second- cousins 0.0156
Self-fertilization (or selfing) ~ Plants 0.5
______________________________________________________________
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Table Inbreeding depression for different components of fitness in wild
species of animals with a 25% increase in inbreeding coefficient. Inbreeding
depression is % reduction of inbred compared to outbred individuals.
Species Character Inbreeding depression %
Deer mice
Litter size 15
Survival to weaning 8
House mice (wild)
Litter size 10
Body wt at 53 days -10
Nesting behaviour 10
Japanese quail
Reproduction and survival 64
Fertility 21
Survival 0-5 wks 10
Body weight 4
Chukar partridges
Reproduction and survival 58
Egg production 16
Body weight 1
Rainbow trout
Hatchability -10, 9, 14
Fry survival 8, 11
Wt at 150 days 12
Zebra fish
Hatchability 89
Survival to 30 days 43
Length at 30 days 11
Channel catfish
Hatchability -11
Body weight at 4 wks 43
Body weight at 12 wks 7
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Sabatia angularis (Gentianaceae)
Male fn
Herkogamy and Dichogamy
Female fn
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1.6
1.8
2
2.2
2.4
2.6
2.8
Greenhouse Garden Field
Environment
Fit
ne
ss
of
ou
tcro
ss
/se
lfInbreeding depression measured in three environments
in Sabatia angularis (Gentianaceae)
Dudash (Evolution 1990) The environment matters!
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Outbreeding depression
Definition:
offspring from crosses between individuals from
different populations have lower fitness than
progeny from crosses between individuals from
the same population
Opposite of inbreeding depression
Optimal outcrossing distance – Price & Waser 1979
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Outbreeding depression
3 Mechanisms proposed to generate outbreeding depression:
1) Chromosomal differences
Post zygotic vs. prezygotic isolating mechanisms
2) Adaptive differentiation among populations
3) Population bottlenecks and genetic drift
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Investigations of inbreeding vs.
outbreeding depression
Inbreeding depression: 4,204 publications
Outbreeding depression: 731 publications
Why are we interested in the expression of outbreeding depression?
March 7, 2012 ISI web of science search
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Genetic Rescue – outcrossing to
reduce effects from inbreeding
• Increases genetic diversity
• Observed subsequent fitness increases of small
isolated inbred populations
Fear of outbreeding depression is preventing
genetic management of fragmented populations
(Frankham et al. 2011)
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Genetic rescue from crosses between different poplns:
Mammals
African elephant, lion and wild dog
Black rhinoceros
Bighorn sheep
Columbia Basin pygmy rabbit
Florida panther
Golden lion tamarin
Mexican wolf
Birds
Greater prairie chicken in Illinois
Red-cockaded woodpecker
Reptile
Swedish adder
Plants
Button wrinklewort
Lakeside daisy population in Illinois
Marsh grass of Parnassus
Mauna Kea silversword (Hawaii)
Brown’s banksia
Round leafed honeysuckle
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Role of inbreeding & drift in Prairie Chickens
https://encrypted-tbn0.google.com/images?q=tbn:ANd9GcRp4RjED7FlH7bkAC-
93Wxj16wsdaYZHOr4B_bJ_zom_3IC1_IU
2009 update – dark green current
and light green pre-settlement
Illinois population decline
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Prairie Chicken populations
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Prairie Chicken populations 2009 update
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Gene flow augmentation plant studies (Frankham et al. 2011)
Button Wrinklewort Rutidosis
leptorrhynchoides
Pickup and Young 2007
Lakeside Daisy Hymenoxys acaulis var.
glabra
DeMauro 1993
Marsh grass of
Parnassus
Parnassia palustris Bossuyt 2007
Mauna Kea silversword Agryoxiphium
sandwicense ssp.
sandwicense
Robichaux et al 1997
Browns’ banksia Banksia brownii Barrett and Jackson
2008
Round leafed
honeysuckle
Lambertia orbifolia Coates et al 1998
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Why is genetic rescue so important?
• What factors are contributing to the
need for genetic rescue?
• Are all local populations (~demes) of a
species similar?
• Examples from our readings?
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3. Gene flow
between
populations in
last 500 yrs?
4. Significant
environmental
differences?
5. Populations
separated
> 20
generations?
Re-establish
gene flow
Keep
populations
separated
Yes
No
No
No
Yes
Decision tree
for considering risk
Outbreeding
Depression 2. Fixed chromosomal differences?
Yes
No
Evaluate risks
in more detail
Yes
1. Is taxonomy resolved?
Yes
Resolve
No
Frankham et al. 2011
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Highlights
• Variability in both plant and animal mating systems
• Evolution of selfing from outcrossing
• Reproductive assurance common in nature
• Transmission bias associated with self matings
• Inbreeding depression - major force preventing evolution of
selfing
• Purging of genetic load
• Inbreeding in nature and factors that influence expression
of inbreeding depression
• Many species can benefit from crossing between
populations with little fear of long term effects of
outbreeding depression ~ Genetic Rescue