Population Genetics and Conservation

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							Population Genetics and
     Conservation
  Is there a relationship between
 genetic diversity and population
           size in nature?
Frankham, R. 1996. Relationship of genetic
   variation to population size in wildlife.
    Conservation Biology 10:1500-1508

For 77 animal, plant, and bacterial species
        with a minimum of 20 loci
 Relationship between Genetic
 Diversity and Population Size

                            r = 0.81
                            p < 0.001

                            r = 0.73 with
He                          E. coli
                            omitted




            log N
Is there a relationship between
    reproductive success and
     genetic diversity in nature?
  Reed, D.H. and R. Frankham. 2003.
Correlation between Fitness and Genetic
Diversity. Conservation Biology 17:230-
                  237
 There are several reasons why there may be a
   weak or nonexistent relationship between
     fitness and levels of genetic diversity
1)   Because molecular markers are neutral, or nearly so, they
     may lose genetic variation more rapidly than loci concerned
     with fitness
2)   Quantitative traits associated with fitness typically have a
     larger proportion of their total genetic variance in the form of
     epistatic and dominance variance than in the form of traits
     less closely associated with fitness – heritabilities can remain
     high in spite of reductions in population size
3)   Selection tends to purge the population of deleterious
     recessive alleles and in theory can create inbred populations
     with a higher fitness than their outbred progenitor
 Positive Relationship between
 Fitness and Genetic Diversity




Overall - X r = 0.432 + 0.058 or 19% of the variation explained
             28 of 34 comparisons were positive
Correlates of Fitness
     Population Size and Fitness
      X = 0.354 + 0.111, N=11
       Heritability and Fitness
       X = 0.509 + 0.134, N=6
 Molecular Heterozygosity and Fitness
      X = 0.447 + 0.081, N=17
Their Conclusion
     Not only does the level of
      heterozygosity relate to
     evolutionary potential, but
    validates its correlation with
     current population fitness
 Most models of the rate of loss of genetic
variation in small populations assume that
      all genotypes have equal fitness
  (i.e., selective neutrality) based on our
        neutral model of genetic drift

                 (1-1/2Ne)t
   where an Ne=50 is necessary to avoid
harmful loss of genetic diversity in the short
                    term
However, some empirical studies
have shown that selection against
homozygotes occurs during early
  stages of growth in natural
      populations of plants
Consequently, models that assume
   selective neutrality may be
           misleading
    Lesica, P. and F.W. Allendorf.
     1991. Are small populations of
       plants worth preserving?
      Conservation Biology 6:135-
                  139.
  They searched the literature for
multilocus studies of plants in which
genotypic frequencies at two or more
    stages in the life cycle were
         reported; found 8
   Data suggest that heterozygotes often
    have a survival advantage that could
   affect the rate of loss of heterozygosity
Three possible explanations
for heterozygote advantage:
1) Inbreeding depression - unmasking
   deleterious recessives
2) Overdominance - greater fitness of
   heterozygotes at the loci examined
3) Associative Overdominance -
   Selection at loci that are linked or
   nonrandomly associated with the loci
   examined
 Whatever the cause, heterozygous
advantage slows the loss of variation
due to drift over what neutral models
                predict

               1 t
     Ht = (1 - 2N ) Ho
 Heterozygosity remaining after 25
generations; Heterozygote fitness is
       1.0, homozygotes 1-s
     100
      90
      80
      70                                  100
      60
%                                         50
      50
                                          25
                                                N
Ht    40
                                          10
      30
      20
      10
       0

       0.00   .02    .05   .10     0.25
           Selection Coefficient
Population size that would lose
genetic variation at a selectively
neutral locus at the same rate as
  observed in the simulations
      ____Population Size_______
  s       10   25       50    100

  0.00    9.7 25.9     49.6   100.4
  0.02   10.4 29.5     63.4   129.6
  0.05   11.4 34.1     81.0   162.6
  0.10   14.8 52.7    116.2   242.9
  0.25   31.3 145.1   361.6   718.7
    Can conclude that small
 populations may be much more
 valuable for conservation than
predicted by models that assume
       selective neutrality
Effective Population Size
Made the assumption that the
 number of males and females
contributing to each subsequent
    generation is the same
If the sex ratio is not 1:1 for each
  generation then the population
   loses genetic variability more
               rapidly
This is because the “effective number” of
   individuals is smaller than the actual
  number of individuals in the population
 The Effective Number (Ne) is the
 number of individuals in an ideal
population that would lose genetic
  variability at the same rate as a
    non-ideal population with N
             individuals
Effective Number can be
calculated as follows:

  Effective Number
                     # breeding
                       females in pop.

    Ne = 4Nm Nf
         Nm + Nf

            # breeding males in pop.
For a sex ratio of 1 male:9 females
  in a population of 100 animals


     Ne = 4(10 X 90)
                     = 36
           10 + 90
Which means that a population of 100
individuals, consisting of 10 breeding
males and 90 breeding females would
lose genetic variability as rapidly as a
   population consisting of only 18
      males and 18 females or 36
              individuals
   Influence of fluctuating
population size on the effective
           number
     e.g., if a population of 100
    individuals drops to only 25 in
       the tenth generation the
    effective number during these
     10 generations would be 77
Recall : Harmonic Mean


          individuals in each generation

  1   1
        ( 1/N1 + 1/N2 +
  N =
        .....1/Nt)
  e   t
Effective # generations
Number
From this example it’s clear
that a single generation with a
low population size has a
large negative influence on
the effective number
Influence of family size on the
effective number
     Actual # of breeding individuals


          4N - 4
     Ne = Vk + 2

   Effective Variance in number of
   number Offspring
Rearrange equation
   Ne/N ~ 4/(Vk + 2)

   Ne/N ~ 4/(2+2) = 1.0 ~ N

   Ne/N ~ 4/(4+2) = 0.67N

   Ne/N ~ 4/(0+2 ) = 2.0N
     Over a range of species,
variation in family sizes reduced
 effective population sizes to an
 average of 54% of census sizes
     Frankham, R. 1995. Effective
    population size/adult population
     size ratios in wildlife: a review.
     Genetic Research 66:95-107.
The influence of generation time
on loss of genetic variability -
loss doesn’t occur per year, per
decade or per century but per
generation!
   Loss of Genetic          Management
   Variability Influenced   Measures
   by                       to Reduce Loss

1. Size of founder           Maximize # of
   population                  founders
2. Rate of growth of         Maximize growth rate,
   population after            especially in the
   foundation                  first generation
3. Sex ratio                 Equalize # males
                               females
4. Generation length         Maximize
5. Family size             Equalize # offspring/
                             breeding individual
6. Fluctuation of          Minimize fluctuations
  population size over
  generations
7. Size of the stable      Maximize carrying
  population (@ carrying    capacity
  capacity)

						
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