So far in this course we have dealt entirely with the evolution of characters that are controlled by simple Mendelian inheritance at a single locus. There are notes on the course website about gametic disequilibrium and how allele frequencies change at two loci simultaneously, but we didn’t discuss them. In every example we’ve considered we’ve imagined that we could understand something about evolution by examining the evolution of a single gene. That’s the domain of classical population genetics. For the next few weeks we’re going to be exploring a field that’s actually older than classical population genetics, although the approach we’ll be taking to it involves the use of population genetic machinery. If you know a little about the history of evolutionary biology, you may know that after the rediscovery of Mendel’s work in 1900 there was a heated debate between the “biometricians” (e.g., Galton and Pearson) and the “Mendelians” (e.g., de Vries, Correns, Bateson, and Morgan). Biometricians asserted that the really important variation in evolution didn’t follow Mendelian rules. Height, weight, skin color, and similar traits seemed to

Human biochemical genetics

The plant allozyme literature was reviewed to: (1) compare genetic diversity in long-lived woody species with species representing other life forms, and (2) to investigate whether the levels and distribution of genetic diversity in woody species are related to life history and ecological characteristics. Data from 322 woody taxa were used to measure genetic diversity within species, and within and among populations of species. Woody species maintain more variation within species and within populations than species with other life forms but have less variation among populations. Woody species with large geographic ranges, outcrossing breeding systems, and wind or animal-ingested seed dispersal have more genetic diversity within species and populations but less variation among populations than woody species with other combinations of traits. Although life history and ecological traits explain a significant proportion (34%) of the variation among species for the genetic parameters measured, a large proportion of the interspecific variation is unexplained. The specific evolutionary history of each species must play an important role in determining the level and distribution of genetic diversity.

Factors influencing levels of genetic diversity in woody plant species

Multilocus DNA markers [random amplified polymorphic DNA (RAPDs), amplified fragment length polymorphism (AFLPs)] are important for population studies because they reveal many polymorphic loci distributed over the genome. The markers are dominant, that is two phenotypes are distinguished at each locus, with a band and with no band. The latter one represents null-homozygotes with unamplified, recessive null-alleles. The frequency of a null-allele can be estimated by taking the square root of the fraction of individuals with no band. Lynch and Milligan (1994) have suggested a modified procedure that reduces bias introduced by the square-root transform. However, the procedure recommends to ignore those samples in which fewer than four null-homozygotes are observed. This may lead to significant bias in estimates of genetic diversity. In this study, I introduce a Bayesian approach to estimation of null-allele frequencies for dominant DNA markers. It follows from computer simulations and data on two conifer species that the Bayesian method gives nearly unbiased estimates of heterozygosity, genetic distances and F-statistics. The influence of a prior distribution and departure from Hardy-Weinberg proportions on the estimates is also considered.

Estimating population structure in diploids with multilocus dominant DNA markers.

Seed dispersal mechanisms should have a direct impact on the genetic structure of populations. Species whose seeds are dispersed near the maternal plant (e.g. gravity or wind dispersal) or species whose seeds are deposited in clumps or patches should have more fine-scale genetic structure than species whose seeds are dispersed singly by mobile animals. Furthermore, due to the overlap of seed shadows, species with high adult densities should have less genetic structure than species with lower densities. Allozyme analyses of three tropical tree species belonging to the moist tropical forest of Barro Colorado Island, Republic of Panama, were used to describe variation in the scale and intensity of genetic structure within their populations. The genetic structure of seedlings and immature trees in the low-density, wind-dispersed species (Platypodium elegans) was the coarsest and strongest whereas genetic structure in a population of Swartzia simplex var. ochnacea (high density, bird-dispersed) was both the finest and the weakest. The genetic structure of Alseis blackiana, a high-density, wind-dispersed species was intermediate in both degree and scale. In P. elegans and A. blackiana, which had ‘J’ shaped size distributions, the significant genetic structure seen in the smaller and intermediate diameter classes disappeared in the largest diameter class. The loss of genetic structure was not observed in S. simplex, a species with a more even size distribution.

The influence of seed dispersal mechanisms on the genetic structure of tropical tree populations

Populus deltoides, P. nigra, and P. trichocarpa are the most important species for poplar breeding programs worldwide. In addition, Populus has become a model for fundamental research on trees. Linkage maps were constructed for these three species by analyzing progeny of two controlled crosses sharing the same female parent, Populus deltoides cv. S9-2 x P. nigra cv. Ghoy and P. deltoides cv. S9-2 x P. trichocarpa cv. V24. The two-way pseudotestcross mapping strategy was used to construct the maps. Amplified fragment length polymorphism (AFLP) markers that segregated 1:1 were used to form the four parental maps. Microsatellites and sequence-tagged sites were used to align homoeologous groups between the maps and to merge linkage groups within the individual maps. Linkage analysis and alignment of the homoeologous groups resulted in 566 markers distributed over 19 groups for P. deltoides covering 86% of the genome, 339 markers distributed over 19 groups for P. trichocarpa covering 73%, and 369 markers distributed over 28 groups for P. nigra covering 61%. Several tests for randomness showed that the AFLP markers were randomly distributed over the genome.

https://www.genetics.org/content/genetics/158/2/787.full.pdf

Dense genetic linkage maps of three Populus species (Populus deltoides, P. nigra and P. trichocarpa) based on AFLP and microsatellite markers.

A genetic map of Pinus radiata plus tree 850.55 was constructed using megagametophytes of S1 seeds. The map contained 19 linkage groups, with 168 RAPD and four microsatellite markers. The total map length was 1116.7 cM (Kosambi’s function) and was estimated to cover 56% of the genome. Of the 172 markers, 59 (34%) were distorted from the expected 1 : 1 ratio in megagametophytes (P<0.05). We show that if the distortion is caused by a single viability gene or by sampling error, the estimate of recombination frequency in megagametophytes of selfed seeds would not be affected.

Genetic analysis of inbreeding depression in plus tree 850.55 of Pinus radiata D. Don. I. Genetic map with distorted markers

When conducting tree breeding experiments, geneticists often assume that individuals from open-pollinated families are halfsibs. The reliability of this assumption was tested using data from enzyme electrophoresis to estimate the genetic relatedness among progeny within 22 open-pollinated families of Robinia pseudoacacia L. (black locust) and 34 open-pollinated families of Gleditsia triacanthos L. (honey locust) from natural stands. An algorithm employing population estimates of fixation indices, pollen allele frequencies, and selfing rates was used to calculate the mean expected number of alleles in common across loci under assumptions of either full-sib (i.e., a single pollen parent) or half-sib (i.e., random mating) relationships. For each open-pollinated family, the average coefficient of relationship among progeny was calculated by linear interpolation from the observed number of alleles in common. For most families of both species, coefficients were significantly higher than 0.25 (half-sib relation), but were significantly lower than 0.50 (full-sib relation). These results suggest that the assumption of a half-sib relationship among progeny of open-pollinated families is violated for these tree species. More critical to the estimation of heritabilities and the prediction of genetic gains was the observation that estimates of the coefficient of relationship varied widely among open-pollinated families (for R. pseudoacacia r0=0.20–0.43, mean=0.34; for G. triacanthos r0=0.29–0.55, mean=0.36).

Genetic relatedness in open-pollinated families of two leguminous tree species, Robinia pseudoacacia L. and Gleditsia triacanthos L.

Seedlings from 23 seed sources representing the natural range of black locust (Robinia pseudoacacia L.) were electrophoretically analyzed at 40 structural loci representing 18 enzyme systems. Genetic diversity was high for an insect-pollinated, early-successional species. On average, 71% of the loci were polymorphic (99% criterion). Average and effective numbers of alleles per locus were 2.61 and 1.58, respectively. Mean expected heterozygosity was 0.291, ranging among the seed sources from 0.250 to 0.322. Most of the genetic diversity (88%) resided within seed sources. Few geographic patterns were evident. The most differentiated sources were located in disjunct sites in Georgia and the Ozark Mountains of Arkansas and Missouri, and in Pennsylvania. The widespread planting of native and European black locust seed sources may have contributed to the lack of geographic patterns. Based on this and other studies, it was concluded that initial selections in black locust breeding programs should concentrate on ...

Allozyme variation in black locust (Robiniapseudoacacia)

When inbred, most outcrossing species show high mortality, manifested at several life stages. The occurrence of homozygotes for deleterious or lethal alleles is believed to be responsible. Here, we report the identification of an allele responsible for the death of selfed Pinus radiata D. Don seedlings in their first month after germination. Among 291 S1 seedlings of plus-tree 850.55, 76 died within 1 month of emergence. Their death appears to be caused by a single recessive lethal allele, SDPr (seedling death in Pinus radiata). SDPr is located in a linkage group with 28 RAPD markers, the closest of which is ai05800a. Of the 76 seedlings that died, megagametophytes of 73 could be genotyped. Of these, 71 had the null (no band) allele of ai05800a; only two had the band allele. Of the 190 surviving S1 diploids that were genotyped, only two individuals were homozygous for the null allele of ai05800a. By two different methods, the map distance between SDPr and ai05800a was estimated to be between 1.0 and 2.7 cM respectively. The frequency of band and null alleles in the combined population of dead and surviving seedlings and in un-sown seeds shows no evidence of selection at this locus prior to germination.

An allele responsible for seedling death in Pinus radiata D. Don

Most coniferous species exhibit severe inbreeding depression. Selfed individuals usually have decreased viability, reduced vigour and morphological defects. The number of filled seeds after selfing Pinus radiata plus tree 850.55 was 48% that of the outcrossing, and 26.1% of the selfed seedlings died at an early stage. The segregation of 172 markers (covering 56% of the genome) in selfed progenies of radiata pine plus tree 850.55 was studied. Based on the segregation ratio of the markers, genes associated with inbreeding depression on viability were identified (P<0.05). Using the Expectation/Conditional Maximization (ECM) algorithm, we estimated the location, degree of dominance and selection coefficient of viability genes. Nine viability genes were discovered. Seven of them appeared to be dominant and one partially dominant (degree of dominance=0.4). The other gene was overdominant or pseudo-overdominant, with selection coefficients for the two homozygotes of 0.4 and 0.42, respectively. Of the genes showing dominance or partial dominance, seven were sub-lethal with selection coefficients ranging from 0.55 to 0.79; one gene (SDPr), which was responsible for seedling death within the first month following germination, was lethal.

B. C. Bongarten

Papers

Genetic analysis of inbreeding depression in plus tree 850.55 of Pinus radiata D. Don. I. Genetic map with distorted markers

Genetic relatedness in open-pollinated families of two leguminous tree species, Robinia pseudoacacia L. and Gleditsia triacanthos L.

Allozyme variation in black locust (Robiniapseudoacacia)

An allele responsible for seedling death in Pinus radiata D. Don

Genetic analysis of inbreeding depression in plus tree 850.55 of Pinus radiata D. Don. II. Genetics of viability genes