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

We redescribe the only previously known species of Myrsidea from bulbuls, M. pycnonoti Eichler. Sixteen new species are described; they and their type hosts are: M. phillipsi ex Pycnonotus goiavier goiavier (Scopoli), M. gieferi ex P. goiavier suluensis Mearns, M. kulpai ex P. flavescens Blyth, M. finlaysoni ex P. finlaysoni Strickland, M. kathleenae ex P. cafer (L.), M. warwicki ex Ixos philippinus (J. R. Forster), M. mcclurei ex Microscelis amaurotis (Temminck), M. zeylanici ex P. zeylanicus (Gmelin), M. plumosi ex P. plumosus Blyth, M. eutiloti ex P. eutilotus (Jardine and Selby), M. adamsae ex P. urostictus (Salvadori), M. ochracei ex Criniger ochraceus F. Moore, M. borbonici ex Hypsipetes borbonicus (J. R. Forster), M. johnsoni ex P. atriceps (Temminck), M. palmai ex C. ochraceus, and M. claytoni ex P. eutilotus. A key is provided for the identification of these 17 species.

A new species

Domestication interests us as the most momentous change in Holocene human history. Why did it operate on so few wild species, in so few geographic areas? Why did people adopt it at all, why did they adopt it when they did, and how did it spread? The answers to these questions determined the remaking of the modern world, as farmers spread at the expense of hunter-gatherers and of other farmers.

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Evolution, consequences and future of plant and animal domestication

Globally, 800 million people are malnourished. Heavily subsidised farmers in rich countries produce sufficient surplus food to feed the hungry, but not at a price the poor can afford. Even donating the rich world's surplus to the poor would not solve the problem. Most poor people earn their living from agriculture, so a deluge of free food would destroy their livelihoods. Thus, the only answer to world hunger is to safeguard and improve the productivity of farmers in poor countries. Diets of subsistence level farmers in Africa and Latin America often contain sufficient carbohydrates (through cassava, corn/maize, rice, wheat, etc.), but are poor in proteins. Dietary proteins can take the form of scarce animal products (eggs, milk, meat, etc.), but are usually derived from legumes (plants of the bean and pea family). Legumes are vital in agriculture as they form associations with bacteria that `fix-nitrogen' from the air. Effectively this amounts to internal fertilisation and is the main reason that legumes are richer in proteins than all other plants. Thousands of legume species exist but more common beans (Phaseolus vulgaris L.) are eaten than any other. In some countries such as Mexico and Brazil, beans are the primary source of protein in human diets. As half the grain legumes consumed worldwide are common beans, they represent the species of choice for the study of grain legume nutrition. Unfortunately, the yields of common beans are low even by the standards of legumes, and the quality of their seed proteins is sub-optimal. Most probably this results from millennia of selection for stable rather than high yield, and as such, is a problem that can be redressed by modern genetic techniques. We have formed an international consortium called `Phaseomics' to establish the necessary framework of knowledge and materials that will result in disease-resistant, stress-tolerant, high-quality protein and high-yielding beans. Phaseomics will be instrumental in improving living conditions in deprived regions of Africa and the Americas. It will contribute to social equity and sustainable development and enhance inter- and intra-cultural understanding, knowledge and relationships. A major goal of Phaseomics is to generate new common bean varieties that are not only suitable for but also desired by the local farmer and consumer communities. Therefore, the socio-economic dimension of improved bean production and the analysis of factors influencing the acceptance of novel varieties will be an integral part of the proposed research (see Figure 1). Here, we give an overview of the economic and nutritional importance of common beans as a food crop. Priorities and targets of current breeding programmes are outlined, along with ongoing efforts in genomics. Recommendations for an international coordinated effort to join knowledge, facilities and expertise in a variety of scientific undertakings that will contribute to the overall goal of better beans are given. To be rapid and effective, plant breeding programmes (i.e., those that involve crossing two different `parents') rely heavily on molecular `markers'. These genetic landmarks are used to position

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Beans (Phaseolus spp.) - model food legumes

Advanced backcross QTL analysis is proposed as a method of combining QTL analysis with variety development. It is tailored for the discovery and transfer of valuable QTL alleles from unadapted donor lines (e.g., land races, wild species) into established elite inbred lines. Following this strategy, QTL analysis is delayed until the BC2 or BC3 generation and, during the development of these populations, negative selection is exercised to reduce the frequency of deleterious donor alleles. Simulations suggest that advanced backcross QTL analysis will be effective in detecting additive, dominant, partially dominant, or overdominant QTLs. Epistatic QTLs or QTLs with gene actions ranging from recessive to additive will be detected with less power than in selfing generations. QTL-NILs can be derived from advanced backcross populations in one or two additional generations and utilized to verify QTL activity. These same QTL-NILs also represent commercial inbreds improved (over the original recurrent inbred line) for one or more quantitative traits. The time lapse from QTL discovery to construction and testing of improved QTL-NILs is minimal (1-2 years). If successfully employed, advanced backcross QTL analysis can open the door to exploiting unadapted and exotic germplasm for the quantitative trait improvement of a number of crop plants.

Advanced backcross QTL analysis: a method for the simultaneous discovery and transfer of valuable QTLs from unadapted germplasm into elite breeding lines.

Seed-crop plants apparently originated from a limited number of mutants in which seed dispersal was changed from that found in nondomesticated populations. Seed nonshattering in cultivated plants may be controlled by a single gene or a small number of genes. Allopolyploid crop plants were derived from a limited number of interspecific hybridizations followed by chromosome doubling. The consequence of this founder effect is a narrow genetic variability in the crop population compared to its wild progenitor. Natural hybridization between the two is prevented by various isolating mechanisms, and gene flow, if it exists, is apparently more effective in the direction from the cultivated to the wild populations. Founder effect in crop-plant evolution indicates the value and the breeding potential of the genetic variability remaining in its wild relatives.

Founder effect in crop-plant evolution

Species relationship between the cultivated chickpea Cicer arietinum and the two newly discovered wild species C echinospermum and C reticulatum were assessed through breeding experiments and cytological examination of the hybrids The two wild species differed from each other by a major reciprocal translocation and their hybrid was completely sterile The wild species C echinospermum also differed from the cultivated species by the same translocation and their hybrid was highly sterile The other wild species, C' reticulatum, was crossed readily with the cultivated chickpea Meiosis of the hybrids, involving 4 different C arietinum lines, was normal, and they were fertile This wild species therefore can be considered as the wild progenitor of the cultivated chickpea

The origin of chickpea Cicer arietinum L.

Preface. 1. Origin of agriculture. 2. Increasing diversity under domestication. 3. The course of reducing and maintaining genetic diversity under domestication. 4. Speciation under domestication. 5. Weeds and their evolution. 6. Evolution of selected crop plants. 7. Genetic resources for future crop evolution. References. Index.

Plant Evolution under Domestication

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Extensive Conservation of Linkage Relationships Between Pea and Lentil Genetic Maps

Genetic relationships between 7 annual species of the genus Cicer, including the cultivated chickpea, have been studied. These species were assigned to 3 crossability groups. In each group interspecific hybrids could be obtained but their fertility differed considerably in the various cross combinations. Crosses between members of different groups yielded no viable seeds. The possibility of gene transfer from the wild species to the cultivated chickpea C. arietinum was also assessed. Only two species could be considered for this purpose, C. reticulatum, which is the wild progenitor of the cultivated species, and C. echinospermum, which is in the secondary gene pool of C. arietinum. A unique postzygotic reproductive barrier mechanism was found between the members of Group II, C. judaicum, C. pinnatifidum and C. bijugum. It is based on a disharmony in the growth rate of the stigma and the anthers at the time of anthesis of the F1 interspecific hybrid so that selfpollination is avoided. It is proposed that this kind of mechanism has been involved only when an effective spatial isolation between the three species had been obtained.

Gideon Ladizinsky

Papers

Founder effect in crop-plant evolution

The origin of chickpea Cicer arietinum L.

Plant Evolution under Domestication

Extensive Conservation of Linkage Relationships Between Pea and Lentil Genetic Maps

Genetic relationships among the annual species of Cicer L.