Variation in the lengths of restriction fragments (RFLPs) of the whole chloroplast DNA molecule was studied in 174 populations of Quercus ilex L. sampled over the entire distribution of this evergreen and mainly Mediterranean oak species. By using five endonucleases, 323 distinct fragments were obtained. From the 29 and 17 cpDNA changes identified as site and length mutations, respectively, 25 distinct chlorotypes were distinguished, mapped and treated cladistically with a parsimony analysis, using as an outgroup Q. alnifolia Poech, a closely related evergreen oak species endemic to Cyprus where Q. ilex does not grow. The predominant role of Q. ilex as maternal parent in hybridization with other species was reflected by the occurrence of a single very specific lineage of related chlorotypes, the most ancestral and recent ones being located in the southeastern and in the northwestern parts of the species' geographical distribution, respectively. The lineage was constituted of two clusters of chlorotypes observed in the 'ilex' morphotyped populations of the Balkan and Italian Peninsulas (including the contiguous French Riviera), respectively. A third cluster was divided into two subclusters identified in the 'rotundifolia' morphotyped populations of North Africa, and of Iberia and the adjacent French regions, respectively. Postglacial colonization probably started from three distinct southerly refugia located in each of the three European peninsulas, and a contact area between the Italian and the Iberian migration routes was identified in the Rhône valley (France). Chlorotypes identical or related to those of the Iberian cluster were identified in the populations from Catalonia and the French Languedoc region, which showed intermediate morphotypes, and in the French Atlantic populations which possessed the 'ilex' morphotype, suggesting the occurrence of adaptive morphological changes in the northern part of the species' distribution.
The results support a Middle-Eastern or a central Mediterranean origin for cork oak with subsequent westward colonization during the Tertiary Period, and suggest that the 'ilex' chlorotype variation does not reflect entirely cytoplasmic introgression by Q. ilex but originated partly in Q. suber.
Summary• Among oak species, Quercus ilex is classified as a monoterpene emitter and Q. suber is mainly known as a nonisoprenoid emitter. The extent and origin of this diversification is unknown.• We examined intra-and interspecific emission variability in two mixed stands which differed in their level of hybridization and reciprocal genetic introgression based on variations in cytoplasmic (chloroplast DNA) and nuclear (allozyme) markers.• At both sites all trees identified as Q. ilex , or as recent descendants from Q. ilex × Q. suber hybrids, emitted monoterpenes. Of Q. suber trees (genetically introgressed or not by Q. ilex ), 91% were also monoterpene emitters, and the remainder nonemitters. One tree identified as a Q. canariensis × Q. ilex hybrid emitted both isoprene and monoterpenes. Compared with Q. ilex , the standard emission rate of Q. suber was higher in summer and lower in autumn. Both species emitted the same monoterpenes, proportions of which showed significant intra-and interspecific variability.• The results suggest that Q. suber populations in the French Mediterranean intrinsically emit monoterpenes, and that gene flow between oak species contributes to diversification of emission signatures.
Maize was first domesticated in a restricted valley in south-central Mexico. It was diffused throughout the Americas over thousands of years, and following the discovery of the New World by Columbus, was introduced into Europe. Trade and colonization introduced it further into all parts of the world to which it could adapt. Repeated introductions, local selection and adaptation, a highly diverse gene pool and outcrossing nature, and global trade in maize led to difficulty understanding exactly where the diversity of many of the local maize landraces originated. This is particularly true in Africa and Asia, where historical accounts are scarce or contradictory. Knowledge of post-domestication movements of maize around the world would assist in germplasm conservation and plant breeding efforts. To this end, we used SSR markers to genotype multiple individuals from hundreds of representative landraces from around the world. Applying a multidisciplinary approach combining genetic, linguistic, and historical data, we reconstructed possible patterns of maize diffusion throughout the world from American "contribution" centers, which we propose reflect the origins of maize worldwide. These results shed new light on introductions of maize into Africa and Asia. By providing a first globally comprehensive genetic characterization of landraces using markers appropriate to this evolutionary time frame, we explore the post-domestication evolutionary history of maize and highlight original diversity sources that may be tapped for plant improvement in different regions of the world.
This study describes the genetic diversity and population structure of 194 native maize populations from 23 countries of Latin America and the Caribbean. The germplasm, representing 131 distinct landraces, was genetically characterized as population bulks using 28 SSR markers. Three main groups of maize germplasm were identified. The first, the Mexico and Southern Andes group, highlights the Pre-Columbian and modern exchange of germplasm between North and South America. The second group, Mesoamerica lowland, supports the hypothesis that two separate human migration events could have contributed to Caribbean maize germplasm. The third, the Andean group, displayed early introduction of maize into the Andes, with little mixing since then, other than a regional interchange zone active in the past. Events and activities in the pre- and post-Columbian Americas including the development and expansion of pre-Columbian cultures and the arrival of Europeans to the Americas are discussed in relation to the history of maize migration from its point of domestication in Mesoamerica to South America and the Caribbean through sea and land routes.
Maize (Zea mays L. ssp. mays) was domesticated from one wild species ancestor, the Balsas teosinte (Zea mays ssp. parviglumis) about 9000 years ago. Higher levels of gene diversity are found in teosinte taxa compared to maize, following domestication and selection bottlenecks. Diversity in maize can be increased via gene flow from teosinte, which has certainly occurred from various taxa, but the rate of flow from different teosinte taxa and the final impact on maize evolution has been difficult to measure. One hundred populations from six Zea taxa, both domesticated (maize) and wild (teosinte), including domesticated landraces from Asia, Africa, and the Americas, were genotyped with 17 SSR markers using 15 individuals per population. Overall levels of diversity were high, and populations could be distinguished based on markers. Relationships between populations followed most published reports, or can now help resolve previously conflicting reports. Gene flow into maize from different teosinte groups, and gene flow between different teosintes, was estimated. Evidence for contributions from the Balsas teosintes and from Chalco teosintes (Z. mays ssp. mexicana) to the maize gene pool was found, as well as from Chalco into ssp. mexicana race ''Durango'' and Z. mays ssp. huehuetenengensis. These contributions are almost certainly the result of post-domestication (and ongoing) exchanges. This information must give more impetus to in situ conservation of teosinte species, and use of these teosintes to continue to direct the evolution of maize, especially in response to new diseases, insect pests, and other biotic and abiotic stresses.
Hybridisation is a potent force in plant evolution, although there are few reported examples of stabilised species that have been created through homoploid hybridisation. We focus here on Quercus afares, an endemic North African species that combines morphological, physiological and ecological traits of both Q. suber and Q. canariensis, two phylogenetically distant species. These two species are sympatric with Q. afares over most of its distribution. We studied two Q. afares populations (one from Algeria and one from Tunisia), as well as several populations of both Q. suber and Q. canariensis sampled both within and outside areas where these species overlap with Q. afares. A genetic analysis was conducted using both nuclear (allozymes) and chloroplastic markers, which shows that Q. afares originates from a Q. suber  Q. canariensis hybridisation. At most loci, Q. afares predominantly possesses alleles from Q. suber, suggesting that the initial cross between Q. suber and Q. canariensis was followed by backcrossing with Q. suber. Other hypotheses that can account for this result, including genetic drift, gene silencing, gene conversion and selection, are discussed. A single Q. suber chlorotype was detected, and all Q. afares individuals displayed this chlorotype, indicating that Q. suber was the maternal parent. Q. afares is genetically, morphologically and ecologically differentiated from its parental species, and can therefore be considered as a stabilised hybrid species.
Interspecific gene flow is common in oaks. In the Mediterranean, this process produced geographical differentiations and new species, which may have contributed to the diversification of the production of volatile terpenes in the oak species of this region. The endemic North African deciduous oak Quercus afares (Pomel) is considered to be a stabilized hybrid between the evergreen Quercus suber (L.) and the deciduous Quercus canariensis (Willd.), presumably being monoterpene and isoprene emitters, respectively. In a common garden experiment, we examined the terpene emission capacities, terpene synthase (TPS) activities and nuclear genetic markers in 52 trees of these three oak species. All but one of the Q. suber and Q. canariensis trees were found to be genetically pure, whereas most Q. afares trees possessed a mixed genotype with a predominance of Q. suber alleles. Analysis of the foliar terpene emissions and TPS activities revealed that all the Q. canariensis trees strongly produced isoprene while all the Q. suber trees were strong monoterpene producers. Quercus afares trees produced monoterpenes as well but at more variable and significantly lower rates, and with a monoterpene pattern different than that observed in Q. suber. Among 17 individuals tested, one Q. afares tree emitted only an insignificant amount of terpenes. No mixed isoprene/monoterpene emitter was detected. Our results suggest that the capacity and pattern of volatile terpene production in Algerian Q. afares populations have strongly diverged from those of its parental species and became quantitatively and qualitatively reduced, including the complete suppression of isoprene production.
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