Role of seed flow on the pattern and dynamics of pearl millet (<Emphasis Type="Italic">Pennisetum glaucum</Emphasis> [L.] R. Br.) genetic diversity assessed by AFLP markers: a study in south-western Niger

We studied the regional genetic diversity and seed exchange dynamics of pearl millet landraces in south-western Niger. The genetic study was based on AFLP markers. We found significant genetic differentiation between landraces in different geographical areas of south-western Niger. However, the degree of differentiation was low insofar as only 1.9% of the total molecular diversity was due to regional differentiation, suggesting a relatively high gene flow. Anthropologic studies on farming practices have suggested that seed exchanges between farmers on a large geographical scale probably make a considerable contribution to this result. In order to test this hypothesis, the effects of seed exchange on the genetic diversity of landraces was analyzed on seed samples from two distant villages in contrasting areas of south-western Niger. Seeds imported by farmers into the southern village of Sina Koara did not differ significantly from locally grown landraces. By contrast, in the northern village of Alzou, several samples were genetically different from locally grown landraces and closer to southern accessions. These data suggest that the seed flow is preferentially from south to north, i.e. from an area with more favorable rainfall conditions. The potential consequences for the genetic diversity and adaptation of northern pearl millet landraces are discussed.     

Bimodal phenotypic structure of two wild pearl millet samples collected in an agricultural area

On the basis of botanical and ecological characters, pearl millet species have been separated into three subspecies:Pennisetum americanum subsp. americanum, cultivated form,P. americanum subsp. monodii, wild form andP. americanum subsp. stenostachyum, weedy form. It has been shown, by enzyme electrophoresis, that these three forms could be discriminated between by their respective enzyme composition even in agricultural areas where they co-exist. Morphological and enzyme analyses of two seed samples collected from monodii plants growing in an agricultural area has shown that the monodii form is not isolated from cultivated plants in its reproduction. The proportion of seeds descending from a cultivated pollen is estimated respectively at 31% and 19%. However, Brunken's typology is corroborated: both samples comprise the two distinct groups of monodii and stenostachyum. This dichotomy is due to the pollen population structure having generated the analysed seeds. Detailedin situ observations on sympatric wild and cultivated populations are essential to explain the absence of recombinant monodii × cultivated pollens and the preservation of monodii millets in spite of their invasion by cultivated pollens.     

Inference of domestication history and differentiation between early‐ and late‐flowering varieties in pearl millet

Pearl millet (Pennisetum glaucum) is a staple crop in Sahelian Africa. Farmers usually grow varieties with different cycle lengths and complementary functions in Sahelian agrosystems. Both the level of genetic differentiation of these varieties and the domestication history of pearl millet have been poorly studied. We investigated the neutral genetic diversity and population genetic structure of early‐ and late‐flowering domesticated and wild pearl millet populations using 18 microsatellite loci and 8 nucleotide sequences. Strikingly, early‐ and late‐flowering domesticated varieties were not differentiated over their whole distribution area, despite a clear difference in their isolation‐by‐distance pattern. Conversely, our data brought evidence for two well‐differentiated genetic pools in wild pearl millet, allowing us to test scenarios with different numbers and origins of domestication using approximate Bayesian computation (ABC). The ABC analysis showed the likely existence of asymmetric migration between wild and domesticated populations. The model choice procedure indicated that a single domestication from the eastern wild populations was the more likely scenario to explain the polymorphism patterns observed in cultivated pearl millet.     

Wild pearl millet population (Pennisetum glaucum,Poaceae) integrity in agricultural Sahelian areas. An example from Keita (Niger)

Morphometric and isozymic analyses of adjacent cultivated and spontaneous populations of pearl millet in Niger revealed in the field a unique continuous distribution of phenotypes ranging from the most cultivated one to a typical cultivated × wild hybrid. The natural population was subdivided into a major wild group and a hybrid wild × cultivated group. Cultivated millet displayed an equilibrium state between recombined domesticated and wild genes. The natural population, in spite of a high rate of immigration by pollen from cultivated plants, retained its structure by apparently reproducing itself exclusively from the major wild group.     

Uptake,Translocation, Distribution and Persistence of 14C-Metalaxyl in Pearl Millet (Pennisetum americanum) [L.] Leeke)1

Time course absorption and desorption of metalaxyl by seeds of pearl millet was analysed by following chemical kinetics equations. Uptake of metalaxyl through roots, leaves and seed, its translocation and distribution in different plant parts and persistence following seed application were studied in pearl millet using ¹⁴C-metalaxyl. Both uptake and efflux of metalaxyl by pearl millet seeds were complex and compartmentalized. Distribution inside the seed was not uniform. A major part of applied fungicide remained within the treated plant part, particularly after seed and foliar applications. Metalaxyl was ambimobile inside the plant and was found to get accumulated at apex and margins of leaf blade. No metalaxyl could be detected in grains, harvested from plants grown from metalaxyl treated seeds.     

Wild to crop introgression and genetic diversity in Lima bean (Phaseolus lunatus L.) in traditional Mayan milpas from Mexico

Despite the evolutionary, ecological and economic importance of introgression between a domesticated species and its wild relatives in centers of diversity and domestication, the role of traditional farmers in this process has received limited attention. In the Yucatan Peninsula, the region of Mexico that has the greatest amount of domesticated varieties of Lima bean, wild populations grow sympatrically with conspecific varieties, allowing the Mayan farmer to act directly on introgressed seed. We used 11 microsatellite loci to assess levels of introgression in three wild-domesticated complexes of Lima bean from the Yucatan Peninsula and analyze its impact on the genetic diversity of this crop. structure and InStruct analyses showed similar results. The Instruct analysis indicated that the complex with the lowest level of introgression was one where the farmer actively selected against wild plants and introgressed seed. In contrast, the complex with the highest level of introgression was one where the farmer has been consciously selecting a weedy morphotype for 15 years and has already incorporated it into his diet. Genetic diversity of the domesticated pool was higher in the complex with the higher level of introgression. This study showed that farmers have an important role in limiting or favoring the wild to crop introgression and influencing the levels of genetic diversity in their domesticated pool. Only when traditional farmers’ knowledge is taken into account can we correctly understand the dynamics, generation and maintenance of genetic diversity of the landraces in the centers of diversity and domestication.     

Phylogeny and origin of pearl millet (Pennisetum glaucum [L.] R. Br) as revealed by microsatellite loci

During the last 12,000 years, different cultures around the world have domesticated cereal crops. Several studies investigated the evolutionary history and domestication of cereals such as wheat in the Middle East, rice in Asia or maize in America. The domestication process in Africa has led to the emergence of important cereal crops like pearl millet in Sahelian Africa. In this study, we used 27 microsatellite loci to analyze 84 wild accessions and 355 cultivated accessions originating from the whole pearl millet distribution area in Africa and Asia. We found significantly higher diversity in the wild pearl millet group. The cultivated pearl millet sample possessed 81% of the alleles and 83% of the genetic diversity of the wild pearl millet sample. Using Bayesian approaches, we identified intermediate genotypes between the cultivated and wild groups. We then analyzed the phylogenetic relationship among accessions not showing introgression and found that a monophyletic origin of cultivated pearl millet in West Africa is the most likely scenario supported by our data set.     

Isozymic classification of pearl millet (Pennisetum glaucum,Poaceae) landraces from Niger (West Africa)

The principal landraces of the pearl millet,Pennisetum glaucum (L.)R. Br., from Niger have been analysed for their genetic structure at eight enzyme systems coded by 12 loci and 46 alleles. Three groups have been identified: (1) early-maturing pearl millets, cultivated between 8° and 13°E longitude, including the oases from Aïr mountains; (2) early-maturing millets situated more to the west (1° and 8°E longitude), and (3) late-maturing millets. Group 1 shows the highest isozyme diversity. The differences between the accessions represent 8.8% of the total diversity and the differences between the three groups 4.5%. The accessions from groups 1 and 3 are the least distant. When considering pearl millets from areas outside Niger, the chadian and sudanese millets are enzymatically close to the Niger group 1. The pearl millets from Niger group 2 are close to millets from east Mali, northern Burkina Faso and Senegal, and the Niger group 3 to the late-maturing millets group from West Africa. This study should help breeders to select the landraces for improvement and parents for crosses from cultivars of Niger and introduced germ plasm.     

A weed-crop complex in sorghum: The dynamics of genetic diversity in a traditional farming system

Despite the major ecological and economic impacts of gene flow between domesticated plants and their wild relatives, many aspects of the process, particularly the relative roles of natural and human selection in facilitating or constraining gene flow, are still poorly understood. We developed a multidisciplinary approach, involving both biologists and social scientists, to investigate the dynamics of genetic diversity of a sorghum weed-crop complex in a village of Duupa farmers in northern Cameroon. Farmers distinguish a gradient from weedy morphotypes (naa baa see, haariya, and genkiya) to domesticated morphotypes; haariya and genkiya have intermediate morphological traits. We investigated the pattern of diversity in this complex using both morphological and genetic data. Our biological results are interpreted in the light of data on farmers' taxonomy and practices such as spatial pattern of planting and plant selection. Both morphological and genetic data are congruent with farmers' taxonomy and confirm the introgressed status of intermediate weedy morphotypes. Farmers actively select against weedy morphotypes, but several practices unconsciously favor gene flow. Furthermore, haariya and genkiya may facilitate introgression between naa baa see and domesticated morphotypes by virtue of their intermediate flowering period and their mode of management by farmers.     

Four thousand years of plant exploitation in the Lake Chad Basin (Nigeria), part III: plant impressions in potsherds from the Final Stone Age Gajiganna Culture*

Late Holocene climatic changes caused a large scale regression of the Lake Chad shoreline followed by an expansion of settlements into previously unexplored territories. Numerous Final Stone Age sites of the Gajiganna Culture (1,800 to 800 b.c.) in the Lake Chad Basin (northeast Nigeria) yielded plant impressions in potsherds. The ceramics of Phase I (1,800–1,400 b.c.) were mineral tempered, and plant impressions, mainly of Paniceae, were caused only by incidental inclusion. In contrast, a considerable number of the sherds from Phase II (1,500–800 b.c.) were intentionally tempered with chaff derived from domesticated pearl millet (Pennisetum glaucum), wild Paniceae and wild rice species (Oryza cf. barthii and O. cf. longistaminata). This plant spectrum suggests the exploitation of the wet wild areas, and also the cultivation of pearl millet on sandy soils. The evidence suggests that agricultural practices were established late and were introduced from elsewhere. During the time of seasonally occupied sites in Phase I, the subsistence strategy was based on herding, fishing, and gathering, while in Phase II there are signs of permanent settlements and agriculture. The evidence from the plant impressions indicates that in the Final Stone Age Gajiganna Culture around 1,000–800 b.c., pearl millet became well established while the gathering of wild millets and rice was still practised.*Klee et al. (2000), Zach and Klee (2003)     

MORPHOLOGICAL DIFFERENTIATION OF PROBOSCIDEA PARVIFLORA SSP. PARVIFLORA (MARTYNIACEAE) UNDER DOMESTICATION

Aboriginal groups in the southwestern United States have grown Proboscidea parviflora ssp. parviflora for basketry fiber. It has been hypothesized that cultivated plants of this subspecies with white seeds and long-clawed fruit (which provide the fiber) have been domesticated. Cluster, multiple discriminant, and unvariate statistical analyses presented here show that the putative domesticate is indeed morphologically distinct from “wild” plants. It has clearly been domesticated, and has diverged from wild populations to the extent that a new variety, var. Hohokamiana, is proposed for the domesticate. Wild and domesticated varieties differ especially in seed color, rostrum, crest, style, and anther lengths. Most characteristics of the wild variety seem to be genetically dominant to those of the domesticate, which may account for absence of a weedy form intermediate between the wild and domesticated varieties. The wild variety is morphologically more variable than the domesticate, probably because of its broader geographical range and its intergradation with another subspecies. Ethnological data and the variational patterns reported here suggest that the domesticate originated in southern Arizonanorthern Sonora, but probably not north of that region.     

Tuareg, a novel miniature-inverted repeat family of pearl millet (Pennisetum glaucum) related to the PIF superfamily of maize

Miniature-inverted repeat transposable elements (MITEs) are abundantly repeated in plant genomes and are especially found in genic regions where they could contribute regulatory elements for gene expression. We describe with molecular and cytological tools the first MITE family reported in pearl millet: Tuareg. It was initially detected in the pearl millet ortholog of Teosinte-branched1, an important developmental gene involved in the domestication of maize. The Tuareg family was amplified recently in the pearl millet genome and elements were found more abundant in wild than in domesticated plants. We found that they shared similarity in their terminal repeats with the previously described mPIF MITEs and that they are also present in other Pennisetum species, in maize and more distantly related grasses. The Tuareg family may be part of MITEs activated by PIF-like transposases and it could have been mobile since pearl millet domestication. Electronic supplementary material Electronic supplementary material is available for this article at and accessible for authorised users. O. Robin contributed the FISH and fiber-FISH hybridizations.     

Evidence from RAPD markers in the evolution ofEchinochloa millets (Poaceae)

Echinochloa (Poaceae) includes two domesticated species,Echinochloa utilis (Japanese barnyard millet) andE. frumentacea (Indian sawa millet) and 20–30 wild species. The two millets are morphologically very variable and overlap in spikelet and inflorescence characteristics. Both species are hexaploids based on x = 9. Cytogenetic studies point to the hexaploid wild speciesE. crusgalli andE. colona as possible progenitors ofE. utilis andE. frumentacea, respectively. The tetraploidE. oryzoides is considered as a possible genome donor to wild and domesticated barnyard millet. Markers from Random Amplified Polymorphic DNA method were used to assess the proposed phylogeny and examine the genetic diversity in both domesticated and wild species. The data were analyzed numerically.Echinochloa utilis andE. frumentacea appear very distinct, but grouped withE. crusgalli andE. colona, respectively. The tetraploidE. oryzoides show strong genetic affinity to theE. utilis—E. crusgalli group. The data are in general agreement with the cytogenetic information; however, some disagreements on the interpretation of some of the cytogenetic information is raised. The variability in DNA markers observed in the domesticated species, particularlyE. frumentacea, points to the feasibility of using RAPD markers in cultivar fingerprinting and breeding programs of these millets.     

Phytolith analysis for differentiating between foxtail millet (Setaria italica) and green foxtail (Setaria viridis)

Foxtail millet (Setaria italica) is one of the oldest domesticated cereal crops in Eurasia, but identifying foxtail millets, especially in charred grains, and differentiating it from its wild ancestor, green foxtail (Setaria viridis), in the archaeobotanical remains, is still problematic. Phytolithic analysis provides a meaningful method for identifying this important crop. In this paper, the silicon structure patterns in the glumes, lemmas, and paleas from inflorescence bracts in 16 modern plants of foxtail millet and green foxtail from China and Europe are examined using light microscopy with phase-contrast and a microscopic interferometer. Our research shows that the silicon structure of ΩIII from upper lemmas and paleas in foxtail millet and green foxtail can be correspondingly divided into two groups. The size of ΩIII type phytolith of foxtail millet is bigger than that from green foxtail. Discriminant function analysis reveals that 78.4% of data on foxtail millet and 76.9% of data on green foxtail are correctly classified. This means certain morphotypes of phytoliths are relatively reliable tools for distinguishing foxtail millet from green foxtail. Our results also revealed that the husk phytolith morphologies of foxtail millets from China and Eastern Europe are markedly different from those from Western Europe. Our research gives a meaningful method of separating foxtail millet and green foxtail. The implications of these findings for understanding the history of foxtail millet domestication and cultivation in ancient civilizations are significant.     

Domestication, cultivation and utilization of two small millets, brachiaria ramosa and Setaria glauca (Poaceae), in South India

Brachiaria ramosa is cultivated in pure stands. Its grains are used in preparing at least nine traditional foods in South India. Setaria glauca is cultivated in mixed stands along with little millet (Panicum sumatrense). In Orissa state and in Southern India the grains are used to make at least six traditional foods as a supplementary ingredient. The weedy forms of these were found growing with upland rice and some millets in diverse agro-ecological niches, and are identified by various vernacular names by local farmers. Explorations were conducted to gather information on the evolutionary trends, domestication and ethnobotany of these minor food crops. These findings substantiate the belief that domestication process may have passed through three distinct phases: (i) an associated mimic weed with upland rice and some millets; (ii) a secondary crop mixed with little millet or kodo millet; and (iii) as an independent crop. Also, this process has been promoted as a means of imparting adaptation to an arid climate and in turn related to the diversification of the agriculture basic complex.     

5-Methylcytosine content and methylation status in six millet DNAs

High performance liquid chromatographic analysis of the total nuclear DNAs of 6 millets plant species indicates that the 5-methylcytosine content ranges from 3% in barn yard millet to 9.6% in great millet while the fraction of cytosines methylated varies between 14% in little millet to 31 % in pearl millet. Digestion of millet DNAs with MspI/HpaII suggests that CpG methylation is more in great millet DNA while CpC methylation is more in the other 5 millet DNAs. Digestion of millet DNAs with MboI, Sau3AI andDpnI indicates that some of the^5’ GATC^3’ sequences are methylated at adenine and/or cytosine residues except in little millet where adenine methylation of the^5’GATC^3’ sequences is insignificant and there is a predominance of cytosine methylation in these sequences.     

Molecular evolution of shattering loci in U.S. weedy rice

Cultivated rice fields worldwide are plagued with weedy rice, a conspecific weed of cultivated rice (Oryza sativa L.). The persistence of weedy rice has been attributed, in part, to its ability to shatter (disperse) seed prior to crop harvesting. In the United States, separately evolved weedy rice groups have been shown to share genomic identity with exotic domesticated cultivars. Here, we investigate the shattering phenotype in a collection of U.S. weedy rice accessions, as well as wild and cultivated relatives. We find that all U.S. weedy rice groups shatter seeds easily, despite multiple origins, and in contrast to a decrease in shattering ability seen in cultivated groups. We assessed allelic identity and diversity at the major shattering locus, sh4, in weedy rice; we find that all cultivated and weedy rice, regardless of population, share similar haplotypes at sh4, and all contain a single derived mutation associated with decreased seed shattering. Our data constitute the strongest evidence to date of an evolution of weeds from domesticated backgrounds. The combination of a shared cultivar sh4 allele and a highly shattering phenotype, suggests that U.S. weedy rice have re‐acquired the shattering trait after divergence from their progenitors through alternative genetic mechanisms.     

Plant management among the Nahua and the Mixtec in the Balsas River Basin,Mexico: An ethnobotanical approach to the study of plant domestication

Different forms of management of wild, weedy, and domesticated plants carried out by the Nahua and the Mixtec in the Balsas River Basin, Mexico, are described. Along with cultivation of domesticated plants, these forms of plant management include gathering from wild populations; in situtolerance of plant individuals during clearings of natural vegetation; in situenhancement and protection of particular plants among populations of some species; as well as sowing or planting of propagules and transplantation of complete individuals of weedy and wild plants in controlled ex situenvironments. Processes of artificial selection and possible routes of domestication occurring in these forms of plant management are discussed.     

Malaysian weedy rice shows its true stripes: wild Oryza and elite rice cultivars shape agricultural weed evolution in Southeast Asia

Weedy rice is a close relative of domesticated rice (Oryza sativa) that competes aggressively with the crop and limits rice productivity worldwide. Most genetic studies of weedy rice have focused on populations in regions where no reproductively compatible wild Oryza species occur (North America, Europe and northern Asia). Here, we examined the population genetics of weedy rice in Malaysia, where wild rice (O. rufipogon) can be found growing in close proximity to cultivated and weedy rice. Using 375 accessions and a combined analysis of 24 neutral SSR loci and two rice domestication genes (sh4, controlling seed shattering, and Bh4, controlling hull colour), we addressed the following questions: (i) What is the relationship of Malaysian weedy rice to domesticated and wild rice, and to weedy rice strains in the USA? (ii) To what extent does the presence of O. rufipogon influence the genetic and phenotypic diversity of Malaysian weeds? (iii) What do the distributions of sh4 and Bh4 alleles and associated phenotypes reveal about the origin and contemporary evolution of Malaysian weedy rice? Our results reveal the following: independent evolutionary origins for Malaysian weeds and US strains, despite their very close phenotypic resemblance; wild‐to‐weed gene flow in Malaysian weed populations, including apparent adaptive introgression of seed‐shattering alleles; and a prominent role for modern Malaysian cultivars in the origin and recent proliferation of Malaysian weeds. These findings suggest that the genetic complexity and adaptability of weedy crop relatives can be profoundly influenced by proximity to reproductively compatible wild and domesticated populations.     

A SYSTEMATIC STUDY OF PENNISETUM SECT. PENNISETUM (GRAMINEAE)

Pennisetum sect. Pennisetum includes two reproductively isolated species. Pennisetum purpureum Schumach. is a tetraploid (2n = 28) perennial species which occurs throughout the wet tropics of the world. Pennisetum americanum (L.) Leeke is a diploid (2n = 14) annual species, native to the semi-arid tropics of Africa and India, and contains three morphologically diverse subspecies. Subspecies americanum includes the wide array of cultivated pearl millets. Subspecies monodii from the Sahel of West Africa is identified as the wild progenitor of pearl millet. Subspecies stenostachyum is morphologically intermediate between subsp. americanum and monodii and includes the mimetic weeds often associated with the cultivation of pearl millet.