GENETIC DIVERSITY IN GOSSYPIUM HIRSUTUM AND THE ORIGIN OF UPLAND COTTON

Gossypium hirsutum has a large indigenous range encompassing most of Mesoamerica and the Caribbean, where it exhibits a diverse array of morphological forms spanning the wild-to-domesticated continuum. Modem, highly improved varieties (“Upland cotton”), which currently account for about 90% of world cotton commerce, are day-length neutral annuals derived from subtropical, perennial antecedents. To assess levels and patterns of genetic variation in the species and to elucidate the origin of Upland cotton, 538 accessions representing the full spectrum of morphological and geographical diversity were analyzed forallozyme variation at 50 loci. Levels of variation are modest overall but are low in Upland cotton. Relationships among accessions reflect pre-Columbian influences of aboriginal peoples and later European colonists superimposed on the preagricultural pattern. In contrast to expectations, two centers of diversity are evident, one in southern Mexico-Guatemala and the other in the Caribbean. Introgression of G. barbadense genes into G. hirsutum has been common in a broad area of sympatry in the Caribbean. The germplasm of present cultivars traces to Mexican highland stocks, which, in turn, were derived from material originally from southern Mexico and Guatemala. Despite the widespread belief that germplasm from several other species has been incorporated into modem Upland stocks through intentional breeding efforts, the 50 Upland cultivars examined contain no unique alleles, suggesting that retention of genes from transspecific sources has been minimal. The most recent infraspecific treatment, which recognizes seven races, does not adequately represent genetic relationships.     

Sequence and RFLP analysis of the ITS2 ribosomal DNA in two Neotropical social bees, <Emphasis>Melipona beecheii</Emphasis> and <Emphasis>Melipona yucatanica</Emphasis> (Apidae,Meliponini)

Two stingless bees species of the genus Melipona, M. beecheii and M. yucatanica, are the only ones reported for the Yucatan Peninsula. The natural distribution of M. beecheii ranges from southern Mexico to Costa Rica, that of M. yucatanica from south Mexico to Guatemala. Colonies of both species occur in a variety of habitats and show adaptations to local conditions denoting the occurrence of ecotypes. The ITS2 of ribosomal DNA has been characterized in both species and its utility to discriminate among colonies has been investigated through RFLP experiments. The ITS2 region is unusually long, 1788 bp in M. beecheii and 1845 bp in M. yucatanica (including the 3′ end of the 5.8S gene and partial 5′ of the 28S gene). Mean nucleotide divergence between both ITS2 sequences is 16% (excluding sites with insertions/deletions) and 20% when the insertions/deletions are taken into account. The G+C content in both sequences is close to 53%. The PCR-RFLP assay was performed with 12 restriction enzymes on colonies of M. beecheii from Mexico (Yucatan, Campeche and Chiapas) Costa Rica, El Salvador and Guatemala, and of M. yucatanica from Mexico (Yucatan) and Guatemala. The restriction patterns obtained allow to discriminating colonies of both species with different origins. Both kinds of data are thus useful for assessing intra and interspecific genetic variability and for developing appropriate conservation strategies for these species. Received 20 June 2007; revised 31 August 2007; accepted 12 September 2007.     

Genetic structure and differentiation of wild and domesticated populations of Capsicum annuum (Solanaceae) from Mexico

 Genetic variation and structure of ten wild, three domesticated and one wild-cultivated populations of pepper (Capsicum annuum) from northwestern Mexico were studied in order to find out if the domestication process has reduced the genetic variation of the modern cultivars of this species. The analysis was based on 12 polymorphic loci from nine isozymes. Wild populations were sampled in different habitats along a latitudinal gradient of ca. 500 km. All populations had high genetic variation (i.e. wild: A = 2.72, P = 90.8%, He = 0.445; wild-cultivated: A = 2.50, P = 92.3%, He = 0.461; domesticated: A = 2.60, P = 84.6%, He = 0.408), indicating little genetic erosion in modern cultivars of pepper. Genetic diversity estimated by Nei's method showed that most genetic variation is found within, rather than among populations. However, genetic differentiation is greater among cultivated (G _ST=0.167) than among wild (G _ST=0.056) populations. Wild populations had an average genetic identity (I) of 0.952, indicating little differentiation and high gene flow (Nm=4.21) among these populations. Average genetic identity between wild and domesticated populations was of I=0.818, revealing that the domestication process has modified the genetic composition of commercial varieties of pepper. Changes in genetic composition among commercial varieties seem to have occurred in different directions, as indicated by the average value of I = 0.817 among these populations. The high level of diversity found in wild populations of C. annuum suggests that the wild relatives of cultivated peppers are a valuable genetic resource which must be conserved. Received May 5, 1999 Accepted October 30, 2000     

Patterns of genetic variation of the genusCapsicum (Solanaceae) in Mexico

The evolutionary relationships of 186 accessions ofCapsicum from Mexico were studied through enzyme electrophoresis. A total of 76 alleles representing 20 genetic loci coding for nine enzyme systems were observed and the allelic variations of enzymes were studied for geographical distribution. Allele frequencies were used to estimate the apportionment of gene diversity within and between populations and to construct a dendrogram based on a similarity matrix containingNei genetic distances. — The gene diversity estimates suggest that the structure ofCapsicum populations in Mexico consists of predominantly homozygous genotypes presumably due to a self-pollinated breeding system and population bottlenecks. Significant genetic differentiation was found mainly between populations of differing geographical regions.—Based on the results of this study, three species of domesticatedCapsicum can be identified in Mexico,C. annuum var.annuum, C. chinense, andC. pubescens. Semidomesticated and wild forms include two species,C. frutescens andC. annuum var.glabriusculum. A sharp geographical division results between the latter species;C. frutescens was collected exclusively in the southeastern states of Oaxaca, Chiapas, and Tabasco; whereas wild and semidomesticated forms from the rest of the country areC. annuum. Based upon the similarity of enzyme genotypes of semidomesticated and wild forms, the primary center of domestication of cultivatedC. annuum was estimated to be the region comprising the states of Tamaulipas, Nuevo Leon, San Luis Potosi, Veracruz, and Hidalgo in eastern Mexico. A possible second center of domestication is suggested to be localized in the state of Nayarit, western Mexico.     

Isozymatic variation and phylogenetic relationships between henequen (Agave fourcroydes) and its wild ancestor A. angustifolia (Agavaceae)

Isozymatic variation and phylogenetic relationships among extant henequén (Agave fourcroydes) germplasm and wild populations of its ancestor A. angustifolia in the Yucatan Peninsula in Mexico were analyzed. Analysis of three isozyme systems using starch gel electrophoresis indicated that while A. angustifolia populations have relatively high levels of variation, within each henequén cultivar all individuals were identical. This result corresponds to previous ethnobotanical and morphological analyses, which indicated severe loss of genetic variation of this domesticated plant as a consequence of the promotion by means of asexual propagation of only one cultivar since the middle of the last century. The three extant cultivars of henequén were distinct from each other. Two of them, Sac Ki (SK) and Yaax Ki (YK), could be matched within the progenitor, but Kitam Ki (KK) has a MDH electrophenotype not found in any of the plants growing inside the Yucatan Peninsula, but found in some A. angustifolia plants growing in the Mexican states of Oaxaca and Veracruz. A parsimony analysis of the morphological data indicated two lineages: that of SK and YK, cultivated cordage plants selected for stronger and longer fibers, whose sister group is the Tropical subdeciduous forest ecotype (SF), and that of all the other wild populations, which also included KK, the cultivated textile plants selected for finer fibers and nearly extinct in Yucatan. These results support the hypothesis of the yucatecan origin of SK and YK from the SF ecotype, as well as the hypothesis of a recent introduction of KK to the Yucatan Peninsula in a domestication trend that probably included also Chelem White (its cultivation being abandoned later).     

Genetic diversity and population structure in the US Upland cotton (Gossypium hirsutum L.)

Key message

Genetic diversity and population structure in the US Upland cotton was established and core sets of allelic richness were identified for developing association mapping populations in cotton.

Abstract

Elite plant breeding programs could likely benefit from the unexploited standing genetic variation of obsolete cultivars without the yield drag typically associated with wild accessions. A set of 381 accessions comprising 378 Upland (Gossypium hirsutum L.) and 3 G. barbadense L. accessions of the United States cotton belt were genotyped using 120 genome-wide SSR markers to establish the genetic diversity and population structure in tetraploid cotton. These accessions represent more than 100 years of Upland cotton breeding in the United States. Genetic diversity analysis identified a total of 546 alleles across 141 marker loci. Twenty-two percent of the alleles in Upland accessions were unique, specific to a single accession. Population structure analysis revealed extensive admixture and identified five subgroups corresponding to Southeastern, Midsouth, Southwest, and Western zones of cotton growing areas in the United States, with the three accessions of G. barbadense forming a separate cluster. Phylogenetic analysis supported the subgroups identified by STRUCTURE. Average genetic distance between G. hirsutum accessions was 0.195 indicating low levels of genetic diversity in Upland cotton germplasm pool. The results from both population structure and phylogenetic analysis were in agreement with pedigree information, although there were a few exceptions. Further, core sets of different sizes representing different levels of allelic richness in Upland cotton were identified. Establishment of genetic diversity, population structure, and identification of core sets from this study could be useful for genetic and genomic analysis and systematic utilization of the standing genetic variation in Upland cotton.     

Morphometric and genetic analyses differentiate Mesoamerican populations of the endangered stingless bee Melipona beecheii (Hymenoptera: Meliponidae) and support their conservation as two separate units

Phenotypic and molecular differences were previously found in populations of the endangered stingless bee Melipona beecheii from two extremes of its geographic range. In this study we combine the use of morphometric and molecular tools, with the aim of investigating patterns of phenotypic and molecular variation in populations across Mesoamerica. Morphometric analyses showed that bees from Mexico have significantly smaller body size compared with populations from Central America, forming two separated groups. Bayesian analysis of the ITS1 spacer of the ribosomal gene also showed the existence of two clusters: one composed by the Mexican populations, and another in which the Central American ones assembled (Guatemala, El Salvador, Nicaragua and Costa Rica). The combined results confirm the presence of two taxonomic units: one distributed in southern Mexico (ranging from the Yucatan peninsula to the north of Guatemala), and a Central American unit found from the southern part of Guatemala down to Costa Rica. These units should be considered separately under conservation programs and therefore, human assisted colony exchange between them should be avoided.     

Genetic mapping and QTL analysis of fiber-related traits in cotton (<Emphasis Type="Italic">Gossypium</Emphasis>)

Cotton, the leading natural fiber crop, is largely produced by two primary cultivated allotetraploid species known as Upland or American cotton (Gossypium hirsutum L.) and Pima or Egyptian cotton (G. barbadense L.). The allotetraploid species diverged from each other and from their diploid progenitors (A or D genome) through selection and domestication after polyploidization. To analyze cotton AD genomes and dissect agronomic traits, we have developed a genetic map in an F₂ population derived from interspecific hybrids between G. hirsutum L. cv. Acala-44 and G. barbadense L. cv. Pima S-7. A total of 392 genetic loci, including 333 amplified fragment length polymorphisms (AFLPs), 47 simple sequence repeats (SSRs), and 12 restriction fragment length polymorphisms (RFLPs), were mapped in 42 linkage groups, which span 3,287 cM and cover approximately 70% of the genome. Using chromosomal aneuploid interspecific hybrids and a set of 29 RFLP and SSR framework markers, we assigned 19 linkage groups involving 223 loci to 12 chromosomes. Comparing four pairs of homoeologous chromosomes, we found that with one exception linkage distances in the A-subgenome chromosomes were larger than those in their D-subgenome homoeologues, reflecting higher recombination frequencies and/or larger chromosomes in the A subgenome. Segregation distortion was observed in 30 out of 392 loci mapped in cotton. Moreover, approximately 29% of the RFLPs behaved as dominant loci, which may result from rapid genomic changes. The cotton genetic map was used for quantitative trait loci (QTL) analysis using composite interval mapping and permutation tests. We detected seven QTLs for six fiber-related traits; five of these were distributed among A-subgenome chromosomes, the genome donor of fiber traits. The detection of QTLs in both the A subgenome in this study and the D subgenome in a previous study suggests that fiber-related traits are controlled by the genes in homoeologous genomes, which are subjected to selection and domestication. Some chromosomes contain clusters of QTLs and presumably contribute to the large amount of phenotypic variation that is present for fiber-related traits.Communicated by J. Dvorak     

Ethnobotany of chia,Salvia hispanica L. (Lamiaceae)

Salvia hispanica L, was an important staple Mesoamerican food and medicinal plant in pre-Columbian times. Unlike other Mesoamerican pseudocereal crops such asAmaranthus andChenopodium, it has received comparatively little research attention. An ethnobotanical review of this Mesoamerican crop plantSalvia hispanica has been undertaken to examine changes in use accompanying Spanish colonization. A comparative analysis of accounts of use from the 16th century codices of Mexico and subsequent publications has revealed subtle changes in medicinal, culinary, artistic, and religious uses. Several hypotheses surrounding changes in use through time and the original use(s) that led to domestication are developed and tested through collection of ethnobotanical data in the highlands of western Mexico and Guatemala. A general decline in ethnobotanical knowledge associated with wild populations coupled with a loss of habitat in some locations has degraded important germplasm and knowledge resources for a species with great economic potential.     

Albumin Mexico (Al Me ) in the Guatemalan highlands

The examination of the sera of two samples of Guatemalans (total n = 386) revealed the presence of albumin Mexico (Al^Me) among unrelated individuals from the western highlands near the Mexican border. The frequency in the putatively less admixed sample, from Quiché, was 0.008. This is the first report of albumin polymorphism among American Indians south of Mexico. The presence of the Al^Me allele among Indians of the Guatemalan highlands argues for a zone of gene flow between contiguous groups of Mexico and Guatemala. The failure to detect albumin polymorphism among lowland populations of the Yucatan peninsula may likewise be significant, but more testing of samples is required before definitive statements can be made with respect to contacts among and within highland and lowland groups.     

Morphometric Analysis of Sunflower (<Emphasis Type="Italic">Helianthus annuus</Emphasis> L.) Achenes from Mexico and Eastern North America<Superscript>1</Superscript>

Morphometric Analysis of Sunflower ( Helianthus annuus L.) Achenes from Mexico and Eastern North America. Sunflower (Helianthus annuus L.) has played a major role in the evolution of agricultural systems in the Americas. The discovery of ancient domesticated remains from archaeological deposits in pre-Columbian Mexico offers new dimensions to widely accepted viewpoints on the domestication pattern of H. annuus. Although American sunflower populations north of Mexico have been examined extensively, Mexican indigenous domesticated landraces have not been studied in any detail. In this study, we morphologically assessed wild and domesticated sunflower achenes from Mexico and compared them to similar datasets from eastern North America. Additionally, we evaluated the utility of four computer-assisted shape measurements in discriminating between wild and domesticated sunflower achenes (fruits) and compared variation in achene size among modern wild and cultivated populations from both Mexico and the U.S. We found that, of the shape parameters tested, none were informative in distinguishing wild achenes from domesticated varieties. Subsequent size analysis, using conventional parameters of length, width, and thickness, showed that modern wild populations from Mexico had smaller achenes compared to modern populations from eastern North America. Domesticated achenes unearthed from Mexican archaeological sites, however, were significantly larger than the early domesticated specimens recovered from eastern North America. Our methodological results indicate that variation in archaeological sunflower achenes is better described by conventional size parameters rather than computerized shape analysis.     

Genetic structure and mating systems in wild and cultivated populations of Phaseolus coccineus and P. vulgaris (Fabaceae)

We present a study of the genetic variation, genetic structure, and outcrossing rate of two wild populations of Phaseolus coccineus ssp. formosus (both from Central Mexico), two cultivated populations of P. coccineus ssp. coccineus (one from Central Mexico, the other from Chiapas), and one cultivated population of P. vulgaris (from Central Mexico), using seven electrophorectic loci. All the populations of P. coccineus showed high and similar levels of genetic variation, without differences among wild and cultivated populations (H ranged between 0.187 to 0.271). In contrast, P. vulgaris has low levels of genetic variation (H = 0.041). The three P. coccineus populations from Central Mexico were genetically similar, while the cultivated P. coccineus from Chiapas and the cultivated P. vulgaris from Central Mexico were very different. In all the loci and all the populations for both species, the fixation indices (F) were positive and differed significantly from random mating expectation in 59% of the cases for P. coccineus. The highest F was the estimated for P. vulgaris. For P. coccineus, the multilocus outcrossing rate estimates ranged from 0.592 to 0.698, and these did not differ significantly among populations. For P. coccineus, we also estimated the outcrossing rate for each of 60 different families, showing great variation, with a majority of families with a t between 0.8 and 1, but a substantial fraction of the families with a t as low as 0.12–0.20. The domestication process has neither eroded the levels of genetic variation of the cultivated populations of P. coccineus nor changed its mating system.     

Ethnobotany and progressive domestication ofjaltomata (solanaceae) in mexico and central america

Members of Jaltomata are herbaceous perennials distributed from southwestern United States into Bolivia, and annuals in the Caribbean and Galapagos Islands. The described species do not fully represent the diversity of this genus. For example, the widespread species, J. procumbens, exhibits morphological variation in several regions of Mexico and Central America. These regions of variation may be in part the result of the plants association with humans. There is a very close match in the geographical distributions of the Tarahumara, Zapotec, Tzeltal, and Chuj cultures and the variant populations of Jaltomata in Chihuahua, Oaxaca, Chiapas (Mexico), and Huehuetenango (Guatemala), respectively. The results of the interaction of Jaltomata and humans are discussed in a general evolutionary context. Observations of Jaltomata among the Tarahumarain Chihuahua are presented as a starting point for further elucidation of progressive domestication. This encouraged weed is recognized as an intermediate between the wild and domesticate. Many vernacular names have been applied to Jaltomata some of these names are also applied to noncongeners (e.g., members of the Solanum nigrum complex).     

Phylogeography of the bark beetle Dendroctonus mexicanus Hopkins (Coleoptera: Curculionidae: Scolytinae)

Dendroctonus mexicanus is polyphagous within the Pinus genus and has a wide geographical distribution in Mexico and Guatemala. We examined the pattern of genetic variation across the range of this species to explore its demographic history and its phylogeographic pattern. Analysis of the mtDNA sequences of 173 individuals from 25 Mexican populations allowed to us identify 53 geographically structured haplotypes. High haplotype and low nucleotide diversities and Tajima’s D indicate that D. mexicanus experienced rapid population expansion during its dispersal across mountain systems within its current range. The nested clade phylogeographic analysis indicates that the phylogeographic pattern of D. mexicanus is explained by continuous dispersion among lineages from the Sierra Madre Occidental, the Sierra Madre Oriental and the Trans-Mexican Volcanic Belt. However, we also observed isolation events among haplotypes from the Cofre de Perote/Trans-Mexican Volcanic Belt/Sierra Madre Oriental and the Trans-Mexican Volcanic Belt/Sierra Madre del Sur, which is consistent with the present conformation of mountain systems in Mexico and the emergence of geographical barriers during the Pleistocene.     

A second spontaneous hybrid in the genusLeucaena (Leguminosae, Mimosoideae)

The detection of hybridity inLeucaena is discussed in relation to: (i) traditional criteria, (ii) molecular criteria and (iii) models to predict hybrid leaf morphology. Morphological, geographical and molecular evidence for the occurrence of interspecific hybrids betweenL. leucocephala andL. diversifolia in south-central Mexico, northern Guatemala, Jamaica, Dominican Republic, the Philippines and Papua New Guinea is presented. Predicted mean hybrid leaf trait values calculated from parent material are compared with data from putative hybrids and shown to be similar. The origin of these hybrids is discussed and shown to be the result of artificial sympatry resulting from indigenous, and recent exotic, domestication of the parent species. The hybrid is described asL. ×spontanea.     

Genetic diversity in and phylogenetic relationships of the Brazilian endemic cotton,Gossypium mustelinum (Malvaceae)

Gossypium mustelinum, one of five tetraploid species in the cotton genus, is geographically restricted to a few states in NE Brazil. Allozyme analysis was used to assess levels and patterns of genetic diversity inG. mustelinum and its relationship to the other tetraploid species. Genetic variation was low, with only 6 of 50 loci examined being polymorphic, a mean of 1.14 alleles per locus and a mean panmictic heterozygosity of 0.08. These estimates are low relative to other tetraploid cotton species, but are typical of island endemics. Interpopulational genetic identities were uniformly high, lending support to the concept of there being only one wild species of Brazilian cotton. The limited allelic diversity observed was correlated with geographical distribution, although variability is so limited in the species that geographically marginal populations are electrophoretically ordinary. Phylogenetic and phenetic analyses demonstrate thatG. mustelinum is isolated among polyploid cotton species, occupying one of the three basal clades resulting from an early radiation of polyploid taxa subsequent to polyploid formation. We suggest thatG. mustelinum represents a paleoendemic that presently exists as a series of widely scattered, relictual populations. Despite several centuries of sympatric cultivation ofG. barbadense andG. hirsutum, there was little evidence of interspecific introgression of alleles from cultivated cottons intoG. mustelinum.     

Patterns of morphological variation,diversity, and domestication of wild and cultivated populations of agave in Yucatan,Mexico

This work presents a statistical and numerical analysis of the patterns of morphological variation of the cultivated variants of henequén (Agave fourcroydes Lem.) presently found in the Mexican state of Yucatan and of the wild populations of A. angustifolia Haw., its putative progenitor. This is the first step in the study of the intrageneric genetic diversity and evolutionary relationships. The study indicated that: (1) There exists a significant discontinuation in morphological variation corresponding to the cultivated variants of traditionally recognized henequén: Sac Ki, Yaax Ki, and Kitam Ki, and to three possible ecotypes of A. angustifolia Haw.: Coastal Dunes, Tropical Deciduous Forest, and Tropical Subdeciduous Forest. (2) Sac Ki and Yaax Ki differ from wild populations in four syndromes of domestication: gigantism, greater fibrosity, less thorniness, and less reproductive capacity. The lower coefficient of variation of their characteristics compared with the wild populations suggests less genetic diversity. This fact, and the disappearance of four out of the seven variants existing early in this century, indicate a dramatic genetic erosion of this crop. (3) Kitam Ki is the cultivated variant more similar to wild ones. Differences with them suggest recent introduction and an artificial selection process with different direction and intensity than the other cultivated variants. (4) A tendency from more to less is observed for characteristics indicating degree of domestication: Sac Ki, Yaax Ki, and Kitam Ki. (5) The differences among the possible wild ecotypes may be associated with the soil conditions and precipitation.     

Morphometric analysis of Triatoma dimidiata populations (Reduviidae:Triatominae) from Mexico and Northern Guatemala

Triatoma dimidiata is one of the major vectors of Chagas disease in Latin America. Its range includes Mexico, all countries of Central America, Colombia, and Ecuador. In light of recent genetic analysis suggesting that the possible origin of this species is the Yucatan peninsula, we have analyzed populations from the state of Yucatan, San Luis Potosi, and Veracruz in Mexico, and a population from the southern region of the Yucatan peninsula located in Northern Guatemala, the region of El Peten. Classical morphometry including principal component, discriminant, sexual dimorphism, and wing asymmetry was analyzed. San Luis Potosi and Veracruz populations were indistinguishable while clearly separate from Yucatan and Peten populations. Despite important genetic differences, Yucatan and Peten populations were highly similar. Yucatan specimens were the smallest in size, while females were larger than males in all populations. Only head characters were necessary to distinguish population level differences, although wing fluctuating asymmetry was present in all populations. These results are discussed in light of recent findings suggesting genetic polymorphism in most populations of Triatoma dimidiata south of Chiapas to Ecuador.