Physiological traits contribute to growth and adaptation of Mexican maize landraces.

Local adaptation of populations results from an interplay between their environment and genetics. If functional trait variation influences plant performance, populations can adapt to their local environment. However, populations may also respond plastically to environmental challenges, altering phenotype without shifting allele frequencies. The level of local adaptation in crop landraces and their capacity for plasticity in response to environmental change may predict their continued utility to farmers facing climate change. Yet we understand little about how physiological traits potentially underlying local adaptation of cultivars influence fitness. Farmers in Mexico-the crop center of origin for maize-manage and rely upon a high diversity of landraces. We studied maize grown in Chiapas, Mexico, where strong elevational gradients cover a relatively small geographic area. We reciprocally transplanted 12 populations sourced from three elevational zones (600, 1550 and 2150 m) back into those elevations for two years using a modified split-split plot design to model effects of environment, genetics, and their interaction. We studied physiological and growth traits, including photosynthetic rate, stomatal conductance, stomatal density, relative growth rate (RGR), and seed production. Maize fitness showed indications of local adaptation with highland and midland types performing poorly at warmer lowland locations, though patterns depended on the year. Several physiological traits, including stomatal conductance, were affected by G x E interactions, some of which indicated non-adaptive plastic responses with potential fitness implications. We discerned a significant positive relationship between fitness and relative growth rate. Growth rates in highland landraces were outperformed by midland and lowland landraces grown in high temperature, lowland garden. Lowland landrace stomatal conductance was diminished compared to that of highland landraces in the cooler highland garden. Thus, both adaptive and non-adaptive physiological responses of maize landraces in southern Mexico may have implications for fitness, as well as responses to climate change.

Crop genetic erosion: understanding and responding to loss of crop diversity.

Crop diversity underpins the productivity, resilience and adaptive capacity of agriculture. Loss of this diversity, termed crop genetic erosion, is therefore concerning. While alarms regarding evident declines in crop diversity have been raised for over a century, the magnitude, trajectory, drivers and significance of these losses remain insufficiently understood. We outline the various definitions, measurements, scales and sources of information on crop genetic erosion. We then provide a synthesis of evidence regarding changes in the diversity of traditional crop landraces on farms, modern crop cultivars in agriculture, crop wild relatives in their natural habitats and crop genetic resources held in conservation repositories. This evidence indicates that marked losses, but also maintenance and increases in diversity, have occurred in all these contexts, the extent depending on species, taxonomic and geographic scale, and region, as well as analytical approach. We discuss steps needed to further advance knowledge around the agricultural and societal significance, as well as conservation implications, of crop genetic erosion. Finally, we propose actions to mitigate, stem and reverse further losses of crop diversity.

Transcriptional differentiation of UV-B protectant genes in maize landraces spanning an elevational gradient in Chiapas, Mexico.

Globally, farmers cultivate and maintain crop landraces (i.e., traditional varieties). Landraces contain unique diversity shaped in part by natural and human-mediated selection and are an indispensable resource for farmers. Since environmental conditions change with elevation, crop landraces grown along elevational gradients have provided ideal locations to explore patterns of local adaptation. To further probe traits underlying this differentiation, transcriptome signatures can help provide a foundation for understanding the ways in which functional genetic diversity may be shaped by environment. In this study, we returned to an elevational gradient in Chiapas, Mexico, to assess transcriptional differentiation of genes underlying UV-B protection in locally adapted maize landraces from multiple elevations. We collected and planted landraces from three elevational zones (lowland, approximately 600 m; midland, approximately 1,550 m; highland approximately 2,100 m) in a common garden at 1,531 m. Using RNA-seq data derived from leaf tissue, we performed differential expression analysis between maize from these distinct elevations. Highland and lowland landraces displayed differential expression in phenylpropanoid and flavonoid biosynthesis genes involved in the production of UV-B protectants and did so at a rate greater than expected based on observed background transcriptional differentiation across the genome. These findings provide evidence for the differentiation of suites of genes involved in complex ecologically relevant pathways. Thus, while neutral evolutionary processes may have played a role in the observed patterns of differentiation, UV-B may have also acted as a selective pressure to differentiate maize landraces in the region. Studies of the distribution of functional crop genetic diversity across variable landscapes can aid us in understanding the response of diversity to abiotic/biotic change and, ultimately, may facilitate its conservation and utilization.

Evolutionary and food supply implications of ongoing maize domestication by Mexican campesinos.

Maize evolution under domestication is a process that continues today. Case studies suggest that Mexican smallholder family farmers, known as campesinos, contribute importantly to this, but their significance has not been explicitly quantified and analysed as a whole. Here, we examine the evolutionary and food security implications of the scale and scope under which campesinos produce maize. We gathered official municipal-level data on maize production under rainfed conditions and identified campesino agriculture as occurring in municipalities with average yields of less than or equal to 3 t ha-1 Environmental conditions vary widely in those municipalities and are associated with a great diversity of maize races, representing 85.3% of native maize samples collected in the country. We estimate that in those municipalities, around 1.38 × 1011 genetically different individual plants are subjected to evolution under domestication each season. This implies that 5.24 × 108 mother plants contribute to the next generation with their standing genetic diversity and rare alleles. Such a large breeding population size also increases the total number of adaptive mutations that may appear and be selected for. We also estimate that campesino agriculture could potentially feed around 54.7 million people in Mexico. These analyses provide insights about the contributions of smallholder agriculture around the world.

Fitness of Mass-Reared Males of Anastrepha obliqua (Diptera: Tephritidae) Resulting From Mating Competition Tests in Field Cages.

The sterile insect technique uses males that have been mass-reared in a controlled environment. The insects, once released in the field, must compete to mate. However, the mass-rearing condition supposes a loss of fitness that will be noticeable by wild females. To compare the fitness of wild males and mass-reared males, three competition settings were established. In setting 1, wild males, mass-reared males and wild females were released in field cages. In setting 2, wild females and wild males were released without competition, and in setting 3, mass-reared males and mass-reared females were also released without competition. Male fitness was based on their mating success, fecundity, weight and longevity. The fitness of the females was measured based on weight and several demographic parameters. The highest percentage of mating was between wild males and wild females between 0800 and 0900 h in the competition condition, while the mass-reared males started one hour later. The successful wild males weighed more and showed longer mating times, greater longevity and a higher number of matings than the mass-reared males. Although the mass-reared males showed the lowest percentage of matings, their fecundity when mating with wild females indicated a high fitness. Since the survival and fecundity of wild females that mated with mass-reared males decreased to become similar to those of mass-reared females that mated with mass-reared males, females seem to be influenced by the type of male (wild or mass-reared).

Differentiated transcriptional signatures in the maize landraces of Chiapas, Mexico.

BACKGROUND: Landrace farmers are the keepers of crops locally adapted to the environments where they are cultivated. Patterns of diversity across the genome can provide signals of past evolution in the face of abiotic and biotic change. Understanding this rich genetic resource is imperative especially since diversity can provide agricultural security as climate continues to shift. RESULTS: Here we employ RNA sequencing (RNA-seq) to understand the role that conditions that vary across a landscape may have played in shaping genetic diversity in the maize landraces of Chiapas, Mexico. We collected landraces from three distinct elevational zones and planted them in a midland common garden. Early season leaf tissue was collected for RNA-seq and we performed weighted gene co-expression network analysis (WGCNA). We then used association analysis between landrace co-expression module expression values and environmental parameters of landrace origin to elucidate genes and gene networks potentially shaped by environmental factors along our study gradient. Elevation of landrace origin affected the transcriptome profiles. Two co-expression modules were highly correlated with temperature parameters of landrace origin and queries into their 'hub' genes suggested that temperature may have led to differentiation among landraces in hormone biosynthesis/signaling and abiotic and biotic stress responses. We identified several 'hub' transcription factors and kinases as candidates for the regulation of these responses. CONCLUSIONS: These findings indicate that natural selection may influence the transcriptomes of crop landraces along an elevational gradient in a major diversity center, and provide a foundation for exploring the genetic basis of local adaptation. While we cannot rule out the role of neutral evolutionary forces in the patterns we have identified, combining whole transcriptome sequencing technologies, established bioinformatics techniques, and common garden experimentation can powerfully elucidate structure of adaptive diversity across a varied landscape. Ultimately, gaining such understanding can facilitate the conservation and strategic utilization of crop genetic diversity in a time of climate change.

Mapping the diversity of maize races in Mexico.

Traditional landraces of maize are cultivated throughout more than one-half of Mexico's cropland. Efforts to organize in situ conservation of this important genetic resource have been limited by the lack of knowledge of regional diversity patterns. We used recent and historic collections of maize classified for race type to determine biogeographic regions and centers of landrace diversity. We also analyzed how diversity has changed over the last sixty years. Based on racial composition of maize we found that Mexico can be divided into 11 biogeographic regions. Six of these biogeographic regions are in the center and west of the country and contain more than 90% of the reported samples for 38 of the 47 races studied; these six regions are also the most diverse. We found no evidence of rapid overall decline in landrace diversity for this period. However, several races are now less frequently reported and two regions seem to support lower diversity than in previous collection periods. Our results are consistent with a previous hypothesis for diversification centers and for migration routes of original maize populations merging in western central Mexico. We provide maps of regional diversity patterns and landrace based biogeographic regions that may guide efforts to conserve maize genetic resources.

Evolutionary response of landraces to climate change in centers of crop diversity.

Landraces cultivated in centers of crop diversity result from past and contemporary patterns of natural and farmer-mediated evolutionary forces. Successful in situ conservation of crop genetic resources depends on continuity of these evolutionary processes. Climate change is projected to affect agricultural production, yet analyses of impacts on in situ conservation of crop genetic diversity and farmers who conserve it have been absent. How will crop landraces respond to alterations in climate? We review the roles that phenotypic plasticity, evolution, and gene flow might play in sustaining production, although we might expect erosion of genetic diversity if landrace populations or entire races lose productivity. For example, highland maize landraces in southern Mexico do not express the plasticity necessary to sustain productivity under climate change, but may evolve in response to altered conditions. The outcome for any given crop in a given region will depend on the distribution of genetic variation that affects fitness and patterns of climate change. Understanding patterns of neutral and adaptive diversity from the population to the landscape scale is essential to clarify how landraces conserved in situ will continue to evolve and how to minimize genetic erosion of this essential natural resource.

Dispersal of transgenes through maize seed systems in Mexico.

OBJECTIVES: Current models of transgene dispersal focus on gene flow via pollen while neglecting seed, a vital vehicle for gene flow in centers of crop origin and diversity. We analyze the dispersal of maize transgenes via seeds in Mexico, the crop's cradle. METHODS: We use immunoassays (ELISA) to screen for the activity of recombinant proteins in a nationwide sample of farmer seed stocks. We estimate critical parameters of seed population dynamics using household survey data and combine these estimates with analytical results to examine presumed sources and mechanisms of dispersal. RESULTS: Recombinant proteins Cry1Ab/Ac and CP4/EPSPS were found in 3.1% and 1.8% of samples, respectively. They are most abundant in southeast Mexico but also present in the west-central region. Diffusion of seed and grain imported from the United States might explain the frequency and distribution of transgenes in west-central Mexico but not in the southeast. CONCLUSIONS: Understanding the potential for transgene survival and dispersal should help design methods to regulate the diffusion of germplasm into local seed stocks. Further research is needed on the interactions between formal and informal seed systems and grain markets in centers of crop origin and diversification.

Asymmetrical local adaptation of maize landraces along an altitudinal gradient.

Crop landraces are managed populations that evolve in response to gene flow and selection. Cross-pollination among fields, seed sharing by farmers, and selection by management and environmental conditions play roles in shaping crop characteristics. We used common gardens to explore the local adaptation of maize (Zea mays ssp. mays) landrace populations from Chiapas, Mexico to altitude. We sowed seeds of 21 populations from three altitudinal ranges in two common gardens and measured two characteristics that estimate fitness: likelihood of producing good quality seed and the total mass of good quality seed per plant. The probability of lowland plants producing good quality seed was invariably high regardless of garden, while highland landraces were especially sensitive to altitude. Their likelihood of producing good seed quadrupled in the highland site. The mass of good quality seed showed a different pattern, with lowland landraces producing 25% less seed mass than the other types at high elevations. Combining these two measures of fitness revealed that the highland landraces were clearly adapted to highland sites, while lowland and midland landraces appear more adapted to the midland site. We discuss this asymmetry in local adaptation in light of climate change and in situ conservation of crop genetic resources.

Maize diversity and ethnolinguistic diversity in Chiapas, Mexico.

The objective of this study is to investigate whether ethnolinguistic diversity influences crop diversity. Factors suggest a correlation between biological diversity of crops and cultural diversity. Although this correlation has been noted, little systematic research has focused on the role of culture in shaping crop diversity. This paper reports on research in the Maya highlands (altitude >1,800 m) of central Chiapas in southern Mexico that examined the distribution of maize (Zea mays) types among communities of two groups, the Tzeltal and Tzotzil. The findings suggest that maize populations are distinct according to ethnolinguistic group. However, a study of isozymes indicates no clear separation of the region's maize into two distinct populations based on ethnolinguistic origin. A reciprocal garden experiment shows that there is adaptation of maize to its environment but that Tzeltal maize sometimes out-yields Tzotzil maize in Tzotzil environments. Because of the proximity of the two groups and selection for yield, we would expect that the superior maize would dominate both groups' maize populations, but we find that such domination is not the case. The role of ethnolinguistic identity in shaping social networks and information exchange is discussed in relation to landrace differentiation.