A B73 x Palomero Toluqueño mapping population reveals local adaptation in Mexican highland maize

Generations of farmer selection in the central Mexican highlands have produced unique maize varieties adapted to the challenges of the local environment. In addition to possessing great agronomic and cultural value, Mexican highland maize represents a good system for the study of local adaptation and acquisition of adaptive phenotypes under cultivation. In this study we characterize a recombinant inbred line population derived from the B73 reference line and the Mexican highland maize variety Palomero Toluqueño. B73 and Palomero Toluqueño showed classic rank-changing differences in performance between lowland and highland field sites, indicative of local adaptation. Quantitative trait mapping identified genomic regions linked to effects on yield components that were conditionally expressed depending on the environment. For the principal genomic regions associated with ear weight and total kernel number, the Palomero Toluqueño allele conferred an advantage specifically in the highland site, consistent with local adaptation. We identified Palomero Toluqueño alleles associated with expression of characteristic highland traits, including reduced tassel branching, increased sheath pigmentation and the presence of sheath macrohairs. The oligogenic architecture of these three morphological traits supports their role in adaptation, suggesting they have arisen from consistent directional selection acting at distinct points across the genome. We discuss these results in the context of the origin of phenotypic novelty during selection, commenting on the role of de novo mutation and the acquisition of adaptive variation by gene flow from endemic wild relatives.

A B73 x Palomero Toluqueo mapping population reveals local adaptation in Mexican highland maize

Generations of farmer selection have produced a unique collection of traditional maize varieties adapted to the environmental challenges of the central Mexican highlands. In addition to agronomic and cultural value, Mexican highland maize represents a good system for the study of local adaptation and acquisition of adaptive phenotypes under cultivation. In this study, we characterized a recombinant inbred line population derived from the cross of the B73 reference line and the Mexican highland maize variety Palomero Toluqueo. Evaluation over multiple years in lowland and highland field sites in Mexico identified genomic regions linked to yield components and putatively adaptive morphological traits. A region on chromosome 7 associated with ear weight showed antagonistic allelic effects in lowland and highland fields, suggesting a trade-off consistent with local adaptation. We identified several alleles of highland origin associated with characteristic highland traits, including reduced tassel branching, increased stem pigmentation and the presence of stem macrohairs. The oligogenic architecture of characteristic morphological traits supports their role in adaptation, suggesting they have arisen from consistent directional selection acting at distinct points across the genome. We discuss these results in the context of the origin of phenotypic novelty during selection, commenting on the role of de novo mutation and the acquisition of adaptive variation by gene flow from endemic wild relatives.

Teosinte introgression modulates phosphatidylcholine levels and induces early maize flowering time

After domestication from lowland teosinte parviglumis (Zea mays ssp.parviglumis) in the warm Mexican southwest, maize (Zea mays ssp. mays) colonized the highlands of México and South America. In the highlands, maize was exposed to lower temperatures that imposed strong selection on flowering time. Phospholipids are important metabolites in plant responses to low-temperature, low phosphorus availability and have also been suggested to influence flowering time.Here, we combined linkage mapping analysis with genome scans to identify High PhosphatidylCholine 1 (HPC1), a gene which encodes a phospholipase A1 enzyme, as a major driver of phospholipid variation in highland maize. Common garden experiments demonstrated strong genotype-by-environment interactions associated with variation at HPC1, with the highland HPC1 allele leading to higher fitness in highlands, possibly by hastening flowering.The highland maize HPC1 variant results in impaired function of the encoded protein due to a polymorphism in a highly conserved sequence. A meta-analysis indicated a strong association between the identity of the amino acid at this position and optimal growth in prokaryotes. Mutagenesis of HPC1 via genome editing validated its role in regulating phospholipid metabolism.Finally, we showed that the highland HPC1 allele entered cultivated maize by introgression from the wild highland teosinte Zea mays ssp. mexicana and has been maintained in maize breeding lines from Northern US, Canada and Europe.Thus, HPC1 introgressed from teosinte mexicana underlies a large metabolic QTL that modulates phosphatidylcholine levels and has an adaptive effect at least in part via induction of early flowering time..

Gene regulatory effects of a large chromosomal inversion in highland maize

Chromosomal inversions play an important role in local adaptation. Inversions can capture multiple locally adaptive functional variants in a linked block by repressing recombination. However, this recombination suppression makes it difficult to identify the genetic mechanisms underlying an inversion’s role in adaptation. In this study, we used large-scale transcriptomic data to dissect the functional importance of a 13 Mb inversion locus (Inv4m) found almost exclusively in highland populations of maize (Zea mays ssp. mays). Inv4m was introgressed into highland maize from the wild relative Zea mays ssp. mexicana, also present in the highlands of Mexico, and is thought to be important for the adaptation of these populations to cultivation in highland environments. However, the specific genetic variants and traits that underlie this adaptation are not known. We created two families segregating for the standard and inverted haplotypes of Inv4m in a common genetic background and measured gene expression effects associated with the inversion across 9 tissues in two experimental conditions. With these data, we quantified both the global transcriptomic effects of the highland Inv4m haplotype, and the local cis-regulatory variation present within the locus. We found diverse physiological effects of Inv4m across the 9 tissues, including a strong effect on the expression of genes involved in photosynthesis and chloroplast physiology. Although we could not confidently identify the causal alleles within Inv4m, this research accelerates progress towards understanding this inversion and will guide future research on these important genomic features.

EFFECTS OF FARMERS’ KNOWLEDGE, ATTITUDE, SKILLS ON HIGHLAND MAIZE ADOPTION IN WESTERN ETHIOPIA

Understanding farmers' indigenous knowledge is vital in hybrid maize popularization. Despite different opportunities, adoption was constrained by the failure of infusing local knowledge in the modern extension system. Hence, farmers' capabilities, preferences, and practices towards adoption were studied. Data were collected from key informants and focus group discussion participants. A cross-sectional survey was to collect data from 154 respondents. Knowledge and attitude were assessed by using a 5-point Likert scale. Descriptive statistics and econometric analyses were run to analyze data accordingly. The results indicated among demographic characteristics; family and land size, owning of ox and experience have positively affected highland maize adoption at 1 per cent significance level; while education, age, and on-farm income have positively affected highland maize adoption at a 1% significance level. However, religion and sex did not affect highland maize adoption at all. Pearson chi-square result indicated, there was a positive and significant relationship of knowledge (χ2=41.49; p=0.000) to adoption. Consequently, an increase in farmers' knowledge of favoured adoption. Finally, poor institutional support, insufficient involvement of resource-poor farmers, and lack of training were major bottlenecks hampering highland maize adoption. Hence, provision of special training, credit services, and farmers-responsive training should be in place for better adoption.

Molecular Parallelism Underlies Convergent Highland Adaptation of Maize Landraces

Convergent phenotypic evolution provides some of the strongest evidence for adaptation. However, the extent to which recurrent phenotypic adaptation has arisen via parallelism at the molecular level remains unresolved, as does the evolutionary origin of alleles underlying such adaptation. Here, we investigate genetic mechanisms of convergent highland adaptation in maize landrace populations and evaluate the genetic sources of recurrently selected alleles. Population branch excess statistics reveal strong evidence of parallel adaptation at the level of individual SNPs, genes and pathways in four independent highland maize populations, even though most SNPs show unique patterns of local adaptation. The majority of selected SNPs originated via migration from a single population, most likely in the Mesoamerican highlands. Polygenic adaptation analyses of quantitative traits reveal that alleles affecting flowering time are significantly associated with elevation, indicating the flowering time pathway was targeted by highland adaptation. In addition, repeatedly selected genes were significantly enriched in the flowering time pathway, indicating their significance in adapting to highland conditions. Overall, our study system represents a promising model to study convergent evolution in plants with potential applications to crop adaptation across environmental gradients.

Development and Release of Two Mid-Late Maturing Maize Hybrids for the Highland Agro-Ecologies of Uganda

The maize yield in the highland areas of Uganda (1500 to 3000 masl), rarely exceeding 2 t ha-1. The use of unimproved maize genotypes, the prevalence of pests and diseases, and low levels of input use has tremendously contributed to the low grain yields. Accordingly, we obtained 100 maize inbred lines majorly from CIMMYT in the year 2013. The inbred lines were evaluated for adaptability and those selected were intercrossed to generate 80 single cross hybrids by January 2015. The 80 single cross hybrids were evaluated in preliminary yield trials in three sites namely, Kachwekano, Kalengyere and Buginyanya in 2015a season. The best combiners were crossed with the selected males to obtain 122 three-way cross hybrids. These were evaluated for adaptability in advanced yield trials in 2016a in Kachwekano, Kalengyere and Bukwo, from where the best five hybrids (BH33, BH31, BH27, BH104 and BH38) were selected for National Performance Trials (NPT) in 2016b and 2017a seasons in Bukwo, Buginyanya, Zombo, Fort portal and Kalengere. Two candidate varieties namely BH31 and BH33 exhibited good agronomic qualities such as earlier maturity, tolerance to MLN and other diseases. The selected hybrids yielded 8.2 t ha-1 and 7.2 t ha-1 for BH31 and BH33, respectively. The two varieties are appropriate for cultivation in major highland and transitional maize growing regions in Uganda and were subsequently released as NAROMAIZE 731 and NAROMAIZE 733, respectively. The newly released varieties are expected to reduce the highland maize cropping season from over six months to five months.

Gene regulatory effects of a large chromosomal inversion in highland maize

Chromosomal inversions play an important role in local adaptation. Inversions can capture multiple locally adaptive functional variants in a linked block by repressing recombination. However, this recombination suppression makes it difficult to identify the genetic mechanisms that underlie an inversion’s role in adaption. In this study, we explore how large-scale transcriptomic data can be used to dissect the functional importance of a 13 Mb inversion locus (Inv4m) found almost exclusively in highland populations of maize (Zea mays ssp. mays). Inv4m introgressed into highland maize from the wild relative Zea mays ssp. mexicana, also present in the highlands of Mexico, and is thought to be important for the adaptation of these populations to cultivation in highland environments. First, using a large publicly available association mapping panel, we confirmed that Inv4m is associated with locally adaptive agronomic phenotypes, but only in highland fields. Second, we created two families segregating for standard and inverted haplotypess of Inv4m in a isogenic B73 background, and measured gene expression variation association with Inv4m across 9 tissues in two experimental conditions. With these data, we quantified both the global transcriptomic effects of the highland Inv4m haplotype, and the local cis-regulatory variation present within the locus. We found diverse physiological effects of Inv4m, and speculate that the genetic basis of its effects on adaptive traits is distributed across many separate functional variants.Author SummaryChromosomal inversions are an important type of genomic structural variant. However, mapping causal alleles within their boundaries is difficult because inversions suppress recombination between homologous chromosomes. This means that inversions, regardless of their size, are inherited as a unit. We leveraged the high-dimensional phenotype of gene expression as a tool to study the genetics of a large chromosomal inversion found in highland maize populations in Mexico - Inv4m. We grew plants carrying multiple versions of Inv4m in a common genetic background, and quantified the transcriptional reprogramming induced by alternative alleles at the locus. Inv4m has been shown in previous studies to have a large effect on flowering, but we show that the functional variation within Inv4m affects many developmental and physiological processes.Author ContributionsT. Crow, R. Rellan-Alvarez, R. Sawers and D. Runcie conceived and designed the experiment. M. Aguilar-Rangel, J. Rodrǵuez, R. Rellan-Alvarez and R. Sawers generated the segregating families. T. Crow, J. Ta, S. Nojoomi, M. Aguilar-Rangel, J. Rodrǵuez D. Gates, D. Runcie performed the experiment. T. Crow, D. Gates, D. Runcie analyzed the data. T. Crow, D. Runcie wrote the original manuscript, and R. Rellan-Alvarez and R. Sawers provided review and editing.