Does millet consumption contribute to raising blood hemoglobin levels compared to regular refined staples?: a systematic review and meta-analysis.

Millets are recognized for their health and nutritional values, and the United Nations declared 2023 the International Year of Millets. Among the several health and nutritional benefits of millets, their impact on hemoglobin concentration is important since anemia is a major public health issue in many countries. To investigate the effect of millet (including sorghum) consumption on hemoglobin concentration in the blood, a systematic review and meta-analysis were conducted. Thirteen published studies featuring randomized control trials involving 590 individuals in the intervention group and 549 control individuals were eligible for the meta-analysis. The difference-in-differences analysis revealed highly significant (p < 0.01) positive effects of millet consumption on hemoglobin concentration, with an effect size of +0.68 standardized mean difference units. The change in hemoglobin concentration observed in the intervention group was +13.6%, which is statistically significant (p < 0.0005), compared to that in the control group, which was +4.8% and not statistically significant (p = 0.1362). In four studies, the consumption of millets in the intervention group demonstrated a change from mild anemia to normal status among children, whereas there was no change in the control group. The findings provide evidence that the consumption of millets can improve blood hemoglobin concentration, likely resulting from increased iron intake. Further research is needed involving the assessment of iron content and bioavailability to better understand the effect variation among millet types and the mechanisms involved.

Genetic and genomic interventions in crop biofortification: Examples in millets.

Micronutrient malnutrition is a serious threat to the developing world's human population, which largely relies on a cereal-based diet that lacks diversity and micronutrients. Besides major cereals, millets represent the key sources of energy, protein, vitamins, and minerals for people residing in the dryland tropics and drought-prone areas of South Asia and sub-Saharan Africa. Millets serve as multi-purpose crops with several salient traits including tolerance to abiotic stresses, adaptation to diverse agro-ecologies, higher productivity in nutrient-poor soils, and rich nutritional characteristics. Considering the potential of millets in empowering smallholder farmers, adapting to changing climate, and transforming agrifood systems, the year 2023 has been declared by the United Nations as the International Year of Millets. In this review, we highlight recent genetic and genomic innovations that can be explored to enhance grain micronutrient density in millets. We summarize the advances made in high-throughput phenotyping to accurately measure grain micronutrient content in cereals. We shed light on genetic diversity in millet germplasm collections existing globally that can be exploited for developing nutrient-dense and high-yielding varieties to address food and nutritional security. Furthermore, we describe the progress made in the fields of genomics, proteomics, metabolomics, and phenomics with an emphasis on enhancing the grain nutritional content for designing competitive biofortified varieties for the future. Considering the close genetic-relatedness within cereals, upcoming research should focus on identifying the genetic and genomic basis of nutritional traits in millets and introgressing them into major cereals through integrated omics approaches. Recent breakthroughs in the genome editing toolbox would be crucial for mainstreaming biofortification in millets.

Biotechnological interventions for improving the seed longevity in cereal crops: progress and prospects.

Seed longevity is a measure of the viability of seeds during long-term storage and is crucial for germplasm conservation and crop improvement programs. Also, longevity is an important trait for ensuring food and nutritional security. Thus, a better understanding of various factors regulating seed longevity is requisite to improve this trait and to minimize the genetic drift during the regeneration of germplasm. In particular, seed deterioration of cereal crops during storage adversely affects agricultural productivity and food security. The irreversible process of seed deterioration involves a complex interplay between different genes and regulatory pathways leading to: loss of DNA integrity, membrane damage, inactivation of storage enzymes and mitochondrial dysfunction. Identifying the genetic determinants of seed longevity and manipulating them using biotechnological tools hold the key to ensuring prolonged seed storage. Genetics and genomics approaches had identified several genomic regions regulating the longevity trait in major cereals such as: rice, wheat, maize and barley. However, very few studies are available in other Poaceae members, including millets. Deploying omics tools, including genomics, proteomics, metabolomics, and phenomics, and integrating the datasets will pinpoint the precise molecular determinants affecting the survivability of seeds. Given this, the present review enumerates the genetic factors regulating longevity and demonstrates the importance of integrated omics strategies to dissect the molecular machinery underlying seed deterioration. Further, the review provides a roadmap for deploying biotechnological approaches to manipulate the genes and genomic regions to develop improved cultivars with prolonged storage potential.

Corrigendum: Grain nutrients variability in pigeonpea genebank collection and its potential for promoting nutritional security in dryland ecologies.

[This corrects the article DOI: 10.3389/fpls.2022.934296.].

Grain Nutrients Variability in Pigeonpea Genebank Collection and Its Potential for Promoting Nutritional Security in Dryland Ecologies.

Pigeonpea, a climate-resilient legume, is nutritionally rich and of great value in Asia, Africa, and Caribbean regions to alleviate malnutrition. Assessing the grain nutrient variability in genebank collections can identify potential sources for biofortification. This study aimed to assess the genetic variability for grain nutrients in a set of 600 pigeonpea germplasms conserved at the RS Paroda Genebank, ICRISAT, India. The field trials conducted during the 2019 and 2020 rainy seasons in augmented design with four checks revealed significant differences among genotypes for all the agronomic traits and grain nutrients studied. The germplasm had a wider variation for agronomic traits like days to 50% flowering (67-166 days), days to maturity (112-213 days), 100-seed weight (1.69-22.17 g), and grain yield per plant (16.54-57.93 g). A good variability was observed for grain nutrients, namely, protein (23.35-29.50%), P (0.36-0.50%), K (1.43-1.63%), Ca (1,042.36-2,099.76 mg/kg), Mg (1,311.01-1,865.65 mg/kg), Fe (29.23-40.98 mg/kg), Zn (24.14-35.68 mg/kg), Mn (8.56-14.01 mg/kg), and Cu (7.72-14.20 mg/kg). The germplasm from the Asian region varied widely for grain nutrients, and the ones from African region had high nutrient density. The significant genotype × environment interaction for most of the grain nutrients (except for P, K, and Ca) indicated the sensitivity of nutrient accumulation to the environment. Days to 50% flowering and days to maturity had significant negative correlation with most of the grain nutrients, while grain yield per plant had significant positive correlation with protein and magnesium, which can benefit simultaneous improvement of agronomic traits with grain nutrients. Clustering of germplasms based on Ward.D2 clustering algorithm revealed the co-clustering of germplasm from different regions. The identified top 10 nutrient-specific and 15 multi-nutrient dense landraces can serve as promising sources for the development of biofortified lines in a superior agronomic background with a broad genetic base to fit the drylands. Furthermore, the large phenotypic data generated in this study can serve as a raw material for conducting SNP/haplotype-based GWAS to identify genetic variants that can accelerate genetic gains in grain nutrient improvement.

Can Feeding a Millet-Based Diet Improve the Growth of Children?-A Systematic Review and Meta-Analysis.

Undernutrition, such as stunting and underweight, is a major public health concern, which requires multi-sectoral attention. Diet plays a key role in growth and should optimally supply all required nutrients to support the growth. While millets (defined broadly to include sorghum) are traditional foods, and climate smart nutritious crops, which are grown across Africa and Asia, they have not been mainstreamed like rice, wheat, and maize. Diversifying staples with millets can potentially provide more macro and micro nutrients, compared to the mainstream crops. However, there is little known scientific evidence to prove millets' efficacy on growth. Therefore, a systematic review and meta-analysis was conducted to collate evidence of the benefits of millets in improving the growth of children. Eight eligible randomized feeding trials were included in the meta-analysis. Results from the randomized effect model showed a significant effect (p < 0.05) of millet-based diets on mean height (+28.2%) (n = 8), weight (n = 9) (+26%), mid upper arm circumference (n = 5) (+39%) and chest circumference (n = 5) (+37%) in comparison to regular rice-based diets over for the period of 3 months to 4.5 years, which was based on largely substituting rice with millets. When an enhanced and diverse diet was served, replacing rice with millet had only minimal growth improvement on chest circumference (p < 0.05). The quality assessment using GRADE shows that the evidence used for this systematic review and meta-analysis had moderate quality, based on eight scoring criteria. These results demonstrate the value of adding millet as the staple for undernourished communities. Further understanding of the efficacy of millets on growth in a wider range of diets is important to develop appropriate dietary programs and improve the nutritional status of various age groups across Africa and Asia.

Millets Can Have a Major Impact on Improving Iron Status, Hemoglobin Level, and in Reducing Iron Deficiency Anemia-A Systematic Review and Meta-Analysis.

The prevalence of iron deficiency anemia is highest among low and middle-income countries. Millets, including sorghum, are a traditional staple in many of these countries and are known to be rich in iron. However, a wide variation in the iron composition of millets has been reported, which needs to be understood in consonance with its bioavailability and roles in reducing anemia. This systematic review and meta-analysis were carried out to analyze the scientific evidence on the bioavailability of iron in different types of millets, processing, and the impact of millet-based food on iron status and anemia. The results indicated that iron levels in the millets used to study iron bioavailability (both in vivo and in vitro) and efficacy varied with the type and variety from 2 mg/100 g to 8 mg/100 g. However, not all the efficacy studies indicated the iron levels in the millets. There were 30 research studies, including 22 human interventions and 8 in vitro studies, included in the meta-analysis which all discussed various outcomes such as hemoglobin level, serum ferritin level, and absorbed iron. The studies included finger millet, pearl millet, teff and sorghum, or a mixture of millets. The results of 19 studies conducted on anaemic individuals showed that there was a significant (p < 0.01) increase in hemoglobin levels by 13.2% following regular consumption (21 days to 4.5 years) of millets either as a meal or drink compared with regular diets where there was only 2.7% increase. Seven studies on adolescents showed increases in hemoglobin levels from 10.8 ± 1.4 (moderate anemia) to 12.2 ± 1.5 g/dl (normal). Two studies conducted on humans demonstrated that consumption of a pearl millet-based meal significantly increased the bioavailable iron (p < 0.01), with the percentage of bioavailability being 7.5 ± 1.6, and provided bioavailable iron of 1 ± 0.4 mg. Four studies conducted on humans showed significant increases in ferritin level (p < 0.05) up to 54.7%. Eight in-vitro studies showed that traditional processing methods such as fermentation and germination can improve bioavailable iron significantly (p < 0.01) by 3.4 and 2.2 times and contributed to 143 and 95% of the physiological requirement of women, respectively. Overall, this study showed that millets can reduce iron deficiency anemia.

Genome-Wide Assessment of Population Structure and Genetic Diversity of the Global Finger Millet Germplasm Panel Conserved at the ICRISAT Genebank.

Finger millet [Eleusine coracana (L.) Gaertn.] is an important climate-resilient nutrient-dense crop grown as a staple food grain in Asia and Africa. Utilizing the full potential of the crop mainly depends on an in-depth exploration of the vast diversity in its germplasm. In this study, the global finger millet germplasm diversity panel of 314 accessions was genotyped, using the DArTseq approach to assess genetic diversity and population structure. We obtained 33,884 high-quality single nucleotide polymorphism (SNP) markers on 306 accessions after filtering. Finger millet germplasm showed considerable genetic diversity, and the mean polymorphic information content, gene diversity, and Shannon Index were 0.110, 0.114, and 0.194, respectively. The average genetic distance of the entire set was 0.301 (range 0.040 - 0.450). The accessions of the race elongata (0.326) showed the highest average genetic distance, and the least was in the race plana (0.275); and higher genetic divergence was observed between elongata and vulgaris (0.320), while the least was between compacta and plana (0.281). An average, landrace accessions had higher gene diversity (0.144) and genetic distance (0.299) than the breeding lines (0.117 and 0.267, respectively). A similar average gene diversity was observed in the accessions of Asia (0.132) and Africa (0.129), but Asia had slightly higher genetic distance (0.286) than African accessions (0.276), and the distance between these two regions was 0.327. This was also confirmed by a model-based STRUCTURE analysis, genetic distance-based clustering, and principal coordinate analysis, which revealed two major populations representing Asia and Africa. Analysis of molecular variance suggests that the significant population differentiation was mainly due to within individuals between regions or between populations while races had a negligible impact on population structure. Finger millet diversity is structured based on a geographical region of origin, while the racial structure made negligible contribution to population structure. The information generated from this study can provide greater insights into the population structure and genetic diversity within and among regions and races, and an understanding of genomic-assisted finger millet improvement.

A Systematic Review and Meta-Analysis of the Potential of Millets for Managing and Reducing the Risk of Developing Diabetes Mellitus.

Millets (including sorghum) are known to be highly nutritious besides having a low carbon footprint and the ability to survive in high temperatures with minimal water. Millets are widely recognised as having a low Glycaemic Index (GI) helping to manage diabetes. This systematic review and meta-analyzes across the different types of millets and different forms of processing/cooking collated all evidences. Of the 65 studies that were collected globally, 39 studies with 111 observations were used to analyze GI outcomes and 56 studies were used to analyze fasting, post-prandial glucose level, insulin index and HbA1c outcomes in a meta-analysis. It is evident from the descriptive statistics that the mean GI of millets is 52.7 ± 10.3, which is about 36% lower than in typical staples of milled rice (71.7 ± 14.4) and refined wheat (74.2 ± 14.9). The descriptive, meta and regression analyses revealed that Job's tears, fonio, foxtail, barnyard, and teff were the millets with low mean GI (<55) that are more effective (35-79%) in reducing dietary GI than the control samples. Millets with intermediate GI (55-69) are pearl millet, finger millet, kodo millet, little millet, and sorghum which have a 13-35% lower GI than the control with high GI (>69). A meta-analysis also showed that all millets had significantly (p < 0.01) lower GI than white rice, refined wheat, standard glucose or white wheat bread except little millet which had inconsistent data. Long term millet consumption lowered fasting and post-prandial blood glucose levels significantly (p < 0.01) by 12 and 15%, respectively, in diabetic subjects. There was a significant reduction in HbA1c level (from 6.65 ± 0.4 to 5.67 ± 0.4%) among pre-diabetic individuals (p < 0.01) who consumed millets for a long period. Minimally processed millets were 30% more effective in lowering GI of a meal compared to milled rice and refined wheat. In conclusion, millets can be beneficial in managing and reducing the risk of developing diabetes and could therefore be used to design appropriate meals for diabetic and pre-diabetic subjects as well as for non-diabetic people for a preventive approach.

Genome-Wide DArTSeq Genotyping and Phenotypic Based Assessment of Within and Among Accessions Diversity and Effective Sample Size in the Diverse Sorghum, Pearl Millet, and Pigeonpea Landraces.

Germplasm should be conserved in such a way that the genetic integrity of a given accession is maintained. In most genebanks, landraces constitute a major portion of collections, wherein the extent of genetic diversity within and among landraces of crops vary depending on the extent of outcrossing and selection intensity infused by farmers. In this study, we assessed the level of diversity within and among 108 diverse landraces and wild accessions using both phenotypic and genotypic characterization. This included 36 accessions in each of sorghum, pearl millet, and pigeonpea, conserved at ICRISAT genebank. We genotyped about 15 to 25 individuals within each accession, totaling 1,980 individuals using the DArTSeq approach. This resulted in 45,249, 19,052, and 8,211 high-quality single nucleotide polymorphisms (SNPs) in pearl millet, sorghum, and pigeonpea, respectively. Sorghum had the lowest average phenotypic (0.090) and genotypic (0.135) within accession distances, while pearl millet had the highest average phenotypic (0.227) and genotypic (0.245) distances. Pigeonpea had an average of 0.203 phenotypic and 0.168 genotypic within accession distances. Analysis of molecular variance also confirms the lowest variability within accessions of sorghum (26.3%) and the highest of 80.2% in pearl millet, while an intermediate in pigeonpea (57.0%). The effective sample size required to capture maximum variability and to retain rare alleles while regeneration ranged from 47 to 101 for sorghum, 155 to 203 for pearl millet, and 77 to 89 for pigeonpea accessions. This study will support genebank curators, in understanding the dynamics of population within and among accessions, in devising appropriate germplasm conservation strategies, and aid in their utilization for crop improvement.

Genome-Wide Population Structure Analyses of Three Minor Millets: Kodo Millet, Little Millet, and Proso Millet.

CORE IDEAS: Developed genome-wide SNP marker data for kodo, proso, and little millet Marker data used to analyze genetic diversity Heritability results of various traits used to validate genetic data Millets are a diverse group of small-seeded grains that are rich in nutrients but have received relatively little advanced plant breeding research. Millets are important to smallholder farmers in Africa and Asia because of their short growing season, good stress tolerance, and high nutritional content. To advance the study and use of these species, we present genome-wide marker datasets and population structure analyses for three minor millets: kodo millet (Paspalum scrobiculatum L.), little millet (Panicum sumatrense Roth), and proso millet (Panicum miliaceum L.).We generated genome-wide marker data sets for 190 accessions of each species with genotyping-by-sequencing (GBS). After filtering, we retained between 161 and 165 accessions of each species, with 3461, 2245, and 1882 single-nucleotide polymorphisms (SNPs) for kodo, little, and proso millet, respectively. Population genetic analysis revealed seven putative subpopulations of kodo millet and eight each of proso millet and little millet. To confirm the accuracy of this genetic data, we used public phenotype data on a subset of these accessions to estimate the heritability of various agronomically relevant phenotypes. Heritability values largely agree with the prior expectation for each phenotype, indicating that these SNPs provide an accurate genome-wide sample of genetic variation. These data represent one of first genome-wide population genetics analyses-and the most extensive-in these species and the first genomic analyses of any sort for little millet and kodo millet. These data will be a valuable resource for researchers and breeders trying to improve these crops for smallholder farmers.

Population Genetics and Structure of a Global Foxtail Millet Germplasm Collection.

Foxtail millet [Setaria italica (L.) P. Beauv.] is one among the most ancient crops of dryland agriculture. It is the second most important crop among millets grown for grains or forage. Foxtail millet germplasm resources provide reservoirs of novel alleles and genes for crop improvement that have remained mostly unexplored. We genotyped a set of 190 foxtail millet germplasm accessions (including 155 accessions of the foxtail millet core collection) using genotyping-by-sequencing (GBS) for rapid single nucleotide polymorphisms (SNP) characterization to study population genetics and structure, which enable allele mining through association mapping approaches. After filtering a total 350,000 raw SNPs identified across 190 germplasm accessions for minor allele frequency (MAF), coverage for samples and coverage for sites, we retained 181 accessions with 17,714 high-quality SNPs with ≥5% MAF. Genetic structure analyses revealed that foxtail millet germplasm accessions are structured along both on the basis of races and geographic origin, and the maximum proportion of variation was due to among individuals within populations. Accessions of race indica were less diverse and are highly differentiated from those of maxima and moharia. Genome-wide linkage disequilibrium (LD) analysis showed on an average LD extends up to ∼150 kbp and varied with individual chromosomes. The utility of the data for performing genome-wide association studies (GWASs) was tested with plant pigmentation and days to flowering and identified significant marker-trait associations. This SNP data provides a foundation for exploration of foxtail millet diversity and for mining novel alleles and mapping genes for economically important traits.