Breeding for provitamin A biofortification of maize (Zea mays L.)

Vitamin A deficiency is widely prevailing in children and women of developing countries. Deficiency of vitamin A causes night blindness, growth retardation, xerophthalmia and increases the susceptibility against epidemic diseases. Among different interventions of overcoming malnutrition, biofortification is the most acceptable and preferred intervention among researchers, growers and consumers. Maize is grown and consumed in those regions where vitamin A deficiency is most prevalent; thus, targeting this crop for provitamin A biofortification is the most appropriate solution. Different breeding strategies including diversity analysis, introduction and stability analysis of exotic germplasm, hybridization, heterosis breeding, mutagenesis and marker‐assisted selection are practised for exploring maize germplasm and development of provitamin A‐enriched cultivars. Genome‐wide association selection and development of transgenic maize genotypes are also being practised, whereas RNA interference and genome editing tools could also be used as potential strategies for provitamin A biofortification of maize genotypes. The use of these breeding strategies for provitamin A biofortification of maize is comprehensively reviewed to provide a working outline for maize breeders.     

磷肥对甜玉米籽粒植酸和锌有效性的影响

为明确磷肥对甜玉米籽粒营养特性的影响,利用不同品种的磷肥定位试验以及磷肥梯度水平试验,探讨甜玉米籽粒植酸、磷(P)、锌(Zn)含量及Zn有效性的时间(不同灌浆期)和空间(果穗上、中、下部;籽粒果皮、胚乳和胚)变化及其对外源磷肥的响应特性。试验结果表明,甜玉米籽粒的P和植酸含量随磷肥用量的增加而增加,Zn含量及其有效性则呈现下降趋势。磷肥投入对吐丝后16～24 d籽粒的植酸、P和Zn的积累(mg plant^–1)的影响最为显著,而对不同灌浆时期Zn有效性均表现出抑制效应。在中磷水平下,果穗不同穗位籽粒植酸(g kg^–1)和Zn有效性的差异并不显著。高磷水平显著提高了中、下部果穗植酸的含量,但对不同穗位籽粒Zn含量和Zn有效性均存在抑制效应。对籽粒内不同部位(果皮、胚乳和胚)进行比较,植酸、P以及Zn含量均以胚中含量最高,积累量占比则以胚乳中最高;而Zn有效性以胚中最高。高磷处理可显著提高籽粒内不同部位植酸和磷含量,降低Zn含量及其有效性。综上,合理磷肥施用可促进籽粒Zn积累,而过量磷肥投入会显著增加不同灌浆时期,以及胚乳和胚中的植酸含量和积累量...     

Identification of quantitative trait locus and prediction of candidate genes for grain mineral concentration in maize across multiple environments

Trace metal elements are essential in daily diets for human health and normal growth. Maize is staple food for people in many countries. However, maize has low mineral concentration which makes it difficult to meet human requirements for micronutrients. The objective of this study was to identify quantitative trait locus (QTL) and predict candidate genes associated with mineral concentration in maize grain. Here, a recombinant inbred line population was used to test phenotype of zinc (Zn), iron (Fe), copper (Cu) and manganese (Mn) concentrations in six environments and then a QTL analysis was conducted using single environment analysis along with multiple environment trial (MET) analysis. These two strategies detected a total of 64 and 67 QTLs for target traits, respectively. Single environment analysis revealed 13 QTL bins distributed on seven chromosomes. We found that five candidate genes associated with mineral concentration were located in the same intervals identified by Comparative mapping of meta-QTLs in our previous study. The genetic and phenotypic correlation coefficients were depended on the nutrient traits and they were significant between Fe and Zn, Fe and Cu, Fe and Mn in all environments. The results of this study illustrated the genetic correlation between maize grain mineral concentrations, and identified some promising genomic regions and candidate genes for further studies on the biofortification of mineral concentration in maize grain.     

Molecular basis of Iron Biofortification in crop plants; A step towards sustainability

Iron is one of the most important micronutrients for crop plants due to its use in important physiological processes such as photosynthesis, mitochondrial respiration, metal homeostasis, and chlorophyll synthesis. Crop plants have adapted different strategies for uptake, transport, accumulation, and storage of iron in tissues and organs which later can be consumed by humans. Estimates indicate that about 2 billion people (33% of human population) are at risk of iron deficiency in which infants, children, and pregnant women are potentially compromised. Biofortification refers to the increase in concentration of micronutrients in edible parts of plants and understanding the pathways for iron accumulation in plants is necessary for breeding iron‐enriched crops. Iron‐biofortified crops are also one of the key factors in achieving multiple United Nations Sustainable Development goals. This review article covers different strategies of iron acquisition and transport in plants, its bioavailability, coping with the iron deficiency as a global perspective, the current status of iron biofortification, and how breeding future biofortified crops could be helpful in combating the said issue in a sustainable manner.     

Aegilops kotschyi and Aegilops tauschii as sources for higher levels of grain Iron and Zinc

Micronutrient malnutrition affects a very large proportion of the world's population. For combating micronutrient malnutrition, biofortification through genetic manipulation has been proposed as an alternative to traditional fortification for increasing the bioavailable nutrient content of food crops. Wheat, being a staple food for a large section of the world's population, is targeted for increasing the Fe and Zn content in the grains. The cultivated germplasm of wheat does not have sufficient variability for grain Fe and Zn content but the wild species of wheat do show wider variation for grain micronutrient density. The analysis of Aegilops kotschyi and A. tauschii for Fe and Zn content in the grains using an atomic absorption spectrophotometer (AAS) indicated that the S and D genome species accumulate significantly higher iron and zinc in the grains than the cultivated wheats. One of the CIMMYT synthetics also had significantly higher Fe and Zn in the grains as compared with the cultivated wheats. Aegilops kotschyi as a promising source for Fe and Zn, is reported for the first time. A systematic programme to identify and utilize the additional sources for high Fe and Zn has been initiated.     

Logistic model-based genetic analysis for kernel filling in a maize RIL population

Kernel filling is an important factor that directly affects kernel yield in maize. Based on a Logistic model, the process of kernel filling in maize can be effectively fitted, and the characteristic parameters with biological significance can be estimated. To clarify the genetic mechanism of characteristic parameters of kernel filling in maize, a recombinant inbred line (RIL) population including 208 lines derived from the maize inbred lines DH1M and T877 were evaluated in Nantong in 2015 and in Yangzhou in 2016, respectively. The kernel dry weights of recombinant inbred lines were measured 10, 15, 20, 25, 30, 35, 40, 43, 46, 49, 52, 55, 58 and 61 days after pollination (DAP). A total of 12 characteristic parameters related to kernel filling were estimated in different environments using the Logistic model. These parameters showed abundant phenotypic variation across two environments in the recombinant inbred line population. Some more ideal genotypes were selected through clustering based on BLUP values of characteristic parameters. Genetic analysis indicated that the 12 characteristic parameters conformed to the “major gene plus polygenes” model. The results of two environments were reproduced well. Most of the characteristic parameters related to kernel filling were controlled by two major genes, and a few characteristic parameters were controlled by three or four major genes. In addition, the genetic models of some characteristic parameters differed in the two environments due to interactions between the genes and environments. This study not only laid a foundation for further clarifying the genetic mechanism of maize kernel filling and mapping the related genes but also suggests a new paradigm for dynamic developing traits.     

Mineral bioavailability in grains of Pakistani bread wheat declines from old to current cultivars

Estimating variation in grain mineral concentration and bioavailability in relation to grain yield and the year of cultivar release is important for breeding wheat with increased content of bioavailable minerals. The grain yield and yield components, grain phytate concentration, and concentration and bioavailability of minerals (zinc Zn, iron Fe and calcium Ca) in wheat grains were estimated in 40 wheat cultivars released in Punjab (Pakistan) during the last five decades. Mean grain Zn and Ca concentrations in current-cultivars were significantly lower (≥14%) than in obsolete cultivars released during the Green Revolution (1965–1976). Much of this variation was related to increased grain weight in current-cultivars. There was a positive correlation among minerals (r = 0.39 or higher, n = 40) and minerals with phytate in wheat grains (r = 0.38 or higher, n = 40). The tested cultivars varied widely in grain yield and grain phytate-to-mineral molar ratios (phytate:mineral). Compared to obsolete cultivars, the current-cultivars had a higher phytate:mineral ratio in grains, indicating poor bioavailability of minerals to humans. The study revealed a non-significant relationship between grain yield and phytate:mineral ratios in grains. Therefore, breeding for lower phytate:mineral ratios in wheat grains can ensure increased mineral bioavailability without significant reduction in the yield potential. Future breeding should be focused on developing new genotypes suitable for mineral biofortification and with increased mineral bioavailability in grains.     

Identification of QTL for zinc and iron concentration in maize kernel and cob

To provide theoretical and applied references for biofortification of maize by increasing Zn and Fe concentration, the correlation and quantitative trait loci (QTL) of four mineral-related traits, i.e. zinc concentration of kernel (ZnK), zinc concentration of cob (ZnC), iron concentration of kernel (FeK) and iron concentration of cob (FeC) were studied for two sets of F_2:3 populations derived from the crosses Mu6 × SDM (MuS) and Mo17 × SDM (MoS) under two different environments (CQ and YN). The parental lines were very different in Zn and Fe concentration of kernels and cobs. A large genetic variation and transgressive segregation of two F_2:3 populations were observed for the four traits. The heritability of FeK was relatively lower (<0.6) than other three traits (>0.7). Analysis for each environment and joint analysis across two environments were used to identify QTL for each population. 16 and 15 QTL were identified in CQ and YN respectively via single environment analysis, some of which were identical in different environments and were also detected in joint analysis. The common regions for same trait at different environments were 3 and 5 in MuS and MoS respectively. Compared with the IBM2 2008 Neighbors Frame6, the distribution and effect of some QTL in two populations were highly consistent and many QTL on chromosome 2, 7 and 9 were detected in both populations. Moreover, several mineral QTL co-localized with each other for both populations such as the QTL for ZnK, ZnC, FeK and FeC on chromosome 2, QTL for Znk, FeK and FeC on chromosome 9 and QTL for ZnK and ZnC on chromosome 7, which probably were closely linked to each other, or were the same pleiotropic QTL.     

12个玉米群体的主要农艺性状与产量、品质的灰色关联度分析

为了明确玉米群体主要农艺性状与产量、营养品质的关系,应用灰色关联度分析方法对12个玉米群体的主要农艺性状、产量、营养品质等进行了灰色关联度分析。结果表明:与产量密切相关的农艺性状是出籽率、行粒数、穗粗、百粒质量、株高等;与粗蛋白、粗脂肪、粗淀粉和赖氨酸含量等营养品质密切相关的农艺性状是穗粒数、百粒质量、行粒数、出籽率、穗行数等。根据玉米群体主要农艺性状间的灰色关联度关系,说明在玉米群体产量、营养品质改良的过程中,在保证适当株高的前提下,要注重选育出籽率、百粒质量较高品种,同时不要忽略对玉米群体穗行数、行粒数、穗粗的选择,为选育高产、优质玉米新品种提供科学依据。     

The genetic architecture of zinc and iron content in maize grains as revealed by QTL mapping and meta-analysis

Micronutrient malnutrition, especially zinc (Zn) and iron (Fe) deficiency in diets, has aroused worldwide attention. Biofortification of food crops has been considered as a promising approach for alleviating this deficiency. Quantitative trait locus (QTL) analysis was performed to dissect the genetic mechanism of Zn and Fe content in maize grains using a total of 218 F_2:3 families derived from a cross between inbred lines 178 and P53. Meta-analysis was used to integrate genetic maps and detect Meta-QTL (MQTL) across several independent QTL researches for traits related to Zn or Fe content. Five significant QTLs and 10 MQTLs were detected. Two informative genomic regions, bins 2.07 and 2.08, showed a great importance for Zn and Fe content QTLs. The correlation between Zn and Fe level in maize grains was proposed by MQTLs as 8 of the 10 involved both traits. The results of this study suggest that QTL mapping and meta-analysis is an effective approach to understand the genetic basis of Zn and Fe accumulation in maize grains.     

磷营养对水稻籽粒锌生物有效性的影响及其与植酸等磷酸肌醇谱含量的关系

作物籽粒锌缺乏所导致的“隐性饥饿”已威胁到全世界1/3人群的健康。为明确磷对水稻籽粒锌的生物有效性的影响,利用大田磷肥长期定位试验,并结合稻穗离体培养技术,探讨了不同外源磷浓度下水稻籽粒锌的生物有效性的变化及其与籽粒植酸等磷酸肌醇谱含量的关系。试验表明高磷处理显著增加水稻籽粒中植酸磷和总磷的含量,以及不同价位磷酸肌醇(InsP1-6)含量,其中以高价磷酸肌醇(InsP4-6)在磷供给下的增幅最为显著。与籽粒植酸的变化趋势相反,高磷水平降低了稻米中锌的含量及其有效性。稻穗离体培养中,高磷(P12)处理相比无磷(P0)对照,籽粒锌的有效性降幅高达81.3%。因此,过量磷肥投入会通过显著提高水稻籽粒中的植酸及高价磷酸肌醇含量,在同步降低籽粒锌含量的同时,进一步显著降低其在水稻籽粒中的生物有效性。     

Genome-wide association study of grain iron and zinc concentration in common bean (Phaseolus vulgaris)

Common bean is one of the widely consumed food security crop in Africa, Asia and South America. It is a rich source of protein, minerals and micronutrients. High genotype by environment interaction is one of the main challenge in breeding for high grain micronutrient concentration. The objective of this study was to estimate SNP markers associated with grain Fe and Zn concentration using 289 common bean genotypes and 11,480 SNP markers. The study revealed that 43 quantitative trait loci (QTLs) were associated with grain Fe and Zn concentration. Five quantitative trait nucleotides (QTNs), that is, QTN Fe_1.1, QTN Fe_6.3, QTN Fe_6.5, QTN Fe_10.3 and QTN Fe_11.6 were detected both at Haramaya and Melkassa locations. Two of the markers, that is, QTN Fe_6.3 and QTN Fe_6.5, were located on chromosome 06 while QTN Fe_1.1, QTN Fe_10.3 and QTN Fe_ 11.6 were residing on chromosomes 01, 10 and 11, respectively. Among these, QTN Fe_11.6 had a large and positive consistent effect across locations. The five stable QTNs along with the potential candidate genes could be used for Fe biofortification through marker assisted selection.     

玉米杂交种农艺性状与籽粒产量的相关和通径分析

玉米杂交种主要农艺性状遗传相关与通径分析结果表明，行粒数、穗行数、百粒重、果穗长在影响籽粒产量诸因素中占主导地位。认为单株籽粒产量改良应在稳定穗数的基础上，着重提高行粒数；适当增加穗行数以利提高百粒重；协调好穗行数、行粒数、百粒重三者关系，实现单株产量的较大幅度增长。     

30%胺鲜酯-乙利水剂对玉米杂交种主要农艺性状的影响

在75000株/hm2的种植密度下,研究了玉米专用增效剂对玉米杂交种主要农艺性状的影响。结果表明:不同剂量的玉米增效剂对玉米杂交种农艺性状的影响存在明显差异;合理使用增效剂可以降低玉米杂交种株高、穗位高、倒伏率、空秆率、秃尖长,增加茎粗和千粒重,稳定穗长、穗粗、行数、行粒数、出籽率,提高玉米籽粒产量。玉米杂交种洛玉5号的适宜施用浓度为150～200mL/hm2。     

Diallel analysis of acid soil tolerant and susceptible maize inbred lines for grain yield under acid and non-acid soil conditions

Maize is not inherently tolerant to soil acidity but due to the ever increasing demand for the crop in the developing world, production of maize on acid soils continues to expand. Breeding for maize acid soil tolerance is the best strategy to improve yield under these conditions. Therefore, the current study was done to determine the general combining ability (GCA) of eight acid-soil tolerant and susceptible inbred lines and the specific combining ability (SCA) of cross combinations of these lines for grain yield under acid and non-acid soils. The eight lines were crossed using a diallel mating design to produce 28 single cross hybrids for evaluation under acid and non-acid soils at four sites for two seasons. Line C2 was the best general combiner under both soil environments while A2/C1 and A1/C2 had the highest desirable SCA effects under optimal conditions. Loss in grain yield and sensitivity to low pH stress was higher among genotypes in light textured soils than heavy soils. Non-additive gene action was more important than additive gene action in conditioning grain yield under both environments. Results revealed that it was feasible to improve grain yield under low pH and optimum soils from the set of genotypes used in the current study.     

不同生态区玉米产量及农艺性状比较

为进一步明晰东北华北春播区和黄淮海夏播区玉米生产的特征,从区域试验对照品种农大108和郑单958入手,对两个区域玉米产量及主要农艺性状进行比较和阐述。结果表明,东华北春玉米平均比黄淮海夏玉米增产1.4－2.0t/hm2,并且其产量优势在生育期、株高和穗位高、产量构成等方面都有着较好的体现。与黄淮海夏玉米相比,东华北春玉米生育期延长30天左右;株高和穗位高分别增加0.15m－0.23m和0.09m－0.15m;穗粒数和千粒重平均增加19.9%－25.4%和9.7％－16.5%。黄淮海区夏玉米粒重和穗粒数都有更大可提升的空间,而东华北春玉米增产主要靠增加穗粒数。     

The potential of lentil (<Emphasis Type="Italic">Lens culinaris</Emphasis> L.) as a whole food for increased selenium,iron, and zinc intake: preliminary results from a 3 year study

Micronutrient malnutrition, especially selenium (Se), iron (Fe), and zinc (Zn) deficiency, is a major global health problem. Previous attempts to prevent micronutrient malnutrition through food fortification, supplementation, and enrichment of staple crops has had limited success. Canadian grown lentils are rich in micronutrients Fe (73–90 mg kg⁻¹), Zn (44–54 mg kg⁻¹), Se (425–673 μg kg⁻¹), and have very low concentrations of phytic acid (2.5–4.4 mg g⁻¹). Our preliminary studies using a Caco-2 cell model show that the uptake of Fe from lentils is relatively greater than that of most other staple food crops. Moreover, preliminary results from our human nutrition study in Sri Lanka show an increased trend in blood Se concentration after lentil consumption. This article briefly overviews our previously published results as well as data from international lentil field trials, and describes the potential for biofortified lentil to provide a whole food solution to combat global human micronutrient malnutrition.     

Varietal Differences in Development of Maize (Zea mays L.) Seedlings on Compacted Soils

Differences among open-pollinated tropical maize ( Zea mays L.) varieties in seedling development and establishment on compacted soils were studied. Soil dry density was used as an index of compaction. Three soil compaction levels and 12 traits associated with development and establishment of maize seedlings were investigated. A control (without compaction) was also included. Varietal differences were observed for most traits measured. Genetic differences for seedling development on compacted soil were detected. Varietal differences contributed about three times the contribution of compaction to total variability in the traits. Better seedling development and performance were obtained in moderately compacted soil than in the control. Shoot length, shoot dry weight and per cent dry matter in roots were good indicators of the tolerance of maize seedlings to compaction. A physiological strategy for maize seedling establishment on compacted soil was proposed. The implications of the results for seed testing were also highlighted. It was concluded that consideration should be given to the genotype of maize destined for use in ecologies prone to high soil densities. All varieties of maize grown in an agroecological zone could be screened to identify genotypes tolerant of higher soil densities. The seeds could then be multiplied and distributed to farmers.     

Low-nitrogen stress tolerance and nitrogen agronomic efficiency among maize inbreds: comparison of multiple indices and evaluation of genetic variation

Limited information is available on genetic variation in low-nitrogen (low-N) stress tolerance and N-use efficiency (NUE) among maize inbreds. To unveil this information, a panel of 189 diverse maize inbred lines was evaluated under contrasting levels of N availability over 2 years. Low-N agronomic efficiency (LNAE), absolute grain yield (GY) at low-N conditions, and the ratio between GY at low-N and optimum-N conditions were taken into account to represent low-N tolerance. Additionally, N-agronomic efficiency (NAE) along with other agronomic traits was also analyzed. Analysis of variance revealed significant effects of genotype on LNAE, NAE, and GY. The estimated broad-sense heritability was 0.38 for LNAE while it was only 0.11 for NAE, implying that selection based on LNAE should be more effective than NAE. LNAE exhibited highly positive genotypic and phenotypic correlations with GY, ear kernel number (EKN), kernel weight, plant height (PH), and chlorophyll content at low-N conditions, while it was negatively correlated with grain-N content and anthesis-silking interval. Path analysis indicated that the EKN at low-N stress had the highest positive effects on LNAE.     

晚粳稻籽粒中As、Cd、Cr、Ni、Pb等重金属含量的基因型与环境效应及其稳定性

选择籽粒Cd、Cr、As、Ni和Pb等重金属含量差异较大的12个晚粳稻基因型，种植于浙江省晚粳稻主产区嘉兴、湖州、杭州、宁波、绍兴等市的6个试点，研究籽粒中5种重金属含量的基因型与环境变异及其稳定性、籽粒和土壤有效态（DTPA提取态）Cd、Cr、As、Ni和Pb等重金属含量之间及与Fe、Zn含量之间的关系，以及土壤pH对籽粒重金属积累的影响。结果表明，环境、基因型及其互作效应对籽粒重金属含量的效应均达极显著水平，表明筛选和选育籽粒重金属含量低的品种以及通过农艺措施减少籽粒重金属含量是可能的。同时，籽粒中这5种重金属含量的稳定性因环境、基因型而有较大差异，且亦因重金属种类而异。因此，为降低籽粒重金属含量，应针对特定重金属污染的环境进行基因型选择，并同时考虑基因型籽粒重金属含量的稳定性。土壤pH不仅影响土壤重金属有效性及水稻籽粒中重金属积累量，还影响籽粒重金属积累的基因型与环境互作效应（即积累稳定性）。此外，土壤中一些重金属常发生复合污染，如Cd和Cr、As或Ni，Cr和As，Pb和As间表现为协同消长，Cd含量较高的稻米往往As和Pb含量也高，Cr和Ni含量以及As和Pb含量之间也呈正相关。