The genetic basis of heterosis for grain yield and its components was investigated at the single- and two-locus levels using molecular markers with an immortalized F(2) (IF(2)) population, which was developed by pair crosses among recombinant inbred lines (RILs) derived from the elite maize hybrid Yuyu22. Mid-parent heterosis of each cross in the IF(2) population was used to map heterotic quantitative trait loci. A total of 13 heterotic loci (HL) were detected. These included three HL for grain yield, seven for ear length, one for ear row number and two for 100-kernel weight. A total of 143 digenic interactions contributing to mid-parent heterosis were detected at the two-locus level involving all three types of interactions (additive x additive = AA, additive x dominance = AD or DA, dominance x dominance = DD). There were 25 digenic interactions for grain yield, 36 for ear length, 31 for ear row number and 51 for 100-kernel weight. Altogether, dominance effects of HL at the single-locus level as well as AA interactions played an important role in the genetic basis of heterosis for grain yield and its components in Yuyu22.

Genetic basis of grain yield heterosis relies on the cumulative effects of dominance, overdominance, and epistasis in maize hybrid Yuyu22. Heterosis, i.e., when F1 hybrid phenotypes are superior to those of the parents, continues to play a critical role in boosting global grain yield. Notwithstanding our limited insight into the genetic and molecular basis of heterosis, it has been exploited extensively using different breeding approaches. In this study, we investigated the genetic underpinnings of grain yield and its components using "immortalized F2" and recombinant inbred line populations derived from the elite hybrid Yuyu22. A high-density linkage map consisting of 3,184 bins was used to assess (1) the additive and additive-by-additive effects determined using recombinant inbred lines; (2) the dominance and dominance-by-dominance effects from a mid-parent heterosis dataset; and (3) the various genetic effects in the "immortalized F2" population. Compared with a low-density simple sequence repeat map, the bin map identified more quantitative trait loci, with higher LOD scores and better accuracy of detecting quantitative trait loci. The bin map showed that, among all traits, dominance was more important to heterosis than other genetic effects. The importance of overdominance/pseudo-overdominance was proportional to the amount of heterosis. In addition, epistasis contributed to heterosis as well. Phenotypic variances explained by the QTLs detected were close to the broad-sense heritabilities of the observed traits. Comparison of the analyzed results obtained for the "immortalized F2" population with those for the mid-parent heterosis dataset indicated identical genetic modes of action for mid-parent heterosis and grain yield performance of the hybrid.

Kernel starch content is an important trait in maize (Zea mays L.) as it accounts for 65–75% of the dry kernel weight and positively correlates with seed yield. A number of starch synthesis-related genes have been identified in maize in recent years. However, many loci underlying variation in starch content among maize inbred lines still remain to be identified. The current study is a genome-wide association study that used a set of 263 maize inbred lines. In this panel, the average kernel starch content was 66.99%, ranging from 60.60 to 71.58% over the three study years. These inbred lines were genotyped with the SNP50 BeadChip maize array, which is comprised of 56,110 evenly spaced, random SNPs. Population structure was controlled by a mixed linear model (MLM) as implemented in the software package TASSEL. After the statistical analyses, four SNPs were identified as significantly associated with starch content (P ≤ 0.0001), among which one each are located on chromosomes 1 and 5 and two are on chromosome 2. Furthermore, 77 candidate genes associated with starch synthesis were found within the 100-kb intervals containing these four QTLs, and four highly associated genes were within 20-kb intervals of the associated SNPs. Among the four genes, Glucose-1-phosphate adenylyltransferase (APS1; Gene ID GRMZM2G163437) is known as an important regulator of kernel starch content. The identified SNPs, QTLs, and candidate genes may not only be readily used for germplasm improvement by marker-assisted selection in breeding, but can also elucidate the genetic basis of starch content. Further studies on these identified candidate genes may help determine the molecular mechanisms regulating kernel starch content in maize and other important cereal crops.

Heterosis is observed for most phenotypic traits and developmental stages in many plants. In this study, the embryos, from germinating seeds after 24 h of soaking, for five elite maize hybrids and their parents were selected to unravel the genetic basis of heterosis using 2-D proteomic method. In total, 257 (80.06%), 363 (58.74%), 351 (79.95%), 242 (54.50%), and 244 (46.30%) nonadditively expressed proteins were identified in hybrids Zhengdan 958, Nongda 108, Yuyu 22, Xundan 20, and Xundan 18, respectively. The nonadditive proteins were divided into above high-parent (++; 811, 55.66%), high-parent (+; 121, 8.30%), partial dominance (+-; 249, 17.09%), low-parent (-; 30, 2.06%), below low-parent (- -; 62, 4.26%), and D (different; 184, 12.63%) expression patterns. The observed patterns indicate the important roles of dominance, partial dominance, and overdominance in regulating seed germination in maize. Additionally, 54 different proteins were identified by mass spectrometry and classified into nine functional groups: metabolism (9), cell detoxification (8), unknown functional proteins (8), chaperones (7), signal transduction (6), development process (5), other (5), transporter (3), and stress response (3). Of these, the most interesting are those involved with germination-related hormone signal transduction and the abscisic acid and gibberellin regulation networks.

Heterosis has been utilized widely in the breeding of maize and other crops, and plays an important role in increasing yield, improving quality and enhancing stresses resistance, but the molecular mechanism responsible for heterosis is far from clear. To illustrate whether miRNA-dependent gene regulation is responsible for heterosis during maize germination, a deep-sequencing technique was applied to germinating embryos of a maize hybrid, Yuyu22, which is cultivated widely in China and its parental inbred lines, Yu87-1 and Zong3. The target genes of several miRNAs showing significant expression in the hybrid and parental lines were predicted and tested using real-time PCR. A total of 107 conserved maize miRNAs were co-detected in the hybrid and parental lines. Most of these miRNAs were expressed non-additively in the hybrid compared to its parental lines. These results indicated that miRNAs might participate in heterosis during maize germination and exert an influence via the decay of their target genes. Novel miRNAs were predicted follow a rigorous criterion and only the miRNAs detected in all three samples were treated as a novel maize miRNA. In total, 34 miRNAs belonged to 20 miRNA families were predicted in germinating maize seeds. Global repression of miRNAs in the hybrid, which might result in enhanced gene expression, might be one reason why the hybrid showed higher embryo germination vigor compared to its parental lines.

Photoperiod sensitivity is an important consideration in maize cultivation. Flowering time is affected by photoperiod and sensitivity to it limits the potential for successful exchange of germplasm across different latitudes. For resolving the genetic basis of photoperiod sensitivity in maize, a set of 207 recombinant inbred lines derived from a temperate and tropical inbred line cross was evaluated for 2 years in a long-day and short-day environment. Genetic linkage maps were constructed using 237 SSR markers with a total length 1,974.3 cM, and an average space between two makers of 8.33 cM. Twenty-nine QTL were detected for the five measured photoperiod sensitivity traits using composite interval mapping and multiple interval mapping. QTL for flowering time, plant height and leaf number, under long-day conditions, were found clustered on chromosome 10, while QTL for short-day conditions resided on chromosome 3. The QTL in the bin 10.04 region of chromosome 10 were detected associated with photoperiod sensitivity and related traits during long days. These results indicated that this region might contain an important photoperiod sensitivity element.

Kernel size and weight are important determinants of grain yield in maize. In this study, multivariate conditional and unconditional quantitative trait loci (QTL), and digenic epistatic analyses were utilized in order to elucidate the genetic basis for these kernel-related traits. Five kernel-related traits, including kernel weight (KW), volume (KV), length (KL), thickness (KT), and width (KWI), were collected from an immortalized F2 (IF2) maize population comprising of 243 crosses performed at two separate locations over a span of two years. A total of 54 unconditional main QTL for these five kernel-related traits were identified, many of which were clustered in chromosomal bins 6.04–6.06, 7.02–7.03, and 10.06–10.07. In addition, qKL3, qKWI6, qKV10a, qKV10b, qKW10a, and qKW7a were detected across multiple environments. Sixteen main QTL were identified for KW conditioned on the other four kernel traits (KL, KWI, KT, and KV). Thirteen main QTL were identified for KV conditioned on three kernel-shape traits. Conditional mapping analysis revealed that KWI and KV had the strongest influence on KW at the individual QTL level, followed by KT, and then KL; KV was mostly strongly influenced by KT, followed by KWI, and was least impacted by KL. Digenic epistatic analysis identified 18 digenic interactions involving 34 loci over the entire genome. However, only a small proportion of them were identical to the main QTL we detected. Additionally, conditional digenic epistatic analysis revealed that the digenic epistasis for KW and KV were entirely determined by their constituent traits. The main QTL identified in this study for determining kernel-related traits with high broad-sense heritability may play important roles during kernel development. Furthermore, digenic interactions were shown to exert relatively large effects on KL (the highest AA and DD effects were 4.6% and 6.7%, respectively) and KT (the highest AA effects were 4.3%).

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