Isozymes as Genetic Markers in Maize Breeding

The major objective of the research is to identify and locate quantitative trait loci (QTL_s) in the Yugoslav maize population. The plants (F2) were selected for the analysis at seedling stage and were selfed to obtain F3 generation. The analysis covered about 15 enzymes controlled by about 30 loci. The seeds of F3 family planted in the greenhouse for measuring some quantitative traits, recorded tasselling and silking during vegetation. At the end of vegetation grain yield, and some other quantitative traits of grain in F3 family were assessed. The relationship between marker loci and the loci for quantitative traits (QTL_s) were estimated by computerized statistical method.     

Quantitative trait loci for yield and correlated traits under high and low soil nitrogen conditions in tropical maize

The first objective of this study was to map and characterize quantitative trait loci (QTL) for grain yield (GY) and for secondary traits under varying nitrogen (N) supply. To achieve this objective, a segregating F_2:3 population previously developed for QTL mapping under water-limited conditions was used. The population was evaluated in Mexico under low N conditions in the dry winter season and under low and high N conditions in the wet summer season. From eight QTLs identified for GY under low N conditions, two were also detected under high N conditions. Five QTLs were stable across the two low N environments and five co-localized with QTLs identified for the anthesis-silking interval (ASI) or for the number of ears per plant (ENO) under low N conditions. The percentage of the phenotypic variance expressed by all QTLs for ASI and ENO was quite different when evaluated under low N conditions during the dry winter (40% for ASI and 22% for ENO) and the wet summer seasons (22% for ASI and 46% for ENO). The results suggest optimizing different breeding strategies based on selection index depending on the growing season. Good QTL colocalization was observed for ASI (four QTLs) and ENO (three QTLs) when looking at QTL identified under low N and water-limited conditions in the same population. The results suggest that that both secondary traits can be used in breeding programs for simultaneous improvement of maize against low N and drought stresses.     

Epistatic interaction is an important genetic basis of grain yield and its components in maize

A population of 294 recombinant inbred lines (RIL) derived from Yuyu22, an elite maize hybrid extending broadly in China, has been constructed to investigate the genetic basis of grain yield, and associated yield components in maize. The main-effect quantitative trait loci (QTL), digenic epistatic interactions, and their interactions with the environment for grain yield and its three components were identified by using the mixed linear model approach. Thirty-two main-effect QTL and forty-four pairs of digenic epistatic interactions were detected for the four measured traits in four environments. Our results suggest that both additive effects and epistasis (additive × additive) effects are important genetic bases of grain yield and its components in the RIL population. Only 30.4% of main-effect QTL for ear length were involved in epistatic interactions. This implies that many loci in epistatic interactions may not have significant effects for traits alone but may affect trait expression by epistatic interaction with the other loci.     

Growth analysis of maize field crops under phosphorus deficiency. II. Radiation-use efficiency,biomass accumulation and yield components

Biomass accumulation by crops depends on both light interception by leaves and on the efficiency with which the intercepted light is used to produce dry matter. Our aim was to identify which of these processes were affected for maize (Zea mays L., cv Volga) field crops grown under phosphorus (P) deficiency. In the preceding paper (Plénet et al., 2000), it was shown that P deficiency severely reduced leaf growth. In this paper, the effect of P deficiency on the radiation-use efficiency (RUE) was investigated. The experimental work was carried out in 1995, 1996 and 1997 on a long-term P fertilisation trial located on a sandy soil in the south-west of France. Three P fertilisation regimes have been applied since 1972: no- P (P0 treatment) and different rates of P fertiliser (P1.5: 1.5 times the grain P export and P3: 3 times the grain P export). These fertilisation regimes have led to contrasted levels of soil P supply. Only slight differences were observed between the P1.5 and P3 treatment for above-ground biomass accumulation and grain yield. Conversely the grain yield was significantly reduced in P0 (–11%). Above-ground biomass production was severely reduced, with the maximum difference between treatment (–60% in P0) occurring between 400 and 600 °C days after sowing. The lower biomass production in P0 was accounted for by the reduced amount of photosynthetically active radiation (PAR) absorbed by the canopy, which was itself the consequence of the reduced leaf area index (see Plénet et al., 2000). The calculated RUE were found to depend on the plant stage, especially during the pre-flowering period, and on the average air temperature. No effect of P deficiency was observed on the calculated RUE, even during the period when above-ground biomass accumulation was the most severely reduced. These results obtained in field crop conditions strengthen the idea that P deficiency affects plant growth, especially leaf growth, earlier and to a greater extent than photosynthesis per unit leaf area.     

In an elite cross of maize a major quantitative trait locus controls one-fourth of the genetic variation for grain yield

Quantitative trait loci (QTLs) for grain yield, dry matter content and test weight were identified in an F₂ segregating population derived from a single cross between two elite maize lines (B73 and A7) and testcrossed to two genetically divergent in breds. Most of the QTLs inferred were consistent across locations, indicating that the expression of the genes influencing the traits under investigation was largely independent of the environment. By using two different tester lines we found that QTLs exhibited by one tester may not necessarily be detected with the second one. Only loci with larger effects were consistent across testers, suggesting that interaction with tester alleles may contribute to the identification of QTLs in a specific fashion. Analysis across both testers revealed four significant QTLs for grain yield that explained more than 35% of the phenotypic variation and showed an overall phenotypic effect of more than 2t/ha. The major QTL for grain yield, located in the proximity of the Nucleolus Organiser Region, accounted for 24.5% of the phenotypic variation for grain yield and showed an average effect of allele substitution of approximately 1 t/ha. Marker-assisted introgression of the superior A7 allele at this locus in the B73 genetic background will not differ from qualitative trait introgression and will eventually lead to new lines having superior testcross performance.     

玉米雌雄开花间隔天数、结穗率与产量的数量性状位点（QTL）分析

采用SSR标记连锁图谱和复合区间作图法在山西灌溉和干旱胁迫条件下,对玉米(Zea mays L.)自交系黄早四×掖107组合的F_3群体雌雄开花间隔天数(ASI)、结穗率和籽粒产量进行了数量性状位点(QTL)定位及基因效应分析。结果表明,在两种水分处理下,ASI、结穗率与籽粒产量的相关性均达到显著水平(P<0.05)。在灌溉和干旱胁迫卜,分别检测到3个和2个控制ASI的QTL,位于第1、2、3和第2、5染色体上。在灌溉条件下,在第3和第6染色体上各检测到1个控制结穗率的QTL,基因作用方式呈加性或部分显性,可解释19.9％的表型变异;在干旱条件下,在第3、7、10染色体上共检测到4个控制结穗率的QTL,基因作用方式为显性或部分显性,可解释60.4％的表型变异。在灌溉和干旱胁迫下,控制产量的QTL分别定位在第3、6、7和第1、2、4、8染色体上,基因作用方式均以加性或部分显性为主,可解释的表型变异为7.3％～22.0％。在干旱条件下,借助连锁分子标记和基因效应分析,可构建包含ASI、结穗率和产量QTL的选择指数,用于分子标记辅助育种。     

Quantitative Changes in Maize Membrane Proteins Induced by Aluminium

Sodium dodecyl sulphate-polyacrylamide gel electrophoresis (SDS-PAGE) was used for monitoring Al-induced changes in polypeptide composition of membrane proteins isolated from 3-d-old maize seedlings subjected to aluminium stress. Analysis of peripheral membrane proteins isolated from maize root showed an Al-induced increase in accumulation of 14 polypeptides with apparent molecular mass from 10 to 135 kDa. Qualitative differences were found between peripheral membrane proteins isolated from root tip (increased accumulation of 4 polypeptides with M_r 42 000 – 135 000) and from root base (increased accumulation of 10 polypeptides with M_r 10 000 – 59 000). On the other hand, no Al-induced changes were observed in peripheral membrane proteins isolated from maize coleoptile and integral membrane proteins isolated either from root or coleoptile. These results indicate that peripheral membrane proteins undergo considerable changes during 24-h Al treatment while integral membrane proteins pattern is stable.     

Quantitative trait loci influencing protein and starch concentration in the Illinois Long Term Selection maize strains

A study was initiated to determine the number, chromosomal location, and magnitude of effect of QTL (quantitative trait loci or locus depending on context) controlling protein and starch concentration in the maize (Zea mays L.) kernel. Restriction fragment length polymorphism (RFLP) analysis was performed on 100 F₃ families derived from a cross of two strains, Illinois High Protein (IHP), X Illinois Low Protein (ILP), which had been divergently selected for protein concentration for 76 generations as part of the Illinois Long Term Selection Experiment. These families were analyzed for kernel protein and starch in replicated field trials during 1990 and 1991. A series of 90 genomic and cDNA clones distributed throughout the maize genome were chosen for their ability to detect RFLP between IHP and ILP. These clones were hybridized with DNA extracted from the 100 F₃ families, revealing 100 polymorphic loci. Single factor analysis of variance revealed significant QTL associations of many loci with both protein and starch concentration (P < 0.05 level). Twenty-two loci distributed on 10 chromosome arms were significantly associated with protein concentration, 19 loci on 9 chromosome arms were significantly associated with starch concentration. Sixteen of these loci were significant for both protein and starch concentration. Clusters of 3 or more significant loci were detected on chromosome arms 3L, 5S, and 7L for protein concentration, suggesting the presence of QTL with large effects at these locations. A QTL with large additive effects on protein and starch concentration was detected on chromosome arm 3L. RFLP alleles at this QTL were found to be linked with RFLP alleles at the Shrunken-2 (Sh2) locus, a structural gene encoding the major subunit of the starch synthetic enzyme ADP-glucose pyrophosphorylase. A multiple linear regression model consisting of 6 significant RFLP loci on different chromosomes explained over 64 % of the total variation for kernel protein concentration. Similar results were detected for starch concentration. Thus, several chromosomal regions with large effects may be responsible for a significant portion of the changes in kernel protein and starch concentration in the Illinois Long Term Selection Experiment.     

Effect of nitrogen form on maize response to drought stress

Two hybrids of maize (Zea mays L.) differing in resistance to drought, were grown in chernozem soil in a greenhouse and were fertilized with two different forms of nitrogen: Ca(NO₃)₂ and (NH₄)₂SO₄ in concentrations corresponding to 100 kg of N ha^-1. After emergence of the 4th leaf, plants were exposed to drought. During the drought period, the parameters of plant water status (water potential, osmotic potential, turgor pressure and relative water content) and chlorophyll a+b concentration were monitored every two days. N and K concentration and accumulation over the drought period were also monitored.Next to differences in adaptability of the two hybrids to drought, the results demonstrate different adaptability of NH₄ and NO₃-treated plants within each hybrid. NH₄-plants of each hybrid maintain higher turgor pressure during the drought by better osmotic adaptation. Especially significant differences appear between chlorophyll (a+b) values of NH₄ and NO₃-treated plants and as affected by drought. Chlorophyll concentrations of NH₄-plants are higher than those of NO₃-plants both in control and droughted plants. NH₄ plants show a characteristic initial chlorophyll increase at the beginning of the drought period while in NO₃ plants chlorophyll constantly decreases throughout the whole drought period. The influence of the nitrogen form on chlorophyll concentration changes during drought does not appear to be affected by regulation of the K concentration.     

Quantitative trait loci for plant height in four maize populations and their associations with qualitative genetic loci

Summary We report that plant height quantitative trait loci (QTLs) identified in a given small population are not consistent with QTLs identified in other small populations, and that most QTLs are in close proximity to mapped qualitative genetic loci. These observations provide evidence to support the hypothesis that qualitative genetic loci are the same loci that affect quantitative traits, and affirm that these modest experiments probably identify real QTLs.     

Genetic analysis of tolerance to low-phosphorus stress in maize using restriction fragment length polymorphisms

Summary An understanding of the genetic nature underlying tolerance to low-phosphorus (low-P) stress could aid in the efficient development of tolerant plant strains. The objective of this study was to identify the number of loci in a maize (Zea mays L.) population segregating for tolerance to low-P stress, their approximate location, and the magnitude of their effect.Seventy-seven restriction fragment length polymorphisms (RFLPs) were identified and scored in a maize F₂ population derived from a cross between line NY821 and line H99. The F₂ individuals were self-pollinated to produce F₃ families. Ninety F₃ families were grown in a sand-alumina system, which simulated diffusion-limited, low-P soil conditions. The F₃ families were evaluated for vegetative growth in a controlled-environment experiment. To identify quantitative trait loci (QTLs) underlying tolerance to low-P stress, the mean phenotypic performances of the F₃ families were contrasted based on genotypic classification at each of 77 RFLP marker loci.Six RFLP marker loci were significantly associated with performance under low-P stress (P<0.01). One marker locus accounted for 25% of the total phenotypic variation. Additive gene action was predominant for all of the QTLs identified. Significant marker loci were located on four separate chromosomes representing five unlinked genomic regions. Two marker loci were associated with an additive by additive epistatic interaction. A multiple regression model including three marker loci and the significant epistatic interaction accounted for 46% of the total phenotypic variation. Heterozygosity per se was not predictive of phenotypic performance.     

Effects of High Temperature on Chlorophyll Fluorescence Induction and the Kinetics of Far Red Radiation-Induced relaxation of apparent F0 in maize leaves

Temperature dependence (25–50 °C) of chlorophyll (Chl) fluorescence induction, far-red radiation (FR)-induced relaxation of the post-irradiation transient increase in apparent F₀, and the trans-thylakoid proton gradients (pH) was examined in maize leaves. Temperatures above 30 °C caused an elevation of F₀ level and an enhancement of F₀ quenching during actinic irradiation. Millisecond delayed light emission (ms-DLE), which reflects the magnitude of pH, decreased strikingly above 35 °C, and almost disappeared at 50 °C. It indicates that the heat-enhanced quenching of F₀ under actinic irradiation could not be attributed mainly to the mechanism of pH-dependent quenching. The relaxation of the post-irradiation transient increase in apparent F₀ upon FR irradiation could be decomposed into two exponential components (₁ = 0.7–1.8 s, ₂ = 2.0–9.9 s). Decay times of both components increased with temperature increasing from 25 to 40–45 °C. The bi-phasic kinetics of FR-induced relaxation of the post-irradiation transient increase in apparent F₀ and its temperature dependence may be related to plastoquinone (PQ) compartmentation in the thylakoid membranes and its re-organisation at elevated temperature.     

Growth,yield, and yield components of ethephon-treated corn

Field studies were conducted during 1985 and 1986 to study the effect of stage and rate of ethephon application on growth, combine-harvested yield, and yield components of three corn (Zea mays L.) hybrids at two densities. Ethephon was applied at four rates from 0 to 560g ha^–1 at three growth stages: tassel elongation (TE 3mm), TE + 6d, and Ear elongation (EE 3mm). The greatest rate of ethephon decreased lodging by 85% in 1985 and 93% in 1986. Reduction in yield at the greatest rate of ethephon was 6% and 2% of the control in 1985 and 1986, respectively. Brace-root rating in 1986 was increased 20%, when comparing the greatest application rate with the control. In the same year, weight per seed was reduced 2%, which was equivalent to the percentage yield reduction. Plant and ear heights generally decreased in a linear fashion with increasing rate of ethephon. The growth stage at the time of ethephon application significantly altered all variables except grain moisture. Grain yield, seed weight, and lodging decreased as ethephon application was delayed. Decrease in seed weight probably caused the yield decrease with delayed application. Stages of application interacted with rate such that ear height was reduced less as ethephon application was delayed. The greatest rate of ethephon applied at the beginning of EE resulted in the best lodging control. However, reduction in lodging did not result in higher yield. Also, ethephon applied at the TE stage mainly affected elongation of internodes below the ear; at the EE stage, elongation of internodes above the ear was affected.     

Identification of QTLs for root characteristics in maize grown in hydroponics and analysis of their overlap with QTLs for grain yield in the field at two water regimes

We investigated the overlap among quantitative trait loci (QTLs) in maize for seminal root traits measured in hydroponics with QTLs for grain yield under well-watered (GY-WW) and water-stressed (GY-WS) field conditions as well as for a drought tolerance index (DTI) computed as GY-WS/GY-WW. In hydroponics, 11, 7, 9, and 10 QTLs were identified for primary root length (R1L), primary root diameter (R1D), primary root weight (R1W), and for the weight of the adventitious seminal roots (R2W), respectively. In the field, 7, 8, and 9 QTLs were identified for GY-WW, GY-WS, and DTI, respectively. Despite the weak correlation of root traits in hydroponics with GY-WW, GY-WS, and DTI, a noticeable overlap between the corresponding QTLs was observed. QTLs for R2W most frequently and consistently overlapped with QTLs for GY-WW, GY-WS, and/or DTI. At four QTL regions, an increase in R2W was positively associated with GY-WW, GY-WS, and/or DTI. A 10 cM interval on chromosome 1 between PGAMCTA205 and php20644 showed the strongest effect on R1L, R1D, R2W, GY-WW, GY-WS, and DTI. These results indicate the feasibility of using hydroponics in maize to identify QTL regions controlling root traits at an early growth stage and also influencing GY in the field. A comparative analysis of the QTL regions herein identified with those described in previous studies investigating root traits in different maize populations revealed a number of QTLs in common.     

Heterogeneity of Maize Root Mitochondria from Plants Grown in the Presence of Ammonium

Mitochondria isolated from root tissue of maize plants grown on a modified Knop solution containing 10.9 mM nitrate ± 7.2 mM ammonium were purified on the discontinuous Percoll density gradient with polyvinylpyrrolidone (PVP) added. The presence of PVP allowed separation of several mitochondrial fractions of a different density. Contrary to mitochondria isolated from plants grown in the presence of nitrate alone, revealing only two fractions, the mitochondria from NH₄⁺/NO₃^–-plants were distributed in four fractions. Total amount of mitochondria, as well as specific activities of some nitrogen metabolism enzymes and tricarboxylic acid (TCA) cycle enzymes of all mitochondrial fractions, and respiratory activities of two lower density fractions isolated from plants grown on mixed nitrogen were higher in comparison to mitochondria from nitrate-grown plants.     

Quantitative trait loci for cold tolerance of rice recombinant inbred lines in low temperature environments

Low temperature is one of the major environmental stresses in rice cultivation in high-altitude and high-latitude regions. In this study, we cultivated a set of recombinant inbred lines (RIL) derived from Dasanbyeo (indica) / TR22183 (japonica) crosses in Yanji (high-latitude area), Kunming (high-altitude area), Chuncheon (cold water irrigation) and Suwon (normal) to evaluate the main effects of quantitative trait loci (QTL) and epistatic QTL (E-QTL) with regard to their interactions with environments for cold-related traits. Six QTLs for spikelet fertility (SF) were identified in three cold treatment locations. Among them, four QTLs on chromosomes 2, 7, 8, and 10 were validated by several near isogenic lines (NILs) under cold treatment in Chuncheon. A total of 57 QTLs and 76 E-QTLs for nine cold-related traits were identified as distributing on all 12 chromosomes; among them, 19 QTLs and E-QTLs showed significant interactions of QTLs and environments (QEIs). The total phenotypic variation explained by each trait ranged from 13.2 to 29.1% in QTLs, 10.6 to 29.0% in EQTLs, 2.2 to 8.8% in QEIs and 1.0% to 7.7% in E-QTL × environment interactions (E-QEIs). These results demonstrate that epistatic effects and QEIs are important properties of QTL parameters for cold tolerance at the reproductive stage. In order to develop cold tolerant varieties adaptable to wide-ranges of cold stress, a strategy facilitating marker-assisted selection (MAS) is being adopted to accumulate QTLs identified from different environments.     

Quantitative trait loci affecting cotton fiber are linked to the t1 locus in upland cotton

Pilose (T ₁), a dominant marker in upland cotton, has been associated with coarse, short fibers. Pilose was, thereby, considered to be pleiotropic on fiber fineness and length. However, a pilose-expressing line with a fiber of average fineness was recently identified. This finding does not support pleiotropy between T ₁ and fiber traits, but is indicative of linkage between pilose and loci influencing fiber characteristics. To understand the relationship between T ₁ and fiber traits, a pilose line with short, coarse fiber was crossed to two t ₁ lines with standard fiber characteristics. One hundred and forty-nine F₂-derived F₃ lines were developed from one cross, and 60 F₂-derived F₃ lines from the other. Seven fiber traits (elongation, maturity, micronaire reading, perimeter, 2.5% span length, strength, and wall thickness) were measured. Segregation was normal, as indicated by allelic frequencies of 0.5 for T ₁ and t ₁, and segregation ratios of 121 for marker genotypes. The association of homozygous T ₁ lines with fibers of average fineness was again observed. Linkage between T ₁ and loci affecting micronaire, perimeter, 2.5% span length, strength, and wall thickness was found in both populations. Significant additive and non-additive gene effects for each of these traits at the marker locus were found as well. The pilose marker accounted for 10–75% of the phenotypic variation associated with each trait. In conclusion, the t ₁ locus is linked to numerous loci that influence fiber traits, and this linkage has previously been misinterpreted as pleiotropy.     

Quantitative trait loci mapping in dairy cattle: review and meta-analysis

From an extensive review of public domain information on dairy cattle quantitative trait loci (QTL), we have prepared a draft online QTL map for dairy production traits. Most publications (45 out of 55 reviewed) reported QTL for the major milk production traits (milk, fat and protein yield, and fat and protein concentration (%)) and somatic cell score. Relatively few QTL studies have been reported for more complex traits such as mastitis, fertility and health. The collated QTL map shows some chromosomal regions with a high density of QTL, as well as a substantial number of QTL at single chromosomal locations. To extract the most information from these published records, a meta-analysis was conducted to obtain consensus on QTL location and allelic substitution effect of these QTL. This required modification and development of statistical methodologies. The meta-analysis indicated a number of consensus regions, the most striking being two distinct regions affecting milk yield on chromosome 6 at 49 cM and 87 cM explaining 4.2 and 3.6 percent of the genetic variance of milk yield, respectively. The first of these regions (near marker BM143) affects five separate milk production traits (protein yield, protein percent, fat yield, fat percent, as well as milk yield).     

Osmotic Stress Increases Alcohol Dehydrogenase Activity in Maize Seedlings

Maize (Zea mays L.) seedlings were exposed to osmotic stress, and alcohol dehydrogenase (ADH) activity and abscisic acid (ABA) concentration were determined. The osmotic stress increased ADH activities in both roots and shoots, whereas the increase was 2-fold greater in roots than the shoots. The stress also increased ABA concentration in both roots and shoots and the increase was greater in the roots than in the shoots.