Defense mechanisms of maize against pink stem borer

The pink stem borer (Sesamia nonagrioides Lef.) is the most important pest of maize (Zea mays L.) in northwestern Spain. The objective of this work was to evaluate defense mechanisms against Sesamia in 10 inbred lines and the 10-parent diallel among these inbreds. Hybrids and inbred lines were tested over 2 yr in northwesternSpain under natural and artificial infestations. Yield of infested and non infested plants per plot were computed to calculate yield loss caused by the pink stem borer attack. Principal component analyses for stem damage traits and for ear damage traits were made for each infestation condition. From the principal component analysis, an index was computed to measure stem damage and ear damage. Damage index under artificial infestation was used to evaluate antibiosis while antixenosis was detected when damage index of each genotype was compared under artificial and natural infestation. The regressions of yield loss on the damage index were used to separate the genotypes into four groups according to the possession of antibiosis and/or tolerance mechanisms. A509, A661, EP31, F7, PB60, and Z77016 showed stem antibiosis. A637, A661, EP31, F7, and PB60 exhibited stem tolerance. A509, A661, and EP31 stood out by their ear antibiosis. A637, A661, F7, and PB60 were tolerant to ear damage. We conclude that the three mechanisms of defense to the pink stem borer attack (antixenosis, antibiosis, and tolerance) were found among inbred lines and hybrids. A multitrait selection scheme using damage traits and yield could improve the defense level against pink stem borer.Research supported by Dep. of Education of the Autonomous Government of Galicia (project XUGA40301B95Xunta de GaliciaN

Combining abilities in maize for the length of the internode basal ring, the entry point of the Mediterranean corn borer larvae

Length of the internode basal ring (LIBR) in maize is a morphological character that has been associated with resistance to Mediterranean corn borer (MCB), Sesamia nonagrioides Lef. The present study is the first research to evaluate the usefulness of this trait in breeding programmes. Six maize hybrids, from a complete diallel set of four inbred lines (two resistant and two susceptible to MCB), were evaluated under early and late sowing conditions at three locations in northwestern Spain. General and specific combining ability (GCA and SCA, respectively) for LIBR were estimated, and LIBR correlations with grain yield and other important agronomic traits were evaluated. Hybrid by environment interactions were not significant for LIBR and the sums of squares partitioning indicated a greater GCA effect (95%), suggesting that this trait is stable and shows important additive effects for this set of hybrids. Correlation coefficients not only indicate that selection for increasing LIBR could enhance grain yield and other related plant traits (height and silking) but also an increase in the MCB susceptibility. Based on the limited number genotypes evaluated, LIBR could be modified by selection; however, if LIBR is used as an indirect selection criterion to improve MCB resistance, then some negative effects on yield may be expected.MICINNEuropean Social FundThis research was supported by the National Plan for Research and Development of Spain (Proje cts Cod. AGL2006-13140, AGL 2009-09611). J. Barros acknowledges a grant from the Ministry of Science and Innovation (Spain). R. Santiago acknowledges a postdoctoral contract “Juan de la Cierva” partially financed by the European Social Fund.Peer Reviewe

Combining abilities for maize stem antibiosis, yield loss and yield under infestation and no infestation with pink stem borer

The pink stem borer (Sesamia nonagrioides Lef.) is the main pest of maize (Zea mays L.) in northwestern Spain. Little is known about combining ability for antibiosis and tolerance to this pest. Therefore, the objectives of this work were to estimate general combining ability (GCA), specific combining ability (SCA), and reciprocal effects for stem damage traits, yield, and yield loss by a complete diallel of 10 inbreds and to determine the most useful trait for evaluating the level of defense to pink stem borer. The diallel design was evaluated for 2 yr for stem damage traits, yield loss, and for yield under two conditions, infestation with Sesamia nonagrioides and non infestation. For all stem damage traits, GCA was significant, while SCA and reciprocal effects were not significant. This indicated that, for this set of inbreds, only additive effects were important for stem antibiosis. GCA and SCA effects were significant, in at least one trial, for yield under both infestation conditions and for yield loss. Reciprocal effects were significant for yield of infested and non infested plants in 1995. The lack of concordance among lines that exhibited the most favorable GCA effects for stem antibiosis, yield loss, and yield under infested conditions and the low correlation coefficient between SCA effects for yield under infested and non infested conditions showed that yield under infested conditions is the best trait for evaluating the level of defense against pink stem borer attack.Research supported by the Committee for Science and Technology of Spain (project AGF92-0161) and by the Department of Education of the Autonomus Government of Galicia (project XUGA 40301B95).MCYTXunta de GaliciaN

Genome-wide association analysis for fumonisin content in maize kernels

Background: Plant breeding has been proposed as one of the most effective and environmentally safe methods to control fungal infection and to reduce fumonisin accumulation. However, conventional breeding can be hampered by the complex genetic architecture of resistance to fumonisin accumulation and marker-assisted selection is proposed as an efficient alternative. In the current study, GWAS has been performed for the first time for detecting high-resolution QTL for resistance to fumonisin accumulation in maize kernels complementing published GWAS results for Fusarium ear rot. Results: Thirty-nine SNPs significantly associated with resistance to fumonisin accumulation in maize kernels were found and clustered into 17 QTL. Novel QTLs for fumonisin content would be at bins 3.02, 5.02, 7.05 and 8.07. Genes with annotated functions probably implicated in resistance to pathogens based on previous studies have been highlighted. Conclusions: Breeding approaches to fix favorable functional variants for genes implicated in maize immune response signaling may be especially useful to reduce kernel contamination with fumonisins without significantly interfering in mycelia development and growth and, consequently, in the beneficial endophytic behavior of Fusarium verticillioides.Xunta de Galicia | Ref. IN607A/013Ministerio de Economía | Ref. AGL2015–67313-C2–1-RMinisterio de Economía | Ref. AGL2015–67313-C2–2-

Environmental factors related to fungal infection and fumonisin accumulation during the development and drying of white maize kernels

In Southern Europe where whole maize kernels are ground and used for making bread and other food products, infection of the kernels by Fusarium verticillioides and subsequent fumonisin contamination pose a serious safety issue. The influence of environmental factors on this fungal infection and mycotoxin accumulation as the kernel develops has not been fully determined, especially in such food grade maize. The objectives of the present study were to determine which environmental factors may contribute to kernel invasion by F. verticillioides and fumonisin accumulation as kernels develop and dry in naturally infected white maize. Three maize hybrids were planted at two different sowing dates and kernel samples were collected 20, 40, 60, 80 and 100. days after silking. The percentage of kernels infected, and ergosterol and fumonisin contents were recorded for each sampling. F. verticillioides was the most prevalent species identified as the kernels developed. Temperature and moisture conditions during the first 80. days after silking favored natural kernel infection by F. verticillioides rather than by Aspergillus or Penicillium species. Fumonisin was found in kernels as early as 20. days after silking however significant fumonisin accumulation above levels acceptable in the EU did not occur until after physiological maturity of the kernel indicating that kernel drying in the field poses a high risk. Our results suggest that this could be due to increasing kernel damage by insects that favor fungal development, such as the damage by the moth Sitotroga cerealella, and to the occurrence of stress conditions for F. verticillioides growth that could trigger fumonisin biosynthesis, such as exposure to suboptimal temperatures for growth simultaneously with low water activity. © 2013 Elsevier B.V.MICINNSpanish Council of ResearchAutonomous Government of GaliciaEuropean Social FundThis research was supported by the National Plan for Research and Development of Spain (AGL2009-12770). A. Cao acknowledges funding from the JAE Program of the Spanish Council of Research. R. Santiago acknowledges postdoctoral contract “Isidro Parga Pondal” supported by the Autonomous Government of Galicia and the European Social Fund.Peer Reviewe

Genome-wide association analysis for maize stem Cell Wall-bound Hydroxycinnamates

Background: The structural reinforcement of cell walls by hydroxycinnamates has a significant role in defense against pests and pathogens, but it also interferes with forage digestibility and biofuel production. Elucidation of maize genetic variations that contribute to variation for stem hydroxycinnamate content could simplify breeding for cell wall strengthening by using markers linked to the most favorable genetic variants in marker-assisted selection or genomic selection approaches. Results: A genome-wide association study was conducted using a subset of 282 inbred lines from a maize diversity panel to identify single nucleotide polymorphisms (SNPs) associated with stem cell wall hydroxycinnamate content. A total of 5, 8, and 2 SNPs were identified as significantly associated to p-coumarate, ferulate, and total diferulate concentrations, respectively in the maize pith. Attending to particular diferulate isomers, 3, 6, 1 and 2 SNPs were related to 8–O–4 diferulate, 5–5 diferulate, 8–5 diferulate and 8–5 linear diferulate contents, respectively. This study has the advantage of being done with direct biochemical determinations instead of using estimates based on Near-infrared spectroscopy (NIRS) predictions. In addition, novel genomic regions involved in hydroxycinnamate content were found, such as those in bins 1.06 (for FA), 4.01 (for PCA and FA), 5.04 (for FA), 8.05 (for PCA), and 10.03 and 3.06 (for DFAT and some dimers). Conclusions: The effect of individual SNPs significantly associated with stem hydroxycinnamate content was low, explaining a low percentage of total phenotypic variability (7 to 10%). Nevertheless, we spotlighted new genomic regions associated with the accumulation of cell-wall-bound hydroxycinnamic acids in the maize stem, and genes involved in cell wall modulation in response to biotic and abiotic stresses have been proposed as candidate genes for those quantitative trait loci (QTL). In addition, we cannot rule out that uncharacterized genes linked to significant SNPs could be implicated in dimer formation and arobinoxylan feruloylation because genes involved in those processes have been poorly characterized. Overall, genomic selection considering markers distributed throughout the whole genome seems to be a more appropriate breeding strategy than marker-assisted selection focused in markers linked to QTL.Xunta de Galicia | Ref. ED431F 2016/014Agencia Estatal de Investigación | Ref. RTI2018–096776-B-C22Agencia Estatal de Investigación | Ref. RTI2018–096776-B-C2

Defensive changes in maize leaves induced by feeding of Mediterranean corn borer larvae

[Background] Plants can respond to insect attack via defense mechanisms that reduce insect performance. In this study, we examined the effects of several treatments applied to two maize genotypes (one resistant, one susceptible) on the subsequent growth and survival of Sesamia nonagrioides Lef. (Mediterranean corn borer, MCB) larvae. The treatments were infestation with MCB larvae, application of MCB regurgitant upon wounding, wounding alone, or exposure to methyl jasmonate, and they were applied at the V6–V8 stage of maize development. We also monitored changes in the concentrations of compounds known to be involved in constitutive resistance, such as cell wall-bound hydroxycinnamates and benzoxazinoids.[Results] In both maize genotypes, the leaves of plants pre-infested with MCB larvae were less suitable for larval development than those from untreated plants. Application of MCB regurgitant upon wounding, and wounding itself, resulted in leaf tissues becoming less suitable for larval growth than those of pre-infested plants, suggesting that there could be herbivore-associated effector molecules that suppress some wounding responses. A single application of MCB regurgitant did not seem to mimic feeding by MCB larvae, although the results suggested that regurgitant deposited during feeding may have enhanced ferulates and diferulates synthesis in infested vs. control plants. Jasmonic acid may play a role in mediating the maize response to MCB attack, but it did not trigger hydroxycinnamate accumulation in the leaves to a level comparable to that induced by larval leaf feeding. The EP39 maize genotype showed an increase in leaf cell wall strength by increasing hemicellulose cross-linking in response to MCB attack, while induced defenses in the EP42 plants appeared to reflect a broader array of resistance mechanisms.[Conclusions] The results indicated that leaf feeding by MCB larvae can increase leaf antibiosis against MCB in two maize genotypes with contrasting levels of resistance against this borer. Also, the larval regurgitant played a positive role in eliciting a defense response. We determined the effects of the plant response on larval growth, and detected defense compounds related to borer resistance.This research was supported by the National Plan for Research and Development of Spain (AGL2012-33415, AGL2015-67313-C2-1-R, and AGL2015-67313-C2-2-R) co-financed by the European Social Fund (FEDER). R. Santiago acknowledges postdoctoral contract “Ramón y Cajal” financed by the Ministry of Economy and Competitiveness (Spain), Vigo University, and the European Social Fund.Peer reviewe

Mapping of QTL for resistance to the Mediterranean corn borer attack using the intermated B73 × Mo17 (IBM) population of maize

The Mediterranean corn borer or pink stem borer (MCB, Sesamia nonagrioides Lefebvre) causes important yield losses as a consequence of stalk tunneling and direct kernel damage. B73 and Mo17 are the source of the most commercial valuable maize inbred lines in temperate zones, while the intermated B73 × Mo17 (IBM) population is an invaluable source for QTL identification. However, no or few experiments have been carried out to detect QTL for corn borer resistance in the B73 × Mo17 population. The objective of this work was to locate QTL for resistance to stem tunneling and kernel damage by MCB in the IBM population. We detected a QTL for kernel damage at bin 8.05, although the effect was small and two QTL for stalk tunneling at bins 1.06 and 9.04 in which the additive effects were 4 cm, approximately. The two QTL detected for MCB resistance were close to other QTL consistently found for European corn borer (ECB, Ostrinia nubilalis Hübner) resistance, indicating mechanisms of resistance common to both pests or gene clusters controlling resistance to different plagues. The precise mapping achieved with the IBM population will facilitate the QTL pyramiding and the positional cloning of the detected QTL.This research was supported by the National Plan for Research and Development of Spain (Project Cod. AGL2006-13140). B Ordas and R Santiago acknowledge a contract from the Spanish National Research Council (I3P Program) and G Sandoya acknowledges a fellowship from the Ministry of Education and Science (Spain).MECCSICPeer reviewe

Genetic factors involved in fumonisin accumulation in maize kernels and their implications in maize agronomic management and breeding

Contamination of maize with fumonisins depends on the environmental conditions; the maize resistance to contamination and the interaction between both factors. Although the effect of environmental factors is a determinant for establishing the risk of kernel contamination in a region, there is sufficient genetic variability among maize to develop resistance to fumonisin contamination and to breed varieties with contamination at safe levels. In addition, ascertaining which environmental factors are the most important in a region will allow the implementation of risk monitoring programs and suitable cultural practices to reduce the impact of such environmental variables. The current paper reviews all works done to address the influence of environmental variables on fumonisin accumulation, the genetics of maize resistance to fumonisin accumulation, and the search for the biochemical and/or structural mechanisms of the maize plant that could be involved in resistance to fumonisin contamination. We also explore the outcomes of breeding programs and risk monitoring of undertaken projects.This research was supported by the National Plan for Research and Development of Spain (AGL2009-12770). R. Santiago acknowledges postdoctoral contracts “Isidro Parga Pondal” supported by the Autonomous Government of Galicia and the European Social Fund and “Ramón y Cajal” supported by the Ministry of Economy and Competitiveness of Spain.We acknowledge support by the CSIC Open Access Publication Initiative through its Unit of Information Resources for Research (URICI).Peer reviewe

Resistencia de variedades de maíz dulce a la plaga de taladro Sesamia nonagrioides Lef. y Ostrinia nubilalis Hbn.

Resumen del trabajo presentado en el Congreso celebrado en Vigo entre el 6 y el 10 de julio de 1998.Peer Reviewe