Drought Tolerance In Tropical Maize Research Articles

La importancia del intervalo de la floración en el mejoramiento para la resistencia a sequía en maíz tropical

The length of the Interval between anthesis and silking (ASI) is increased by drought which coincides with flowering. Four elite CIMMYT lowland tropical populations are undergoing recurrent selection (S1 or full-sib) for improved grain yield and several other traits under drought and well watered conditions. Data collected from more than 200 families per population grown In single row plots under three water stress levels (pre and post-flowering stress; post-flowering stress; normal irrigation, all in the absence of rain) showed weak or no correlation between grain yield and traits related to plant water status, such as leaf rolling and senescence, photooxidation, leaf chlorophyll concentration, shoot elongation rate, canopy temperature and predawn water potential. Yield under all levels of stress was significantly negatively correlated with AST, and as AST increased due to drought, kernels and ears per plant were significantly reduced. In all populations yield decreased by approximately 10 % per day increase in AST up to 8 days. In several stress situations broad-sense heritability of AST was greater than that of grain yield and the genetic correlation between grain yield and AST approached -1.00. Synthetics formed from one population following bidirectional selection and tested under drought showed adaptive advantage of cool canopy temperature, delayed leaf senescence, reduced AST and erect leaves, especially when all were combined with grain yield in a single index during selection. Selection for reduced AST and high grain yield under drought can be an effective means of improving drought tolerance in tropical maize.

Molecular mapping across three populations reveals a QTL hotspot region on chromosome 3 for secondary traits associated with drought tolerance in tropical maize

Identifying quantitative trait loci (QTL) of sizeable effects that are expressed in diverse genetic backgrounds across contrasting water regimes particularly for secondary traits can significantly complement the conventional drought tolerance breeding efforts. We evaluated three tropical maize biparental populations under water-stressed and well-watered regimes for drought-related morpho-physiological traits, such as anthesis-silking interval (ASI), ears per plant (EPP), stay-green (SG) and plant-to-ear height ratio (PEH). In general, drought stress reduced the genetic variance of grain yield (GY), while that of morpho-physiological traits remained stable or even increased under drought conditions. We detected consistent genomic regions across different genetic backgrounds that could be target regions for marker-assisted introgression for drought tolerance in maize. A total of 203 QTL for ASI, EPP, SG and PEH were identified under both the water regimes. Meta-QTL analysis across the three populations identified six constitutive genomic regions with a minimum of two overlapping traits. Clusters of QTL were observed on chromosomes 1.06, 3.06, 4.09, 5.05, 7.03 and 10.04/06. Interestingly, a ~8-Mb region delimited in 3.06 harboured QTL for most of the morpho-physiological traits considered in the current study. This region contained two important candidate genes viz., zmm16 (MADS-domain transcription factor) and psbs1 (photosystem II unit) that are responsible for reproductive organ development and photosynthate accumulation, respectively. The genomic regions identified in this study partially explained the association of secondary traits with GY. Flanking single nucleotide polymorphism markers reported herein may be useful in marker-assisted introgression of drought tolerance in tropical maize.Electronic supplementary materialThe online version of this article (doi:10.1007/s11032-014-0068-5) contains supplementary material, which is available to authorized users.

Early vertical distribution of roots and its association with drought tolerance in tropical maize

Selection for deep roots to improve drought tolerance of maize (Zea mays L.) requires presence of genetic variation and suitable screening methods. We examined a diverse set of 33 tropical maize inbred lines that were grown in growth columns in the greenhouse up to the 2-, 4-, and 6-leaf stage and in the field in Mexico. To determine length of roots from different depths at high throughput, we tested an approach based on staining roots with methylene blue and measuring the amount of absorbed dye as proxy measure for root length. Staining provided no advantage over root weights that are much easier to measure and therefore preferable. We found significant genotypic variation for all traits at the 6-leaf stage. For development rates between the 2-leaf and the 6-leaf stage, genotypes only differed for rooting depth and the number of crown roots. Positive correlations of leaf area with root length and rooting depth indicated a common effect of plant vigor. However, leaf area in growth columns was negatively related to grain yield under drought (r = −0.50). The selection for deeper roots by an increase in plant vigor likely results in a poorer performance under drought conditions. The proportion of deep roots was independent of other traits but showed a low heritability and was not correlated to field performance. An improved screening protocol is proposed to increase throughput and heritability for this trait.

Validation of the use of secondary traits and selection indices for drought tolerance in tropical maize (Zea mays L.)

Breeding progress for drought tolerance in maize has been slow since drought tolerance is a complex trait controlled by many genes. Breeders improving maize for drought tolerance have therefore been using secondary traits and selection indices for selecting the best genotypes under drought stress. The objectives of this study were to determine the combining ability of the inbred lines in stress and non stress environments as well as compare the use and efficacy of secondary traits and selection indices in selecting for drought tolerant genotypes. Fifty hybrids formed through a North Carolina Design II and four checks were evaluated using the 0.1 alpha lattice planting design under optimum and drought environments. General combining ability (GCA), specific combining ability (SCA) and seven drought indices (SDI), stress susceptibility index (SSI), tolerance (TOL), yield index (YI), yield stability index (YSI), mean productivity index (MPI), stress tolerance index (STI) and geometric productivity index (GMP) were used in the computation of results. Results showed that under drought conditions, GCA was highly significant (P<0.001) for grain yield, anthesis-silking interval (ASI) and ears per plant (EPP) while SCA was significant for grain yield and EPP. Selection indices STI and GMP had positive and significant correlation (P<0.01) with grain yield under both drought stress (Ys) and optimum (Yp) conditions. EPP also had significant (P<0.01) and positive correlation with Ys and Yp. ASI had significant and negative correlation with Ys (P<0.01) and TOL (P <0.05). In addition EPP had a positive and significant correlation with STI (P<0.05) and GMP (P<0.01). The results indicated that ASI, EPP, STI and GMP are effective in identifying high yielding genotypes under different moisture regimes. Narrow sense heritability showed that phenotypic variation attributed to genetic effects increased under stress conditions for EPP and ASI and making them more reliable parameters for use in selecting for genotypes under stress conditions. The two secondary traits (ASI and EPP) together with two drought indices; (STI and GMP) can therefore be used in tandem for increased efficiency in selecting for genotypes under stress environments. Key words: Combining ability drought stress, drought index, heritability.

Gains from improvement for mid-season drought tolerance in tropical maize ( Zea mays L.)

Drought and low soil fertility are major factors limiting maize production in the tropics. We examined the performance of hybrid progenies of drought-tolerant populations (DTP) in stressed (drought and low-N) and unstressed environments. A set of high-yielding normal single cross hybrids, developed using inbred lines improved with main emphasis on yield per se under optimal input conditions, was evaluated and compared with DTP c9 S 3 topcrosses across environments. Performance of normal hybrids was slightly better than DTP-topcrosses under unstressed conditions. However, under stress normal hybrids performed poorly with an average yield of 3.3–4.8% under drought and 34.8–36.2% under low-N stress. Hybrid progenies from DTP yielded up to 31.8–42.4% under drought and 48.9–63.6% under low-N as compared with their yields with out stress. Estimation of gains with selection for mid-season drought stress in DTPs over selection for improved yields under optimal input conditions showed that the gains were remarkable for drought (89.6%), and also for low-N stress (39.3%). The improved performance of DTP hybrids across environments was due to improvements in secondary traits such as reduced anthesis–silking interval, increased ears per plant, delayed senescence and relatively high leaf chlorophyll during late grain filling. Correlations between the performance of genotypes under drought and low-N stresses were positive and fairly high for DTP progenies, both white ( r 2=0.45 ∗,r=0.56 ∗ ) and yellow ( r 2=0.42 ∗,r=0.52 ∗ ), while it was not significant with normal hybrids. Selection for mid-season drought tolerance resulted in morpho-physiological changes that proved advantageous under both drought and low-N stress, without significant yield penalties under optimal input conditions.

Genotype by environment effects and selection for drought tolerance in tropical maize. II. Three-mode pattern analysis

A selection program in three tropical maize populations aimed to improve tolerance of mid-season to late season drought environments while maintaining grain yield (GY) potential. The selection process employed other attributes that included maintaining a constant anthesis date (AD) and, under drought, shortening the anthesis-silking interval (ASI) and increasing ear number per plant (EPP). Three-mode (genotypes x environments x attributes) pattern analysis, which consists of clustering and ordination, should be able to collectively interpret these changes from ten evaluation trials. Mixture maximum likelihood clustering identified four groups that indicated the populations' performance had changed with selection. Groups containing the advanced cycles of selection were higher yielding in most environments and had lower ASI and higher EPP, particularly in drought environments. Check entries with no selection for drought tolerance remained grouped with the initial cycles of selection. A 3 x 2 x 3 (genotypes by environments by attributes) principal component model explained 70% of the variation. For the first environmental component, ASI was shown to be highly negatively correlated with both GY and EPP while anthesis date (AD) was virtually uncorrelated with other traits. The second environmental component (explaining 10% of the variation) contrasted droughted and well-watered environments and showed that EPP and GY were better indicators of this contrast (in terms of changes in population performance) than were AD or ASI. Three-mode analysis demonstrated that improvements with selection occurred in both droughted and well-watered environments and clearly summarised the overall success of the breeding program.

Genotype by environment effects and selection for drought tolerance in tropical maize. I. Two mode pattern analysis of yield

Ten trials evaluated the performance of several late tropical maize populations (La Posta Sequia, Pool 26 Sequia and Tuxpeno Sequia) selected for tolerance to drought during flowering and grain filling and also for yield potential. Families (S1 or full-sib) had been selected recurrently for six to eight years on an index of traits. Pattern (clustering and ordination) analysis was used to analyse the relative performance of entries that included cycles of selection for drought tolerance in the populations and non-drought tolerant checks. Mean environment (E) yields ranged from 1.0 to 10.4 t ha-1. Analysis of variance showed that 97.9% of the total sums of squares was accounted for by E, and that, of the remaining sums of squares the G × E (genotype by environment interaction) was almost 3 times that of the contribution of G alone. Cluster analysis separated the checks, the earlier maturing drought tolerant entries and the later maturing drought tolerant entries. This was verified by principal component (PC) analysis of the G × E matrix. Grouping of the environments (i.e. based on entry performance), resulted in the separation of different types of droughts, and of medium and high yielding well-watered environments. The patterns of discrimination observed indicated that the yield gains under drought would have been unlikely to occur if selection had been done only in well-watered environments. Within each population, selection improved broad adaptation (higher mean yield) to both drought and well-watered environments and cycles of selection ‘jumped’ from non-drought-tolerant to drought-tolerant groups as their specific adaptation to drought environments increased.

The importance of the anthesis-silking interval in breeding for drought tolerance in tropical maize

Selection for improved performance under drought based on grain yield alone has often been considered inefficient, but the use of secondary traits of adaptive value whose genetic variability increases under drought can increase selection efficiency. In the course of recurrent selection for drought tolerance in six tropical maize ( Zea mays L.) populations, a total of 3509 inbred progenies (S 1 to S 3 level) were evaluated in 50 separate yield trials under two or three water regimes during the dry winter seasons of 1986–1990 at Tlaltizapán, México. In over 90% of the trials, ears plant −1, kernels plant −1, weight kernel −1, anthesis-silking interval (ASI), tassel branch number and visual scores for leaf angle, leaf rolling and leaf senescence were determined. Low scores indicated erect, unrolled or green leaves. Canopy temperature, leaf chlorophyll concentration and stem-leaf extension rate were measured in 20–50% of the trials. Across all trials, linear phenotypic correlations ( P < 0.01) between grain yield under drought and these traits, in order listed, were 0.77, 0.90, 0.46, −0.53, −0.16, 0.06 NS, −0.18, −0.11, −0.27, 0.17 and 0.10. Genetic correlations were generally similar in size and sign. None of physiological or morphological traits indicative of improved water status correlated with grain yield under drought, although some had relatively high heritabilities. Genetic variances for grain yield, kernels ear −1, kernels plant −1 and weight kernel −1 decreased with increasing drought, but those for ASI and ears plant −1 increased. Broad-sense heritability for grain yield averaged around 0.6, but fell to values near 0.4 at very low grain yield levels. Genetic correlations between grain yield and ASI or ears plant −1 were weak under well-watered conditions, but approached −0.6 and 0.9, respectively, under severe moisture stress. These results show that secondary traits are not lacking genetic variability within elite maize populations. Their low correlation with grain yield may indicate that variation in grain yield under moisture stress is dominated by variation in ear-setting processes related to biomass partitioning at flowering, and much less by factors putatively linked to crop water status. Field-based selection programs for drought tolerance should consider these results.

Arbuscular mycorrhizas and water relations in maize under drought stress at tasselling

summaryA greenhouse experiment was conducted to investigate the influence of an arbuscular mycorrhizal (AM) fungus (Glomus intraradices Schenck &amp; Smith) on drought tolerance in tropical maize. Freshly regenerated seeds of selection cycles 0 (cv. C0, drought‐sensitive) and 8 (cv. C8, drought‐resistant) of the lowland tropical population ‘Tuxpeño sequia’ were used for this study. Maize plants were subjected to drought stress for three weeks following tasselling (75–95 days after sowing). During the drought stress period, midday leaf water potential (LWP), stomatal resistance (SR) and transpiration rate (TR) were monitored daily, and green leaf area (GLA) determined at alternate days to assess the effects of mycorrhizal inoculation. Mycorrhizal plants of the cultivars CO and C8 had higher (less negative) LWP and TR, and lower SR throughout the experiment. The GLA was 27.5 % higher in mycorrhizal than non‐mycorrhizal CO plants under drought conditions. Results indicate that mycorrhizal significantly improve tolerance to moderate drought stress imposed at tasselling, especially for the sensitive CO maize cultivar.