One of the biggest challenges for genetic studies on natural or unstructured populations is the unbalanced datasets where individuals are measured at different times and environments. This problem is ...also common in crop and animal breeding where many individuals are only evaluated for a single year and large but unbalanced datasets can be generated over multiple years. Many wheat breeding programs have focused on increasing bread wheat (Triticum aestivum L.) yield, but processing and end-use quality are critical components when considering its use in feeding the rising population of the next century. The challenges with end-use quality trait improvements are high cost and seed amounts for testing, the latter making selection in early breeding populations impossible. Here we describe a novel approach to identify marker-trait associations within a breeding program using a meta-genome wide association study (GWAS), which combines GWAS analysis from multi-year unbalanced breeding nurseries, in a manner reflecting meta-GWAS in humans. This method facilitated mapping of processing and end-use quality phenotypes from advanced breeding lines (n = 4,095) of the CIMMYT bread wheat breeding program from 2009 to 2014. Using the meta-GWAS we identified marker-trait associations, allele effects, candidate genes, and can select using markers generated in this process. Finally, the scope of this approach can be broadly applied in 'breeding-assisted genomics' across many crops to greatly increase our functional understanding of plant genomes.
Pre-harvest sprouting (PHS) in wheat can cause substantial reduction in grain yield and end-use quality. Grain color (GC) together with other components affect PHS resistance. Several quantitative ...trait loci (QTL) have been reported for PHS resistance, and two of them on chromosome 3AS (TaPHS1) and 4A have been cloned.
To determine genetic architecture of PHS and GC and genetic relationships of the two traits, a genome-wide association study (GWAS) was conducted by evaluating a panel of 185 U.S. elite breeding lines and cultivars for sprouting rates of wheat spikes and GC in both greenhouse and field experiments. The panel was genotyped using the wheat 9K and 90K single nucleotide polymorphism (SNP) arrays.
Four QTL for GC on four chromosomes and 12 QTL for PHS resistance on 10 chromosomes were identified in at least two experiments. QTL for PHS resistance showed varied effects under different environments, and those on chromosomes 3AS, 3AL, 3B, 4AL and 7A were the more frequently identified QTL. The common QTL for GC and PHS resistance were identified on the long arms of the chromosome 3A and 3D.
Wheat grain color is regulated by the three known genes on group 3 chromosomes and additional genes from other chromosomes. These grain color genes showed significant effects on PHS resistance in some environments. However, several other QTL that did not affect grain color also played a significant role on PHS resistance. Therefore, it is possible to breed PHS-resistant white wheat by pyramiding these non-color related QTL.
Wheat (
L.) grain yield response to plant density is inconsistent, and the mechanisms driving this response are unclear. A better understanding of the factors governing this relationship could ...improve plant density recommendations according to specific environmental and genetics characteristics. Therefore, the aims of this paper were to: i) execute a synthesis-analysis of existing literature related to yield-plant density relationship to provide an indication of the need for different agronomic optimum plant density (AOPD) in different yield environments (YEs), and ii) explore a data set of field research studies conducted in Kansas (USA) on yield response to plant density to determine the AOPD at different YEs, evaluate the effect of tillering potential (TP) on the AOPD, and explain changes in AOPD
variations in wheat yield components. Major findings of this study are: i) the synthesis-analysis portrayed new insights of differences in AOPD at varying YEs, reducing the AOPD as the attainable yield increases (with AOPD moving from 397 pl m
for the low YE to 191 pl m
for the high YE); ii) the field dataset confirmed the trend observed in the synthesis-analysis but expanded on the physiological mechanisms underpinning the yield response to plant density for wheat, mainly highlighting the following points: a) high TP reduces the AOPD mainly in high and low YEs, b) at canopy-scale, both final number of heads and kernels per square meter were the main factors improving yield response to plant density under high TP, c) under varying YEs, at per-plant-scale, a compensation between heads per plant and kernels per head was the main factor contributing to yield with different TP.
Plant breeding has increased the yield of winter wheat (
L.) over decades, and the rate of genetic gain has been faster under high fertility in some countries. However, this response is not ...universal, and limited information exists on the physiological traits underlying the interaction between varieties and fertilization. Thus, our objectives were to identify the key shifts in crop phenotype in response to selection for yield and quality, and to determine whether historical and modern winter wheat varieties respond differently to in-furrow fertilizer. Factorial field experiments combined eight winter wheat varieties released between 1920 and 2016, and two fertilizer practices control
112 kg ha
in-furrow 12 -40-0-10-1 (N-P-K-S-Zn) in four Kansas environments. Grain yield and grain N-removal increased nonlinearly with year of release, with greater increases between 1966 and 2000. In-furrow fertilizer increased yield by ~300 kg ha
with no variety × fertility interaction. Grain protein concentration related negatively to yield, and the residuals of this relationship were unrelated to year of release. Yield increase was associated with changes in thermal time to critical growth stages, as modern varieties had shorter vegetative period and longer grain filling period. Yield gains also derived from more kernels m
resultant from more kernels head
. Historical varieties were taller, had thinner stems, and allocated more biomass to the stem than semidwarf varieties. Yield gains resulted from increases in harvest index and not in biomass accumulation at grain filling and maturity, as shoot biomass was similar among varieties. The allometric exponent (i.e., the slope between log of organ biomass and log of shoot biomass) for grain increased with, and for leaves was unrelated to, year of release. The ability of modern varieties to allocate more biomass to the kernels coupled to an early maturity increased grain yield and grain N-removal over time. However, increases in grain yield were greater than increases in grain N-removal, reducing grain protein concentration in modern varieties.
A great majority of dryland wheat producers are reluctant to intensify management due to the assumption that lack of water availability is the most critical factor limiting yield and thus, the ...response to management intensification would be limited. We conducted on-farm field experiments across three locations and two growing seasons in Kansas using 21 modern winter wheat genotypes grown under either standard (SM) or intensified management (IM) systems. The goals of this study were to (i) determine whether the SM adopted is adequate to reach achievable yields by farmers in the region and (ii) identify differences in responsiveness to IM among a range of modern genotypes. Across all sites-years and genotypes, the IM increased yield by 0.9 Mg ha
, outyielding the SM system even in the lowest yielding conditions. As expected, the yield response to IM increased with the achievable yield of the environment and genotype. Across all sources of variation, the yield responsiveness to IM was related to increased biomass rather than harvest index, strongly driven by improvements in grain number (and independent of changes in grain weight), and by improvements in N uptake which resulted from greater biomass and shoot N concentration. The IM system generally also increased grain N concentration and decreased the grain N dilution effect from increased yield. Genotypes varied in their response to IM, with major response patterns resulting from the combination of response magnitude (large vs. small) and consistency (variable vs. consistent). Genotypes with high mean response and high variability in the response to IM across years could offer greater opportunities for producers to maximize yield as those genotypes showed greater yield gain from IM when conditions favored their response. For the background conditions evaluated, intensifying management could improve wheat yield in between c. 0.2 and 1.5 Mg ha
depending on genotype.
Wheat (Triticum aestivum L.) is a temperate cereal with an optimum temperature range of 15–22°C during the grain filling stage. Heat stress is one of the major environmental constraints for wheat ...production worldwide. Temperatures above 25°C during the grain filling stage significantly reduced wheat yield and quality. This reduction was reported due to the inactivation of the soluble starch synthase, a key heat-labile enzyme in starch transformation of wheat endosperm. To improve wheat productivity under heat stress, the rice soluble starch synthase I, under the control of either a constitutive promoter or an endosperm-specific promoter, was expressed in wheat and the transgenic lines were monitored for expression and the effects on yield-related traits. The results showed that the transgenic wheat events expressed rice soluble starch synthase I at a high level after four generations, and transgenic plants produced grains of greater weight during heat stress. Under heat stress conditions, the thousand kernel weight increased 21–34% in T2 and T3 transgenic plants compared to the non-transgenic control plants. In addition, the photosynthetic duration of transgenic wheat was longer than in non-transgenic controls. This study demonstrated that the engineering of a heat tolerant soluble starch synthase gene can be a potential strategy to improve wheat yield under heat stress conditions.
Core Ideas
Genomic selection applied for wheat quality in CIMMYT spring bread wheat breeding program.
All wheat quality traits predicted and validated using forward genomic selection.
Dough and loaf ...traits have moderately high predictive ability in CIMMYT breeding program.
Genomic selection genetic gain 1.4 to 2.7 times higher than phenotypic selection.
Wheat (Triticum aestivum L.) cultivars must possess suitable end‐use quality for release and consumer acceptability. However, breeding for quality traits is often considered a secondary target relative to yield largely because of amount of seed needed and expense. Without testing and selection, many undesirable materials are advanced, expending additional resources. Here, we develop and validate whole‐genome prediction models for end‐use quality phenotypes in the CIMMYT bread wheat breeding program. Model accuracy was tested using forward prediction on breeding lines (n = 5520) tested in unbalanced yield trials from 2009 to 2015 at Ciudad Obregon, Sonora, Mexico. Quality parameters included test weight, 1000‐kernel weight, hardness, grain and flour protein, flour yield, sodium dodecyl sulfate sedimentation, Mixograph and Alveograph performance, and loaf volume. In general, prediction accuracy substantially increased over time as more data was available to train the model. Reflecting practical implementation of genomic selection (GS) in the breeding program, forward prediction accuracies (r) for quality parameters were assessed in 2015 and ranged from 0.32 (grain hardness) to 0.62 (mixing time). Increased selection intensity was possible with GS since more entries can be genotyped than phenotyped and expected genetic gain was 1.4 to 2.7 times higher across all traits than phenotypic selection. Given the limitations in measuring many lines for quality, we conclude that GS is a powerful tool to facilitate early generation selection for end‐use quality in wheat, leaving larger populations for selection on yield during advanced testing and leading to better gain for both quality and yield in bread wheat breeding programs.
Senescence is a genetically programmed and environmentally influenced process resulting in the destruction of chlorophyll and remobilization of nutrients to younger or reproductive parts of plants. ...Delayed senescence, or stay-green, contributes to a long grain-filling period and stable yield under stress. To model senescence and identify quantitative trait loci (QTL) for the trait, a population of recombinant inbred lines (RIL) from a cross between winter wheat cultivars, ‘Ventnor' and ‘Karl 92' was evaluated for heat tolerance under optimum temperature of 20/15°C (day/night) and continuous heat stress of 30/25°C from 10 days after anthesis (DAA) until maturity. Ventnor is a heat-tolerant cultivar and Karl 92 is a relatively heat-susceptible cultivar. Green leaf area was measured and used to model percent greenness retained over the reproductive period. Chlorophyll content and chlorophyll fluorescence were recorded on flag leaves. Senescence was converted to a quantitative trait using the model. Based on the modeled parameters, the RILs were categorized into three groups. When senescence-related traits were evaluated, nine QTL for heat tolerance were found on chromosome 2A, two each on chromosomes 6A and 6B and one each on chromosome 3A, 3B, and 7A. Both parents contributed favorable alleles for most of the senescence-related traits. Microsatellite markers Xgwm356 and Xgwm5 prominently linked to the senescence-related traits may be useful in marker-assisted breeding. These and the linked AFLP (amplified fragment length polymorphism) markers XCGT.TGCG-349, XCGT.GTG-343, and XCGT.CTCG-406, if converted to STS (sequence tagged sites), can be used for further molecular dissection of the QTL for post-anthesis heat tolerance.
Increasing temperatures can severely affect wheat (Triticum aestivum L.) production, particularly when it coincides with the grain‐filling period. Heat stress induces rapid senescence resulting in ...early maturity and shortened grain‐filling period. In this study, the applicability of in vivo chlorophyll fluorescence (Chl‐F) and chlorophyll index to track rate of senescence in flag leaves and spikes exposed to heat stress were investigated. Seven winter wheat varieties were exposed to post‐flowering heat stress using growth chambers 35/15 °C (heat stress) and 25/15 °C (control) day/night and unique field‐based heat tents (imposed +6 °C higher than ambient). Effective quantum yield of photosystem II (PSII) (QY) was recorded temporally in flag leaves and spikes, and compared with in vitro chlorophyll‐a (Chl‐a) concentration and non‐invasive estimation of chlorophyll and flavonoids index. Time point indicating the start of senescence (change‐point, CP) for QY was advanced by 0–8 and 0–6 d in flag leaves and spikes, respectively, under heat stress. In the chamber experiment, sustained heat stress induced accelerated decline of QY, particularly in wheat cultivars Larry and WB4458. Stronger positive relationship between days to senescence in spikes and thousand kernel weight indicated an extended period of assimilate supply from sink compared to the source tissue, during grain filling. Capturing heat stress‐induced changes in photosynthetic pigments and QY at high temporal frequency presents an effective phenotyping approach for testing genetic diversity in large‐scale field experiments involving different crops.
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