Abstract
High grain yield (GY) is the primary selection target in commercial hard winter wheat (Triticum aestivum L.) breeding programs, with milling and bread-making quality as important secondary selection targets. Grain protein concentration (GPRO) is strongly correlated with important dough rheology and bread-making characteristics. Simultaneous improvement is difficult given the strong negative relationship of GY and GPRO in cereal crops. Nitrogen use efficiency (NUE), defined as the amount of grain produced per unit of N supply, promotes high GY through the component traits N uptake (NUpE) and N utilization (NUtE) efficiencies. Grain protein accumulation relies on N uptake from the soil and remobilization from plant tissue reserves. One study was conducted to characterize variation for NUE among a set of 20 breeding lines and varieties adapted to the west central Great Plains of the United States. Path analysis was applied to characterize the NUE component structure during the 2010-2011 growing season and then for two newly released varieties in the 2011-2012 growing season. Nitrogen use efficiency ranged from 39.9 kg kg-1 for 'RonL' to 46.7 kg kg-1 for 'Byrd'. By path analysis, we determined that variation in NUE depended on NUpE under N sufficiency and on NUtE under limiting N. Additionally, strategies for simultaneous improvement of GY and GPRO were explored. Analysis of standardized residuals of the linear regression of GPRO on GY, or 'grain protein deviation', identified one cultivar ('Brawl CL Plus') that had 6.7 g kg-1 higher GPRO than the average for all 20 genotypes. In a second study, selection strategies based on protein-yield selection indices for a set of 775 breeding lines and varieties representing the Colorado State University hard winter wheat breeding program were evaluated based on field data obtained during the 2012-2015 growing seasons. Selection based on high values for a particular index delivered a characteristic emphasis on GY or GPRO. Correlation analysis between index values and GY or GPRO showed that each simultaneous selection strategy focused to differing extents on the primary traits. Genomic selection applied to index values in univariate models provided forward prediction accuracy ranging between r = .21 to .44 for the 2013 validation set, but approached zero for the 2014 validation set. Index values were also calculated from genomic estimated breeding values obtained in bivariate genomic selection models. Prediction accuracy for individual trait values was not substantially improved in the bivariate model. Protein-yield indices calculated from bivariate genomic estimated breeding values showed similar relationships to GY and GPRO as for the genomic estimated breeding values for indices calculated in the univariate models. A set of selection strategies generate sufficient predictive ability in phenotypic or genomic selection to be effective tools for simultaneous selection for GY and GPRO.
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Claims 10
Claim · model-written summary
Nitrogen use efficiency ranged from 39.9 kg kg-1 for 'RonL' to 46.7 kg kg-1 for 'Byrd' among 20 breeding lines and varieties.
- Taxon
- Triticum aestivum
Triticum aestivumt0 - Outcome
- Nitrogen use efficiencye1
- Value
- 39.9 kg kg-1 for 'RonL' to 46.7 kg kg-1e1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Nitrogen use efficiency ranged from 39.9 kg kg-1 for 'RonL' to 46.7 kg kg-1 for 'Byrd'”
t0“hard winter wheat (Triticum aestivum L.)”
Claim · model-written summary
Variation in NUE depended on N uptake efficiency (NUpE) under N sufficiency and on N utilization efficiency (NUtE) under limiting N.
- Taxon
- Triticum aestivum
Triticum aestivumt0 - Treatment
- N sufficiencye2
- Outcome
- variation in NUEe2
- Qualifier
- NUpE under N sufficiency and on NUtE under limiting Ne2
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e2“By path analysis, we determined that variation in NUE depended on NUpE under N sufficiency and on NUtE under limiting N”
t0“hard winter wheat (Triticum aestivum L.)”
Claim · model-written summary
The cultivar 'Brawl CL Plus' had 6.7 g kg-1 higher GPRO than the average for all 20 genotypes.
- Taxon
- Triticum aestivum
Triticum aestivumt0 - Material
- Brawl CL Pluse3
- Outcome
- GPROe3
- Direction
- highere3
- Value
- 6.7 g kg-1e3
- Comparator
- the average for all 20 genotypese3
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e3“identified one cultivar ('Brawl CL Plus') that had 6.7 g kg-1 higher GPRO than the average for all 20 genotypes”
t0“hard winter wheat (Triticum aestivum L.)”
Claim · model-written summary
Selection based on high values for a particular index gave a characteristic emphasis on GY or GPRO.
- Taxon
- Triticum aestivum
Triticum aestivumt0 - Treatment
- Selection based on high values for a particular indexe4
- Outcome
- emphasis on GY or GPROe4
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e4“Selection based on high values for a particular index delivered a characteristic emphasis on GY or GPRO”
t0“hard winter wheat (Triticum aestivum L.)”
Claim · model-written summary
Each simultaneous selection strategy focused to differing extents on the primary traits, as shown by correlation of index values with GY or GPRO.
- Taxon
- Triticum aestivum
Triticum aestivumt0 - Treatment
- each simultaneous selection strategye5
- Outcome
- Correlation analysis between index values and GY or GPROe5
Not reportedDirection
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e5“Correlation analysis between index values and GY or GPRO showed that each simultaneous selection strategy focused to differing extents on the primary traits”
t0“hard winter wheat (Triticum aestivum L.)”
Claim · model-written summary
Genomic selection on index values in univariate models gave forward prediction accuracy of r = .21 to .44 for the 2013 validation set.
- Taxon
- Triticum aestivum
Triticum aestivumt0 - Outcome
- forward prediction accuracye6
- Value
- r = .21 to .44e6
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e6“Genomic selection applied to index values in univariate models provided forward prediction accuracy ranging between r = .21 to .44 for the 2013 validation set”
t0“hard winter wheat (Triticum aestivum L.)”
Claim · model-written summary
Forward prediction accuracy approached zero for the 2014 validation set.
- Taxon
- Triticum aestivum
Triticum aestivumt0 - Outcome
- forward prediction accuracye7
- Value
- approached zeroe7
- Qualifier
- 2014 validation sete7
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e7“forward prediction accuracy ranging between r = .21 to .44 for the 2013 validation set, but approached zero for the 2014 validation set”
t0“hard winter wheat (Triticum aestivum L.)”
Claim · model-written summary
Bivariate genomic selection did not substantially improve prediction accuracy for individual trait values compared with univariate models.
- Taxon
- Triticum aestivum
Triticum aestivumt0 - Outcome
- Prediction accuracy for individual trait valuese1
- Direction
- not substantially improvede1
- Comparator
- univariate modelse2
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Prediction accuracy for individual trait values was not substantially improved in the bivariate model”
e2“applied to index values in univariate models”
t0“hard winter wheat (Triticum aestivum L.)”
Claim · model-written summary
Protein-yield indices from bivariate genomic estimated breeding values showed relationships to GY and GPRO similar to those of the univariate-model indices.
- Taxon
- Triticum aestivum
Triticum aestivumt0 - Treatment
- bivariate genomic estimated breeding valuese1
- Outcome
- relationships to GY and GPROe1
- Direction
- similare1
- Comparator
- genomic estimated breeding values for indices calculated in the univariate modelse1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“Protein-yield indices calculated from bivariate genomic estimated breeding values showed similar relationships to GY and GPRO as for the genomic estimated breeding values for indices calculated in the univariate models”
t0“hard winter wheat (Triticum aestivum L.)”
Claim · model-written summary
The authors recommend selection strategies with sufficient predictive ability as effective tools for simultaneous selection for GY and GPRO.
- Taxon
- Triticum aestivum
Triticum aestivumt0 - Outcome
- effective tools for simultaneous selection for GY and GPROe1
Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Verbatim
e1“A set of selection strategies generate sufficient predictive ability in phenotypic or genomic selection to be effective tools for simultaneous selection for GY and GPRO”
t0“hard winter wheat (Triticum aestivum L.)”