Abstract:
Development of nitrogen (N) efficient bread wheat cultivars better suited to limiting nitrogen
condition is one of the prime objectives in plant breeding research. Three years (2013/14,
2014/15 and 2015/16) study was carried out to investigate genetic variability, identify traits
influencing N use efficiency (NUE) and their pattern of inheritance using Hayman and Griffing
approaches at The University of Agriculture, Peshawar Pakistan. During 2013/14, 30 wheat
varieties released during 1981 till 2013 were preliminary screened under high nitrogen
(Recommended dose, N+) and low nitrogen (N0) conditions as independent experiments.
Significant G×E for all traits justified independent analysis. Independent analysis revealed
significant genotypic variation for all traits under each level of nitrogen. Significant reductions
under N0 condition for all traits signified the role of nitrogen fertilization in crop production.
Path coefficient analysis identified biological yield, grain nitrogen content, harvest index and
grain growth rate as direct contributors, whereas, tillers m-2 and 1000 grains weight, biological
growth rate, total nitrogen uptake and nitrogen harvest index as indirect contributors for the
improvement of NUE in bread wheat. Based on stress selection indices, cluster analysis,
principal components analysis and 3D biplot analysis Shahkar-13 (Shk), Pirsabak-05 (PS) and
Tatara (Tat) were classified as N-efficient, while Hashim-08 (Hsm), Inqilab-91(Inq) and Pak-81
(Pak) were found as N-inefficient varieties. During 2014/15, an objectives-based hybridization
program was initiated by crossing two contrasting groups of varieties to generate a 6 × 6
straight diallel. In the following year, 15 F1 hybrids along with six parent cultivars were
evaluated under N+ and N0 conditions. Significant GEI for all traits necessitated independent
analysis under each nitrogen level, which exhibited significant variation among genotypes
justifying genetic analysis for all traits. Diallel analysis of variance exhibited preponderance of
non-additive gene action in the inheritance of all traits under both N+ and N0 conditions.
However, non-additive genetic effects for days to emergence, anthesis, maturity and grain
filling duration were changed to additive genetic control under N0 condition. Pre-dominant
role of non-additive genetic effects was further validated by the greater proportion of
dominance than additive genetic variance, higher magnitude of H1 than D genetic component,
proportion of dominant genes, average degrees of dominance (>1), Wr/Vr graphs and low
narrow sense heritability. Position of parents along the regression line expressed that Tatara20
96 and Hashim-08 possessed more dominant and recessive genes, respectively for most of
the traits. Heritability estimates both in narrow and broad sense were greater under N+ than
N0 condition for most of the traits. Low to moderate ratio of narrow and broad sense
heritability for most of the traits exhibited that non-additive variance was predominant than
additive variance in the expression of these traits. Combining ability analysis revealed the
significance of both GCA and SCA effects for almost all traits. Traits with GCA/SCA ratio less
than 1 indicated importance of non-additive genetics effects in phenotypic expression of
these traits. Tatara appeared as best general combiner for all important NUE and yield
associated traits. Relative ranking of crosses for SCA effects were not the same under both N+
and N0 conditions. The F1 hybrid, Shk × Hsm expressed maximum SCA effects for most of the
important traits under both N+ and N0 conditions, whereas, PS × Shk was ranked as top
specific combination for most of the traits only under N0 condition. Correlation analysis
suggested that mean values of both parents and F1 hybrids predict their suitability as general
or specific combiner. Better parent heterosis was exhibited by cross combinations Shk × Hsm
and PS × Tat for most of the important traits under N+ and N0 conditions, respectively. Both
Hayman and Griffing analyses revealed the involvement of non-additive gene actions in the
phenotypic expression of most of the traits. Non-additive genetic variance of traits warranted
improvement through delayed selection. Besides exploring genetics of N-efficient lines, this
study also identified some indices for selecting N-efficient wheat lines. Cross combinations
Shk × Hsm, PS × Shk and PS × Tat with higher SCA and heterobiltiosis may be useful in evolving
wheat cultivars with enhanced NUE.