Pattern of genetic inheritance of grain yield and grain related traits in rabi sorghum Sorghum bicolor (L.) Moench

doi:10.21203/rs.3.rs-1563283/v1

Grain related and yield contributing traits greatly influences both grain size and grain yield. In present experiment four crosses viz., BJV 44 × DSMR-8, BJV 44 × DSMR-4, PKV Kranti × DSMR-8 and PKV Kranti × DSMR-4 were studied for inheritance pattern during rabi-2020. Both additive and dominance gene actions were found to be predominant for all the traits in the cross BJV 44 × DSMR-8, whereas rest of the crosses exhibited additive gene action for some traits and dominance for other traits. The majority of the crosses exhibited duplicate gene interactions for most of the traits and were found suitable for population improvement program whereas some crosses, showed complementary gene interaction and were found suitable for heterosis breeding. While two crosses viz., BJV 44 × DSMR-4 and PKV Kranti × DSMR-4 were found suitable for heterosis breeding for grain yield/plant. It is the first report on inheritance pattern of grain size viz., grain length, width, thickness and other grain related traits such as 100 grain volume and density studied using generation analysis.

Sorghum

grain size

grain yield

genetic inheritance

generation mean analysis

A C₄ plant with higher photosynthetic efficiency and abiotic stress tolerance (Nagy et al., 1995; Reddy et al., 2009), with center of origin in Africa, sorghum (Sorghum bicolor L. Moench) is adapted to a diverse set of environments ranging from arid and semiarid to tropical and temperate regions throughout the world. On world basis, sorghum is the fifth major cereal crop in the extent of production following wheat, rice, maize and barley. In addition to its significance as a food and fodder crop, sorghum plays an important role in supplying micronutrients at low cost (Rao et al., 2006; 2010). As a species, sorghum bicolor with 2n = 2x = 20 chromosome number belong to family Poaceae, sub family panicoideae, tribe Andropogene and is classified into five distinct races; bicolor, caudatum, durra, guinea and kafir, and an array of intermediate classes (Harlan and de Wet, 1972). Sorghum is an often cross pollinated crop with 5 to 10 per cent cross pollination and may exceed up to 40 per cent out crossing (Barnaud et al., 2008).

World’s sorghum production of 57.50 million tones is produced from the area of 40.28 mha with the productivity of 1.43 tons per hectare in 2019-20 (Foreign Agricultural Service/USDA, 2020). India shares about 8 per cent of world sorghum production after USA and Nigeria and ranks third in area after Sudan and Nigeria.

In India, sorghum is primarily grown for food and fodder purposes in both rainy and post-rainy seasons. The total production and productivity of sorghum is of 4.63 million tons and around 1 ton per hectare, respectively from an area of 4.66 m ha (Indiastat, 2019–2020).

The grain related traits viz., grain length, grain width, grain thickness, 100 grain weight, 100 grain volume, grain density, grains/panicle and days to flowering showed positive and significant correlation with grain yield while path analysis also exhibited positive and significant direct & indirect effect on grain yield (Verma and Biradar, 2021). So, the genetic study is prerequisite for the improvement of these traits.

Knowledge of inheritance of these traits is critical to exploit the large variability identified in sorghum. Generation mean analysis provides information on the relative importance of additive effects, dominance deviations and effects due to non-allelic gene interactions, in determining genotypic values of the individuals and, consequently, means genotypic values of families and generations. Therefore, an attempt was made to determine the inheritance pattern of grain yield and grain related traits using the generation mean analysis, trait heritability and type of gene action prevailing in rabi sorghum.

Two promising varieties viz., BJV 44 & PKV Kranti and two strong restorers such as DSMR-8 & DSMR-4 (Verma et al., 2022) were crossed to develop four crosses viz., BJV 44 × DSMR-8, BJV 44 x DSMR-4, PKV Kranti × DSMR-8 and PKV Kranti × DSMR-4. Six generations, i e., P₁, P₂, F₁, F₂, B₁ and B₂ were developed during summer and kharif seasons of 2020. In summer-2020 the F₁’s of all the crosses generated and in kharif-2020 the B₁ and B₂ populations were generated by crossing the F₁’s with their parents. Simultaneously, The F₁’s were advanced to generate F₂ population. Since development of generations involve crossing and backcrossing in four different cross combinations, the seeds were not sufficient to evaluate in multiple seasons/year. Therefore only one season evaluation was carried out in post rainy season-2020. All the six generations were evaluated during post rainy season of 2020 at the Botany Garden, College of Agriculture, University of Agricultural Sciences, Dharwad, state of Karnataka, India. The experimental site is located at an altitude of 750 m above mean sea level, latitude of 15.46^oN and longitude of 75.01^oE.

To assess the six generations of all the four crosses, a trial was conducted without replications; the population size and characteristics of the parents is given in the Table 1. All the recommended package of practices were followed to raise a healthy crop. Data were recorded on days to flowering, grain yield/plant (g), grains/panicle, grain related traits viz., grain width (mm), grain length (mm), grain thickness (mm), 100 grain weight (g), 100 grain volume (cc), grain density and plant height (cm). The grain size related traits were measured using the varnier caliper while, the volume of the 100 grains were measured based on the water displacement method.

Table 1

Characteristics of parents used and population size of the different generations of different crosses
Cross-1: BJV 44 × DSMR-8			Cross-2: BJV 44 x DSMR-4
Sl. No	Generation	Population Size	Generation	Population Size
1	P₁ (BJV-44- High grain yield and quality)	20	P₁ (BJV-44- High grain yield and quality)	20
2	P₂ (DSMR-4- Strong restorer and bold seeds)	20	P₂ (DSMR-4- Strong restorer and bold seeds)	20
3	F₁	20	F₁	20
4	F₂	453	F₂	453
5	B₁	158	B₁	158
6	B₂	157	B₂	157
Cross-3: PKV Kranti × DSMR-8			Cross-4: PKV Kranti × DSMR-4
1	P₁ (PKV Kranti- High grain yield and quality)	20	P₁ (PKV Kranti- High grain yield and quality)		30
2	P₂ (DSMR-8- Strong restorer and bold seeds)	22	P₂ (DSMR-4- Strong restorer and bold seeds)		24
3	F₁	19	F₁		22
4	F₂	455	F₂		453
5	B₁	156	B₁		156
6	B₂	154	B₂		158

Statistical analysis

The data obtained on different characters under study were subjected to generation mean analysis by following Mather (1949), scaling test (Hayman and Mather, 1955) for associated scaling test and (Cavalli, 1952) for joint scaling test and Hayman (1958) approach to find the significant non allelic interaction. The validity of this model for the scaling tests and non allelic interactions was examined using Indostat 9.1 software.

Means and variances

The mean and variances for different traits in different generations are presented in Table 2. The experiment exhibited adequate variance for all the traits in all the crosses.

Adequacy of additive-dominance model

The scaling tests were performed to determine the efficacy of additive dominance model for the inheritance of grain yield and grain related traits. The scaling test values were found significantly differed from zero for all the studied traits in all the crosses except for grain density in PKV Kranti × DSMR-4, indicating the presence of non allelic gene interaction and dominance additive model is inadequate for explaining the inheritance of these traits (Table 3). Whereas, the additive dominant model found to be adequate for the grain density in PKV Kranti × DSMR-4. The joint scaling test also showed significant values and supported the inadequacy of the simple additive dominance model for all the traits in all the crosses except PKV Kranti × DSMR-4 for the grain density. These results are in harmony with earlier reports of Rao et al. (1994), Singh et al. (2007), Kumar et al. (2011), Salmi et al. (2019), Koubisy (2019) and Feltaous et al. (2020).

Gene effects

The six parameter model was followed to determine the nature and magnitude of gene effects involved in the genetic control of studied traits. The estimated mean effect parameter [m], which reflects the contribution due to the overall mean plus the locus effects and interactions of the fixed loci, was found to be significant for all the traits in all the crosses except for the grain density in PKV Kranti × DSMR-4, indicating that these characters are quantitatively inherited and showed the importance of non-allelic interactions.

The additive [d] gene effect was recorded negative and significant for grain length in BJV 44 x DSMR-4, for grain density in BJV 44 × DSMR-8 and for grain width in BJV 44 × DSMR-4 & PKV Kranti × DSMR-4. While, it was found to be non significant for grain volume in PKV Kranti × DSMR-4, for 100 grain weight in BJV 44 × DSMR-8 and PKV Kranti × DSMR-4, for grain width in PKV Kranti × DSMR-8 and for grain density in BJV 44 x DSMR-4. Rest of the traits of the crosses mentioned above showed significant additive effect. Interestingly, the traits, viz., days to flowering, grain yield/plant, grain thickness, grains/panicle and plant height showed significant additive effects in all the crosses.

The dominance effect [h] was recorded significant for days to flowering, grain yield/plant, 100 grain weight, grain width, grain length, grain thickness, grain volume, grains/panicle and plant height in all the crosses. Whereas, it was non significant for grain width, and grain thickness in PKV Kranti × DSMR-4 and for grain density in PKV Kranti × DSMR-8.

Additive × additiveinteraction [i] was found to be significant in all the crosses for days to flowering, grain yield/plant, grain length, grain thickness, grains/panicle and plant height. Whereas, it was significant for 100 grain weight and volume in all the crosses except PKV Kranti × DSMR-4. The crosses viz., PKV Kranti × DSMR-8 and BJV 44 × DSMR-8 were recorded significant while, rest of the crosses were non significant for grain density. In the crosses viz., BJV 44 × DSMR-8 and BJV 44 × DSMR-4 the additive × additiveinteraction [i] was found to be significant for grain width whereas, it was non significant in rest of the crosses.

In all the crosses except PKV Kranti × DSMR-4 for days to flowering and 100 grain volume, in BJV 44 × DSMR-4 & PKV Kranti × DSMR-8 for 100 grain weight and grain length, in BJV 44 × DSMR-4 & PKV Kranti × DSMR-8 for 100 grain weight & grain length and in BJV 44 × DSMR-8 & BJV 44 × DSMR-4 for grain density and plant height the additive × dominance interaction [j] was found to be significant. Interestingly, it was significant in all the crosses for grain yield/plant and grains/panicle.

The dominance × dominance interaction [l] effect was noticed significant in all the studied crosses for days to flowering, grain yield/plant, 100 grain weight, grain width, grain thickness, 100 grain volume, grains/panicle and plant height. Whereas for grain length, it was significant in all the crosses except PKV Kranti × DSMR-8, while for grain thickness and grain density, it was significant in all the crosses except PKV Kranti × DSMR-4. Dominance × dominance interaction [l] effects were found to be higher than additive × dominance and additive × additiveinteractions for most of the crosses. These results are in harmony with the earlier reports of Vasudev Rao and Goud (1977), Hoshmand and Rezai (1997), Biradar et al. (2000), Pahuja (2003), Narain (2007), Singh et al. (2007), Prakash et al. (2010), Kumar et al. (2011), Shivani and Sreelakshmi (2013), Vyas (2014) and Aruna et al. (2015) Gaddameedi et al. (2018) and Rokade et al. (2021).

Types of epistasis

The type of epistasis is determined based on the signs of dominance [h] and dominance × dominance [l] interaction. When these effects are in same direction, the epistasis is complementary type if not it is duplicate type. The different type of epistasis observed in the present study for different characters in the different crosses and details are furnished in Table 4.

Most of the traits of all the crosses were controlled by duplicate type of interactions. Interestingly, all the traits of the cross BJV 44 × DSMR-8 exhibited duplicate type of gene interaction. These results are in agreement with the reports of Singh et al. (2007), Narain et al. (2007), Shivani and Sreelakshmi (2013), Vyas et al. (2014), Gaddameedi et al. (2018) and Mohammed et al. (2018).

Some traits viz., days to flowering, grain yield/plant, grain/panicle in BJV 44 x DSMR-4, grain length, grain width and plant height in PKV Kranti × DSMR-8 and days to flowering, 100 grain weight, grain width, grain thickness & plant height in PKV Kranti × DSMR-4 exhibited complementary type of epistasis. Earlier workers such as Kumar et al. (2011) and Singh et al. (2007) reported the similar results.

Components of genetic variation

It can be seen from Table 5 that the additive variance (σ²a) was recorded greater than the dominance variance (σ²d) in all the crosses for days to flowering, grain yield/plant, 100 grain weight, grain width, grain length, grain thickness, grain density, grains/panicle and plant height. Whereas for 100 grain volume all the crosses except PKV Kranti × DSMR-4 exhibited higher additive variance than the dominance variance.

The degree of dominance was recorded lower than unity in all the crosses except BJV 44 × DSMR-8 for days to flowering and grain width. For grain yield/plant it was greater than unity in all the crosses except PKV Kranti × DSMR-8. For 100 grain weight, grain thickness, 100 grain volume and grains/panicle all the crosses except PKV Kranti × DSMR-4 showed degree of dominance greater than unity. All the crosses except BJV 44 × DSMR-4 recorded degree of dominance greater than unity for grain length. For grain density two crosses viz., BJV 44 × DSMR-8 and PKV Kranti × DSMR-8 showed degree of dominance greater than unity whereas for rest of the crosses it was lower than unity. The degree of dominance was found to be lower than unity for all the crosses except PKV Kranti × DSMR-4 for plant height.

Heritability

The traits viz., days to flowering and plant height exhibited moderate broad sense heritability whereas it was high for rest of the traits in BJV 44 × DSMR-8 (Table 5). In BJV 44 × DSMR-4 the traits viz., 100 grain weight, grain thickness and grain density exhibited moderate heritability whereas for grain width it was low. Rest of the traits in the cross showed high heritability. All the traits in PKV Kranti × DSMR-8 showed high heritability while the traits viz., grain thickness and plant height exhibited moderate heritability. Four traits such as grain width, grain length, grain thickness and plant height showed moderate heritability while rest of the traits showed high heritability. On the other hand, the narrow sense heritability was found to be high for all the traits in BJV 44 × DSMR-8 and PKV Kranti × DSMR-8. In BJV 44 x DSMR-4, all the traits expressed high narrow sense heritability except for grain width where it was moderate. In the cross PKV Kranti × DSMR-4 the 100 grain volume showed low heritability whereas rest of the traits showed high heritability.

For all the traits in all the crosses except for 100 grain volume in PKV Kranti × DSMR-4 the narrow sense heritability was in the range of moderate to high whereas it was found to be low for 100 grain volume. These results are in agreement with those reported previously by Abd El-Hamid and Ghareeb (2018) and Koubisy (2019).

The information of inheritance pattern, heritability and gene action helps considerably for the improvement of traits and ultimately yield. Hayman’s generation mean analysis provides information on the relative importance of additive effect, dominance effect and non-allelic gene interactions, to determining genotypic values of the individuals and consequently, mean genotypic values of families and generations. This experiment showed considerable variance in the different families of different crosses, revealing the existence of genetic diversity in the parental material used for the experiment (Table 2). The present experiment based on the significant values of different scales and joint scale test indicated the presence of gene interaction for all the traits in all the crosses except in PKV Kranti × DSMR-4 for the grain density. The gene effect such as [m] indicated that all the traits in all the crosses are quantitatively inherited and showed the importance of non allelic interactions. The significant [d] additive effects for most of the traits in majority of the crosses indicting the possibility of improvement through selection in early segregating generations. Similarly, significant dominance effect [h] suggesting to postpone the selection for late generations to increase homozygosity. The results indicated that both additive and dominance effects have important contributions in the inheritance of the studied traits, suggesting that selection for desirable traits might be effective in early generations. However, it is better to postpone it to later segregating generations. The significance of additive × dominance [j] and dominance × dominance [l] gene interactions indicates that the traits are greatly affected by additive × dominance, dominance and non allelic interactions. Therefore, it is advisable to delay selection for later segregating generation till the populations attain of homozygosity. The present results suggesting that all type of epistasis viz., additive × additive, additive × dominance and dominance × dominance playing role for the expression of all the studied traits.

Based on the results of present study, it is advisable to delay selection for most of the traits in most of the crosses and interestingly for all the traits in the cross BJV 44 × DSMR-8 as these were showing duplicate type of epistasis. This study reveals the presence of duplicate type of epistasis for most of the traits and it would limit the range of variability (Kearsy and Pooni, 1996) and delay in the selection until the attainment of homozygosity would be better to exploit the transgressive segregants. Whereas, it is advisable to go for heterosis breeding for the traits showing complementary gene interaction (Table 4). It would be rewarding to exploit heterosis in the crosses viz., BJV 44 × DSMR-4 and PKV Kranti × DSMR-4 for grain yield/plant. For the traits in the crosses such as for days to flowering in BJV 44 × DSMR-4 & PKV Kranti × DSMR-4, For grain yield in BJV 44 x DSMR-4, for grain width in PKV Kranti × DSMR-8 & PKV Kranti × DSMR-4, for grain length in PKV Kranti × DSMR-8, for grains/panicle in BJV 44 × DSMR-4 and for plant height in PKV Kranti × DSMR-8 the complementary epistasis was in desirable direction and this increases the heterosis in the desirable direction. Thus, heterosis breeding would be worthwhile for these crosses for the concerned traits. While, the crosses exhibiting duplicate gene interaction (Table 4) can not be used directly for heterosis exploitation as this gene action reduces the heterosis. Therefore, for the characters to improve the concerned cross should be subjected to recurrent selection (Comstock et al., 1949) or bi parental matting to accumulate favorable genes followed by selection in the subsequent generations. Liang and Walter, 1968 suggested delaying selection until the genotypes are stabilized in homogenous dominance condition, so that superior dominant allelic combinations can be selected.

The results of the present experiment reveals that all the traits in all the crosses except for 100 grain volume in PKV Kranti × DSMR-4 is under the control of additive gene action (Table 6) and selection in the early generation would be rewarding. Whereas for the 100 grain volume in PKV Kranti × DSMR-4 delayed selection would be good. The traits of the crosses showing degree of dominance less than unity reveals the role of partial dominance gene effects and selection in the later segregating generation would be meaningful. The degree of dominance greater than unity for most of traits in most of the crosses revealing the role of over dominance. These results suggest that the delayed selection to later generation would be rewarding.

Table 6

Identified gene actions for the different traits in different crosses
6. 100 grain volume
σ²g		0.10	0.251		0.157			0.132
σ²a		0.14	0.360		0.275			0.029
σ²d		-0.03	-0.109		-0.118			0.104
σ²p		0.14	0.284		0.205			0.157
σ²b		0.71	0.884		0.769			0.841
σ²h		0.94	1.267		1.345			0.182
Degree of dominance		2.40	2.208		1.666			-4.283
7. Grain density
σ²g	0.03		0.011			0.007	0.009
σ²a	0.06		0.025			0.011	0.014
σ²d	-0.03		-0.014			-0.004	-0.005
σ²p	0.04		0.021			0.010	0.010
σ²b	0.81		0.537			0.702	0.836
σ²h	1.50		1.176			1.066	1.338
Degree of dominance	1.38		-5.025			1.155	-0.950
8. Number of grains per panicle
σ²g	8472.09		63738.36	8375.67			7170.84
σ²a	14081.05		124523.90	14692.34			9052.41
σ²d	-5608.97		-60785.54	-6316.66			-1881.57
σ²p	12303.59		100894.52	11970.35			11469.14
σ²b	0.69		0.63	0.70			0.63
σ²h	1.14		1.23	1.23			0.79
Degree of dominance	2.58		2.19	-2.22			1.75
10. Plant height
σ²g	19.59		44.52	22.27			25.25
σ²a	23.47		55.06	30.69			29.31
σ²d	-3.89		-10.55	-8.42			-4.06
σ²p	34.98		73.81	42.53			48.47
σ²b	0.56		0.60	0.52			0.52
σ²h	0.67		0.75	0.72			0.61
Degree of dominance	-1.26		-0.70	-0.80			1.10

Traits	BJV 44 × DSMR-8	BJV 44 × DSMR-4	PKV Kranti × DSMR-8	PKV Kranti × DSMR-4
Days to 50 per cent flowering	A & D	D	D	D
Grain yield per plant	A & D	A & D	D	A & D
100 grain weight	A & D	A & D	A & D	D
Grain width	A & D	D	D	D
Grain length	A & D	D	A & D	A & D
Grain thickness	A & D	A & D	A & D	D
100 Grain volume	A & D	A & D	A & D	D
Grain density	A & D	D	A & D	*
Number of grains per panicle	A & D	A & D	D	A & D
Plant height	D	D	D	D
* Trait showed absence of non allelic interaction.
Where,
A- Additive gene action, D- Dominance gene action
and A&D- both Additive and Dominance gene action.

The estimates of heritability in the broad sense suggests a high portion of the phenotypic variation and is attribute to genetic variation for all the crosses except for grain width in BJV 44 × DSMR-8 as the broad sense heritability observed between moderate to high. The magnitude of narrow sense heritability shows that the additive effects have a higher role than the dominance effect in controlling all the traits except 100 grain volume. The present study indicates that the selection process will be effective, which result in a higher response but delaying selection would be more rewarding.

The current investigation indicated the significant influence of all the two types of gene actions viz., additive, dominance and interaction components in the expression of almost all the traits. The crosses showing complementary gene interaction can be improved through heterosis breeding while reciprocal and biparental matting strategies can be adopted to accumulate favorable genes in crosses for concerned traits showing duplicate gene interaction. This is a first report on using generation mean analysis to study the inheritance pattern of grain width, grain length, grain thickness, grain volume and grain density in sorghum. The information from the present study could be helpful for further study of the grain yield and grain related traits.

Acknowledgements

The author is thankful to Sorghum scheme of Dr. PDKV, Akola, India for providing the promising verity PKV Kranti and Dr. B. D. Biradar for providing the promising variety BJV 44.

Conflict of Interest

The authors have no conflict of interest

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Table 2: Mean and variance of four crosses for different characters under study

Cross	BJV 44 × DSMR-8		BJV 44 × DSMR-4		PKV Kranti × DSMR-8		PKV Kranti × DSMR-4
Pedigree	Mean	Variance	Mean	Variance	Mean	Variance	Mean	Variance
*Days to 50 per cent* flowering**
P₁	70.45	1.88	70.40	1.73	70.10	1.78	70.57	1.63
P₂	67.15	2.03	60.30	1.91	67.45	2.93	60.88	2.03
F₁	67.41	1.87	61.65	1.19	68.79	1.06	62.55	1.12
F₂	67.22	3.48	64.40	7.04	69.55	6.69	66.31	9.05
BC₁	70.18	2.08	66.16	2.75	70.22	1.81	68.21	3.07
BC₂	67.20	2.38	63.66	2.71	65.26	3.15	63.34	3.82
2. Grain yield per plant
P₁	81.37	10.49	77.15	11.23	91.94	12.87	94.83	10.28
P₂	80.64	11.49	50.70	12.97	81.85	11.23	60.47	11.10
F₁	96.54	11.82	94.20	11.89	124.84	12.82	105.19	10.47
F₂	92.52	42.06	71.18	37.14	118.56	47.20	86.81	36.25
BC₁	116.53	20.07	78.11	12.24	120.02	20.54	103.12	17.23
BC₂	104.74	18.14	69.58	16.52	88.36	18.52	82.11	19.10
3. 100 grain weight
P₁	3.942	0.106	3.887	0.116	4.440	0.108	4.412	0.112
P₂	3.830	0.112	4.034	0.120	4.073	0.111	4.242	0.110
F₁	3.613	0.037	3.477	0.103	3.721	0.012	3.878	0.005
F₂	3.578	0.348	3.681	0.243	3.946	0.279	4.311	0.197
BC₁	4.072	0.110	4.061	0.118	4.984	0.113	4.325	0.129
BC₂	4.031	0.117	3.894	0.121	4.325	0.118	4.284	0.122
4. Grain width
P₁	3.843	0.018	3.984	0.019	4.101	0.019	4.182	0.018
P₂	3.980	0.014	4.187	0.024	4.06	0.014	4.298	0.015
F₁	3.723	0.016	4.339	0.015	4.300	0.014	4.304	0.018
F₂	3.862	0.057	4.228	0.026	4.110	0.051	4.199	0.043
BC₁	4.110	0.022	4.216	0.022	4.108	0.025	4.188	0.025
BC₂	4.051	0.021	4.314	0.017	4.117	0.022	4.227	0.021
5. Grain length
P₁	4.813	0.028	4.836	0.056	4.985	0.030	4.979	0.027
P₂	4.661	0.026	5.035	0.051	4.758	0.027	4.884	0.030
F₁	4.831	0.024	5.052	0.055	5.382	0.019	4.987	0.029
F₂	4.692	0.094	4.887	0.232	4.883	0.087	4.599	0.070
BC₁	5.070	0.035	5.021	0.063	5.069	0.041	4.949	0.034
BC₂	4.832	0.033	5.106	0.057	4.890	0.033	4.885	0.037
6. Grain thickness
P₁	2.846	0.016	2.808	0.021	3.088	0.016	3.103	0.018
P₂	2.781	0.015	2.863	0.027	2.834	0.018	2.854	0.025
F₁	2.644	0.016	2.803	0.022	2.953	0.016	2.758	0.021
F₂	2.753	0.041	2.758	0.053	2.851	0.031	2.841	0.046
BC₁	2.973	0.019	2.934	0.024	3.14	0.021	2.960	0.022
BC₂	2.812	0.019	2.833	0.030	2.841	0.023	2.793	0.029

Contd….

Cross	BJV 44 × DSMR-8		BJV 44 × DSMR-4		PKV Kranti × DSMR-8		PKV Kranti × DSMR-4
Pedigree	Mean	Variance	Mean	Variance	Mean	Variance	Mean	Variance
7. 100 Grain volume
P₁	3.215	0.048	3.043	0.042	3.720	0.059	4.402	0.056
P₂	3.112	0.043	3.404	0.036	3.448	0.045	4.242	0.039
F₁	2.982	0.038	3.078	0.027	3.561	0.043	3.253	0.003
F₂	2.962	0.144	3.108	0.284	3.429	0.205	3.608	0.157
BC₁	3.634	0.084	3.415	0.108	3.994	0.070	3.608	0.101
BC₂	3.311	0.070	3.258	0.100	3.525	0.064	3.543	0.185
8. Grain density
P₁	1.2106	0.0029	1.2377	0.0081	1.1937	0.0033	1.1984	0.0012
P₂	1.2241	0.0025	1.8549	0.0179	1.1876	0.0032	1.2073	0.0018
F₁	1.1928	0.0116	1.1302	0.0067	1.0448	0.0029	1.1890	0.0019
F₂	1.2123	0.0380	1.2059	0.0213	1.1937	0.0103	1.1981	0.0105
BC₁	1.1224	0.0094	1.2134	0.0109	1.2466	0.0052	1.1987	0.0037
BC₂	1.2212	0.0096	1.1968	0.0067	1.2268	0.0044	1.2092	0.0033
9. Number of grains per panicle
P₁	2075.20	4528.14	2002.43	45899.13	2082.55	4374.91	2163.48	4354.68
P₂	2119.09	4118.54	1264.55	17311.53	2016.90	3941.52	1481.63	4510.71
F₁	2683.28	3339.66	2731.26	42706.99	3356.93	3031.14	2713.51	4163.90
F₂	2467.63	12303.58	1968.45	100894.50	3058.92	11970.35	2035.32	11469.14
BC₁	2891.16	5602.36	1959.02	36390.93	2439.54	4894.45	2398.59	6605.38
BC₂	2627.30	4923.76	1802.79	40874.21	2041.13	4353.92	1936.55	7280.49
10. plant height
P₁	281.06	16.57	284.56	32.26	298.40	21.43	299.47	24.39
P₂	241.81	17.81	235.70	39.35	251.23	19.66	235.61	31.15
F₁	250.20	13.60	268.91	22.78	253.46	19.98	276.29	18.66
F₂	270.38	34.98	262.27	73.81	270.44	42.53	253.41	48.47
BC₁	276.55	23.17	267.74	43.37	291.59	28.71	277.76	29.64
BC₂	244.16	23.32	247.35	49.18	245.25	25.66	244.76	37.99

Table 3: Scaling test and gene effects for grain yield and grain related traits

Cross	BJV 44 × DSMR-8	BJV 44 × DSMR-4	PKV Kranti × DSMR-8	PKV Kranti × DSMR-4
*Days to 50 per cent* flowering**
Scaling test
A	2.51**±0.48	0.28±0.46	1.55*±0.44	3.30**±0.43
B	-0.16±0.50	5.36**±0.47	-5.73**±0.52	3.26**±0.48
C	-3.55**±0.81	3.58**±0.82	3.07**±0.83	8.72**±0.81
D	-2.95**±0.25	-1.03**±0.31	3.63**±0.30	1.08**±0.35
Joint Scaling test	237.31**	171.29**	136.42**	270.88**
Genetic parameters
m	67.28**±0.09	64.40**±0.13	69.55**±0.12	66.31**±0.14
d	2.99**±0.17	2.51**±0.19	4.96**±0.18	4.86**±0.21
h	4.50**±0.61	-1.64*±0.70	-7.24**±0.69	-5.34**±0.76
i	5.89**±0.49	2.06**±0.62	-7.25**±0.60	-2.16**±0.70
j	2.67**±0.55	-5.08**±0.57	7.27**±0.59	0.04±0.56
l	-8.23**±1.06	-7.70**±1.11	11.43**±1.09	-4.40**±1.17
2. Grain yield per plant (g)
Scaling test
A	55.15**±1.24	-15.14**±1.21	23.28**±1.36	6.22**±1.12
B	32.30**±1.26	-5.73**±1.29	-29.97**±1.29	-1.45±1.19
C	14.99**±2.17	-31.51**±2.21	50.78**±2.35	-18.43**±2.00
D	-36.24**±0.79	-5.32**±0.72	28.74**±0.82	-11.60**±0.74
Joint Scaling test	3240.99**	255.16**	2626.16**	287.83**
Genetic parameters
m	92.52**±0.31	71.19**±0.29	118.56**±0.32	86.81**±0.28
d	11.79**±0.50	8.52**±0.43	31.66**±0.50	21.01**±0.48
h	88.01**±1.82	40.92**±1.71	-19.53**±1.91	50.75**±1.70
i	72.47**±1.58	10.64**±1.43	-57.48**±1.63	23.21**±1.49
j	22.85**±1.43	-9.41**±1.39	53.24**±1.47	7.66**±1.32
l	-159.93**±2.95	10.22**±2.80	64.17**±3.09	-27.98**±2.77
3. 100 grain weight (g)
Scaling test
A	0.589**±0.097	0.758**±0.118	1.807**±0.094	0.359**±0.085
B	0.620**±0.102	0.276*±0.119	0.857**±0.093	0.449**±0.089
C	-0.686**±0.172	-0.151±0.202	-0.169±0.151	0.836**±0.127
D	-0.947**±0.068	-0.592**±0.061	-1.417**±0.063	0.014±0.058
Joint Scaling test	214.96**	117.75**	717.07**	56.14**
Genetic parameters
m	3.578**±0.028	3.681**±0.023	3.946**±0.025	4.311**±0.021
d	0.041±0.039	0.167**±0.039	0.659**±0.039	0.040±0.040
h	1.621**±0.150	0.701**±0.151	2.297**±0.138	-0.476**±0.125
i	1.894**±0.135	1.185**±0.121	2.833**±0.126	-0.027±0.115
j	-0.03±0.13	0.48**±0.13	0.95**±0.13	-0.09±0.12
l	-3.103**±0.231	-2.219**±0.255	-5.498**±0.216	-0.781**±0.204

Contd….

Cross	BJV 44 × DSMR-8	BJV 44 × DSMR-4	PKV Kranti × DSMR-8	PKV Kranti × DSMR-4
4. Grain width (mm)
Scaling test
A	0.654**±0.046	0.110*±0.048	-0.184**±0.048	-0.111*±0.045
B	0.398**±0.045	0.102*±0.049	-0.128**±0.044	-0.147**±0.045
C	0.180*±0.080	0.063±0.078	-0.326**±0.079	-0.292**±0.078
D	-0.436**±0.028	-0.074**±0.022	-0.007±0.027	-0.017±0.026
Joint Scaling test	367.25**	14.67**	20.11**	15.34**
Genetic parameters
m	3.862**±0.011	4.228**±0.008	4.109**±0.011	4.199**±0.010
d	0.060**±0.017	-0.098**±0.016	-0.009±0.017	-0.040*±0.017
h	0.683**±0.065	0.402**±0.057	0.231**±0.064	0.098±0.062
i	0.872**±0.056	0.148**±0.044	0.013±0.055	0.034±0.052
j	0.26**±0.05	0.01±0.06	-0.05±0.05	0.04±0.05
l	-1.925**±0.105	-0.360**±0.100	0.299**±0.105	0.224*±0.104
5. Grain length (mm)
Scaling test
A	0.497**±0.057	0.154*±0.085	-0.229**±0.059	-0.067±0.055
B	0.173**±0.057	0.127±0.082	-0.360**±0.055	-0.101*±0.059
C	-0.369**±0.101	-0.424**±0.157	-0.974**±0.098	-1.440**±0.099
D	-0.519**±0.036	-0.352**±0.053	-0.193**±0.035	-0.636**±0.033
Joint Scaling test	243.09**	44.63**	104.72**	440.79**
Genetic parameters
m	4.692**±0.014	4.887**±0.023	4.883**±0.014	4.599**±0.012
d	0.238**±0.021	-0.086**±0.028	0.179**±0.022	0.064**±0.021
h	1.132**±0.083	0.821**±0.124	0.895**±0.081	1.328**±0.078
i	1.038**±0.072	0.704**±0.106	0.385**±0.070	1.272**±0.065
j	0.32**±0.07	0.027±0.09	0.14*±0.07	0.03±0.06
l	-1.707**±0.132	-0.985**±0.192	0.203±0.132	-1.103**±0.131
6. Grain thickness
Scaling test
A	0.457**±0.044	0.259**±0.052	0.239**±0.046	0.058±0.046
B	0.199**±0.044	0.001±0.057	-0.105*±0.047	-0.026±0.053
C	0.098±0.076	-0.243*±0.094	-0.425**±0.078	-0.107±0.084
D	-0.279**±0.025	-0.251**±0.028	-0.280**±0.024	-0.070*±0.027
Joint Scaling test	190.87**	100.38**	192.69**	9.67*
Genetic parameters
m	2.753**±0.010	2.758**±0.011	2.851**±0.008	2.841**±0.010
d	0.161**±0.016	0.101**±0.018	0.299**±0.017	0.167**±0.018
h	0.389**±0.060	0.470**±0.070	0.551**±0.059	-0.081±0.065
i	0.558**±0.050	0.503**±0.057	0.559**±0.047	0.140*±0.054
j	0.26**±0.05	0.26**±0.06	0.34**±0.05	0.08±0.05
l	-1.214**±0.099	-0.762**±0.119	-0.693**±0.103	-0.172±0.111

Contd….

Cross	BJV 44 × DSMR-8	BJV 44 × DSMR-4		PKV Kranti × DSMR-8		PKV Kranti × DSMR-4
7. 100 grain volume
Scaling test
A	1.072**±0.078		0.710**±0.079		0.706**±0.083		-0.438**±0.068
B	0.529**±0.076		0.034±0.075		0.042±0.077		-0.409**±0.080
C	-0.441**±0.128		-0.169±0.139		-0.575**±0.146		-0.719**±0.098
D	-1.021**±0.048		-0.456**±0.062		-0.661**±0.052		0.064±0.057
Joint Scaling test	515.50**		117.39**		218.06**		90.65**
Genetic parameters
m	2.962**±0.018		3.108**±0.025		3.429**±0.021		3.608**±0.019
d	0.323**±0.032		0.157**±0.036		0.468**±0.029		0.065±0.043
h	1.861**±0.109		0.766**±0.133		1.299**±0.119		-1.197**±0.118
i	2.043**±0.096		0.912**±0.124		1.322**±0.103		-0.128±0.113
j	0.54**±0.09		0.68**±0.01		0.66**±0.09		-0.03±0.10
l	-3.644**±0.180		-1.655**±0.201		-2.070**±0.187		0.975**±0.197
8. Grain density
Scaling test
A	-0.159**±0.030		0.059*±0.032		0.255**±0.021		0.010±0.015
B	0.026±0.030		-0.591**±0.037		0.221**±0.020		0.022±0.016
C	0.029±0.061		-0.530**±0.058		0.304**±0.036		0.009±0.029
D	0.081**±0.022		0.001±0.017		-0.086**±0.012		-0.012±0.012
Joint Scaling test	62.12**		292.94**		207.02**		6.15^NS
Genetic parameters
m	1.212**±0.009		1.206**±0.007		1.194**±0.005
d	-0.099**±0.011		0.017±0.011		0.020*±0.008
h	-0.187**±0.049		-0.419**±0.043		0.026±0.029
i	-0.162**±0.043		-0.003±0.035		0.172**±0.025
j	-0.18**±0.03		0.65**±0.04		0.03±0.02
l	0.295**±0.076		0.536**±0.072		-0.648**±0.048
9. Number of grains per panicle
Scaling test
A	1023.84**±22.47		-815.65**±73.16		-560.40**±22.45		-79.82**±22.45
B	452.24**±22.08		-390.23**±63.58		-1291.58**±21.26		-322.04**±23.70
C	309.68**±38.15		-855.67**±123.55		1422.39**±38.17		-930.88**±38.67
D	-583.20**±13.36		175.10**±37.17		1637.19**±12.84		-264.51**±13.77
Joint Scaling test	3200.74**		132.99**		17689.97**		772.81**
Genetic parameters
m	2467.63**±5.22		1968.45**±14.92		3058.92**±5.129		2035.32**±5.03
d	263.86**±8.32		156.23**22.15		398.41**±7.72		462.04**±9.40
h	1752.53**±31.12		747.56**±91.93		-1967.17**±30.31		1419.97**±32.11
i	1166.40**±26.71		-350.21**±74.34		-3274.37**±25.68		529.02**±27.55
j	571.60**±26.24		-425.41**±71.58		731.17**±25.23		242.22**±26.20
l	-2642.47**±50.63		1556.09**±152.04		5126.35**±49.11		-127.15*±53.94

Contd….

Cross	BJV 44 × DSMR-8	BJV 44 × DSMR-4		PKV Kranti × DSMR-8		PKV Kranti × DSMR-4
10. Plant height
Scaling test
A	21.84**±1.41		-17.99**±1.96		31.31**±1.69		-20.25**±1.56
B	-3.70*±1.46		-9.92**±2.09		-14.20**±162		-22.38**±1.76
C	58.25**±2.32		-9.00**±3.28		25.21**±2.77		-74.04**±2.69
D	20.06**±0.79		9.45**±1.11		4.05**±0.85		-15.71**±0.93
Joint Scaling test	1205.93**		125.73**		690.91**		823.51**
Genetic parameters
m	270.38**±0.28		262.27**±0.40		270.44**±0.31		253.41**±0.33
d	32.39**±0.55		20.39**±0.77		46.34**±0.59		33.00**±0.66
h	-51.35**±1.87		-10.12**±2.64		-29.45**±2.11		40.17**±2.19
i	-40.11**±1.57		-18.90**±2.23		-8.10**±1.70		31.41**±1.85
j	25.54**±1.69		-8.07**±2.43		45.51±1.84		2.13±1.96
l	21.98**±3.20		46.80**±4.49		-9.02*±3.64		11.22**±3.76

Table 4. Identified different gene interaction for the traits under study in four different crosses

Sl. No.	Crosses	BJV 44 × DSMR-8	BJV 44 × DSMR-4	PKV Kranti × DSMR-8	PKV Kranti × DSMR-4
Sl. No.	Characters	BJV 44 × DSMR-8	BJV 44 × DSMR-4	PKV Kranti × DSMR-8	PKV Kranti × DSMR-4
1.	Days to 50 % flowering	Duplicate	Complementary	Duplicate	Complementary
2.	Grain yield / plant (g)	Duplicate	Complementary	Duplicate	Duplicate
3.	100 grain weight (g)	Duplicate	Duplicate	Duplicate	Complementary
4.	Grains/ panicle	Duplicate	Complementary	Duplicate	Duplicate
5.	100 grains volume (cc)	Duplicate	Duplicate	Duplicate	Duplicate
6.	Grain density	Duplicate	Duplicate	Duplicate	No interaction
7.	Grain length (mm)	Duplicate	Duplicate	Complementary	Duplicate
8.	Grain width (mm)	Duplicate	Duplicate	Complementary	Complementary
9.	Grain thickness (mm)	Duplicate	Duplicate	Duplicate	Complementary
10.	Plant height (cm)	Duplicate	Duplicate	Complementary	complementary

Table 5. Estimates of various genetic parameters for the traits under study

Cross	BJV 44 × DSMR-8	BJV 44 × DSMR-4	PKV Kranti × DSMR-8	PKV Kranti × DSMR-4
*Days to 50 per cent* flowering**
σ²g	1.56	5.54	4.98	7.58
σ²a	2.49	8.61	8.43	11.22
σ²d	-0.92	-3.07	-3.44	-3.64
σ²p	3.48	7.04	6.69	9.05
σ²b	0.45	0.79	0.75	0.84
σ²h	0.72	1.22	1.23	1.24
Degree of dominance	1.23	-0.81	-1.21	-1.05
2. Grain yield per plant (g)
σ²g	30.65	25.14	34.77	25.67
σ²a	45.91	45.52	55.34	36.18
σ²d	-15.25	-20.38	-20.57	-10.51
σ²p	42.06	37.14	47.20	36.25
σ²b	0.73	0.68	0.74	0.71
σ²h	1.09	1.23	1.17	1.00
Degree of dominance	2.73	2.19	-0.79	1.55
100 grain weight (g)
σ²g	0.28	0.13	0.22	0.139
σ²a	0.47	0.25	0.33	0.143
σ²d	-0.19	-0.11	-0.11	-0.004
σ²p	0.35	0.24	0.28	0.197
σ²b	0.79	0.55	0.78	0.705
σ²h	1.35	1.02	1.17	0.727
Degree of dominance	6.32	2.05	1.87	-3.436
Grain width (mm)
σ²g	0.04	0.007	0.036	0.026
σ²a	0.07	0.011	0.055	0.040
σ²d	-0.03	-0.004	-0.019	-0.015
σ²p	0.06	0.026	0.051	0.043
σ²b	0.72	0.285	0.710	0.595
σ²h	1.24	0.449	1.082	0.938
Degree of dominance	3.39	-0.029	-5.059	-1.572
Grain length (mm)
σ²g	0.07	0.178	0.063	0.041
σ²a	0.12	0.344	0.099	0.068
σ²d	-0.05	-0.166	-0.036	-0.027
σ²p	0.09	0.232	0.087	0.070
σ²b	0.73	0.766	0.729	0.588
σ²h	1.27	1.483	1.144	0.980
Degree of dominance	2.18	-3.091	2.236	4.544
Grain thickness
σ²g	0.03	0.030	0.015	0.024
σ²a	0.04	0.053	0.019	0.040
σ²d	-0.01	-0.023	-0.004	-0.016
σ²p	0.04	0.053	0.031	0.046
σ²b	0.62	0.565	0.471	0.534
σ²h	1.07	0.999	0.601	0.877
Degree of dominance	1.55	2.152	1.356	-0.696

Contd….

100 grain volume
σ²g		0.10	0.251		0.157			0.132
σ²a		0.14	0.360		0.275			0.029
σ²d		-0.03	-0.109		-0.118			0.104
σ²p		0.14	0.284		0.205			0.157
σ²b		0.71	0.884		0.769			0.841
σ²h		0.94	1.267		1.345			0.182
Degree of dominance		2.40	2.208		1.666			-4.283
Grain density
σ²g	0.03		0.011			0.007	0.009
σ²a	0.06		0.025			0.011	0.014
σ²d	-0.03		-0.014			-0.004	-0.005
σ²p	0.04		0.021			0.010	0.010
σ²b	0.81		0.537			0.702	0.836
σ²h	1.50		1.176			1.066	1.338
Degree of dominance	1.38		-5.025			1.155	-0.950
Number of grains per panicle
σ²g	8472.09		63738.36	8375.67			7170.84
σ²a	14081.05		124523.90	14692.34			9052.41
σ²d	-5608.97		-60785.54	-6316.66			-1881.57
σ²p	12303.59		100894.52	11970.35			11469.14
σ²b	0.69		0.63	0.70			0.63
σ²h	1.14		1.23	1.23			0.79
Degree of dominance	2.58		2.19	-2.22			1.75
10. Plant height
σ²g	19.59		44.52	22.27			25.25
σ²a	23.47		55.06	30.69			29.31
σ²d	-3.89		-10.55	-8.42			-4.06
σ²p	34.98		73.81	42.53			48.47
σ²b	0.56		0.60	0.52			0.52
σ²h	0.67		0.75	0.72			0.61
Degree of dominance	-1.26		-0.70	-0.80			1.10

Pattern of genetic inheritance of grain yield and grain related traits in rabi sorghum Sorghum bicolor (L.) Moench

Status:

Version 1

Abstract

Introduction

Material And Methods

Statistical analysis

Results

Discussion

Conclusion

Declarations

References

Tables

Status:

Version 1