Genetic parameters estimation for egg quality traits of four chicken breeds using a full diallel cross

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A total of 480 eggs from 4 purebreds and 12 crossbreds were evaluated for traits like egg weight ( EWT ), egg length ( EL ), egg width ( EW ), shell weight ( SWT ), shell thickness ( ST ), shell ratio ( SR ), albumen height ( AH ), albumen width ( AW ), yolk height ( YH ), yolk width ( YW ), yolk weight ( YWT ), yolk ratio ( YR ), and hugh unit ( HU ). Results showed positive H e for most traits. GCA were significant (P < 0.001) for EWT, EL, EW, AW, YW, YWT, and YR. Variation due to SCA was high (P < 0.001) for all traits. K×S cross showed positive RE for EWT and AW. D×S cross had the highest positive RE for ST. Both additive and non-additive gene actions play significant roles in controlling egg quality traits. EWT and YWT were primarily influenced by additive genetic effects, while EL, EW, ST, and YR were dominated by non-additive effects. M e exhibited significant variation across genotypes for most traits. In conclusion, breeding strategies should be tailored to the genetic basis of each trait, focusing on economically important traits and using molecular tools for efficiency. Heterosis effect Combining ability Reciprocal effect Maternal effects Diallel cross Introduction Poultry eggs are highly nutritious and relatively affordable food sources, offering high-quality protein complete with all essential amino acids, essential fatty acids, and a wide range of vitamins and minerals (Chambers et al. 2017 ; Morris et al. 2018 ). They play a vital role in supporting early growth and development due to their rich nutritional content (Omer et al. 2022 ). Despite the nutritional significance, egg production in Ethiopia remains notably low (Tadelle 2003 ). One of the key factors contributing to this issue is the poor production performance of indigenous breeds, largely stemming from their limited genetic potential (Kassa et al. 2021 ). Conversely, various efforts have been made to enhance the overall productivity of the poultry sector by introducing exotic chicken breeds (Tadelle et al. 2000 ). However, these initiatives have achieved only limited success, primarily due to several challenges. These include the breeds poor adaptation to harsh production environments, their heightened vulnerability to diseases, and poor on-farm management practices (Tadelle et al. 2000 ; Fikrineh et al. 2023 ). Among poultry traits, egg quality stands as one of the most important, holding immense value for both producers and consumers alike (Ademola et al. 2023 ). In today’s world, consumers are more discerning than ever, paying close attention to what they purchase and consume. This increased focus on egg quality is a big change, directly affecting the market demand and economic value of both fertile and table eggs (Hanusova et al. 2015 ; Ademola et al. 2023 ). Most importantly, the quality of eggs also significantly influences the reproductive success of the parents (Islam et al. 2001 ). So, examining the attributes that define egg quality characteristics is essential for boosting productivity and addressing the rising demand for chicken products (Beyene et al. 2022 ). Genetic factors play a crucial role in determining egg quality characteristics, and estimating the genetic parameters for targeted traits can determine the degree to which these traits are influenced by genetics (Hanusova et al. 2015 ; Zhang et al. 2024 ). Improving poultry genetics usually involves two main strategies: crossbreeding and artificial selection (Duenk et al. 2021 ). Both methods have different merits in poultry breeding, and the choice depends on the specific goals of the breeder. Crossbreeding in poultry is often more beneficial because it allows us to combine the favorable traits of two distinct breeds or genetic lines to produce offspring with enhanced qualities (Mokoena et al. 2024 ). Therefore, this study was planned to estimate crossbreeding parameters of various egg quality traits by crossing four chicken breeds: one locally improved breed (Improved Horro), two exotic breeds (Sasso and Potchefstroom Koekoek), and one locally developed composite breed (Dz-White feathered), utilizing a 4×4 full diallel mating system. The diallel cross design facilitates a detailed analysis of inheritance mechanisms, such as general combining ability (GCA), specific combining ability (SCA), and the heritability of traits that are either qualitative or complex in nature (Wearden et al. 1965 ). This approach offers critical insights into the genetic potential of each breed and their hybrid offspring. Materials and Methods Study area This study was carried out at the poultry farm of Werer Agricultural Research Center, situated in the Afar Regional State of Ethiopia, approximately 280 km from the capital city, Addis Ababa. The farm is located at an elevation of 750 meters above sea level (Jemal et al. 2019 ). The region experiences a mean annual rainfall of less than 590 mm, with May and June being the driest months. The area is characterized by high temperatures, ranging from 19.3°C to 45°C (Atsbaha et al. 2022 ). Breeding plan and management Crossbred genetic groups were produced by using random mating, and the sex ratio was maintained at 1:5 for all breed groups. For Horro dams, due to their small size, artificial insemination was used by collecting semen from sires of S, K, and D. Fertile eggs were collected, cleaned, and recorded daily according to breeds and crossbreds. After seven days collection the eggs were weighed and set for incubation separately according to their different breeds category. Accordingly, from the diallel cross, 4 purebred (H, S, K, D), 6 crossbred (H×S, H×K, H×D, S×K, S×D, K×D), and six reciprocal crossbred (S×H, K×H, D×H, K×S, D×S, D×K) progenies were obtained. From purebreds and crossbreds of each genotype a total of 800 female chicks were randomly distributed into 48 pens (16 progenies by 3 replications) under a completely randomized design (CRD). All experimental chickens were vaccinated against Newcastle disease (HB1 and Lasota), Gumboro (Infectious Bursal disease), fowl typhoid, and fowl pox based on the producer’s manual. Starting from hatch chickens were provided with a starter ration (20% CP and 3000 kcal/kg ME) until 8 week (WK) of age, followed by grower ration (18% CP and 2950 kcal/kg of ME) from 9 to 20 WK, and then the ration was changed into layer ration (16% CP and 2800 kcal/kg of ME) from 21 to 40 WK. Feed was purchased from Alema Feeds Co., Ltd., Debrezeit, Ethiopia, and clean water were always available in a drinker. During the chicks growing stage (from hatch upto WK 8) fluorescent lamps were placed to provide lighting for the chickens. Due to the hot temperature of the study area for the first 5 days 16–18 hours of light was provided, afterwards 12 hours of light up to the 8th WK. Traits recorded A total of 480 eggs (10 from each replication) were randomly chosen at the pick production period (at 32 WK). Sampled eggs were evaluated for egg weight ( EWT ), egg length ( EL ), egg width ( EW ), shell weight ( SWT ), shell thickness ( ST ), shell ratio ( SR ), albumen height ( AH ), albumen width ( AW ), yolk height ( YH ), yolk width ( YW ), yolk weight ( YWT ), yolk ratio ( YR ), and hugh unit ( HU ). Eggs were weighed using a sensitive balance then length and width of each egg were measured using a compass. Then, the eggs were broken within 24 hours on a table glass cover and YH and AH were measured using a spherometer. YW and AW were measured using a compass. Then yolk of each egg was separated from the albumen, then weighed in grams and expressed as a percentage relative to the EWT. The ST were measured at 3 different points of the eggshell and the calculated average of the three points were used. The shell and membrane were weighed together in grams for each egg and expressed as a percentage relative to the egg weight. Haugh unit was calculated according to the equation of (Haugh 1937 ): \(\:HU\:=\:100\:\text{log}\left(AH-1.7E{W}^{0.37}+7.6\right)\) , where; H = observed height of the albumen (mm) and W = weight of the eggs in grams. Statistical analysis Egg quality data were analyzed for variation between the genotypes using the general linear model procedure of R (Version 4.4.2) (R Core Team 2023 ). Differences considered to be significant (P < 0.05) were compared by Duncan’s Multiple Range Test (Duncan 1955 ). Genetic parameters estimation Diallel analysis was carried out only when the differences among the genotypes were significant, fixed-effect model of (Griffing 1956 ) Model 1, Method 1, a commonly used approach for assessing the genetic potential of breeding lines, especially in diallel crossing experiments, was used. The mathematical model for the combing ability analysis used was: $$\:{Y}_{ij}=\mu\:+{G}_{i}+{G}_{j}+{S}_{ij}+{r}_{ij}+{e}_{ij}$$ Where, Y ij = the observed performance of the i th genotype crossed with the j th genotype, µ = the overall mean of the population, G i and G j = general combining abilities of the i th and j th genotypes (represents the additive genetic effect of genotype i and j), S ij = specific combining ability for the i th and j th cross (represents the non-additive genetic interaction between i and j genotypes), r ij = is the reciprocal effect involving the reciprocal crosses between the i th and j th parents, and 𝑒 𝑖𝑗 = random error term, which accounts for any variation not explained by the fixed effects. Heterosis was calculated on percentage of mid-parents by application of the following formula (Williams et al. 2002 ): $$\:H\%=F1\:-\left\{\frac{\frac{\left(P1+P2\right)}{2}}{\frac{\left(P1+P2\right)}{2}}\right\}\times\:100$$ , Where F1 = the first filial and P1 and P2 are parents in diallel and reciprocal crosses. Maternal effects were estimated using a set of linear contrasts of the genetic group means (Dickerson 1969 ) as follows: \(\:{M}^{e}=\frac{1}{2}\left[\:\left(A\times\:B\right)-\left(B\times\:A\right)\:\right]\) . Results Egg quality traits The least square means for EWT, EL, EW, SWT, ST, and SR, across different genotypes are detailed in Table 1 . Differences (P < 0.05) were observed for all traits. From the result, K×S crossbred demonstrated the heaviest EWT at 55.10 g, followed by D×S at 53.91 g. S×K, S×H, D×K, and S, also displayed notable EWT, while the purebred H recorded the lightest EWT at 46.08 g. The purebred D stood out with the longest EL at 53.89 mm, trailed by S at 53.50 mm. Conversely, the D×S crossbred exhibited the shortest EL at 42.79 mm. For EW, S achieved the largest measurement at 41.82 mm, followed by D×K and D. On the lower end, the D×S crossbred recorded the narrowest EW at 31.76 mm. SWT was highest in the purebred S at 5.97 g, while the D×H crossbred displayed the lightest shells. ST varied significantly, with the S×D crossbred showing the thickest shells at 0.47 mm, whereas the purebred K and D exhibited the thinnest shells at 0.32 mm. The SR was highest in the purebred S at 11.44%, with the crossbreds K×H and D×K recording the lowest at 9.36% and 9.35%, respectively. The least square means of AH, AW, YH, YW, YWT, YR, and HU are detailed in Table 2 . The analysis revealed differences (P < 0.05) across all traits. The crossbred K×H (7.28 mm) and its reciprocal H×K (7.00 mm) achieved the highest AH, surpassing other genotypes. Followed by H×D (6.93 mm) and its reciprocal D×H (6.83 mm), while the lowest AH values were recorded in S×D (5.72 mm), D (5.55 mm), and K (5.17 mm). In terms of AW, the H×D crossbred stood out with the highest measurement, trailed by D×K, D, and K×S. Conversely, the H×K crossbred exhibited the narrowest AW. The purebred D (19.44 mm), crossbred K×S (19.33 mm), and purebred S (19.11 mm) demonstrated the highest YH. On the other hand, D×S (42.69 mm), S (42.33 mm), and S×D (41.22 mm) recorded the highest YW. In contrast, the H×K crossbred displayed the smallest YH (15.94 mm) and YW (34.98 mm) values. The purebred S (16.36 g) emerged with the heaviest yolk, outperforming all other genetic groups. For YR, the purebred H (34.26%) and crossbred H×S (32.78%) achieved the highest percentages, with H×D (24.30%) and D×K (22.80%) showing the lowest values. The K×H crossbred (87.67) exhibited the highest HU score. This was closely followed by D×H (86.21) and its reciprocal H×K (85.97). Other genetic groups displayed moderate HU values, while K×S (77.12) and K (73.87) recorded the lowest scores. Table 1 Least square means (± SE) for external egg quality traits Genotype EWT(g) EL (mm) EW (mm) SWT (g) ST (mm) SR (%) Purebred H 46.08 j ±0.28 51.14 ab ± 0.17 39.57 abc ± 0.41 5.23 a − d ±0.05 0.33 e ± 0.03 11.36 a ± 0.06 S 52.15 de ± 0.11 53.50 a ± 0.39 41.82 a ± 0.43 5.97 a ± 0.04 0.35 de ± 0.00 11.44 a ± 0.06 K 50.44 g ± 0.34 51.29 ab ± 0.48 38.93 abc ± 0.14 5.05 bcd ± 0.02 0.32 e ± 0.01 10.02 abc ± 0.11 D 49.13 h ± 0.36 53.89 a ± 0.47 41.46 ab ± 0.40 5.52 abc ± 0.21 0.32 e ± 0.01 11.24 ab ± 0.50 Crosses H×S 48.80 hi ± 0.42 53.17 a ± 0.85 39.55 abc ± 0.90 5.06 bcd ± 0.23 0.43 ab ± 0.02 10.39 abc ± 0.57 H×K 51.07 fg ± 0.22 53.17 a ± 1.08 39.47 abc ± 0.41 5.38 a − d ±0.26 0.42 abc ± 0.01 10.53 abc ± 0.50 H×D 50.38 g ± 0.26 53.29 a ± 0.36 40.94 abc ± 0.52 4.85 cd ± 0.32 0.37 cde ± 0.01 9.61 c ± 0.59 S×K 53.46 bc ± 0.57 51.66 ab ± 1.67 40.04 abc ± 0.54 5.42 a − d ±0.30 0.33 e ± 0.02 10.16 abc ± 0.66 S×D 50.85 fg ± 0.06 45.63 c ± 2.13 34.05 d ± 2.28 5.72 ab ± 0.14 0.47 a ± 0.01 11.26 ab ± 0.28 K×D 50.11 g ± 0.19 49.30 b ± 1.98 37.74 c ± 2.04 5.25 a − d ±0.18 0.43 ab ± 0.02 10.48 abc ± 0.34 RE S×H 53.31 bc ± 0.28 51.07 ab ± 0.87 39.93 abc ± 0.75 5.18 a − d ±0.30 0.40 ± bcd 0.01 9.71 bc ± 0.52 K×H 51.63 ef ± 0.22 52.66 ab ± 0.51 38.92 abc ± 0.44 4.83 cd ± 0.08 0.45 a ± 0.02 9.36 c ± 0.19 D×H 48.05 i ± 0.14 50.63 ab ± 0.62 38.34 bc ± 0.42 4.66 d ± 0.05 0.40 bcd ± 0.01 9.69 bc ± 0.08 K×S 55.10 a ± 0.13 52.59 ab ± 0.81 41.42 ab ± 0.28 5.33 a − d ±0.05 0.33 e ± 0.01 9.68 bc ± 0.07 D×S 53.91 b ± 0.32 42.79 c ± 0.72 31.76 d ± 0.63 5.56 abc ± 0.51 0.37 cde ± 0.03 10.32 abc ± 1.00 D×K 52.61 cd ± 0.50 52.59 ab ± 0.46 41.50 ab ± 1.08 4.92 bcd ± 0.35 0.42 ab ± 0.01 9.35 c ± 0.65 P-values < 0.0001 < 0.0001 < 0.0001 0.0400 < 0.0001 0.0300 CV 1.05 3.56 4.27 7.93 7.12 8.07 a−j Means with the same letter in a column are not significantly different (P < 0.05) Table 2 Least square means (± SE) for internal egg quality traits Genotype AH (mm) AW (mm) YH (mm) YW (mm) YWT (g) YR (%) HU Purebred H 6.33 a − d ±0.51 62.40 efg ± 2.58 18.67 ab ± 0.33 40.51 abc ± 0.86 15.79 abc ± 0.47 34.26 a ± 1.00 83.57 a − d ±3.13 S 6.33 a − d ±0.67 61.06 fgh ± 2.14 19.11 a ± 0.40 42.33 ab ± 0.63 16.36 a ± 0.76 31.36 abc ± 1.48 81.21 a − e ±4.68 K 5.17 d ± 0.17 66.93 c − f ±2.56 18.22 abc ± 0.11 37.63 de ± 0.74 13.87 c − f ±0.30 27.49 cd ± 0.49 73.87 e ± 1.30 D 5.55 cd ± 0.22 70.74 abc ± 3.10 19.44 a ± 0.29 39.02 cd ± 1.06 14.60 a − d ±0.65 29.74 bc ± 1.50 77.32 cde ± 1.74 Crosses H×S 5.83 bcd ± 0.44 67.03 c − f ±1.92 16.89 cd ± 0.78 35.923 e ± 0.36 15.99 ab ± 0.39 32.78 ab ± 1.07 79.28 a − e ±3.25 H×K 7.00 ab ± 0.58 55.91 h ± 1.09 15.94 d ± 0.44 34.98 e ± 0.34 14.03 b − e ±0.56 27.47 cd ± 1.08 85.97 abc ± 3.40 H×D 6.93 ab ± 0.47 75.97 a ± 0.53 15.97 d ± 0.41 35.43 e ± 1.14 12.24 ef ± 0.26 24.30 de ± 0.49 85.88 a − d ±2.85 S×K 6.22 a − d ±0.59 67.53 cde ± 1.71 18.33 abc ± 0.33 39.88 bcd ± 0.16 14.77 a − d ±0.95 27.67 cd ± 2.06 80.11 a − e ±3.97 S×D 5.72 bcd ± 0.15 66.20 c − f ±1.75 18.89 a ± 0.40 41.22 abc ± 0.75 15.41 abc ± 0.65 30.31 bc ± 1.31 77.93 b − e ±1.07 K×D 6.10 a − d ±0.10 68.32 cde ± 0.97 16.89 cd ± 0.22 35.59 e ± 1.43 12.55 ef ± 0.43 25.05 de ± 0.77 80.85 a − e ±0.73 RE S×H 6.28 a − d ±0.50 63.94 d − g ±0.77 17.27 bcd ± 0.06 36.13 e ± 0.60 13.13 def ± 0.72 24.62 de ± 1.25 80.67 a − e ±3.33 K×H 7.28 a ± 0.15 62.22 efg ± 1.73 16.94 cd ± 0.53 35.06 e ± 0.37 13.21 def ± 1.11 25.60 de ± 2.25 87.67 a ± 0.90 D×H 6.83 abc ± 0.10 59.84 gh ± 1.88 16.56 d ± 0.87 35.36 e ± 1.27 12.30 ef ± 0.46 25.61 de ± 0.99 86.21 ab ± 0.54 K×S 5.83 bcd ± 0.17 69.33 bcd ± 1.26 19.33 a ± 0.33 40.85 abc ± 0.45 15.39 abc ± 0.47 27.93 cd ± 0.79 77.12 de ± 1.27 D×S 6.06 a − d ±0.47 64.46 d − g ±1.16 18.78 ab ± 0.49 42.69 a ± 0.66 15.05 a − d ±0.23 27.93 cd ± 0.59 78.94 a − e ±3.32 D×K 6.22 a − d ±0.11 74.08 ab ± 0.84 17.00 cd ± 0.88 35.53 e ± 0.29 11.99 f ± 0.45 22.80 e ± 0.95 80.77 a − e ±0.72 P-values 0.0361 < 0.0001 < 0.0001 < 0.0001 < 0.0001 < 0.0001 0.0287 CV 10.69 4.80 4.78 3.62 7.37 7.66 5.50 a−g Means with the same letter in a column are not significantly different (P < 0.05) Crossbreeding effects Heterosis effect (H e ) H e varied significantly across different traits Table 3 . EWT displayed positive H e for most crosses and their reciprocals, ranging from 0.42–8.54%, with the exception of H×S cross. For EL, the majority of H e estimates were negative, except for H×S (1.64%), H×K (3.82%), H×D (1.49%), K×H (2.83%), and K×S (0.37%). Similarly, EW predominantly exhibited negative H e , with the exception of H×K (0.56%), H×D (1.04%), K×S (2.58%), and D×K (3.24%). In terms of SWT, H e was negative across all genetic groups except for H×K (4.54%). Conversely, ST showed high and positive H e for most groups, excluding S×K (-2.97%) and its reciprocal K×S (-1.98%). However, SR demonstrated negative H e across all crosses and reciprocals. AH exhibited strong positive H e in the majority of crosses, except for H×S, S×D, and S×H. Similarly, AW also displayed positive H e in most crosses. In contrast, YH showed negative H e across all crosses and reciprocals, with the exception of K×S. YW and YWT also had negative H e , except for S×D, K×S, and D×S. For HU, most H e were positive but relatively low, except for the crossbreds H×S, S×D, S×H, K×S, and D×S. on the other hand, YR predominantly exhibited high and negative H e . Table 3 Estimation of heterosis percentage for egg quality traits Traits Genotypes Crosses Reciprocals H×S H×K H×D S×K S×D K×D S×H K×H D×H K×S D×S D×K EWT -0.65 5.82 5.83 4.22 0.42 0.65 8.54 6.98 0.93 7.41 6.45 5.68 EL 1.64 3.82 1.49 -1.40 -15.02 -6.25 -2.38 2.83 -3.59 0.37 -20.31 0.00 EW -2.81 0.56 1.04 -0.84 -18.23 -6.12 -1.88 -0.85 -5.38 2.58 -23.73 3.24 SWT -9.58 4.54 -9.86 -1.57 -0.35 -0.63 -7.56 -6.03 -13.39 -3.21 -3.25 -6.94 ST 27.12 27.55 13.89 -2.97 40.10 35.18 16.67 39.12 22.81 -1.98 9.82 32.41 SR -8.88 -1.51 -14.92 -5.35 -0.76 -1.39 -14.85 -12.40 -14.23 -9.80 -9.01 -12.05 AH -7.87 21.77 16.69 8.17 -3.70 13.81 -0.87 26.59 15.01 1.45 1.91 16.04 AW 8.59 -13.53 14.13 5.52 0.46 -0.74 3.57 -3.78 -10.11 8.33 -2.19 7.63 YH -10.58 -13.56 -16.17 -1.79 -2.01 -10.31 -8.55 -8.13 -13.11 3.58 -2.58 -9.74 YW -13.26 -10.46 -10.91 -0.24 1.33 -7.14 -12.78 -10.26 -11.09 2.18 4.95 -7.29 YWT -0.53 -5.40 -19.42 -2.28 -0.42 -11.80 -18.28 -10.90 -19.02 1.84 -2.75 -15.74 YR -0.10 -11.04 -24.06 -5.99 -0.78 -12.46 -24.95 -17.07 -19.97 -5.09 -8.57 -20.30 HU -3.78 9.21 6.75 3.32 -1.69 6.94 -2.09 11.37 7.17 -0.54 -0.41 6.84 H Improved Horro, S Sasso, K Potchefstroom Koekoek, D Dz-white; Males are listed first in the cross Combining ability and reciprocal effects Estimates of GCA, SCA, and RE for egg quality traits are presented in Table 4 . GCA effects were highly significant (P < 0.001) for EWT, EL, EW, AW, YW, YWT, and YR, but not for ST. The S breed showed the highest positive GCA estimates for EWT, followed by K, while D and H breeds recorded negative values. Positive GCA estimate for EL was observed in H and K breeds, while S and D breeds showed negative estimates. The D breed exhibited the highest positive GCA value for AW, followed by K, with S and H breeds showing negative values. Only the S breed recorded positive GCA values for YW and YWT, while D, K, and H breeds had negative estimates. For YR, the S breed showed the highest positive GCA estimates, followed by H, while D and K breeds had small negative estimates. SCA variations were highly significant (P < 0.001) for all traits. The S×D crossbred had the highest positive SCA estimates for EWT, followed by K×D, H×K, S×K, and H×S, while H×D recorded a negative SCA estimates. The K×D crossbred showed the highest positive SCA value for EL, followed by H×S, S×K, and H×K, while H×D and S×D had negative estimates. For ST, the H×K crossbred had the highest positive SCA estimate, followed by H×S, H×D, and S×K, while K×D and S×D exhibited small negative SCA values. Positive SCA estimates for AW were observed in K×D, H×S, and S×K, while S×D, H×K, and H×D showed negative estimates. Most SCA estimates for YW were negative, except for S×D, S×K. The S×K crossbred showed the highest SCA values for YWT, while H×S recorded the highest negative estimate. For YR, only H×D and S×K crossbreds had positive SCA estimates, while the remaining crosses showed negative values. RE were significant (P < 0.005) for EWT, EW, ST, AW, and YR, but not for EL, YW, and YWT. The K×S crossbred had the highest positive RE for EWT, followed by D×S, D×H, and D×K, while K×H and S×H showed negative values. For EW, RE was positive for most crosses, except S×H and D×S. However, D×S had the highest positive estimate for ST. The K×S crossbred recorded the highest positive estimate for AW but a negative estimate for YR. The S×H crossbred exhibited the highest positive estimate for YR. From the results, EL had the highest GCA/SCA ratio, while AW and YW showed moderate ratios. Conversely, YR, EL, EW, and ST exhibited the lowest ratios. Table 4 Estimates (± SE) of combining ability for external and internal egg quality traits Genotype EWT (g) EL (mm) EW (mm) ST (mm) AW (mm) YW (mm) YWT (g) YR (%) GCA < 0.0001 0.0026 0.0479 NS < 0.0001 < 0.0001 < 0.0001 0.0002 g H -1.64 hi ± 0.27 0.89 a ± 0.36 0.45 a ± 0.36 0.01 a − e ±0.01 -2.29 abc ± 0.12 -1.27 cd ± 0.58 -0.11 abc ± 0.58 0.80 ab ± 1.25 g S 1.40 ab ± 0.13 -0.66 ab ± 0.18 -0.29 ab ± 0.18 -0.01 b − f ±0.01 -0.92 ab ± 0.06 2.16 a ± 0.29 1.14 ab ± 0.29 1.44 ab ± 0.62 g K 0.79 a − d ±0.27 0.67 ab ± 0.36 0.53 a ± 0.36 0.00 b − f ±0.01 0.41 ab ± 0.12 -0.87 bcd ± 0.58 -0.46 abc ± 0.58 -1.37 b ± 1.25 g D -0.55 fg ± 0.08 -0.90 ab ± 0.11 -0.68 ab ± 0.11 0.00 a − e ±0.00 2.80 a ± 0.04 -0.03 abc ± 0.18 -0.57 abc ± 0.18 -0.87 b ± 0.38 SCA < 0.0001 < 0.0001 < 0.0001 < 0.0001 0.0003 < 0.0001 0.0005 < 0.0001 s H×S 0.23 c − f ±0.23 0.75 a ± 0.31 0.50 a ± 0.31 0.03 a − d ±0.01 2.69 a ± 0.10 -2.87 d ± 0.50 -0.64 abc ± 0.50 -1.35 b ± 1.08 s H×K 1.13 abc ± 0.30 0.21 ab ± 0.40 -0.87 ab ± 0.40 0.05 a ± 0.01 -5.06 bc ± 0.13 -0.85 bcd ± 0.65 0.02 abc ± 0.65 -0.71 b ± 1.40 s H×D -1.17 gh ± 0.13 -1.33 ab ± 0.18 -1.30 ab ± 0.18 0.01 a − d ±0.01 -8.07 c ± 0.06 -0.04 abc ± 0.29 0.03 abc ± 0.29 0.65 ab ± 0.62 s S×K 0.82 a − d ±0.19 0.46 ab ± 0.25 0.69 a ± 0.25 0.00 a − f ±0.01 0.90 ab ± 0.08 0.48 abc ± 0.41 0.31 abc ± 0.41 0.13 ab ± 0.88 s S×D 1.53 a ± 0.20 -1.42 ab ± 0.27 -1.15 ab ± 0.27 -0.05 f ± 0.01 -0.87 ab ± 0.09 0.74 abc ± 0.44 -0.18 abc ± 0.44 -1.19 b ± 0.94 s K×D 1.25 abc ± 0.23 1.64 a ± 0.31 1.88 a ± 0.31 -0.01 b − f ±0.01 2.88 a ± 0.10 -0.03 abc ± 0.50 -0.28 abc ± 0.50 -1.12 b ± 1.08 RE < 0.0001 NS 0.0310 0.0018 < 0.0001 NS NS 0.0021 r S×H -2.26 i ± 0.15 1.05 a ± 0.20 -0.19 ab ± 0.20 0.02 a − d ±0.01 1.55 a ± 0.06 -0.10 abc ± 0.33 1.43 a ± 0.33 4.08 a ± 0.70 r K×H -0.28 efg ± 0.19 0.25 ab ± 0.25 0.28 ab ± 60.25 -0.02 def ± 0.01 -3.15 abc ± 0.08 -0.04 abc ± 0.41 0.41 abc ± 0.41 0.93 ab ± 0.88 r D×H 0.34 c − f ±0.20 0.82 a ± 0.27 0.79 a ± 0.27 -0.01 c − f ±0.01 1.40 a ± 0.09 -1.32 cd ± 0.44 -1.21 c ± 0.44 -2.79 b ± 0.94 r K×S 1.02 a − d ±0.08 0.96 a ± 0.11 1.40 a ± 0.11 -0.04 ef ± 0.00 2.94 a ± 0.04 1.06 abc ± 0.18 0.23 abc ± 0.18 -0.08 b ± 0.38 r D×S 0.46 b − e ±0.19 -5.38 b ± 0.25 -5.21 b ± 0.25 0.04 abc ± 0.01 -2.54 abc ± 0.08 1.81 ab ± 0.41 0.50 abc ± 0.41 0.75 ab ± 0.88 r D×K 0.05 def ± 0.15 0.02 ab ± 0.20 0.68 a ± 0.20 0.05 ab ± 0.01 2.00 a ± 0.06 -1.56 cd ± 0.33 -0.86 bc ± 0.33 -1.64 b ± 0.70 GCA/SCA 0.681 0.044 0.016 0.002 0.259 0.227 0.345 0.158 a−i Means with the same letter in a column are not significantly different (P < 0.05); g : general combining ability, s : specific combining ability, r : reciprocal effect Maternal effect (M e ) The M e estimates for the studied traits varied, displaying both positive and negative values with low to moderate to high magnitudes Table 5 . With the exception of the H×D, all other crosses displayed negative M e on EWT. In terms of EL, S×K and K×D crosses showed negative estimates, while the remaining crosses exhibited positive estimates. For EW, the estimates were positive for the H×K, H×S, and S×D, whereas the other three crosses showed negative values. Regarding SWT and SR, the H×S and H×D crosses demonstrated negative M e , while the H×K, S×K, S×D, and K×D showed positive estimates. However, for ST, the H×K, H×D, and S×K crosses exhibited negative estimates, whereas the H×S, S×D, and K×D displayed positive estimates. For AH, all crosses except H×D and S×D exhibited negative estimates. In contrast, only the S×D and K×D showed negative estimates for AW. Similarly, YH displayed negative estimates for all crosses except S×K. For YW, only the H×K and H×S showed negative estimates, while the others exhibited positive values. Conversely, YWT and YR displayed positive M e for all crosses except H×D and S×D.</p Discussion Egg quality traits EWT exhibited significant variation across genotypes, highlighting the strong influence of genetics on this trait. Purebred S had the highest EWT at 32 WK of age, followed by K and D. The EWT of the S in this study was consistent with findings of Yonas et al. ( 2019 ) and Wolde et al. ( 2021 ). In contrast, Beyene et al. ( 2022 ) reported a higher EWT, however their observed EWT for the K breed aligned with the present result. The EWT for the D breed reported by Samrawit ( 2020 ) was slightly higher than that found in this study. However, the same author reported similar EWT values for H with the present findings. Table 1 showed that the H×K, H×D, S×H, K×H, S×K, K×S, D×S, and D×K crossbreds exhibited higher EWT compared to their purebred counterparts, suggesting hybrid vigor. This implies that crossing K and D with S improves EWT compared to purebreds. H recorded the lowest EWT, indicating possible genetic constraints for this trait within the breed. However, crossbreeding H with S and K led to notable improvements in EWT. These findings align with (Kedija et al. 2019 ; Atsbaha et al. 2023 ), they reported significant gains in EWT through crossbreeding H with exotic breeds, further supporting the heterosis effects observed in this study, particularly in H×S, S×H, H×K, and K×H crossbreds. However, the reciprocal cross D×H performed poorly, suggesting that the maternal or paternal influence of H may negatively impact EWT. Purebreds D and S had the longest eggs, indicating that these breeds inherently contribute to increased EL. The EL observed in this study for D and H agreed with findings reported by Samrawit ( 2020 ). In contrast, Beyene et al. ( 2022 ) and Kefelegn and Ashenafi ( 2024 ), reported slightly longer EL for S breed. The EL of the K breed in this study was consistent with finding of Yesihak et al. ( 2024 ). Crossbred H×D also performed well, suggesting that combining H and D can enhance this trait. D×S had the smallest EL, given that both D and S purebreds performed well, this suggests a potential negative interaction or dominance effect when S is used as the dam in this cross. EW exhibited differences (P < 0.05) among genotypes. Significant differences for EW in consensus to this study had also been reported by (Kedija et al. 2019 ; Udoh et al. 2020 ; Fikrineh et al. 2023 ). S showed the widest eggs, followed by the D×K and D, suggesting that both S and D breeds positively influence EW. This observation was consistent with the result reported by Wolde et al. ( 2021 ). Conversely, the D×S crossbred had the smallest EW, which corresponds to its poor performance in EL. This further reinforces the notion of a negative interaction in the D×S cross. Significant differences (P < 0.04) in SWT were observed across groups, though the variations were minimal. This suggests that genetic influences on this trait are limited or that environmental factors play a more substantial role. S purebred exhibited the heaviest shell, followed by S×D and D×S, indicating that breed S positively contributes to SWT. Similar SWT value for the S was reported by Wolde et al. ( 2021 ), though Samrawit ( 2020 ) recorded higher SWT values for D and H. The D×H crossbred had the lowest SWT, consistent with its poor performance in other traits. In contrast, the S×D crossbred displayed the thickest shell, followed by K×H and K×D, suggesting that specific combinations can enhance ST, potentially due to complementary genetic effects. These findings align with previous studies (Sola-Ojo and Ayorinde, 2011 ; Khalil et al. 2013 ), which also demonstrated that crossbreeding improved ST. The K and D had the thinnest shells, indicating possible genetic limitations for this trait. Similarly, Samrawit ( 2020 ) observed that breed D exhibited lower performance in terms of ST compared to H. Differences (P < 0.03) were noted in SR across groups, although the variations were minimal. S demonstrated the highest SR, followed by H and the S×D crossbred, underscoring the positive influence of S and H on SR. Conversely, the K×H and D×K recorded the lowest SR, indicating that these combinations may not be effective for enhancing shell quality. Differences in AH (P < 0.0360) and AW (P < 0.0001) were observed across genotypes. K×H and H×K demonstrated the highest AH, reflecting strong heterosis in these combinations. Table 2 showed, H×K, H×D, K×D, K×H, D×H, and D×K crossbreds displayed higher AH compared to their purebred counterparts, indicating hybrid vigor. The superiority of crossbreds over purebreds has been supported by previous studies (Sola-Ojo and Ayorinde, 2011 ; Sinha et al. 2018 ; Kedija et al. 2019 ). Purebred K and crossbred S×D recorded the lowest AH, while H×K had the narrowest AW, implying that certain combinations, particularly those involving K and S, may not be optimal for enhancing albumen quality. The H breed performed well in AH and AW when crossed with K and D, highlighting its potential to positively influence albumen quality in crossbreds. The S breed showed moderate performance in AH and AW, suggesting that while S is a robust breed overall, its contribution to albumen quality may be limited in specific crosses. Udoh et al. ( 2020 ) reported results consistent with the present study regarding AH for the S, although their reported AW value was lower. K breed underperformed in AH, but showed promise in crosses such as K×H. In contrast, Fikrineh et al. ( 2023 ) reported a slightly higher AH value for the K breed. However, the same author noted that K performed moderately relative to Fayoumi and White Leghorn but underperformed compared to its reciprocal cross with Fayoumi, aligning with the current findings. The purebred D and crossbred K×S exhibited the highest YH, while D×S and S demonstrated the widest YW. This suggests that both D and S breeds positively influence yolk size. In contrast, the crossbred H×K had the smallest YH and YW, indicating that this combination may not be optimal for enhancing yolk characteristics. These findings align with Samrawit ( 2020 ), who reported similar values for the D in terms of YH and YWT, though the YWT value was lower than that reported by Tewodros et al. ( 2022 ). However, H showed mixed results, with some crosses (H×D) performing well in YH but others (H×K) underperforming. The K breed underperformed in YH and YW, particularly in its purebred form. The YH and YW values for the S breed in this study were higher than those reported by Yonas et al. ( 2019 ). The S breed consistently excelled in YWT and YR, both in purebred form and in crosses, while the H breed showed strong performance in YR, particularly in its purebred form. For YR, H, H×S, and S achieved the highest percentages, indicating that both H and S positively influence yolk size and proportion. Similar findings were reported by Wolde et al. ( 2021 ), where S crossed with local chickens, S outperformed the local breed and their F1 cross in YWT, while the local breed and the F1 cross outperformed S in YR. The D breed underperformed in YWT and YR, particularly in crosses like H×D and D×K.Samrawit ( 2020 ) also noted that D underperformed in YR compared to H. Meanwhile, the K breed showed mixed results, with some crosses (K×S) performing well in YWT but others (K×D) underperforming. The crossbred K×H had the highest HU, indicating superior egg quality, closely followed by D×H and H×K. This suggests that crosses involving H and K, or D can enhance egg quality. These results are consistent with Tewodros et al. ( 2022 ), who reported similar HU values for the D breed. Crossbreeding effects Heterosis effect (H e ) Crossbreeding is a commonly used approach in the poultry to improve favorable traits (Dzungwe et al. 2024 ). Most crosses and their reciprocals exhibited positive H e for EWT, except for H×S. However, its reciprocal, S×H, displayed the highest positive H e . The negative H e observed in H×S suggests that this particular cross may not be ideal for improving EWT. In agreement,Saadey et al. ( 2008 ) reported positive H e for EWT in a direct cross between White Leghorn and Rhode Island Red but negative H e in its reciprocal cross. Crossbreds H×S, H×K, H×D, K×H, K×S, and D×K demonstrated positive H e values for EL and EW. Similarly, Atsbaha et al. ( 2023 ) reported positive H e for EL and EW. The occurrence of both positive and negative H e highlights the role of non-additive genetic effects in determining egg dimensions. Specific combinations, such as H×K and K×S, which exhibit positive H e , suggest that certain parental lines (H, K, S) could possess complementary alleles or genetic backgrounds that interact favorably to enhance egg size. These crosses are particularly promising for breeding programs focused on improving egg dimensions, as they demonstrate potential hybrid vigor. Negative H e was observed for SWT in all crosses except H×K. Conversely, positive H e was noted ST in most crosses, except S×K and K×S. In agreement, Santosh and Raj ( 2023 ) reported the highest positive H e values for ST. Similarly, Atsbaha et al. ( 2023 ) also documented positive H e for ST. However, in contrast, the same authors reported positive H e for SWT. The positive H e for ST in most crosses is advantageous for enhancing shell quality, which is critical for reducing breakage and improving hatchability. The positive H e for AH and AW observed in most crosses indicates that crossbreeding can improve albumen quality, which is important for egg grading and consumer preference. These results are supported by Atsbaha et al. ( 2023 ), who reported positive H e values for AH. For YH, negative H e was observed in all crosses except K×S. Similarly, for YW and YWT negative H e was observed in most crosses, except S×D, K×S, and D×S. However, the positive H e in specific crosses indicates potential for targeted improvement. Most crosses showed positive H e for HU, but values were relatively low. Similarly, Khalil et al. ( 2013 ) also recorded negative H e for YWT. However, in contrast, the same author reported negative H e for HU. Conversely, Santosh and Raj ( 2023 ) reported positive H e for YWT in most crossbreds, which disagreed with the present study. The positive H e for HU is beneficial for improving egg quality, as HU is a key indicator of freshness. Combining ability GCA is a key metric in breeding programs, used to estimate the average performance of offspring based on the genetic potential inherited from that individual Adebambo et al. ( 2011 ). GCA effects were significant (P < 0.001) across purebreds (Table 4 ). The significance of GCA underscores the critical role of additive genetic effects in the inheritance of egg quality traits (Khalil et al. 2018 ). S breed showed the highest positive GCA for EWT, YW, and YWT, making it a strong contributor to egg quality. K had positive GCA for EWT, EL, EW, ST and AW, indicating its potential for improving these traits, however the breed exhibited negative value for YWT, YW, and YR. In agreement, positive GCA value for K on EWT and EW was reported by Fikrineh et al. ( 2023 ). However, in contrast, the same author reported negative value for EL and ST and positive value for YWT. SCA refers to situations where certain genetic combinations perform either better or worse than predicted based on the average performance of the parent lines involved (Emad et al. 2017 ). It highlights the non-additive genetic interactions that influence the performance of specific crosses. Table 4 indicated that variation due to SCA was high (P < 0.001) for all traits. The S×D cross exhibited the highest positive SCA for EWT, demonstrating significant heterosis. Meanwhile, the K×D cross displayed the highest positive SCA for EL, EW, and AW, indicating its potential to enhance these characteristics. The H×K cross showed the highest positive SCA for ST and YR, positioning it as a promising combination for improving shell and yolk quality. Similar findings were reported by Singh et al. ( 2016 ) and Santosh and Raj ( 2023 ) in their studies. RE showed significant variations for various traits, except EL, YW, and YWT. K×S showed the highest positive RE for EWT and AW, indicating that using K as the maternal parent and S as the paternal parent improves EWT and AW. D×S cross had the highest positive RE for ST, suggesting that using D as the maternal parent and S as the paternal parent improves ST. Crosses like S×D and K×D should be prioritized for improving EWT and EL, while H×K can be used to enhance ST. The direction of crossing should be considered to optimize egg quality traits, as seen in the positive RE for K×S and D×S. Similarly, Fikrineh et al. ( 2023 ) reported, except for EL, other studied traits aligned with this study showed variation in RE. Singh et al. ( 2016 ) also reported significant RE for EWT. The GCA/SCA ratio is used to assess the relative importance of additive genetic effects versus non-additive genetic effects in the expression of a trait. A high GCA/SCA ratio suggests that additive genetic effects are more important, while a low ratio indicates that non-additive genetic effects (dominance or epistasis) play a more significant role. Traits like EWT (0.681) and YWT (0.345) were primarily influenced by additive genetic effects, suggesting selection-based breeding (such as recurrent selection) would be more effective. Conversely, traits like EL (0.044), EW (0.016), ST (0.002), and YR (0.158) were dominated by non-additive effects, making hybrid breeding or specific crosses to exploit heterosis more suitable. For traits with moderate ratios like AW (0.259) and YW (0.227), a combined approach of selection and hybrid breeding is recommended. Maternal effect (M e ) The M e exhibited significant variation across most traits Table 5 . Significant M e observed on EWT in crosses H×K, H×S, H×D, S×D, and K×D (P < 0.05). Notably, H×S displayed a strong negative M e , whereas H×D exhibited a positive effect. This highlights the substantial impact of maternal breed selection on EWT. Crossbreeding also significantly influenced traits such as AW and YWT, with H×D showing positive M e for EWT and AW, and H×S for YWT, reflecting enhanced performance. However, negative M e were observed in H×S, S×D, and K×D for EWT, as well as in K×D for AW, indicating these combinations may be less favorable. Traits like EL and SWT showed no significant M e , suggesting limited crossbreeding influence. These findings underscore the need for strategic crossbreeding to harness heterosis and improve key traits, while avoiding combinations with detrimental M e in targeted breeding programs. Declarations Acknowledgements Many thanks to Werer Agricultural Research Center (WARC), for allowing the poultry farm and providing the necessary facilities for this experiment. The Animal and Human Health Program of the International Livestock Research Institute (ILRI) and Ethiopian Institute of Agricultural Research (EIAR) are highly appreciated for financing the study. Author contributions Philimon Teshome was responsible for the study's design, data collection, analysis, and interpretation. He also wrote the first draft of the manuscript. All authors reviewed, provided feedback on earlier versions, and approved the final manuscript. Funding This work was manly funded by the International Livestock Research Institute (ILRI), Ethiopia, under the Animal and Human Health Program (grant number: 04/GF/23/6443, January 2023). Under the grant conditions of the Foundation, a Creative Commons Attribution 4.0 Generic License has already been assigned to the Author Accepted Manuscript version that might arise from this submission. The findings and conclusions contained within are those of the authors and do not necessarily reflect the positions or policies of ILRI. Data availability Data associated with this article are included in the manuscript but are available from the corresponding author on reasonable request. Ethics approval The protocol for the conduct of animal experiment was approved by the animal research ethical review committee of Collage of Veterinary Medicine and Agriculture of Addis Ababa University under the certificate reference number VM/ERC/01/06/15/2024. The birds were treated humanely in accordance with all international, national, and institutional guidelines for the care of animals used in research and teaching. Competing interests The authors declare that they have no known competing interests that could have appeared to influence the work reported in this paper. References Adebambo, A. O., Ikeobi, C. O. N., Ozoje, M. O., Oduguwa, O. O., and Adebambo Olufunmilayo, A. (2011). Combining abilities of growth traits among pure and crossbred meat type chickens. Arch. 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Diallel crossing analysis for body weight and egg production traits of two native Egyptian and two exotic chicken breeds. Int. J. Poult. Sci., 7(1), 64–71. https://doi.org/10.3923/ijps.2008.64.71 Samrawit, M. (2020). Performance evaluation of Dz-white synthetic and Improved indigenous Horro chicken breeds under on-farm and on-station management in different agro-ecological zones of Ethiopia [Ph.D. Thesis]. Addis Ababa University College of Veterinary Medicine and Agriculture. Santosh, M., and Raj, N. (2023). Estimation of egg production and quality parameters through complete diallel crosses of pure and crossbred Japanese quails. Pharma Innov. J., 12(12), 400–405. Singh, V., Narayan, R., Saxena, V. K., Tyagi, P. K., and Chandrahas. (2016). Analysis of heterosis and combining ability for production and egg quality traits in complete diallel cross of three chicken breeds. Indian J. Poult. Sci., 51(2), 132. https://doi.org/10.5958/0974-8180.2016.00038.6 Sinha, B., Mandal, K., Kumari, R., and Kumari, T. (2018). Estimate and effect of breeds on egg quality traits of poultry: A Review. Int. J. Livest. Res., 1. https://doi.org/10.5455/ijlr.20170812102444 Sola-Ojo, F. E., and Ayorinde, K. L. (2011). Evaluation of Reproductive Performance and Egg quality Traits in Progenies of Dominant Black Strain Crossed with Fulani Ecotype Chicken. Journal of Agricultural Science, 3(1). https://doi.org/10.5539/jas.v3n1p258 Tadelle, D. (2003). Phenotypic and genetic characterization of chicken ecotypes in Ethiopia. [Ph.D. Thesis]. Humboldt University. Tadelle, D., Alemu, Y., and Peters, K. J. (2000). Indigenous chickens in Ethiopia: genetic potential and attempts at improvement. World’s Poult. Sci. J., 56(1), 45–54. https://doi.org/10.1079/WPS20000005 Tewodros, F., Gebreyohannes, B., Mammo, M., and Misba, A. (2022). Effect of dietary metabolizable energy and crude protein on the laying performance, egg quality, hatchability, and fertility of DZ-White chickens. Int. J. Livest. Res., 12(7). Udoh, J. E., Udoh, U. H., and Adeoye, A. A. (2020). Comparative analysis of fertile egg quality traits in exotic, two Nigerian ecotype chickens and their crosses. Niger. J. Anim. Sci., 22(1), 50–59. Wearden, S., Tindell, D., and Craig, J. V. (1965). Use of a full diallel cross to estimate general and specific combining ability in chickens. Poult. Sci., 44(4), 1043–1053. https://doi.org/10.3382/ps.0441043 Williams, S., Price, S., and Siegel, P. (2002). Heterosis of growth and reproductive traits in fowl. Poult. Sci., 81(8), 1109–1112. https://doi.org/10.1093/ps/81.8.1109 Wolde, S., Mirkena, T., Melesse, A., Dessie, T., and Abegaz, S. (2021). Egg production and quality traits of Sasso-RIR, Normal Feathered local and their F1 cross chickens managed under on-station condition in Southern Ethiopia. Trop. Subtrop. Agroecosyst., 24(3). https://doi.org/10.56369/tsaes.3828 Yesihak, M., Ashenafi, G., Kefelegn, K., and Mengistu Urge. (2024). Comparative evaluation of egg quality traits in two exotic chicken breeds. J. Food Chem. Nanotechnol., 10(1). https://doi.org/10.17756/jfcn.2024-169 Yonas, K., Sandip, B., and Mestawet, T. (2019). Some internal and external egg quality characteristics of local and exotic chickens reared in Yirgalem and Hawassa towns, Ethiopia. Int. J. Livest. Prod., 10(5), 135–142. https://doi.org/10.5897/IJLP2018.0547 Zhang, X., Li, Y., Li, Q., Zhang, T., Sun, Y., Shi, F., and Chen, J. (2024). Research Note: Genetic parameters estimation of egg quality traits in Rhode Island Red and White Leghorn chickens. Poult. Sci., 103(12), 104263. https://doi.org/10.1016/j.psj.2024.104263 Table 5 Table 5 is available in the Supplementary Files section. Supplementary Files Table5.docx Cite Share Download PDF Status: Under Review Version 1 posted Reviewers agreed at journal 08 Apr, 2025 Reviewers invited by journal 27 Mar, 2025 Editor assigned by journal 12 Mar, 2025 First submitted to journal 07 Mar, 2025 You are reading this latest preprint version Research Square lets you share your work early, gain feedback from the community, and start making changes to your manuscript prior to peer review in a journal. As a division of Research Square Company, we’re committed to making research communication faster, fairer, and more useful. We do this by developing innovative software and high quality services for the global research community. Our growing team is made up of researchers and industry professionals working together to solve the most critical problems facing scientific publishing. Also discoverable on Platform About Our Team In Review Editorial Policies Advisory Board Help Center Resources Author Services Accessibility API Access RSS feed Manage Cookie Preferences © Research Square 2026 | ISSN 2693-5015 (online) Privacy Policy Terms of Service Do Not Sell My Personal Information {"props":{"pageProps":{"initialData":{"identity":"rs-6173323","acceptedTermsAndConditions":true,"allowDirectSubmit":false,"archivedVersions":[],"articleType":"Research Article","associatedPublications":[],"authors":[{"id":435022889,"identity":"6cca40f6-91a5-4076-a268-e1ba9d7f51c8","order_by":0,"name":"Philimon Teshome Feyisa","email":"data:image/png;base64,iVBORw0KGgoAAAANSUhEUgAAAZAAAAAyAQMAAABI0h/eAAAABlBMVEX///8AAABVwtN+AAAACXBIWXMAAA7EAAAOxAGVKw4bAAAAxklEQVRIiWNgGAWjYJACZhDBz8DARqR6NqgWyQaStRgcIFaLuXzz4c8FFdvkjG8kP3vwoYJBnl/sAH4tlm1sadIzztw2NruRZm444wyD4czZCfi1GBzjMWPmbbuduO1Ggpk0bxtDgsFtglr4P38GaqnfPCP9G7FaeBiAKm8nGEjkEGmLZVuamTTPmdtAb7wpk5xxRoKwX8yZDz/+zFNxW56/PX2bxIcKG3l+aUIOg7MEwCol8CtH1cJ/gLDqUTAKRsEoGJkAAOESP/Bu/E7sAAAAAElFTkSuQmCC","orcid":"https://orcid.org/0009-0000-9458-4729","institution":"Ethiopian Institute of Agricultural Research","correspondingAuthor":true,"prefix":"","firstName":"Philimon","middleName":"Teshome","lastName":"Feyisa","suffix":""},{"id":435022890,"identity":"fbf4d982-d7b2-426c-8bc0-54a831a37666","order_by":1,"name":"Gebeyehu Goshu","email":"","orcid":"","institution":"Addis Ababa University","correspondingAuthor":false,"prefix":"","firstName":"Gebeyehu","middleName":"","lastName":"Goshu","suffix":""},{"id":435022891,"identity":"e2df7642-2620-448e-ab47-59bddaaa6e28","order_by":2,"name":"Wondmeneh Esatu","email":"","orcid":"","institution":"International Livestock Research Institute Ethiopia","correspondingAuthor":false,"prefix":"","firstName":"Wondmeneh","middleName":"","lastName":"Esatu","suffix":""},{"id":435022892,"identity":"4c470a0e-818a-4127-aa85-868078cab0d3","order_by":3,"name":"Tadelle Dessie","email":"","orcid":"","institution":"International Livestock Research Institute Ethiopia","correspondingAuthor":false,"prefix":"","firstName":"Tadelle","middleName":"","lastName":"Dessie","suffix":""}],"badges":[],"createdAt":"2025-03-06 20:34:31","currentVersionCode":1,"declarations":"","doi":"10.21203/rs.3.rs-6173323/v1","doiUrl":"https://doi.org/10.21203/rs.3.rs-6173323/v1","draftVersion":[],"editorialEvents":[],"editorialNote":"","failedWorkflow":false,"files":[{"id":80752358,"identity":"926ce64b-5793-4678-ae08-fccd90cc0d7c","added_by":"auto","created_at":"2025-04-16 16:46:36","extension":"pdf","order_by":0,"title":"","display":"","copyAsset":false,"role":"manuscript-pdf","size":1442136,"visible":true,"origin":"","legend":"","description":"","filename":"manuscript.pdf","url":"https://assets-eu.researchsquare.com/files/rs-6173323/v1/21273eca-6f62-452e-8054-8dea6e10a84a.pdf"},{"id":80751825,"identity":"3139e95b-a919-431a-ad91-2fa822e94afa","added_by":"auto","created_at":"2025-04-16 16:38:35","extension":"docx","order_by":1,"title":"","display":"","copyAsset":false,"role":"supplement","size":216961,"visible":true,"origin":"","legend":"","description":"","filename":"Table5.docx","url":"https://assets-eu.researchsquare.com/files/rs-6173323/v1/7d22b481c9aee7a4cc455778.docx"}],"financialInterests":"","formattedTitle":"Genetic parameters estimation for egg quality traits of four chicken breeds using a full diallel cross","fulltext":[{"header":"Introduction","content":"\u003cp\u003ePoultry eggs are highly nutritious and relatively affordable food sources, offering high-quality protein complete with all essential amino acids, essential fatty acids, and a wide range of vitamins and minerals (Chambers et al. \u003cspan citationid=\"CR6\" class=\"CitationRef\"\u003e2017\u003c/span\u003e; Morris et al. \u003cspan citationid=\"CR24\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). They play a vital role in supporting early growth and development due to their rich nutritional content (Omer et al. \u003cspan citationid=\"CR25\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). Despite the nutritional significance, egg production in Ethiopia remains notably low (Tadelle \u003cspan citationid=\"CR33\" class=\"CitationRef\"\u003e2003\u003c/span\u003e). One of the key factors contributing to this issue is the poor production performance of indigenous breeds, largely stemming from their limited genetic potential (Kassa et al. \u003cspan citationid=\"CR18\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Conversely, various efforts have been made to enhance the overall productivity of the poultry sector by introducing exotic chicken breeds (Tadelle et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2000\u003c/span\u003e). However, these initiatives have achieved only limited success, primarily due to several challenges. These include the breeds poor adaptation to harsh production environments, their heightened vulnerability to diseases, and poor on-farm management practices (Tadelle et al. \u003cspan citationid=\"CR34\" class=\"CitationRef\"\u003e2000\u003c/span\u003e; Fikrineh et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2023\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eAmong poultry traits, egg quality stands as one of the most important, holding immense value for both producers and consumers alike (Ademola et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). In today\u0026rsquo;s world, consumers are more discerning than ever, paying close attention to what they purchase and consume. This increased focus on egg quality is a big change, directly affecting the market demand and economic value of both fertile and table eggs (Hanusova et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Ademola et al. \u003cspan citationid=\"CR2\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Most importantly, the quality of eggs also significantly influences the reproductive success of the parents (Islam et al. \u003cspan citationid=\"CR16\" class=\"CitationRef\"\u003e2001\u003c/span\u003e). So, examining the attributes that define egg quality characteristics is essential for boosting productivity and addressing the rising demand for chicken products (Beyene et al. \u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eGenetic factors play a crucial role in determining egg quality characteristics, and estimating the genetic parameters for targeted traits can determine the degree to which these traits are influenced by genetics (Hanusova et al. \u003cspan citationid=\"CR14\" class=\"CitationRef\"\u003e2015\u003c/span\u003e; Zhang et al. \u003cspan citationid=\"CR42\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Improving poultry genetics usually involves two main strategies: crossbreeding and artificial selection (Duenk et al. \u003cspan citationid=\"CR8\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Both methods have different merits in poultry breeding, and the choice depends on the specific goals of the breeder. Crossbreeding in poultry is often more beneficial because it allows us to combine the favorable traits of two distinct breeds or genetic lines to produce offspring with enhanced qualities (Mokoena et al. \u003cspan citationid=\"CR23\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Therefore, this study was planned to estimate crossbreeding parameters of various egg quality traits by crossing four chicken breeds: one locally improved breed (Improved Horro), two exotic breeds (Sasso and Potchefstroom Koekoek), and one locally developed composite breed (Dz-White feathered), utilizing a 4\u0026times;4 full diallel mating system. The diallel cross design facilitates a detailed analysis of inheritance mechanisms, such as general combining ability (GCA), specific combining ability (SCA), and the heritability of traits that are either qualitative or complex in nature (Wearden et al. \u003cspan citationid=\"CR37\" class=\"CitationRef\"\u003e1965\u003c/span\u003e). This approach offers critical insights into the genetic potential of each breed and their hybrid offspring.\u003c/p\u003e"},{"header":"Materials and Methods","content":"\u003cdiv id=\"Sec3\" class=\"Section2\"\u003e \u003ch2\u003eStudy area\u003c/h2\u003e \u003cp\u003eThis study was carried out at the poultry farm of Werer Agricultural Research Center, situated in the Afar Regional State of Ethiopia, approximately 280 km from the capital city, Addis Ababa. The farm is located at an elevation of 750 meters above sea level (Jemal et al. \u003cspan citationid=\"CR17\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). The region experiences a mean annual rainfall of less than 590 mm, with May and June being the driest months. The area is characterized by high temperatures, ranging from 19.3\u0026deg;C to 45\u0026deg;C (Atsbaha et al. \u003cspan citationid=\"CR3\" class=\"CitationRef\"\u003e2022\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eBreeding plan and management\u003c/h3\u003e\n\u003cp\u003eCrossbred genetic groups were produced by using random mating, and the sex ratio was maintained at 1:5 for all breed groups. For Horro dams, due to their small size, artificial insemination was used by collecting semen from sires of S, K, and D. Fertile eggs were collected, cleaned, and recorded daily according to breeds and crossbreds. After seven days collection the eggs were weighed and set for incubation separately according to their different breeds category. Accordingly, from the diallel cross, 4 purebred (H, S, K, D), 6 crossbred (H\u0026times;S, H\u0026times;K, H\u0026times;D, S\u0026times;K, S\u0026times;D, K\u0026times;D), and six reciprocal crossbred (S\u0026times;H, K\u0026times;H, D\u0026times;H, K\u0026times;S, D\u0026times;S, D\u0026times;K) progenies were obtained. From purebreds and crossbreds of each genotype a total of 800 female chicks were randomly distributed into 48 pens (16 progenies by 3 replications) under a completely randomized design (CRD).\u003c/p\u003e \u003cp\u003eAll experimental chickens were vaccinated against Newcastle disease (HB1 and Lasota), Gumboro (Infectious Bursal disease), fowl typhoid, and fowl pox based on the producer\u0026rsquo;s manual. Starting from hatch chickens were provided with a starter ration (20% CP and 3000 kcal/kg ME) until 8 week (WK) of age, followed by grower ration (18% CP and 2950 kcal/kg of ME) from 9 to 20 WK, and then the ration was changed into layer ration (16% CP and 2800 kcal/kg of ME) from 21 to 40 WK. Feed was purchased from Alema Feeds Co., Ltd., Debrezeit, Ethiopia, and clean water were always available in a drinker. During the chicks growing stage (from hatch upto WK 8) fluorescent lamps were placed to provide lighting for the chickens. Due to the hot temperature of the study area for the first 5 days 16\u0026ndash;18 hours of light was provided, afterwards 12 hours of light up to the 8th WK.\u003c/p\u003e\n\u003ch3\u003eTraits recorded\u003c/h3\u003e\n\u003cp\u003eA total of 480 eggs (10 from each replication) were randomly chosen at the pick production period (at 32 WK). Sampled eggs were evaluated for egg weight (\u003cb\u003eEWT\u003c/b\u003e), egg length (\u003cb\u003eEL\u003c/b\u003e), egg width (\u003cb\u003eEW\u003c/b\u003e), shell weight (\u003cb\u003eSWT\u003c/b\u003e), shell thickness (\u003cb\u003eST\u003c/b\u003e), shell ratio (\u003cb\u003eSR\u003c/b\u003e), albumen height (\u003cb\u003eAH\u003c/b\u003e), albumen width (\u003cb\u003eAW\u003c/b\u003e), yolk height (\u003cb\u003eYH\u003c/b\u003e), yolk width (\u003cb\u003eYW\u003c/b\u003e), yolk weight (\u003cb\u003eYWT\u003c/b\u003e), yolk ratio (\u003cb\u003eYR\u003c/b\u003e), and hugh unit (\u003cb\u003eHU\u003c/b\u003e).\u003c/p\u003e \u003cp\u003eEggs were weighed using a sensitive balance then length and width of each egg were measured using a compass. Then, the eggs were broken within 24 hours on a table glass cover and YH and AH were measured using a spherometer. YW and AW were measured using a compass. Then yolk of each egg was separated from the albumen, then weighed in grams and expressed as a percentage relative to the EWT. The ST were measured at 3 different points of the eggshell and the calculated average of the three points were used. The shell and membrane were weighed together in grams for each egg and expressed as a percentage relative to the egg weight. Haugh unit was calculated according to the equation of (Haugh \u003cspan citationid=\"CR15\" class=\"CitationRef\"\u003e1937\u003c/span\u003e): \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:HU\\:=\\:100\\:\\text{log}\\left(AH-1.7E{W}^{0.37}+7.6\\right)\\)\u003c/span\u003e\u003c/span\u003e, where; H\u0026thinsp;=\u0026thinsp;observed height of the albumen (mm) and W\u0026thinsp;=\u0026thinsp;weight of the eggs in grams.\u003c/p\u003e \u003cdiv id=\"Sec6\" class=\"Section2\"\u003e \u003ch2\u003eStatistical analysis\u003c/h2\u003e \u003cp\u003eEgg quality data were analyzed for variation between the genotypes using the general linear model procedure of R (Version 4.4.2) (R Core Team \u003cspan citationid=\"CR26\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). Differences considered to be significant (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05) were compared by Duncan\u0026rsquo;s Multiple Range Test (Duncan \u003cspan citationid=\"CR9\" class=\"CitationRef\"\u003e1955\u003c/span\u003e).\u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eGenetic parameters estimation\u003c/h3\u003e\n\u003cp\u003eDiallel analysis was carried out only when the differences among the genotypes were significant, fixed-effect model of (Griffing \u003cspan citationid=\"CR13\" class=\"CitationRef\"\u003e1956\u003c/span\u003e) Model 1, Method 1, a commonly used approach for assessing the genetic potential of breeding lines, especially in diallel crossing experiments, was used. The mathematical model for the combing ability analysis used was:\u003cdiv id=\"Equa\" class=\"Equation\"\u003e\u003cdiv format=\"TEX\" class=\"mathdisplay\" id=\"FileID_Equa\" name=\"EquationSource\"\u003e\n$$\\:{Y}_{ij}=\\mu\\:+{G}_{i}+{G}_{j}+{S}_{ij}+{r}_{ij}+{e}_{ij}$$\u003c/div\u003e\u003c/div\u003e\u003c/p\u003e \u003cp\u003eWhere, Y\u003csub\u003eij\u003c/sub\u003e = the observed performance of the i\u003csup\u003eth\u003c/sup\u003e genotype crossed with the j\u003csup\u003eth\u003c/sup\u003e genotype, \u0026micro;\u0026thinsp;=\u0026thinsp;the overall mean of the population, G\u003csub\u003ei\u003c/sub\u003e and G\u003csub\u003ej\u003c/sub\u003e = general combining abilities of the i\u003csup\u003eth\u003c/sup\u003e and j\u003csup\u003eth\u003c/sup\u003e genotypes (represents the additive genetic effect of genotype i and j), S\u003csub\u003eij\u003c/sub\u003e = specific combining ability for the i\u003csup\u003eth\u003c/sup\u003e and j\u003csup\u003eth\u003c/sup\u003e cross (represents the non-additive genetic interaction between i and j genotypes), r\u003csub\u003eij =\u003c/sub\u003e is the reciprocal effect involving the reciprocal crosses between the i\u003csup\u003eth\u003c/sup\u003e and j\u003csup\u003eth\u003c/sup\u003e parents, and \u0026#119890;\u003csub\u003e\u0026#119894;\u0026#119895;\u003c/sub\u003e = random error term, which accounts for any variation not explained by the fixed effects.\u003c/p\u003e \u003cp\u003eHeterosis was calculated on percentage of mid-parents by application of the following formula (Williams et al. \u003cspan citationid=\"CR38\" class=\"CitationRef\"\u003e2002\u003c/span\u003e):\u003cdiv id=\"Equb\" class=\"Equation\"\u003e\u003cdiv format=\"TEX\" class=\"mathdisplay\" id=\"FileID_Equb\" name=\"EquationSource\"\u003e\n$$\\:H\\%=F1\\:-\\left\\{\\frac{\\frac{\\left(P1+P2\\right)}{2}}{\\frac{\\left(P1+P2\\right)}{2}}\\right\\}\\times\\:100$$\u003c/div\u003e\u003c/div\u003e,\u003c/p\u003e \u003cp\u003eWhere F1\u0026thinsp;=\u0026thinsp;the first filial and P1 and P2 are parents in diallel and reciprocal crosses.\u003c/p\u003e \u003cp\u003eMaternal effects were estimated using a set of linear contrasts of the genetic group means (Dickerson \u003cspan citationid=\"CR7\" class=\"CitationRef\"\u003e1969\u003c/span\u003e) as follows: \u003cspan class=\"InlineEquation\"\u003e\u003cspan class=\"mathinline\"\u003e\\(\\:{M}^{e}=\\frac{1}{2}\\left[\\:\\left(A\\times\\:B\\right)-\\left(B\\times\\:A\\right)\\:\\right]\\)\u003c/span\u003e\u003c/span\u003e.\u003c/p\u003e"},{"header":"Results","content":"\u003cdiv id=\"Sec9\" class=\"Section2\"\u003e \u003ch2\u003eEgg quality traits\u003c/h2\u003e \u003cp\u003eThe least square means for EWT, EL, EW, SWT, ST, and SR, across different genotypes are detailed in Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e. Differences (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05) were observed for all traits. From the result, K\u0026times;S crossbred demonstrated the heaviest EWT at 55.10 g, followed by D\u0026times;S at 53.91 g. S\u0026times;K, S\u0026times;H, D\u0026times;K, and S, also displayed notable EWT, while the purebred H recorded the lightest EWT at 46.08 g. The purebred D stood out with the longest EL at 53.89 mm, trailed by S at 53.50 mm. Conversely, the D\u0026times;S crossbred exhibited the shortest EL at 42.79 mm. For EW, S achieved the largest measurement at 41.82 mm, followed by D\u0026times;K and D. On the lower end, the D\u0026times;S crossbred recorded the narrowest EW at 31.76 mm. SWT was highest in the purebred S at 5.97 g, while the D\u0026times;H crossbred displayed the lightest shells. ST varied significantly, with the S\u0026times;D crossbred showing the thickest shells at 0.47 mm, whereas the purebred K and D exhibited the thinnest shells at 0.32 mm. The SR was highest in the purebred S at 11.44%, with the crossbreds K\u0026times;H and D\u0026times;K recording the lowest at 9.36% and 9.35%, respectively.\u003c/p\u003e \u003cp\u003eThe least square means of AH, AW, YH, YW, YWT, YR, and HU are detailed in Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e. The analysis revealed differences (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05) across all traits. The crossbred K\u0026times;H (7.28 mm) and its reciprocal H\u0026times;K (7.00 mm) achieved the highest AH, surpassing other genotypes. Followed by H\u0026times;D (6.93 mm) and its reciprocal D\u0026times;H (6.83 mm), while the lowest AH values were recorded in S\u0026times;D (5.72 mm), D (5.55 mm), and K (5.17 mm). In terms of AW, the H\u0026times;D crossbred stood out with the highest measurement, trailed by D\u0026times;K, D, and K\u0026times;S. Conversely, the H\u0026times;K crossbred exhibited the narrowest AW. The purebred D (19.44 mm), crossbred K\u0026times;S (19.33 mm), and purebred S (19.11 mm) demonstrated the highest YH. On the other hand, D\u0026times;S (42.69 mm), S (42.33 mm), and S\u0026times;D (41.22 mm) recorded the highest YW. In contrast, the H\u0026times;K crossbred displayed the smallest YH (15.94 mm) and YW (34.98 mm) values. The purebred S (16.36 g) emerged with the heaviest yolk, outperforming all other genetic groups. For YR, the purebred H (34.26%) and crossbred H\u0026times;S (32.78%) achieved the highest percentages, with H\u0026times;D (24.30%) and D\u0026times;K (22.80%) showing the lowest values. The K\u0026times;H crossbred (87.67) exhibited the highest HU score. This was closely followed by D\u0026times;H (86.21) and its reciprocal H\u0026times;K (85.97). Other genetic groups displayed moderate HU values, while K\u0026times;S (77.12) and K (73.87) recorded the lowest scores.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab1\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 1\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eLeast square means (\u0026plusmn;\u0026thinsp;SE) for external egg quality traits\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"7\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eGenotype\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eEWT(g)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eEL (mm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eEW (mm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eSWT (g)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eST (mm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eSR (%)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePurebred\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eH\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e46.08 \u003csup\u003ej\u003c/sup\u003e\u0026plusmn;0.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e51.14\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e39.57\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.41\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e5.23\u003csup\u003ea\u0026thinsp;\u0026minus;\u0026thinsp;d\u003c/sup\u003e\u0026plusmn;0.05\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.33\u003csup\u003ee\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e11.36\u003csup\u003ea\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e52.15\u003csup\u003ede\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e53.50\u003csup\u003ea\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.39\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e41.82\u003csup\u003ea\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.43\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e5.97\u003csup\u003ea\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.35\u003csup\u003ede\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.00\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e11.44\u003csup\u003ea\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eK\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e50.44\u003csup\u003eg\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.34\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e51.29\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.48\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e38.93\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e5.05\u003csup\u003ebcd\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.32\u003csup\u003ee\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e10.02\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.11\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eD\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e49.13\u003csup\u003eh\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.36\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e53.89\u003csup\u003ea\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.47\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e41.46\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e5.52\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.32\u003csup\u003ee\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e11.24\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.50\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCrosses\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eH\u0026times;S\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e48.80\u003csup\u003ehi\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.42\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e53.17\u003csup\u003ea\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.85\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e39.55\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.90\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e5.06\u003csup\u003ebcd\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.23\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.43\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e10.39\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.57\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eH\u0026times;K\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e51.07\u003csup\u003efg\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.22\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e53.17\u003csup\u003ea\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;1.08\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e39.47\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.41\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e5.38\u003csup\u003ea\u0026thinsp;\u0026minus;\u0026thinsp;d\u003c/sup\u003e\u0026plusmn;0.26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.42\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e10.53\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.50\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eH\u0026times;D\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e50.38\u003csup\u003eg\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e53.29\u003csup\u003ea\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.36\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e40.94\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e4.85\u003csup\u003ecd\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.32\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.37\u003csup\u003ecde\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e9.61\u003csup\u003ec\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.59\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eS\u0026times;K\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e53.46\u003csup\u003ebc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.57\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e51.66\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;1.67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e40.04\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.54\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e5.42\u003csup\u003ea\u0026thinsp;\u0026minus;\u0026thinsp;d\u003c/sup\u003e\u0026plusmn;0.30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.33\u003csup\u003ee\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e10.16\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.66\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eS\u0026times;D\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e50.85\u003csup\u003efg\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e45.63\u003csup\u003ec\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;2.13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e34.05\u003csup\u003ed\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;2.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e5.72\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.47\u003csup\u003ea\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e11.26\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.28\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eK\u0026times;D\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e50.11\u003csup\u003eg\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e49.30\u003csup\u003eb\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;1.98\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e37.74\u003csup\u003ec\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;2.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e5.25\u003csup\u003ea\u0026thinsp;\u0026minus;\u0026thinsp;d\u003c/sup\u003e\u0026plusmn;0.18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.43\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e10.48\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.34\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRE\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eS\u0026times;H\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e53.31\u003csup\u003ebc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e51.07\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.87\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e39.93\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.75\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e5.18\u003csup\u003ea\u0026thinsp;\u0026minus;\u0026thinsp;d\u003c/sup\u003e\u0026plusmn;0.30\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.40\u0026thinsp;\u0026plusmn;\u0026thinsp;\u003csup\u003ebcd\u003c/sup\u003e0.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e9.71\u003csup\u003ebc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.52\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eK\u0026times;H\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e51.63\u003csup\u003eef\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.22\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e52.66\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.51\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e38.92\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.44\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e4.83\u003csup\u003ecd\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.45\u003csup\u003ea\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.02\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e9.36\u003csup\u003ec\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.19\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eD\u0026times;H\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e48.05\u003csup\u003ei\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e50.63\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.62\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e38.34\u003csup\u003ebc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.42\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e4.66\u003csup\u003ed\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.05\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.40\u003csup\u003ebcd\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e9.69\u003csup\u003ebc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eK\u0026times;S\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e55.10\u003csup\u003ea\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e52.59\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.81\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e41.42\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e5.33\u003csup\u003ea\u0026thinsp;\u0026minus;\u0026thinsp;d\u003c/sup\u003e\u0026plusmn;0.05\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.33\u003csup\u003ee\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e9.68\u003csup\u003ebc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.07\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eD\u0026times;S\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e53.91\u003csup\u003eb\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.32\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e42.79\u003csup\u003ec\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.72\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e31.76\u003csup\u003ed\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.63\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e5.56\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.51\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.37\u003csup\u003ecde\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.03\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e10.32\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;1.00\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eD\u0026times;K\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e52.61\u003csup\u003ecd\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.50\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e52.59\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.46\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e41.50\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;1.08\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e4.92\u003csup\u003ebcd\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.42\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e9.35\u003csup\u003ec\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.65\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eP-values\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.0001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.0001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.0001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.0400\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.0001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.0300\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e1.05\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e3.56\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e4.27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e7.93\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e7.12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e8.07\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"7\"\u003e\u003csup\u003ea\u0026minus;j\u003c/sup\u003eMeans with the same letter in a column are not significantly different (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05)\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab2\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 2\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eLeast square means (\u0026plusmn;\u0026thinsp;SE) for internal egg quality traits\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"8\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eGenotype\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eAH (mm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eAW (mm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eYH (mm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eYW (mm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eYWT (g)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eYR (%)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eHU\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003ePurebred\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eH\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e 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\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e77.32\u003csup\u003ecde\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;1.74\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCrosses\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eH\u0026times;S\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e 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\u003cp\u003e27.67\u003csup\u003ecd\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;2.06\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e80.11\u003csup\u003ea\u0026thinsp;\u0026minus;\u0026thinsp;e\u003c/sup\u003e\u0026plusmn;3.97\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eS\u0026times;D\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e5.72\u003csup\u003ebcd\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e66.20\u003csup\u003ec\u0026thinsp;\u0026minus;\u0026thinsp;f\u003c/sup\u003e\u0026plusmn;1.75\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e18.89\u003csup\u003ea\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e41.22\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.75\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e15.41\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.65\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e30.31\u003csup\u003ebc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;1.31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e77.93\u003csup\u003eb\u0026thinsp;\u0026minus;\u0026thinsp;e\u003c/sup\u003e\u0026plusmn;1.07\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eK\u0026times;D\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e6.10\u003csup\u003ea\u0026thinsp;\u0026minus;\u0026thinsp;d\u003c/sup\u003e\u0026plusmn;0.10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e68.32\u003csup\u003ecde\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.97\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e16.89\u003csup\u003ecd\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.22\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e35.59\u003csup\u003ee\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;1.43\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e12.55\u003csup\u003eef\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.43\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e25.05\u003csup\u003ede\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.77\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e80.85\u003csup\u003ea\u0026thinsp;\u0026minus;\u0026thinsp;e\u003c/sup\u003e\u0026plusmn;0.73\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eRE\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eS\u0026times;H\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e6.28\u003csup\u003ea\u0026thinsp;\u0026minus;\u0026thinsp;d\u003c/sup\u003e\u0026plusmn;0.50\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e63.94\u003csup\u003ed\u0026thinsp;\u0026minus;\u0026thinsp;g\u003c/sup\u003e\u0026plusmn;0.77\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e17.27\u003csup\u003ebcd\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e36.13\u003csup\u003ee\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.60\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e13.13\u003csup\u003edef\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.72\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e24.62\u003csup\u003ede\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;1.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e80.67\u003csup\u003ea\u0026thinsp;\u0026minus;\u0026thinsp;e\u003c/sup\u003e\u0026plusmn;3.33\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eK\u0026times;H\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e7.28\u003csup\u003ea\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e62.22\u003csup\u003eefg\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;1.73\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e16.94\u003csup\u003ecd\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.53\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e35.06\u003csup\u003ee\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.37\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e13.21\u003csup\u003edef\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;1.11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e25.60\u003csup\u003ede\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;2.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e87.67\u003csup\u003ea\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.90\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eD\u0026times;H\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e6.83\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e59.84\u003csup\u003egh\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;1.88\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e16.56\u003csup\u003ed\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.87\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e35.36\u003csup\u003ee\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;1.27\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e12.30\u003csup\u003eef\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.46\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e25.61\u003csup\u003ede\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.99\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e86.21\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.54\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eK\u0026times;S\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e5.83\u003csup\u003ebcd\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e69.33\u003csup\u003ebcd\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;1.26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e19.33\u003csup\u003ea\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e40.85\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.45\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e15.39\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.47\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e27.93\u003csup\u003ecd\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.79\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e77.12\u003csup\u003ede\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;1.27\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eD\u0026times;S\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e6.06\u003csup\u003ea\u0026thinsp;\u0026minus;\u0026thinsp;d\u003c/sup\u003e\u0026plusmn;0.47\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e64.46\u003csup\u003ed\u0026thinsp;\u0026minus;\u0026thinsp;g\u003c/sup\u003e\u0026plusmn;1.16\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e18.78\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.49\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e42.69\u003csup\u003ea\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.66\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e15.05\u003csup\u003ea\u0026thinsp;\u0026minus;\u0026thinsp;d\u003c/sup\u003e\u0026plusmn;0.23\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e27.93\u003csup\u003ecd\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.59\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e78.94\u003csup\u003ea\u0026thinsp;\u0026minus;\u0026thinsp;e\u003c/sup\u003e\u0026plusmn;3.32\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eD\u0026times;K\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e6.22\u003csup\u003ea\u0026thinsp;\u0026minus;\u0026thinsp;d\u003c/sup\u003e\u0026plusmn;0.11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e74.08\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.84\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e17.00\u003csup\u003ecd\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.88\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e35.53\u003csup\u003ee\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.29\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e11.99\u003csup\u003ef\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.45\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e22.80\u003csup\u003ee\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.95\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e80.77\u003csup\u003ea\u0026thinsp;\u0026minus;\u0026thinsp;e\u003c/sup\u003e\u0026plusmn;0.72\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eP-values\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.0361\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.0001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.0001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.0001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.0001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.0001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.0287\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eCV\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e10.69\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e4.80\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e4.78\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e3.62\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e7.37\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e7.66\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e5.50\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"8\"\u003e\u003csup\u003ea\u0026minus;g\u003c/sup\u003eMeans with the same letter in a column are not significantly different (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05)\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e\n\u003ch3\u003eCrossbreeding effects\u003c/h3\u003e\n\u003cdiv id=\"Sec11\" class=\"Section2\"\u003e \u003ch2\u003eHeterosis effect (H\u003csup\u003ee\u003c/sup\u003e)\u003c/h2\u003e \u003cp\u003eH\u003csup\u003ee\u003c/sup\u003e varied significantly across different traits Table\u0026nbsp;\u003cspan refid=\"Tab3\" class=\"InternalRef\"\u003e3\u003c/span\u003e. EWT displayed positive H\u003csup\u003ee\u003c/sup\u003e for most crosses and their reciprocals, ranging from 0.42\u0026ndash;8.54%, with the exception of H\u0026times;S cross. For EL, the majority of H\u003csup\u003ee\u003c/sup\u003e estimates were negative, except for H\u0026times;S (1.64%), H\u0026times;K (3.82%), H\u0026times;D (1.49%), K\u0026times;H (2.83%), and K\u0026times;S (0.37%). Similarly, EW predominantly exhibited negative H\u003csup\u003ee\u003c/sup\u003e, with the exception of H\u0026times;K (0.56%), H\u0026times;D (1.04%), K\u0026times;S (2.58%), and D\u0026times;K (3.24%). In terms of SWT, H\u003csup\u003ee\u003c/sup\u003e was negative across all genetic groups except for H\u0026times;K (4.54%). Conversely, ST showed high and positive H\u003csup\u003ee\u003c/sup\u003e for most groups, excluding S\u0026times;K (-2.97%) and its reciprocal K\u0026times;S (-1.98%). However, SR demonstrated negative H\u003csup\u003ee\u003c/sup\u003e across all crosses and reciprocals. AH exhibited strong positive H\u003csup\u003ee\u003c/sup\u003e in the majority of crosses, except for H\u0026times;S, S\u0026times;D, and S\u0026times;H. Similarly, AW also displayed positive H\u003csup\u003ee\u003c/sup\u003e in most crosses. In contrast, YH showed negative H\u003csup\u003ee\u003c/sup\u003e across all crosses and reciprocals, with the exception of K\u0026times;S. YW and YWT also had negative H\u003csup\u003ee\u003c/sup\u003e, except for S\u0026times;D, K\u0026times;S, and D\u0026times;S. For HU, most H\u003csup\u003ee\u003c/sup\u003e were positive but relatively low, except for the crossbreds H\u0026times;S, S\u0026times;D, S\u0026times;H, K\u0026times;S, and D\u0026times;S. on the other hand, YR predominantly exhibited high and negative H\u003csup\u003ee\u003c/sup\u003e.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab3\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 3\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eEstimation of heterosis percentage for egg quality traits\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"14\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c10\" colnum=\"10\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c11\" colnum=\"11\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c12\" colnum=\"12\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c13\" colnum=\"13\"\u003e\u003c/div\u003e \u003cdiv align=\"char\" char=\".\" class=\"colspec\" colname=\"c14\" colnum=\"14\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\" morerows=\"2\" rowspan=\"3\"\u003e \u003cp\u003eTraits\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colspan=\"13\" nameend=\"c14\" namest=\"c2\"\u003e \u003cp\u003eGenotypes\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colspan=\"6\" nameend=\"c7\" namest=\"c2\"\u003e \u003cp\u003eCrosses\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colspan=\"6\" nameend=\"c14\" namest=\"c9\"\u003e \u003cp\u003eReciprocals\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eH\u0026times;S\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eH\u0026times;K\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eH\u0026times;D\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eS\u0026times;K\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eS\u0026times;D\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eK\u0026times;D\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eS\u0026times;H\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c10\"\u003e \u003cp\u003eK\u0026times;H\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c11\"\u003e \u003cp\u003eD\u0026times;H\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c12\"\u003e \u003cp\u003eK\u0026times;S\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c13\"\u003e \u003cp\u003eD\u0026times;S\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c14\"\u003e \u003cp\u003eD\u0026times;K\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEWT\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e-0.65\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e5.82\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e5.83\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e4.22\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.42\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e0.65\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e8.54\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e6.98\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e0.93\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e7.41\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e6.45\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e5.68\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEL\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e1.64\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e3.82\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1.49\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e-1.40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e-15.02\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e-6.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e-2.38\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e2.83\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e-3.59\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e0.37\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e-20.31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e0.00\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eEW\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e-2.81\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e0.56\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e1.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e-0.84\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e-18.23\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e-6.12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e-1.88\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e-0.85\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e-5.38\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e2.58\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e-23.73\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e3.24\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSWT\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e-9.58\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e4.54\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e-9.86\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e-1.57\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e-0.35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e-0.63\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e-7.56\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e-6.03\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e-13.39\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e-3.21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e-3.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e-6.94\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eST\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e27.12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e27.55\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e13.89\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e-2.97\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e40.10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e35.18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e16.67\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e39.12\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e22.81\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e-1.98\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e9.82\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e32.41\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eSR\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e-8.88\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-1.51\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e-14.92\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e-5.35\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e-0.76\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e-1.39\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e-14.85\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e-12.40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e-14.23\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e-9.80\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e-9.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e-12.05\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAH\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e-7.87\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e21.77\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e16.69\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e8.17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e-3.70\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e13.81\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e-0.87\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e26.59\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e15.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e1.45\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e1.91\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e16.04\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eAW\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e8.59\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-13.53\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e14.13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e5.52\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e0.46\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e-0.74\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e3.57\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e-3.78\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e-10.11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e8.33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e-2.19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e7.63\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eYH\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e-10.58\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-13.56\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e-16.17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e-1.79\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e-2.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e-10.31\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e-8.55\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e-8.13\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e-13.11\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e3.58\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e-2.58\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e-9.74\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eYW\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e-13.26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-10.46\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e-10.91\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e-0.24\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e1.33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e-7.14\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e-12.78\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e-10.26\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e-11.09\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e2.18\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e4.95\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e-7.29\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eYWT\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e-0.53\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-5.40\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e-19.42\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e-2.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e-0.42\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e-11.80\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e-18.28\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e-10.90\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e-19.02\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e1.84\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e-2.75\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e-15.74\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eYR\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e-0.10\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e-11.04\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e-24.06\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e-5.99\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e-0.78\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e-12.46\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e-24.95\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e-17.07\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e-19.97\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e-5.09\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e-8.57\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e-20.30\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eHU\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c2\"\u003e \u003cp\u003e-3.78\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c3\"\u003e \u003cp\u003e9.21\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c4\"\u003e \u003cp\u003e6.75\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c5\"\u003e \u003cp\u003e3.32\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c6\"\u003e \u003cp\u003e-1.69\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c7\"\u003e \u003cp\u003e6.94\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e\u0026nbsp;\u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c9\"\u003e \u003cp\u003e-2.09\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c10\"\u003e \u003cp\u003e11.37\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c11\"\u003e \u003cp\u003e7.17\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c12\"\u003e \u003cp\u003e-0.54\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c13\"\u003e \u003cp\u003e-0.41\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"char\" char=\".\" colname=\"c14\"\u003e \u003cp\u003e6.84\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003cp\u003e \u003cem\u003eH\u003c/em\u003e Improved Horro, \u003cem\u003eS\u003c/em\u003e Sasso, \u003cem\u003eK\u003c/em\u003e Potchefstroom Koekoek, \u003cem\u003eD\u003c/em\u003e Dz-white; Males are listed first in the cross\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec12\" class=\"Section2\"\u003e \u003ch2\u003eCombining ability and reciprocal effects\u003c/h2\u003e \u003cp\u003eEstimates of GCA, SCA, and RE for egg quality traits are presented in Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e. GCA effects were highly significant (P\u0026thinsp;\u0026lt;\u0026thinsp;0.001) for EWT, EL, EW, AW, YW, YWT, and YR, but not for ST. The S breed showed the highest positive GCA estimates for EWT, followed by K, while D and H breeds recorded negative values. Positive GCA estimate for EL was observed in H and K breeds, while S and D breeds showed negative estimates. The D breed exhibited the highest positive GCA value for AW, followed by K, with S and H breeds showing negative values. Only the S breed recorded positive GCA values for YW and YWT, while D, K, and H breeds had negative estimates. For YR, the S breed showed the highest positive GCA estimates, followed by H, while D and K breeds had small negative estimates.\u003c/p\u003e \u003cp\u003eSCA variations were highly significant (P\u0026thinsp;\u0026lt;\u0026thinsp;0.001) for all traits. The S\u0026times;D crossbred had the highest positive SCA estimates for EWT, followed by K\u0026times;D, H\u0026times;K, S\u0026times;K, and H\u0026times;S, while H\u0026times;D recorded a negative SCA estimates. The K\u0026times;D crossbred showed the highest positive SCA value for EL, followed by H\u0026times;S, S\u0026times;K, and H\u0026times;K, while H\u0026times;D and S\u0026times;D had negative estimates. For ST, the H\u0026times;K crossbred had the highest positive SCA estimate, followed by H\u0026times;S, H\u0026times;D, and S\u0026times;K, while K\u0026times;D and S\u0026times;D exhibited small negative SCA values. Positive SCA estimates for AW were observed in K\u0026times;D, H\u0026times;S, and S\u0026times;K, while S\u0026times;D, H\u0026times;K, and H\u0026times;D showed negative estimates. Most SCA estimates for YW were negative, except for S\u0026times;D, S\u0026times;K. The S\u0026times;K crossbred showed the highest SCA values for YWT, while H\u0026times;S recorded the highest negative estimate. For YR, only H\u0026times;D and S\u0026times;K crossbreds had positive SCA estimates, while the remaining crosses showed negative values.\u003c/p\u003e \u003cp\u003eRE were significant (P\u0026thinsp;\u0026lt;\u0026thinsp;0.005) for EWT, EW, ST, AW, and YR, but not for EL, YW, and YWT. The K\u0026times;S crossbred had the highest positive RE for EWT, followed by D\u0026times;S, D\u0026times;H, and D\u0026times;K, while K\u0026times;H and S\u0026times;H showed negative values. For EW, RE was positive for most crosses, except S\u0026times;H and D\u0026times;S. However, D\u0026times;S had the highest positive estimate for ST. The K\u0026times;S crossbred recorded the highest positive estimate for AW but a negative estimate for YR. The S\u0026times;H crossbred exhibited the highest positive estimate for YR. From the results, EL had the highest GCA/SCA ratio, while AW and YW showed moderate ratios. Conversely, YR, EL, EW, and ST exhibited the lowest ratios.\u003c/p\u003e \u003cp\u003e \u003cdiv class=\"gridtable\"\u003e\u003ctable float=\"Yes\" id=\"Tab4\" border=\"1\"\u003e \u003ccaption language=\"En\"\u003e \u003cdiv class=\"CaptionNumber\"\u003eTable 4\u003c/div\u003e \u003cdiv class=\"CaptionContent\"\u003e \u003cp\u003eEstimates (\u0026plusmn;\u0026thinsp;SE) of combining ability for external and internal egg quality traits\u003c/p\u003e \u003c/div\u003e \u003c/caption\u003e \u003ccolgroup cols=\"9\"\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c1\" colnum=\"1\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c2\" colnum=\"2\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c3\" colnum=\"3\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c4\" colnum=\"4\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c5\" colnum=\"5\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c6\" colnum=\"6\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c7\" colnum=\"7\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c8\" colnum=\"8\"\u003e\u003c/div\u003e \u003cdiv align=\"left\" class=\"colspec\" colname=\"c9\" colnum=\"9\"\u003e\u003c/div\u003e \u003cthead\u003e \u003ctr\u003e \u003cth align=\"left\" colname=\"c1\"\u003e \u003cp\u003eGenotype\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c2\"\u003e \u003cp\u003eEWT (g)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c3\"\u003e \u003cp\u003eEL (mm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c4\"\u003e \u003cp\u003eEW (mm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c5\"\u003e \u003cp\u003eST (mm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c6\"\u003e \u003cp\u003eAW (mm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c7\"\u003e \u003cp\u003eYW (mm)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c8\"\u003e \u003cp\u003eYWT (g)\u003c/p\u003e \u003c/th\u003e \u003cth align=\"left\" colname=\"c9\"\u003e \u003cp\u003eYR (%)\u003c/p\u003e \u003c/th\u003e \u003c/tr\u003e \u003c/thead\u003e \u003ctbody\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eGCA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.0001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.0026\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.0479\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003eNS\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e\u0026lt;\u0026thinsp;0.0001\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e 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\u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-0.08\u003csup\u003eb\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.38\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003er\u003csup\u003eD\u0026times;S\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.46\u003csup\u003eb\u0026thinsp;\u0026minus;\u0026thinsp;e\u003c/sup\u003e\u0026plusmn;0.19\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e-5.38\u003csup\u003eb\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e-5.21\u003csup\u003eb\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.25\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.04\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e-2.54\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.08\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e1.81\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.41\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.50\u003csup\u003eabc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.41\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.75\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.88\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003er\u003csup\u003eD\u0026times;K\u003c/sup\u003e\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.05\u003csup\u003edef\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.15\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.02\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.68\u003csup\u003ea\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.20\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.05\u003csup\u003eab\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.01\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e2.00\u003csup\u003ea\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.06\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e-1.56\u003csup\u003ecd\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e-0.86\u003csup\u003ebc\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.33\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e-1.64\u003csup\u003eb\u003c/sup\u003e\u0026thinsp;\u0026plusmn;\u0026thinsp;0.70\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003ctr\u003e \u003ctd align=\"left\" colname=\"c1\"\u003e \u003cp\u003eGCA/SCA\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c2\"\u003e \u003cp\u003e0.681\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c3\"\u003e \u003cp\u003e0.044\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c4\"\u003e \u003cp\u003e0.016\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c5\"\u003e \u003cp\u003e0.002\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c6\"\u003e \u003cp\u003e0.259\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c7\"\u003e \u003cp\u003e0.227\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c8\"\u003e \u003cp\u003e0.345\u003c/p\u003e \u003c/td\u003e \u003ctd align=\"left\" colname=\"c9\"\u003e \u003cp\u003e0.158\u003c/p\u003e \u003c/td\u003e \u003c/tr\u003e \u003c/tbody\u003e \u003c/colgroup\u003e \u003ctfoot\u003e \u003ctr\u003e\u003ctd colspan=\"9\"\u003e\u003csup\u003ea\u0026minus;i\u003c/sup\u003eMeans with the same letter in a column are not significantly different (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05); \u003cem\u003eg\u003c/em\u003e: general combining ability, \u003cem\u003es\u003c/em\u003e: specific combining ability,\u003c/td\u003e\u003c/tr\u003e \u003ctr\u003e\u003ctd colspan=\"9\"\u003e\u003cem\u003er\u003c/em\u003e: reciprocal effect\u003c/td\u003e\u003c/tr\u003e \u003c/tfoot\u003e \u003c/table\u003e\u003c/div\u003e \u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec13\" class=\"Section2\"\u003e \u003ch2\u003eMaternal effect (M\u003csup\u003ee\u003c/sup\u003e)\u003c/h2\u003e \u003cp\u003eThe M\u003csup\u003ee\u003c/sup\u003e estimates for the studied traits varied, displaying both positive and negative values with low to moderate to high magnitudes Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e. With the exception of the H\u0026times;D, all other crosses displayed negative M\u003csup\u003ee\u003c/sup\u003e on EWT. In terms of EL, S\u0026times;K and K\u0026times;D crosses showed negative estimates, while the remaining crosses exhibited positive estimates. For EW, the estimates were positive for the H\u0026times;K, H\u0026times;S, and S\u0026times;D, whereas the other three crosses showed negative values. Regarding SWT and SR, the H\u0026times;S and H\u0026times;D crosses demonstrated negative M\u003csup\u003ee\u003c/sup\u003e, while the H\u0026times;K, S\u0026times;K, S\u0026times;D, and K\u0026times;D showed positive estimates. However, for ST, the H\u0026times;K, H\u0026times;D, and S\u0026times;K crosses exhibited negative estimates, whereas the H\u0026times;S, S\u0026times;D, and K\u0026times;D displayed positive estimates.\u003c/p\u003e \u003cp\u003eFor AH, all crosses except H\u0026times;D and S\u0026times;D exhibited negative estimates. In contrast, only the S\u0026times;D and K\u0026times;D showed negative estimates for AW. Similarly, YH displayed negative estimates for all crosses except S\u0026times;K. For YW, only the H\u0026times;K and H\u0026times;S showed negative estimates, while the others exhibited positive values. Conversely, YWT and YR displayed positive M\u003csup\u003ee\u003c/sup\u003e for all crosses except H\u0026times;D and S\u0026times;D.\u003c/p"},{"header":"Discussion","content":"\u003cdiv id=\"Sec15\" class=\"Section2\"\u003e \u003ch2\u003eEgg quality traits\u003c/h2\u003e \u003cp\u003eEWT exhibited significant variation across genotypes, highlighting the strong influence of genetics on this trait. Purebred S had the highest EWT at 32 WK of age, followed by K and D. The EWT of the S in this study was consistent with findings of Yonas et al. (\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2019\u003c/span\u003e) and Wolde et al. (\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). In contrast, Beyene et al. (\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) reported a higher EWT, however their observed EWT for the K breed aligned with the present result. The EWT for the D breed reported by Samrawit (\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) was slightly higher than that found in this study. However, the same author reported similar EWT values for H with the present findings. Table\u0026nbsp;\u003cspan refid=\"Tab1\" class=\"InternalRef\"\u003e1\u003c/span\u003e showed that the H\u0026times;K, H\u0026times;D, S\u0026times;H, K\u0026times;H, S\u0026times;K, K\u0026times;S, D\u0026times;S, and D\u0026times;K crossbreds exhibited higher EWT compared to their purebred counterparts, suggesting hybrid vigor. This implies that crossing K and D with S improves EWT compared to purebreds. H recorded the lowest EWT, indicating possible genetic constraints for this trait within the breed. However, crossbreeding H with S and K led to notable improvements in EWT. These findings align with (Kedija et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Atsbaha et al. \u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), they reported significant gains in EWT through crossbreeding H with exotic breeds, further supporting the heterosis effects observed in this study, particularly in H\u0026times;S, S\u0026times;H, H\u0026times;K, and K\u0026times;H crossbreds. However, the reciprocal cross D\u0026times;H performed poorly, suggesting that the maternal or paternal influence of H may negatively impact EWT.\u003c/p\u003e \u003cp\u003ePurebreds D and S had the longest eggs, indicating that these breeds inherently contribute to increased EL. The EL observed in this study for D and H agreed with findings reported by Samrawit (\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2020\u003c/span\u003e). In contrast, Beyene et al. (\u003cspan citationid=\"CR5\" class=\"CitationRef\"\u003e2022\u003c/span\u003e) and Kefelegn and Ashenafi (\u003cspan citationid=\"CR20\" class=\"CitationRef\"\u003e2024\u003c/span\u003e), reported slightly longer EL for S breed. The EL of the K breed in this study was consistent with finding of Yesihak et al. (\u003cspan citationid=\"CR40\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Crossbred H\u0026times;D also performed well, suggesting that combining H and D can enhance this trait. D\u0026times;S had the smallest EL, given that both D and S purebreds performed well, this suggests a potential negative interaction or dominance effect when S is used as the dam in this cross.\u003c/p\u003e \u003cp\u003eEW exhibited differences (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05) among genotypes. Significant differences for EW in consensus to this study had also been reported by (Kedija et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2019\u003c/span\u003e; Udoh et al. \u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2020\u003c/span\u003e; Fikrineh et al. \u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). S showed the widest eggs, followed by the D\u0026times;K and D, suggesting that both S and D breeds positively influence EW. This observation was consistent with the result reported by Wolde et al. (\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2021\u003c/span\u003e). Conversely, the D\u0026times;S crossbred had the smallest EW, which corresponds to its poor performance in EL. This further reinforces the notion of a negative interaction in the D\u0026times;S cross.\u003c/p\u003e \u003cp\u003eSignificant differences (P\u0026thinsp;\u0026lt;\u0026thinsp;0.04) in SWT were observed across groups, though the variations were minimal. This suggests that genetic influences on this trait are limited or that environmental factors play a more substantial role. S purebred exhibited the heaviest shell, followed by S\u0026times;D and D\u0026times;S, indicating that breed S positively contributes to SWT. Similar SWT value for the S was reported by Wolde et al. (\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), though Samrawit (\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) recorded higher SWT values for D and H. The D\u0026times;H crossbred had the lowest SWT, consistent with its poor performance in other traits. In contrast, the S\u0026times;D crossbred displayed the thickest shell, followed by K\u0026times;H and K\u0026times;D, suggesting that specific combinations can enhance ST, potentially due to complementary genetic effects. These findings align with previous studies (Sola-Ojo and Ayorinde, \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Khalil et al. \u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2013\u003c/span\u003e), which also demonstrated that crossbreeding improved ST. The K and D had the thinnest shells, indicating possible genetic limitations for this trait. Similarly, Samrawit (\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) observed that breed D exhibited lower performance in terms of ST compared to H. Differences (P\u0026thinsp;\u0026lt;\u0026thinsp;0.03) were noted in SR across groups, although the variations were minimal. S demonstrated the highest SR, followed by H and the S\u0026times;D crossbred, underscoring the positive influence of S and H on SR. Conversely, the K\u0026times;H and D\u0026times;K recorded the lowest SR, indicating that these combinations may not be effective for enhancing shell quality.\u003c/p\u003e \u003cp\u003eDifferences in AH (P\u0026thinsp;\u0026lt;\u0026thinsp;0.0360) and AW (P\u0026thinsp;\u0026lt;\u0026thinsp;0.0001) were observed across genotypes. K\u0026times;H and H\u0026times;K demonstrated the highest AH, reflecting strong heterosis in these combinations. Table\u0026nbsp;\u003cspan refid=\"Tab2\" class=\"InternalRef\"\u003e2\u003c/span\u003e showed, H\u0026times;K, H\u0026times;D, K\u0026times;D, K\u0026times;H, D\u0026times;H, and D\u0026times;K crossbreds displayed higher AH compared to their purebred counterparts, indicating hybrid vigor. The superiority of crossbreds over purebreds has been supported by previous studies (Sola-Ojo and Ayorinde, \u003cspan citationid=\"CR32\" class=\"CitationRef\"\u003e2011\u003c/span\u003e; Sinha et al. \u003cspan citationid=\"CR31\" class=\"CitationRef\"\u003e2018\u003c/span\u003e; Kedija et al. \u003cspan citationid=\"CR19\" class=\"CitationRef\"\u003e2019\u003c/span\u003e). Purebred K and crossbred S\u0026times;D recorded the lowest AH, while H\u0026times;K had the narrowest AW, implying that certain combinations, particularly those involving K and S, may not be optimal for enhancing albumen quality. The H breed performed well in AH and AW when crossed with K and D, highlighting its potential to positively influence albumen quality in crossbreds. The S breed showed moderate performance in AH and AW, suggesting that while S is a robust breed overall, its contribution to albumen quality may be limited in specific crosses. Udoh et al. (\u003cspan citationid=\"CR36\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) reported results consistent with the present study regarding AH for the S, although their reported AW value was lower. K breed underperformed in AH, but showed promise in crosses such as K\u0026times;H. In contrast, Fikrineh et al. (\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) reported a slightly higher AH value for the K breed. However, the same author noted that K performed moderately relative to Fayoumi and White Leghorn but underperformed compared to its reciprocal cross with Fayoumi, aligning with the current findings.\u003c/p\u003e \u003cp\u003eThe purebred D and crossbred K\u0026times;S exhibited the highest YH, while D\u0026times;S and S demonstrated the widest YW. This suggests that both D and S breeds positively influence yolk size. In contrast, the crossbred H\u0026times;K had the smallest YH and YW, indicating that this combination may not be optimal for enhancing yolk characteristics. These findings align with Samrawit (\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2020\u003c/span\u003e), who reported similar values for the D in terms of YH and YWT, though the YWT value was lower than that reported by Tewodros et al. (\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2022\u003c/span\u003e). However, H showed mixed results, with some crosses (H\u0026times;D) performing well in YH but others (H\u0026times;K) underperforming. The K breed underperformed in YH and YW, particularly in its purebred form. The YH and YW values for the S breed in this study were higher than those reported by Yonas et al. (\u003cspan citationid=\"CR41\" class=\"CitationRef\"\u003e2019\u003c/span\u003e).\u003c/p\u003e \u003cp\u003eThe S breed consistently excelled in YWT and YR, both in purebred form and in crosses, while the H breed showed strong performance in YR, particularly in its purebred form. For YR, H, H\u0026times;S, and S achieved the highest percentages, indicating that both H and S positively influence yolk size and proportion. Similar findings were reported by Wolde et al. (\u003cspan citationid=\"CR39\" class=\"CitationRef\"\u003e2021\u003c/span\u003e), where S crossed with local chickens, S outperformed the local breed and their F1 cross in YWT, while the local breed and the F1 cross outperformed S in YR. The D breed underperformed in YWT and YR, particularly in crosses like H\u0026times;D and D\u0026times;K.Samrawit (\u003cspan citationid=\"CR28\" class=\"CitationRef\"\u003e2020\u003c/span\u003e) also noted that D underperformed in YR compared to H. Meanwhile, the K breed showed mixed results, with some crosses (K\u0026times;S) performing well in YWT but others (K\u0026times;D) underperforming. The crossbred K\u0026times;H had the highest HU, indicating superior egg quality, closely followed by D\u0026times;H and H\u0026times;K. This suggests that crosses involving H and K, or D can enhance egg quality. These results are consistent with Tewodros et al. (\u003cspan citationid=\"CR35\" class=\"CitationRef\"\u003e2022\u003c/span\u003e), who reported similar HU values for the D breed.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec16\" class=\"Section2\"\u003e \u003ch2\u003eCrossbreeding effects\u003c/h2\u003e \u003cdiv id=\"Sec17\" class=\"Section3\"\u003e \u003ch2\u003eHeterosis effect (H\u003csup\u003ee\u003c/sup\u003e)\u003c/h2\u003e \u003cp\u003eCrossbreeding is a commonly used approach in the poultry to improve favorable traits (Dzungwe et al. \u003cspan citationid=\"CR10\" class=\"CitationRef\"\u003e2024\u003c/span\u003e). Most crosses and their reciprocals exhibited positive H\u003csup\u003ee\u003c/sup\u003e for EWT, except for H\u0026times;S. However, its reciprocal, S\u0026times;H, displayed the highest positive H\u003csup\u003ee\u003c/sup\u003e. The negative H\u003csup\u003ee\u003c/sup\u003e observed in H\u0026times;S suggests that this particular cross may not be ideal for improving EWT. In agreement,Saadey et al. (\u003cspan citationid=\"CR27\" class=\"CitationRef\"\u003e2008\u003c/span\u003e) reported positive H\u003csup\u003ee\u003c/sup\u003e for EWT in a direct cross between White Leghorn and Rhode Island Red but negative H\u003csup\u003ee\u003c/sup\u003e in its reciprocal cross.\u003c/p\u003e \u003cp\u003eCrossbreds H\u0026times;S, H\u0026times;K, H\u0026times;D, K\u0026times;H, K\u0026times;S, and D\u0026times;K demonstrated positive H\u003csup\u003ee\u003c/sup\u003e values for EL and EW. Similarly, Atsbaha et al. (\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) reported positive H\u003csup\u003ee\u003c/sup\u003e for EL and EW. The occurrence of both positive and negative H\u003csup\u003ee\u003c/sup\u003e highlights the role of non-additive genetic effects in determining egg dimensions. Specific combinations, such as H\u0026times;K and K\u0026times;S, which exhibit positive H\u003csup\u003ee\u003c/sup\u003e, suggest that certain parental lines (H, K, S) could possess complementary alleles or genetic backgrounds that interact favorably to enhance egg size. These crosses are particularly promising for breeding programs focused on improving egg dimensions, as they demonstrate potential hybrid vigor. Negative H\u003csup\u003ee\u003c/sup\u003e was observed for SWT in all crosses except H\u0026times;K. Conversely, positive H\u003csup\u003ee\u003c/sup\u003e was noted ST in most crosses, except S\u0026times;K and K\u0026times;S. In agreement, Santosh and Raj (\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) reported the highest positive H\u003csup\u003ee\u003c/sup\u003e values for ST. Similarly, Atsbaha et al. (\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) also documented positive H\u003csup\u003ee\u003c/sup\u003e for ST. However, in contrast, the same authors reported positive H\u003csup\u003ee\u003c/sup\u003e for SWT. The positive H\u003csup\u003ee\u003c/sup\u003e for ST in most crosses is advantageous for enhancing shell quality, which is critical for reducing breakage and improving hatchability. The positive H\u003csup\u003ee\u003c/sup\u003e for AH and AW observed in most crosses indicates that crossbreeding can improve albumen quality, which is important for egg grading and consumer preference. These results are supported by Atsbaha et al. (\u003cspan citationid=\"CR4\" class=\"CitationRef\"\u003e2023\u003c/span\u003e), who reported positive H\u003csup\u003ee\u003c/sup\u003e values for AH.\u003c/p\u003e \u003cp\u003eFor YH, negative H\u003csup\u003ee\u003c/sup\u003e was observed in all crosses except K\u0026times;S. Similarly, for YW and YWT negative H\u003csup\u003ee\u003c/sup\u003e was observed in most crosses, except S\u0026times;D, K\u0026times;S, and D\u0026times;S. However, the positive H\u003csup\u003ee\u003c/sup\u003e in specific crosses indicates potential for targeted improvement. Most crosses showed positive H\u003csup\u003ee\u003c/sup\u003e for HU, but values were relatively low. Similarly, Khalil et al. (\u003cspan citationid=\"CR22\" class=\"CitationRef\"\u003e2013\u003c/span\u003e) also recorded negative H\u003csup\u003ee\u003c/sup\u003e for YWT. However, in contrast, the same author reported negative H\u003csup\u003ee\u003c/sup\u003e for HU. Conversely, Santosh and Raj (\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) reported positive H\u003csup\u003ee\u003c/sup\u003e for YWT in most crossbreds, which disagreed with the present study. The positive H\u003csup\u003ee\u003c/sup\u003e for HU is beneficial for improving egg quality, as HU is a key indicator of freshness.\u003c/p\u003e \u003c/div\u003e \u003c/div\u003e \u003cdiv id=\"Sec18\" class=\"Section2\"\u003e \u003ch2\u003eCombining ability\u003c/h2\u003e \u003cp\u003eGCA is a key metric in breeding programs, used to estimate the average performance of offspring based on the genetic potential inherited from that individual Adebambo et al. (\u003cspan citationid=\"CR1\" class=\"CitationRef\"\u003e2011\u003c/span\u003e). GCA effects were significant (P\u0026thinsp;\u0026lt;\u0026thinsp;0.001) across purebreds (Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e). The significance of GCA underscores the critical role of additive genetic effects in the inheritance of egg quality traits (Khalil et al. \u003cspan citationid=\"CR21\" class=\"CitationRef\"\u003e2018\u003c/span\u003e). S breed showed the highest positive GCA for EWT, YW, and YWT, making it a strong contributor to egg quality. K had positive GCA for EWT, EL, EW, ST and AW, indicating its potential for improving these traits, however the breed exhibited negative value for YWT, YW, and YR. In agreement, positive GCA value for K on EWT and EW was reported by Fikrineh et al. (\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2023\u003c/span\u003e). However, in contrast, the same author reported negative value for EL and ST and positive value for YWT.\u003c/p\u003e \u003cp\u003eSCA refers to situations where certain genetic combinations perform either better or worse than predicted based on the average performance of the parent lines involved (Emad et al. \u003cspan citationid=\"CR11\" class=\"CitationRef\"\u003e2017\u003c/span\u003e). It highlights the non-additive genetic interactions that influence the performance of specific crosses. Table\u0026nbsp;\u003cspan refid=\"Tab4\" class=\"InternalRef\"\u003e4\u003c/span\u003e indicated that variation due to SCA was high (P\u0026thinsp;\u0026lt;\u0026thinsp;0.001) for all traits. The S\u0026times;D cross exhibited the highest positive SCA for EWT, demonstrating significant heterosis. Meanwhile, the K\u0026times;D cross displayed the highest positive SCA for EL, EW, and AW, indicating its potential to enhance these characteristics. The H\u0026times;K cross showed the highest positive SCA for ST and YR, positioning it as a promising combination for improving shell and yolk quality. Similar findings were reported by Singh et al. (\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) and Santosh and Raj (\u003cspan citationid=\"CR29\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) in their studies.\u003c/p\u003e \u003cp\u003eRE showed significant variations for various traits, except EL, YW, and YWT. K\u0026times;S showed the highest positive RE for EWT and AW, indicating that using K as the maternal parent and S as the paternal parent improves EWT and AW. D\u0026times;S cross had the highest positive RE for ST, suggesting that using D as the maternal parent and S as the paternal parent improves ST. Crosses like S\u0026times;D and K\u0026times;D should be prioritized for improving EWT and EL, while H\u0026times;K can be used to enhance ST. The direction of crossing should be considered to optimize egg quality traits, as seen in the positive RE for K\u0026times;S and D\u0026times;S. Similarly, Fikrineh et al. (\u003cspan citationid=\"CR12\" class=\"CitationRef\"\u003e2023\u003c/span\u003e) reported, except for EL, other studied traits aligned with this study showed variation in RE. Singh et al. (\u003cspan citationid=\"CR30\" class=\"CitationRef\"\u003e2016\u003c/span\u003e) also reported significant RE for EWT.\u003c/p\u003e \u003cp\u003eThe GCA/SCA ratio is used to assess the relative importance of additive genetic effects versus non-additive genetic effects in the expression of a trait. A high GCA/SCA ratio suggests that additive genetic effects are more important, while a low ratio indicates that non-additive genetic effects (dominance or epistasis) play a more significant role. Traits like EWT (0.681) and YWT (0.345) were primarily influenced by additive genetic effects, suggesting selection-based breeding (such as recurrent selection) would be more effective. Conversely, traits like EL (0.044), EW (0.016), ST (0.002), and YR (0.158) were dominated by non-additive effects, making hybrid breeding or specific crosses to exploit heterosis more suitable. For traits with moderate ratios like AW (0.259) and YW (0.227), a combined approach of selection and hybrid breeding is recommended.\u003c/p\u003e \u003c/div\u003e \u003cdiv id=\"Sec19\" class=\"Section2\"\u003e \u003ch2\u003eMaternal effect (M\u003csup\u003ee\u003c/sup\u003e)\u003c/h2\u003e \u003cp\u003eThe M\u003csup\u003ee\u003c/sup\u003e exhibited significant variation across most traits Table\u0026nbsp;\u003cspan refid=\"Tab5\" class=\"InternalRef\"\u003e5\u003c/span\u003e. Significant M\u003csup\u003ee\u003c/sup\u003e observed on EWT in crosses H\u0026times;K, H\u0026times;S, H\u0026times;D, S\u0026times;D, and K\u0026times;D (P\u0026thinsp;\u0026lt;\u0026thinsp;0.05). Notably, H\u0026times;S displayed a strong negative M\u003csup\u003ee\u003c/sup\u003e, whereas H\u0026times;D exhibited a positive effect. This highlights the substantial impact of maternal breed selection on EWT. Crossbreeding also significantly influenced traits such as AW and YWT, with H\u0026times;D showing positive M\u003csup\u003ee\u003c/sup\u003e for EWT and AW, and H\u0026times;S for YWT, reflecting enhanced performance. However, negative M\u003csup\u003ee\u003c/sup\u003e were observed in H\u0026times;S, S\u0026times;D, and K\u0026times;D for EWT, as well as in K\u0026times;D for AW, indicating these combinations may be less favorable. Traits like EL and SWT showed no significant M\u003csup\u003ee\u003c/sup\u003e, suggesting limited crossbreeding influence. These findings underscore the need for strategic crossbreeding to harness heterosis and improve key traits, while avoiding combinations with detrimental M\u003csup\u003ee\u003c/sup\u003e in targeted breeding programs.\u003c/p\u003e \u003c/div\u003e"},{"header":"Declarations","content":"\u003cp\u003e\u003cstrong\u003eAcknowledgements\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eMany thanks to Werer Agricultural Research Center (WARC), for allowing the poultry farm and providing the necessary facilities for this experiment. The Animal and Human Health Program of the International Livestock Research Institute (ILRI) and Ethiopian Institute of Agricultural Research (EIAR) are highly appreciated for financing the study.\u0026nbsp;\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eAuthor contributions\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003ePhilimon Teshome was responsible for the study\u0026apos;s design, data collection, analysis, and interpretation. He also wrote the first draft of the manuscript. All authors reviewed, provided feedback on earlier versions, and approved the final manuscript.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eFunding\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThis work was manly funded by the International Livestock Research Institute (ILRI), Ethiopia, under the Animal and Human Health Program (grant number: 04/GF/23/6443, January 2023). Under the grant conditions of the Foundation, a Creative Commons Attribution 4.0 Generic License has already been assigned to the Author Accepted Manuscript version that might arise from this submission. The findings and conclusions contained within are those of the authors and do not necessarily reflect the positions or policies of ILRI.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eData availability\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eData associated with this article are included in the manuscript but are available from the corresponding author on reasonable request.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eEthics approval\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe protocol for the conduct of animal experiment was approved by the animal research ethical review committee of Collage of Veterinary Medicine and Agriculture of Addis Ababa University under the certificate reference number VM/ERC/01/06/15/2024. The birds were treated humanely in accordance with all international, national, and institutional guidelines for the care of animals used in research and teaching.\u003c/p\u003e\n\u003cp\u003e\u003cstrong\u003eCompeting interests\u0026nbsp;\u003c/strong\u003e\u003c/p\u003e\n\u003cp\u003eThe authors declare that they have no known competing interests that could have appeared to influence the work reported in this paper.\u003c/p\u003e"},{"header":"References","content":"\u003col\u003e\n\u003cli\u003eAdebambo, A. O., Ikeobi, C. O. N., Ozoje, M. O., Oduguwa, O. O., and Adebambo Olufunmilayo, A. (2011). Combining abilities of growth traits among pure and crossbred meat type chickens. Arch. Zootec., 60(232), 953\u0026ndash;963. https://doi.org/10.4321/S0004-05922011000400012 \u003c/li\u003e\n\u003cli\u003eAdemola, A. A., Fayeye, T. R., Akintunde, A. O., and Chimezie, V. O. (2023). 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Sci., 8(1), 1\u0026ndash;9.\u003c/li\u003e\n\u003cli\u003eKefelegn, K., and Ashenafi, G. M. (2024). Differentiation of exotic chicken breeds based on egg quality traits using classification tree algorithm. J. Food Technol. Nutr. Sci., 6(4), 1\u0026ndash;9.\u003c/li\u003e\n\u003cli\u003eKhalil, M. H., Debes, A. A., and Shebl, M. K. (2018). Estimation of heterosis, combining ability and reciprocal effects for growth traits in chickens from a full diallel cross. Int. J. Res. Agric. Sci., 5(6), 291\u0026ndash;295.\u003c/li\u003e\n\u003cli\u003eKhalil, M. H., Iraqi, M. M., and El-Atrouny, M. M. (2013). Effects on egg quality traits of crossing Egyptian Golden Montazah with White Leghorn chickens. Livest. Res. Rural Dev., 25(6).\u003c/li\u003e\n\u003cli\u003eMokoena, K., Mbazima, V., and Tyasi, T. L. (2024). A literature review on effect of chicken crossbreeding on egg production traits. World\u0026rsquo;s Poult. Sci. J., 1\u0026ndash;17. https://doi.org/10.1080/00439339.2024.2443184 \u003c/li\u003e\n\u003cli\u003eMorris, S. S., Beesabathuni, K., and Headey, D. (2018). An egg for everyone: Pathways to universal access to one of nature\u0026rsquo;s most nutritious foods. Matern. Child Nutr., 14(S3). https://doi.org/10.1111/mcn.12679 \u003c/li\u003e\n\u003cli\u003eOmer, A., Hailu, D., and Whiting, S. J. (2022). Egg consumption of children under two years of age through a child-owned poultry and nutrition intervention in rural Ethiopia: A community-based randomized controlled trial. J. Agric. Food Res., 9, 100354. https://doi.org/10.1016/j.jafr.2022.100354 \u003c/li\u003e\n\u003cli\u003eR Core Team. (2023). R: A language and environment for statistical computing (4.4.2). R Foundation for Statistical Computing. https://www.R-project.org/ \u003c/li\u003e\n\u003cli\u003eSaadey, S., Mekky, A., Galal, Z. H. I., and El-Dein, A. Z. (2008). 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Indian J. Poult. Sci., 51(2), 132. https://doi.org/10.5958/0974-8180.2016.00038.6 \u003c/li\u003e\n\u003cli\u003eSinha, B., Mandal, K., Kumari, R., and Kumari, T. (2018). Estimate and effect of breeds on egg quality traits of poultry: A Review. Int. J. Livest. Res., 1. https://doi.org/10.5455/ijlr.20170812102444 \u003c/li\u003e\n\u003cli\u003eSola-Ojo, F. E., and Ayorinde, K. L. (2011). Evaluation of Reproductive Performance and Egg quality Traits in Progenies of Dominant Black Strain Crossed with Fulani Ecotype Chicken. Journal of Agricultural Science, 3(1). https://doi.org/10.5539/jas.v3n1p258 \u003c/li\u003e\n\u003cli\u003eTadelle, D. (2003). Phenotypic and genetic characterization of chicken ecotypes in Ethiopia. [Ph.D. Thesis]. Humboldt University.\u003c/li\u003e\n\u003cli\u003eTadelle, D., Alemu, Y., and Peters, K. J. (2000). Indigenous chickens in Ethiopia: genetic potential and attempts at improvement. World\u0026rsquo;s Poult. Sci. J., 56(1), 45\u0026ndash;54. https://doi.org/10.1079/WPS20000005 \u003c/li\u003e\n\u003cli\u003eTewodros, F., Gebreyohannes, B., Mammo, M., and Misba, A. (2022). Effect of dietary metabolizable energy and crude protein on the laying performance, egg quality, hatchability, and fertility of DZ-White chickens. Int. J. Livest. Res., 12(7).\u003c/li\u003e\n\u003cli\u003eUdoh, J. E., Udoh, U. H., and Adeoye, A. A. (2020). Comparative analysis of fertile egg quality traits in exotic, two Nigerian ecotype chickens and their crosses. Niger. J. Anim. Sci., 22(1), 50\u0026ndash;59.\u003c/li\u003e\n\u003cli\u003eWearden, S., Tindell, D., and Craig, J. V. (1965). Use of a full diallel cross to estimate general and specific combining ability in chickens. Poult. Sci., 44(4), 1043\u0026ndash;1053. https://doi.org/10.3382/ps.0441043 \u003c/li\u003e\n\u003cli\u003eWilliams, S., Price, S., and Siegel, P. (2002). Heterosis of growth and reproductive traits in fowl. Poult. Sci., 81(8), 1109\u0026ndash;1112. https://doi.org/10.1093/ps/81.8.1109 \u003c/li\u003e\n\u003cli\u003eWolde, S., Mirkena, T., Melesse, A., Dessie, T., and Abegaz, S. (2021). Egg production and quality traits of Sasso-RIR, Normal Feathered local and their F1 cross chickens managed under on-station condition in Southern Ethiopia. Trop. Subtrop. Agroecosyst., 24(3). https://doi.org/10.56369/tsaes.3828 \u003c/li\u003e\n\u003cli\u003eYesihak, M., Ashenafi, G., Kefelegn, K., and Mengistu Urge. (2024). Comparative evaluation of egg quality traits in two exotic chicken breeds. J. Food Chem. Nanotechnol., 10(1). https://doi.org/10.17756/jfcn.2024-169 \u003c/li\u003e\n\u003cli\u003eYonas, K., Sandip, B., and Mestawet, T. (2019). Some internal and external egg quality characteristics of local and exotic chickens reared in Yirgalem and Hawassa towns, Ethiopia. Int. J. Livest. Prod., 10(5), 135\u0026ndash;142. https://doi.org/10.5897/IJLP2018.0547 \u003c/li\u003e\n\u003cli\u003eZhang, X., Li, Y., Li, Q., Zhang, T., Sun, Y., Shi, F., and Chen, J. (2024). Research Note: Genetic parameters estimation of egg quality traits in Rhode Island Red and White Leghorn chickens. Poult. Sci., 103(12), 104263. https://doi.org/10.1016/j.psj.2024.104263 \u003c/li\u003e\n\u003c/ol\u003e"},{"header":"Table 5","content":"\u003cp\u003eTable 5 is available in the Supplementary Files section.\u003c/p\u003e\n"}],"fulltextSource":"","fullText":"","funders":[],"hasAdminPriorityOnWorkflow":false,"hasManuscriptDocX":true,"hasOptedInToPreprint":true,"hasPassedJournalQc":"","hasAnyPriority":false,"hideJournal":false,"highlight":"","institution":"","isAcceptedByJournal":true,"isAuthorSuppliedPdf":false,"isDeskRejected":"","isHiddenFromSearch":false,"isInQc":false,"isInWorkflow":true,"isPdf":false,"isPdfUpToDate":true,"isWithdrawnOrRetracted":false,"journal":{"display":true,"email":"[email protected]","identity":"tropical-animal-health-and-production","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"trop","sideBox":"Learn more about [Tropical Animal Health and Production](https://www.springer.com/journal/11250)","snPcode":"11250","submissionUrl":"https://submission.nature.com/new-submission/11250/3","title":"Tropical Animal Health and Production","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false},"keywords":"Heterosis effect, Combining ability, Reciprocal effect, Maternal effects, Diallel cross","lastPublishedDoi":"10.21203/rs.3.rs-6173323/v1","lastPublishedDoiUrl":"https://doi.org/10.21203/rs.3.rs-6173323/v1","license":{"name":"CC BY 4.0","url":"https://creativecommons.org/licenses/by/4.0/"},"manuscriptAbstract":"\u003cp\u003eThis study aimed to estimate heterosis effects (\u003cb\u003eH\u003c/b\u003e\u003csup\u003e\u003cb\u003ee\u003c/b\u003e\u003c/sup\u003e), general combining ability (\u003cb\u003eGCA\u003c/b\u003e), specific combining ability (\u003cb\u003eSCA\u003c/b\u003e), reciprocal effects (\u003cb\u003eRE\u003c/b\u003e), and maternal effects (\u003cb\u003eM\u003c/b\u003e\u003csup\u003e\u003cb\u003ee\u003c/b\u003e\u003c/sup\u003e) for egg quality traits in a 4\u0026times;4 diallel cross involving four chicken breeds: Improved Horro (\u003cb\u003eH\u003c/b\u003e), Sasso (\u003cb\u003eS\u003c/b\u003e), Potchefstroom Koekoek (\u003cb\u003eK\u003c/b\u003e), and Dz-white feathered (\u003cb\u003eD\u003c/b\u003e). A total of 480 eggs from 4 purebreds and 12 crossbreds were evaluated for traits like egg weight (\u003cb\u003eEWT\u003c/b\u003e), egg length (\u003cb\u003eEL\u003c/b\u003e), egg width (\u003cb\u003eEW\u003c/b\u003e), shell weight (\u003cb\u003eSWT\u003c/b\u003e), shell thickness (\u003cb\u003eST\u003c/b\u003e), shell ratio (\u003cb\u003eSR\u003c/b\u003e), albumen height (\u003cb\u003eAH\u003c/b\u003e), albumen width (\u003cb\u003eAW\u003c/b\u003e), yolk height (\u003cb\u003eYH\u003c/b\u003e), yolk width (\u003cb\u003eYW\u003c/b\u003e), yolk weight (\u003cb\u003eYWT\u003c/b\u003e), yolk ratio (\u003cb\u003eYR\u003c/b\u003e), and hugh unit (\u003cb\u003eHU\u003c/b\u003e). Results showed positive H\u003csup\u003ee\u003c/sup\u003e for most traits. GCA were significant (P\u0026thinsp;\u0026lt;\u0026thinsp;0.001) for EWT, EL, EW, AW, YW, YWT, and YR. Variation due to SCA was high (P\u0026thinsp;\u0026lt;\u0026thinsp;0.001) for all traits. K\u0026times;S cross showed positive RE for EWT and AW. D\u0026times;S cross had the highest positive RE for ST. Both additive and non-additive gene actions play significant roles in controlling egg quality traits. EWT and YWT were primarily influenced by additive genetic effects, while EL, EW, ST, and YR were dominated by non-additive effects. M\u003csup\u003ee\u003c/sup\u003e exhibited significant variation across genotypes for most traits. In conclusion, breeding strategies should be tailored to the genetic basis of each trait, focusing on economically important traits and using molecular tools for efficiency.\u003c/p\u003e","manuscriptTitle":"Genetic parameters estimation for egg quality traits of four chicken breeds using a full diallel cross","msid":"","msnumber":"","nonDraftVersions":[{"code":1,"date":"2025-04-16 16:30:30","doi":"10.21203/rs.3.rs-6173323/v1","editorialEvents":[{"type":"communityComments","content":0},{"type":"reviewerAgreed","content":"","date":"2025-04-08T06:43:14+00:00","index":0,"fulltext":""},{"type":"reviewersInvited","content":"","date":"2025-03-27T20:34:38+00:00","index":"","fulltext":""},{"type":"editorAssigned","content":"","date":"2025-03-12T04:09:16+00:00","index":"","fulltext":""},{"type":"submitted","content":"Tropical Animal Health and Production","date":"2025-03-07T10:14:36+00:00","index":"","fulltext":""}],"status":"published","journal":{"display":true,"email":"[email protected]","identity":"tropical-animal-health-and-production","isNatureJournal":false,"hasQc":true,"allowDirectSubmit":false,"externalIdentity":"trop","sideBox":"Learn more about [Tropical Animal Health and Production](https://www.springer.com/journal/11250)","snPcode":"11250","submissionUrl":"https://submission.nature.com/new-submission/11250/3","title":"Tropical Animal Health and Production","twitterHandle":"","acdcEnabled":true,"dfaEnabled":true,"editorialSystem":"em","reportingPortfolio":"Springer Hybrid","inReviewEnabled":true,"inReviewRevisionsEnabled":false}}],"origin":"","ownerIdentity":"0974471c-b6f5-4786-ac27-6fad5b1d1240","owner":[],"postedDate":"April 16th, 2025","published":true,"recentEditorialEvents":[],"rejectedJournal":[],"revision":"","amendment":"","status":"under-review","subjectAreas":[],"tags":[],"updatedAt":"2025-07-30T14:23:08+00:00","versionOfRecord":[],"versionCreatedAt":"2025-04-16 16:30:30","video":"","vorDoi":"","vorDoiUrl":"","workflowStages":[]},"version":"v1","identity":"rs-6173323","journalConfig":"researchsquare"},"__N_SSP":true},"page":"/article/[identity]/[[...version]]","query":{"redirect":"/article/rs-6173323","identity":"rs-6173323","version":["v1"]},"buildId":"8U1c8b4HqxoKbykW_rLl7","isFallback":false,"isExperimentalCompile":false,"dynamicIds":[84888],"gssp":true,"scriptLoader":[]}