Heritability and correlation coefficient analysis for yield and its components in rice (Oryza sativa L.)
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1 African Journal of Plant Science Vol. 5 (3), pp , March 2011 Available online at ISSN Academic Journals Full Length Research Paper Heritability and correlation coefficient analysis for yield and its components in rice (Oryza sativa L.) M. G. Akinwale 1, G. Gregorio 2, F. Nwilene 1, B. O. Akinyele 4 *, S. A. Ogunbayo 3 and A. C. Odiyi 4 1 Africa Rice Center, Ibadan, Oyo State, Nigeria. 2 International Rice Research Institute (IRRI), Philippines. 3 Africa Rice Center, Cotonou, Benin Republic. 4 Federal University of Technology, Akure, Ondo State, Nigeria. Accepted 8 February, 2011 The objectives of this study were to estimate the phenotypic and genotypic coefficients of variation, broad sense heritability, genetic gain and correlations in rice (Oryza sativa L.). The experiment was laid out in a randomized complete block design (RCBD) with three replications in the International Institute of Tropical Agriculture, Ibadan, Nigeria during 2008/2009 cropping season. Genotypes differed significantly at (p>0.001) for all the traits studied, which implies that the genotypes constitute a pool of germplasm with adequate genetic variability. Genotypic coefficients of variation were lower than the corresponding phenotypic coefficients in all the traits studied, indicating considerable influence of the environment on the expression of the traits. High to medium broad sense heritability estimates observed on days to heading, days to maturity, plant height, grain yield and number of grains per panicle, panicle weight, number of panicles per m 2 and panicle length suggests high component of heritable portion of variation, which is the portion exploited by breeder and that selection for these traits can be achieved directly based on their phenotypic performance. The low broad sense heritability observed for the number of tillers per plant and 1000 grain weight is indicative of the influence of the environment on these traits. Low heritability of these traits indicates the ineffectiveness of direct selection for these traits. High to medium heritability and genetic advance were recorded for the number of grains per panicle, grain yield, panicle weight and the number of panicles per plant. This suggests that these traits are primarily under genetic control and selection for them can be achieved through their phenotypic performance. Grain yield exhibited significantly positive correlation with the number of tillers per plant (r = 0.58**), panicle weight (r =0.60*) and number of grains per panicle (r= 0.52*). Therefore, the results suggest that these traits can be used for grain yield selection. Key words: Broad sense heritability, rice, genotypic coefficient of variation, phenotypic coefficient of variation, genetic gain, rice yield, yield components. INTRODUCTION The world population is expected to reach 8 billion by 2030 and rice production must be increased by 50% in order to meet the growing demand (Khush and Brar, 2002). Hence, rice breeders are interested in developing cultivars with improved yield and other desirable agronomic characters. Genetic variability for agronomic traits is the key component of breeding programmes for *Corresponding author. [email protected]. broadening the gene pool of rice. Plant breeders commonly select for yield components which indirectly increase yield. Heritability (h 2 ) of a trait is important in determining its response to selection. It was found out earlier that genetic improvement of plants for quantitative traits requires reliable estimates of heritability in order to plan an efficient breeding program. El-Malky et al. (2003) observed high broad sense heritability estimate of 98.89% for days to maturity, 75.20% for the number of tillers per plant, 41.74% for the number of panicles per plant, 98.97% for 1000 grain weight and 90.87% for
2 208 Afr. J. Plant Sci. panicle weight. Babar et al. (2007) also reported high heritability of 0.74 for panicle length, 0.75 for plant height, 0.63 for the number of panicles per plant, 0.64 for days to heading and 0.81 for grain yield. On the other hand, Kato (1997) estimated low broad sense heritability of 0.16 for the number of panicles per plant and 0.20 to 0.33 for number of spikelet per panicle. Sürek and Korkut (1998) estimated high narrow sense heritability for grain weight, moderate for the number of spikelets per panicle and low for the number of panicles per plant. The relationship between rice yield and yield components has been studied extensively at phenotypic level; Sharma and Choubey (1985) and Dhanraj and Jagadish (1987) reported that yield per plant was positively correlated with the number of productive tillers per plant, the number of panicles per plant and spikelets per plant and 1000 grain weight while Prasad et al. (1988) observed positive correlations between grain yield per plant and the number of spikelets per panicle, the number of fertile grains per panicle and 1000 grain weight. Bai et al. (1992) reported that grain yield is positively correlated with the number of productive tillers per plant, and the number of grains per panicle. Sürek and korkut (1998) reported that grain yield per plant was significantly correlated with the number of panicles per plant and 1000 grain weight. Yield component breeding to increase grain yield would be most effective, if the components involved are highly heritable and genetically independent or positively correlated with grain yield. However, it is very difficult to judge whether observed variability is highly heritable or not. Moreover, knowledge of heritability is essential for selection based improvement as it indicates the extent of transmissibility of a character into future generations (Sabesan et al., 2009). Therefore, the objectives of this study were (i) to estimate the heritability of yield and yield components, and (ii) to compute the correlation of grain yield with yield components among the rice genotypes. MATERIALS AND METHODS Twenty rice genotypes were laid out in a randomized complete block design with three replications in the International Institute of Tropical Agriculture, Ibadan, Nigeria during 2008/2009 cropping season. Seeds sown in the wet nursery were transplanted to the puddled field after 21 days. Two seedlings were transplanted per each hill at a spacing of 20 cm between rows and between hills in 12 rows of 6 m 2. Inorganic fertilizer (NPK ) was applied in three split applications, a basal of 200 kg/ha before transplanting and top dressed with urea at the rate of 65 kg/ha at the tillering stage and at the rate of 35 kg/ha at booting stage. Approximately 5 cm of standing water was maintained in the field until drainage before harvest. Weeds were controlled by application of postemergence herbicide (Oritzoplus) 14 days after transplanting (DAT) and hand weeding. Data collection In all the plots, Data were collected for plants for days to 50% flowering, days to maturity, plant height, number of tillers per plant, number of panicles per plant, panicle length, 1000 grain weight, panicle weight, number of grains per panicle, and grain yield. Statistical analysis Analysis of variance (ANOVA) was carried out on the data to assess the genotypic effects and their interaction using general linear model (GLM) procedure for randomized complete blocks design in SAS (9.2 version). Estimates of variance components were generated. Broad-sense heritability (h 2 ) was calculated as the ratio of the genotypic variance to the phenotypic variance using the formula according to Allard (1960): h 2 = Ơ 2 g/ Ơ 2 ph x 100 where h 2 = broad sense heritability (%), Ơ 2 g = genotypic variance and Ơ 2 ph = phenotypic variance. Genetic advance was calculated at 20% selection intensity (i=1.4). Phenotypic coefficients of correlation were computed using Pearson s linear correlation outlined by Steel and Torrie (1984). RESULTS AND DISCUSSION The analysis of variance revealed significant variations among the genotypes for all the character examined (Table 1). The phenotypic variance was partitioned into genotypic and environmental variances for a clear understanding of the pattern of variation. Number of grains per panicle exhibited the highest genotypic variance (322) and also the highest environmental variance (139). However, panicle weight had the lowest magnitude of genotypic, environmental and phenotypic variance of 0.15, 0.1 and 0.2 respectively. Genotypic coefficient of variation measures the variability of any character. The extent of the environmental influence on any character is indicated by the magnitude of the differences between the genotypic and phenotypic coefficients of variation. Large differences reflect high environmental influence, while small differences reveal high genetic influence. Phenotypic coefficients of variation were slightly higher than the genotypic coefficients of variation for all the traits studied. This indicates the presence of environmental influence to some degree in the phenotypic expression of the characters. Iftekharuddeula et al. (2001) observed similar result. Moderate to low genotypic and phenotypic coefficients of variation obtained for days to heading, days to maturity, plant height and panicle length may be due to the presence of both positive and negative alleles in the population. The findings were supported by Iftekharuddeula et al. (2001). They obtained moderate genotypic and phenotypic coefficients of variation for plant height, days to maturity and panicle length when nineteen rice hybrids were evaluated. The small differences observed between genotypic and phenotypic
3 Akinwale et al. 209 Table 1. Genetic parameters of variation for agronomic characters. Traits Mean square 2 g S 2 e S 2 P GCV% PCV% h 2 B GA heading 43.98*** maturity 46.12*** Plant height (cm) *** panicles/ plant 11.63*** length(cm) 6.13*** tillers /plant 3.08* weight(gm) 0.52* Grains/panicle *** grain weight 16.51* Yield(t/ha) 0.85* Genetic variance(s 2 g), error variance(s 2 e), phenotypic variance (S 2 P),genotypic coefficient of variation (GCV), phenotypic coefficient of variation (PCV), broad sense heritability(h2b) and genetic advance (GA)*, ** and *** significant P= 0.05, P=0.01, and P= respectively. coefficients of variation on days to heading, days to maturity, plant height and panicle length indicate the presence of sufficient genetic variability for the traits which may facilitate selection (Yadav, 2000). The high to moderate differences between genotypic and phenotypic coefficients of variation observed in number of panicles per square meter, number of tillers per plant, panicle weight, number of grains per panicle, 1000 grain weight and grain yield indicate high influence of the environment on the traits. The broad sense heritability is the relative magnitude of genotypic and phenotypic variances for the traits and it is used as a predictive role in selection procedures (Allard, 1960). This gives an idea of the total variation ascribable to genotypic effects, which are exploitable portion of variation. The heritability estimates obtained for the traits studied ranged between 19.2 to 95.1% (Table 1). High to medium broad sense heritability estimates observed for days to heading, days to maturity, plant height, grain yield and the number of grains per panicle, panicle weight, the number of panicles per square meter and panicle length. It suggests high component of heritable portion of variation, it is the portion which is exploited by breeders. On the other hand, Bhatti et al. (1998) reported high heritability for the number of spikelets per panicle, 1000 grain weight and the number of panicles per plant in rice. The low broad sense heritability observed for the number of tillers per plant and 1000 grain weight indicates the influence of the environment on these traits. The low heritability recorded for these traits indicates that direct selection for these traits will be ineffective. Since high heritability do not always indicate high genetic gain, heritability with genetic advance considered together should be used in predicting the ultimate effect for selecting superior varieties (Ali et al., 2002). High to medium heritability and genetic advance were recorded for the number of grains per panicle, grain yield, panicle weight and the number of panicles per plant. This suggests that these traits are primarily under genetic control and selection for them can be achieved through their phenotypic performance. High heritability estimates with low genetic advance observed for days to heading, days to maturity, plant height and panicle length indicates non additive type of gene action and that genotype x environment interaction plays a significant role in the expression of the traits. The results further revealed that most of the characters exhibited wide range of Variability (Table 2). 50% flowering shows significant differences among genotypes at (p < 0.001) with genotype IR B-93-B-3 taking the shortest time (74.33 days) to attained 50% flowering while genotype PSBRC 14 took the longest period (91.66 days). maturity follows the same trend with days to 50% flowering which ranges between 104 and days with genotype IR B-93-B-3 taking the shortest period (104 days) to mature while genotype PSBRC 14 takes the longest period ( days) to mature. Plant height shows significant difference (p<0.001) among the genotypes at maturity. Genotype NERICA -L-40 is significantly taller with a height of cm while genotype IR is the shortest (86.06 cm). Number of tillers per plant varied significantly among the genotypes. The number of tillers per plant ranges between 9.50 and with genotypes IR , IR B , IR B and NERICA -L-2 recording the highest number (12.25). The number of panicles per m2 is significantly affected by genotypes. It varies from 6.06 in IR B-44-4 to in genotype IR Number of grains per panicle is one of the most important components of yield and probably this character will be helpful in breaking the yield plateau. Number of grains per panicle differs significantly among the tested genotypes. The highest grain number (164.88) is recorded in genotype IR followed by FARO 37 (ITA 306) (154.66). The highest grain yield (5.16t/ha) is observed in IR (p<0.001) while the lowest grain yield (2.83 t/ha) is observed in genotype
4 210 Afr. J. Plant Sci. Table 2. Mean performance of 20 rice genotypes for 10 agronomic traits. Designation Heading maturity Plant height(cm) tillers/plant panicles/ plant length(cm) weight (g) grains/pa nicle 1000grain weight (g) Grain yield at 14% t/ha FARO 35 (ITA 212) 88.33dc ab abcdef 9.58defg 10.66bcdef 26.26ab 1.76fgh def 25.67bc 2.93f FARO 37 (ITA 306) 87.00de cde 95.33fghij 11.25def 9.16efg 24.90abcde 2.92ab ab 25.93bc 4.20bcde IR B 87.66d 118bcd 91.83ghijkl 9.58defg 12.25abcd 24.23bcdef 2.91ab ed 29.86abc 3.97cde IR B fg efg 97.2defghi 10.25gh 12.32abc 23.10defg 2.29cde abc 31.66ab 4.42bc IR B fgh fgh 90.83ijkl 11.42ab 12.50abc 24.10bcdef 2.76abc fg 25.00c 4.70ab IR B fg 116def abcd 12.25a 10.90bcdef 23.60dcef 2.06defg f 34.66a 4.05cde IR B i 114fgh abcde 11.33ab 9.25efg 24.96abc 2.46abcd ef 28.33bc 4.02cde IR B ghi gh 99.26bcdefg 10.58ab 8.91fg 26.66a 2.03defg cde 28.33bc 4.11cde IR B ij hi 93.96fghij 12.25a 11.91bcde 22.23fg 2.37cde def 29.66abc 3.84cde IR B j 111i 94.30fghij 9.67def 11.17bcdef 26.06ab 2.16def 131cde 29.33abc 3.92cde IR ab a 90.33ijkl 12.25a 10.11bcdefg 24.33bcdef 2.23def bcd 27.00bc 4.39bc IR B-93-B k 104j ab 9.75abc 9.75cdefg 21.10g 1.43h 85.67g 26.00bc 2.83f IR B de 117cde abcde 9.50abcd 6.08h 24.00bcdef 2.43bcde cde 25.66bc 3.65de IR B ef 116def 98.26cdefgh 10.25ab 10.66bcdef 23.66dcef 2.49abcd abc 26.33bc 4.24bcd IR ef fgh 88.20jkl 10.91ab 10.25bcdef 24.43abcdef 2.44bcde def 28.66bc 3.86cde IR j 111i 96.06efhhij 10.67ab 11.58bcdef 22.43fg 2.95a ef 27.00bc 3.68de IR bc 119abc 86.06l 12.00a 14.70a 25.63abc 2.96a a 29.33abc 5.16a NERICA -L i hi 90.23ijkl 12.25a 12.75ab 24.23bcdef 2.39cde ef 28.67bc 4.29bc NERICA -L hi 113hi hi 11.75ab 9.36defg 22.73efg 2.28cde edf 27.66bc 4.34bc PSBRC a a 86.63a 10.25ab 7.41h 25.26bcda 2.29cde cde 30.00abc 3.60e Means SE CV R Pr > F * * * * * * * * * * * * * * * * * * * * * * * * * * * = significant at P<0.05; ** = significant at P<0.01; *** = significant at P<0.001; ns = non significant. IR B-93-B-3. The high grain yield performance of genotype IR could be attributed to the high number of grains per panicle, high panicle weight and large number of panicles per plant. Correlation The degree of correlation among the characters is an important factor especially in economic and complex character as yield. Steel and Torrie (1984) stated that correlations are measures of the intensity of association between traits. The selection for one trait results in progress for all characters that are positively correlated and retrogress for traits that are negatively correlated.
5 Akinwale et al. 211 Table 3. The phenotypic correlations among the traits. Traits heading maturity Plant height (cm) panicles/plant length tillers/plant weight grains/panicle 1000 grain weight heading 1 maturity 0.98*** 1 Plant Height (cm) -0.49* -0.38ns 1 s/plant -0.06ns -0.06ns -0.44* 1 length(cm) 0.55* 0.55* -0.32ns -0.07ns 1 No. of tillers/plant 0.12ns 0.08ns -0.23ns 0.35ns 0.13ns 1 weight(gm) 0.38* 0.34ns -0.50* 0.32ns 0.01ns 0.23ns 1 grains/panicle. 0.60** 0.58** -0.39ns 0.1ns 0.41ns 0.05ns 0.53* Gwt 0.10ns 0.10ns -0.11ns 0.26ns 0.01ns 0.26ns 0.25ns 0.07ns 1 Yield(t/ha) 0.37ns 0.32ns -0.42ns 0.47ns 0.18ns 0.58** 0.60* 0.52* 0.20ns 1 * = significant at P<0.05; ** = significant at P<0.01; *** = significant at P<0.001; ns = non significant. Grain yield at 14% t/ha The result of correlation analysis as shown by their coefficients of correlation (Table 3) reveals that grain yield exhibits significantly positive correlation with the number of tillers per plant (r = 0.58**), panicle weight (r =0.60*) and the number of grains per panicle (r= 0.52*). The results are in conformity with Mohammed.et al. (2007) for the number of panicles per plant and Ramakrishman et al. (2006) for number of grains per panicle. The number of grains per panicle shows significantly positive correlation with days to 50% heading (r=o.60**), days to maturity (r=0.58**) and panicle weight (0.53*). weight correlated positively with days to heading (r=0.38*) and negatively correlated with plan height (r=0.50*). Conclusion The overall results indicated that there is adequate genetic variability present in the material studied. The broad sense heritability, genetic advance and correlation analysis of the study revealed that the number of panicles per plant, panicle weight and the number of grains per panicle were the most important yield components. These characters also showed moderate to high heritability and genetic advance in percentage of mean. Therefore, the results suggest that the number of panicles per plant, panicle weight and the number of grains per panicle are important yield contributing traits and selection based on these traits would be most effective. REFERENCES Ali A, Khan S, Asad MA (2002). Drought tolerance in wheat: Genetic variation and heritability for growth and ion relations. Asian J. Plant Sci., 1: Allard RW (1960). Principles of Plant Breeding. John Wiley and Sons, Inc. New York, p Bai NR, Devika R, Regina A, Joseph CA (1992). Correlation of yield and yield components in medium duration rice cultivars. Environ. Ecol., 10: Bhatti MA, Khan AM, Sadagat H A, Khan AA (1998). Path coefficient analysis in coarse rice. Anim. Plant Sci., 8: Dhanraj A, Jagadish CA (1987). Studies on character association in f2 generation of ten selected crosses in rice (Oryza sativa L.) J. Res.-A FAU, 15: El-Malky M, El-Habashy M, Abdelkhalik AF (2008). Rice germplasm evaluation for agronomic traits and their influence on stem borer (Chilo agamemnon bles.) resistance. J. Agric. Res., 46(3): 206. Iftekharuddeula M, Hassan S, Islam MJ, Badshah MA, Islam MR, Khaleda A (2001). Genetic evaluation and selection criteria of Hybrid rice in irrigated ecosystem of Bangladesh. Pak. J. Bio. Sci., 4(7): Kato T (1997). Selection response for the characters related to yield sink capacity of rice. Crop Sci., 37: Khush GS, Brar DS (2002). Biotechnology for rice breeding: progress and impact. In: Sustainable rice production for food security. Proceedings of the 20th Session of theinternational Rice Commission (23-26 July, 2002). Bangkok, Thailand. Muhammed B, Asif AK, Anjuman A, Yusuf Z, Muhammad A (2007). Path analysis of some leaf and panicle traits affecting grain yield in doubled haploid lines of rice (oryza sativa l.). J. Agric. Res., 45(4): Prasad GSV, Prasad ASR, Sastry MVS, Srinivasan TE (1988). Genetic relationship among yield components in rice (Oryza sativa L.). Indian J. Agric. Sci., 58(6):
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