HETEROSIS AND DEGREES OF DOMINANCE OF QUANTITATIVE TRAITS IN HYBRIDS (PISUM SATIVUM L.)

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1 HETEROSIS AND DEGREES OF DOMINANCE OF QUANTITATIVE TRAITS IN HYBRIDS (PISUM SATIVUM L.) Valentin Ivanov KOSEV DOI: / V(9) 23 Institute of Forage Crops Pleven, Gen.Vladimir Vazov 89 Street 5800 Pleven, BULGARIA, e mail:valkosev@hotmail.com Abstract. The experimental study was conducted during period in second experimental field of Institute of Forage Crops, Pleven. The hybridization scheme including direct and back crosses covered four varieties of forage pea (Pisum sativum L.), spring Rezonator and Harkovskii etalon from Ukrainian, winter Bulgarian varieties Vesela and Mir. Inheritance of quantitative traits was analysed plant height, seed number per plant, pod number per plant, seed number per pod and seed weight per plant of parental components (Р 1 and Р 2 ) and crosses of first and second hybrid generation (F 1 and F 2 ). Heterosis effect in F 1 hypothetical and true and degree of dominance in F 1 (h p1 ) and in F 2 (h p2 ), depression, degree (Tn) and frequency of transgression in F 2 and variability (VC %) were determined for each studied trait. Highest real heterosis effect was found in crosses Mir x Rezonator for plant height (60.24%) and seed number per pod ( 0.05%), Harkovskii etalon x Mir for pod number per plant (58.88%) and seed weight per plant (51.92%), Rezonator x Mir for seed number per plant (7.39%). In F 2 with strongest depression are plants from cross Mir x Vesela for number seeds (43.53%), number pod (47.03%) and seed weight per plant (46.55%) and Mir x Rezonator for plant height (28.71%) and seed number per pod (18.59%). Cross Harkovskii etalon x Mir is with highest degree of transgression for traits plant height (24.10%), pod number per plant (64.49%) and seed weight per plant (69.62%). Low variability in F 1 and F 2 was found at plant height ( %). Highest is variability for seed weight per plant in F 1 ( %). Key words: degrees of dominance forage pea, heterosis, transgression Introduction Hybridization is one of main methods of combining different characters [MIHOV et and qualities of parental varieties al., 2002]. The study of heterotic manifestations is of great importance to reveal regularities in inheritance of [POPOVA and SACHANSKI, 1988; POPOVA, characters 1991]. The manifestations of heterosis and inheritance of characters associated with plant productivity is an object of a number [KALAPCHIEVA, 2008; HENAUT et al., of investigations 1992] [МАКАSHEVA, 1979], indicated that inheritance of longer stem in F 1 was a result of effect of dominant genes. According to [SNOAD and ARTHUR, 1973], flower number per flower stalk and seed number per pod are under control of a polygenic system of an additive nature. [CEYHAN, 2006; Many other authors BURSTIN et al., 2007; BEECK et al., 2008; CEYHAN et al., 2008] found both additive and no additive gene 23 effect in inheritance of some quantitative and qualitative characters in pea. The objective of this study was to determine manifestations of heterosis, degrees of dominance, inbreeding depression, degree and frequency of transgression and variation of a group of characters conditioning productivity of F 1 and F 2 hybrid plants, which supplements breeding evaluation of studied materials and allows their purposeful inclusion in breeding process. Material and methods The experimental study was conducted during period in second experimental field of Institute of Forage Crops, Pleven. The parent partners used for crossing were chosen according to observation of variety collection in previous years. As parental component are using following varieties forage pea: spring

2 Banat s University of Agricultural Sciences and Veterinary Medicine Regele Mihai I al României from Timisoara, Contact: web: bjb@usab tm.ro forms (Pisum sativum ssp. sativum) Rezonator and Harkovskii etalon (Ukraine republic), winter types pea Mir (Pisum Genotype Vesela Rezonator Harkovskii etalon Mir Pea parental components used in research Distinctive features sativum ssp. arvense) and Vesela (Pisum sativum ssp. sativum) from Bulgaria. The characteristics of these varieties are reported in Table 1. Table 1. Clear seed, winter, normal leaves, few branches, white flowered, long vine Clear seed, spring, normal leaves, without branshes, white flowered, long vine Clear seed, spring, leafless (afila), without branshes, white flowered, short vine Dark seed, winter, normal leaves, few branshes, purple flowered, long vine, These forms were crossed by hand in The parental forms (P 1 and P 2 ) and first and second generation (F 1 and F 2 ) are sown at scheme P 1 F 1 F 2 P 2 on a row spacing 70 cm and distance in row 5 cm. Hand planting was applied with depth of sowing 5 cm. The forage pea is grown by approved technology of Institute of forage Crops Pleven. In hybridization were included next direct and reciprocals crosses: Mir х Vesela, Vesela х Mir, Mir х Rezonator, Rezonator х Mir, Mir х Harkovskii etalon, Harkovskii etalon х Mir. Inheritance of quantitative traits was analyzed plant height, seed number per plant, pod number per plant, seed number per pod and seed weight per plant of parental components (Р 1 and Р 2 ) and crosses of first and second hybrid generation (F 1 and F 2 ). The individual number measurement of each character on one plant is respectively: from F 1 all plants, from Р 1, Р 2 and F 2 20 plants. For each studied trait were determined: mean (x) and standard error [DIMOVA and (Se) and variability (VC, %) MARINKOV, 1999], heterosis effect in F 1 hypothetical and true and depression [ОMAROV, 1975], degree of dominance in F 1 (h p1 ) and in F 2 (h p2 ) ROMERO and FREY, 1973 ; degree (Tn) and frequency of [VOSKRESENSKAYA and transgression (T f ) in F 2 SHPOTA, 1967]. All experimental data were processed statistically with using computer software MS Excel for Windows XP. Results and discussion It is evident from biometric data on plant height (Table 2) that in hybrid Mir х Rezonator inheritance was overdominant (hp 1 >1) with a strongly expressed true heterotic effect (60.24%), but also with highest degree of depression (28.71%). Negative dominance was found for Mir х Vesela ( 0.72), in which true heterotic effect had also lowest value ( 12.07%). Epistatic gene effects (hp 2 > hp 1 ) were observed in other studied hybrids for inheritance of this character. A negative degree of transgression ( 19.52% and 8.58%) was found in Mir х Vesela and Mir х Harkovskii etalon. The other crosses showed positive transgression and homozygous genotypes could be expected for them, in which gene recombinations should be available resulting in an increase of plant height. Harkovskii etalon х Mir (20.10%) had most strongly expressed degree. The character of transgression frequency recorded in four of studied hybrids varied greatly (0% 95.00%), Rezonator х Mir (95.00%) having greatest frequency. The seed number per plant (Table 2) was inherited in an overdominant way in Vesela x Mir with a true heterotic effect (3.07%), but with highest depression as well (43.53%). In reciprocal cross, inheritance was of the type of negative overdominance with the lowest value of true heterotic effect ( 22.65%). 24

3 Biometrical data related to the quantitative traits in the forage pea crosses Table 2. Hybrids Degrees of Degree of Freq. of P 1 P 2 F 1 F 2 Heterosis F 1 (%) Depression dominance transgress. transgress F 2 (%) х ± SE Hypothetical Real in F 1h p1 in F 2h p2 Tn (%) T f (%) Height of the plants, cm Mir х Vesela ± ± ± ± Vesela х Mir ± ± ± ± Mir х Rezonator ± ± ± ± Rezonator х Mir 45.25± ± ± ± Mir х Harkovskii etalon ± ± ± ± Harkovskii etalon х Mir 36.25± ± ± ± Number of seeds per plant Mir х Vesela 71.70± ± ± ± Vesela х Mir 91.15± ± ± ± Mir х Rezonator 71.70± ± ± ± Rezonator х Mir 20.65± ± ± ± Mir х Harkovskii etalon 71.70± ± ± ± Harkovskii etalon х Mir 8.10± ± ± ± Number of pods per plant Mir х Vesela 16.05± ± ± ± Vesela х Mir 17.45± ± ± ± Mir х Rezonator 16.05± ± ± ± Rezonator х Mir 5.50± ± ± ± Mir х Harkovskii etalon 16.05± ± ± ± Harkovskii etalon х Mir 3.15± ± ± ± Numbe of seeds per pod Mir х Vesela 4.51± ± ± ± Vesela х Mir 5.24± ± ± ± Mir х Rezonator 4.51± ± ± ± Rezonator х Mir 3.92± ± ± ± Mir х Harkovskii etalon 4.51± ± ± ± Harkovskii etalon х Mir 2.96± ± ± ± Seed weight per plant, g Mir х Vesela 9.63± ± ± ± Vesela х Mir 17.40± ± ± ± Mir х Rezonator 9.63± ± ± ± Rezonator х Mir 5.66± ± ± ± Mir х Harkovskii etalon 9.63± ± ± ± Harkovskii etalon х Mir 2.52± ± ± ±

4 Banat s University of Agricultural Sciences and Veterinary Medicine Regele Mihai I al României from Timisoara, Contact: web: bjb@usab tm.ro Epistatic manifestations in this character were observed for other hybrids. The strongest positive degree and frequency of transgression were observed in Mir х Rezonator (30.87%, 25.00%). The transgression degree was negative for crosses between varieties Mir and Vesela, irrespective of direction of crossing Mir х Vesela ( 36.20%), Vesela х Mir ( 19.59%). With exception of these hybrids, transgression frequency was found for rest, though within a narrow range (5.00% 25.00%). The trait of pod number per plant exerted a strong influence on plant productivity [BILGILI et al., 2001]. In crosses from F 1 (Table 2), highest values were observed for MirхVesela (27.13) with a very well expressed hypothetical and true heterotic effect (61.97% and 55.47%), with positive over dominant inheritance of character and with strong depression of plants in F 2, reaching to 47.03%. In hybrids, maternal parental components of which are Vesela, Rezonator and Harkovskii etalon, pod number was inherited in an epistatic way. The hybrids Harkovskii etalonхmir (64.49% and 75.00%), Mir х Rezonator (58.06% and 50.00%) and Rezonator х Mir (9.68% and 30.00%) were characterized by high frequency and degree of transgression with relatively close values of character in F 1 and F 2. For crosses from F 1 greatest seed number per pod per plant (Table 2) was observed in Mir х Rezonator (4.51), in which inheritance was positive dominant. The plants of same cross in F 2 were depressed to greatest extent (18.59%). Negative overdominance was found in Mir x Vesela and its reciprocal cross Hybrids and an epistatic type of inheritance was found in other hybrids. In F 2 cross Mir х Harkovskii etalon differed significantly from parental forms, as well as from other hybrids, having shown highest values (5.90) and lowest depression ( 46.04%). Negative true heterosis was observed for all hybrids reaching to 43.51% in Vesela х Mir. Positive degree and frequency of transgression existed only in Mir х Harkovskii etalon, being expressed to a great extent (29.91% and 65.00%). Relatively equalized values of grain weight per plant (Table 2) in F 1 and F 2 were recorded for crosses Mir х Harkovskii etalon and Harkovskii etalon х Mir (9.69, and 16.57, 15.45), in which most strongly expressed heterotic effect was found (51.92%). Epistasis in inheritance of this character and low depression existed in most of hybrids. In Mir x Vesela and Vesela x Mir inheritance was overdominant and negative dominant, respectively, depression had a higher value (46.55% and 6.13%), with manifestations of negative transgression ( 33.57% and 42.69%). The hybrids Mir х Rezonator (104.69% and 70.00%) and Harkovskii etalon х Mir (69.62% and 80.00%) were characterized by greatest frequency and degree of transgression. The studied hybrids were analyzed also with regard to variability of tested traits (Table 3) through variation coefficient (VC, %), which characterized the phenotypic diversity. Variability (VC %) of the investigated traits in the hybrids F 1 and F 2 Plants height Seeds number per plant Pods number per plant Seeds number per pod Table 3. Seeds weight per plant F 1 F 2 F 1 F 2 F 1 F 2 F 1 F 2 F 1 F 2 Mir х Vesela Vesela х Mir Mir х Rezonator Rezonator х Mir MirхHarkovskii etalon Harkovskii etalon хmir

5 The variation is considered low, medium or high at VC of 10%, 10 20% and over 20%, respectively. It was found that it varied within different limits for each character. It was most slightly expressed for plant height of crosses Rezonator х Mir (3.74%) and Mir х Harkovskii etalon (4.46%) from F 1 and Rezonator х Mir from F 2 (3.06%). With regard to trait of seed number per plant in F 1 variation was high over 20%, except for Mir х Rezonator (19.08%). For F 2 variation was medium to high (20.63% %) in hybrids, in which variety Mir participates as a maternal component. The level of variation was similar for pod number per plant and in hybrids Mir х Rezonator from F 1 and Rezonator х Mir from F 2 variability was more slightly expressed, 26.21% and 11.26%, respectively. Low to high variation was found with regard to seed number per pod. In F 1 highest variation was found for Mir х Harkovskii etalon (46.06%), which is evidence of greater genetic instability of this character. The hybrids from F 2 were characterized by low variation in Vesela х Mir (6.53%) tо medium variation in other crosses. The variance analysis in F 1 and F 2 with regard to grain weight per plant showed that this trait, though being genetically conditioned, was greatly influenced by environmental conditions. In F 1 and F 2 lowest variation was found for Rezonator х Mir (21.48% and 15.70%). The obtained experimental data are in according with published results of [SHARMA et al., 1998; SHARMA et al., other authors 2000; BORAH, 2009] which announce for manifestation as positive so negative heterosis effect for plant height, number of seeds per pod and seed weight per [LEJEUNE HENAUT et al., 1992; MISHRA et al., 1993; plant. CEYHAN, 2003] established non additive genes actions and overdominance for number pod per plant. When dominant gene actions predominate in these traits there is small probability selection in early crossed 27 generations to be successful [BHARDWAJ and VIKRAM, 2004; KUMAR et al., 2003; SORPHI et al., 2006] reported for high coefficient of variation for yield, plant height, number of primary branches per plant and pod weight. [SHARMA et al., 2003] also reported high coefficient of variation for seed yield per plant followed by pod per plant. Highest real heterosis effect was found in crosses Mir x Rezonator for plant height (60.24%) and seed number per pod ( 0.05%), Harkovskii etalon x Mir for pod number per plant (58.88%) and seed weight per plant (51.92%), Rezonator x Mir for seed number per plant (7.39%). %). In F 2 with strongest depression are plants from cross Mir x Vesela for number seeds (43.53%), number pod (47.03%) and seed weight per plant (46.55%) and Mir x Rezonator for plant height (28.71%) and seed number per pod (18.59%). The cross Harkovskii etalon x Mir is with highest degree of transgression for traits plant height (24.10%), pod number per plant (64.49%) and seed weight per plant (69.62%). Determinated is stability of the characters. Low variability in F 1 and F 2 was found at plant height ( %). Highest is variability for seed weight per plant in F 1 ( %). References 1. Beeck, C.P.; Wroth, J.; Cowling, W.A.; Additive genetic variance for stem strength in field pea (Pisum sativum), Australian Journal of Agricultural Research, : Bhardwaj, R.K.; Vikram, A.; Genetics of yield and other'characters in a cross of garden pea (Pisum Sativum 'L.). Indian J, Agric, Res., (2): Bilgili, U.; Uzun, M.A.S.; Acikgoz, E.; The effects of supplemental lighting and light density on plant growing of pea (Pisum sativum L.) in greenhouse conditions. 4th Turkey Field Crops Congress, Sep., Grassland and Forage Crops, Tekirdag, PAYMAS Press, Istanbul, Turkey pp. 4. Borah, H.K.; Studies on combining ability and heterosis in field pea, Legume Res (4):

6 Banat s University of Agricultural Sciences and Veterinary Medicine Regele Mihai I al României from Timisoara, Contact: web: bjb@usab tm.ro 5. Burstin, J.; Marget, P.M.; Huart, An.; Moessner, Br.; Mangin, Ch.; Duchene, Br.D.; Jolain, N.M.; Duc, G.; Developmental Genes Have Pleiotropic Effects on Plant Morphology and Source Capacity, Eventually Impacting on Seed Protein Content and Productivity in Pea, Plant Physiology, : Ceyhan, E.; Combining abilities for grain yield and yield leaf characters in pea parents and crosses, S. Ü., Ziraat Fakültesi Dergisi (40): Ceyhan, E.; Determination of some agricultural characters and their heredity through line x tester method in pea parents and crosses. Selcuk Univ., Graduate School Nat. Appl. Sci Ceyhan, Er.; Avci, M.A.; Karadas, S.; Line X tester analysis in pea (Pisum sativum L.): entification of superior parents for seed yield and its components, African Journal of Biotechnology (16): Dimova, D.; Marinkov, E.; Opitno delo s biometriya, Sofia: Zemizdat. 10. Henaut, L.; Fouilloux, G.; Ambrose, M.J.; Dumoulin, V.; Eteve, G.; Analysis of 5 parent half diallel in dried pea (Pisum sativum L.). I. Seed yield heterosis, Agronomie, : Kalapchieva, S.; Selektsiona otsenka na obrastsi gradinski grah, PhD Thesis, Plovdiv: IZK Maritsa. 12. Kumar, A.; Jain, B.P.; Kumar, A.; Genetic variability in pea (Pisum sativum L.). J. Res. Birsa Agric. Univ., (1): Lejeune Henaut, I.; Fouilloux, G.; Ambrose, M.J.; Dumoulin, V.; Eteve, G.; Analysis of a 5 parent half diallel in dried pea (Pisum sativum L.). I. Seed yield heterosis. Agronomie : Makasheva, RKh.; Flora of Cultivated Plants IV, Grain Legumes 1, Pea, Kolos, St. Petersburg, Russia, pp Mehandjiev, A.; Kosturkova, G.; Mihov, M.; Enrichment of Pisum sativum gene resources through combined use of physical and chemical mutagens. Israel Plant Science, : Mehandjiev, A.; Mihov, M.; Noveva, S.; Rodeva, R.; Kosturkova, G.; Some 28 results from the investigation on genetic improvement of pea (Pisum sativum L.). Field Crops Studies, : Mehandjiev, A.; Mihov, M.; Stoyanova, M.; Todorova, J.; Enrichment of P. Sativum genetic variation through experimental mutagenesis. Plant Science (Bg), : Mihov, M.; Mehandjiev, A.; Stoyanova, M.; Opportunities for enlarging of germplasm in spring pea. Proc. of Balkan Symposium on Vegetables and Potatoes, Novi Sad, : Mihov, M.; Stoyanova, M.; Agrobiological and economic characterization of some introduced pea varieties. Proc. of Balkan Symposium on Vegetables and Potatoes, Novi Sad, pp Mikic, A.; Cupina, B.; Katic, S.; Karagic, D.; Importance of annual forage legumes in supplying plant proteins. A Periodical of Scientific Research on Field and Vegetable Crops, (1): Mishra, S.P.; Asthana, A.N.; Yadav, L.G.S.; Heterosis for yield and yield components in field pea. Heterosis Breeding in Crop Plants Theory and Application: Short Communications: Symposium Ludhiana, February pp Omarov, D.S.; K metodike ucheta i otsenki geterozisa u rasteniy, Sel skohozyaystvennaya biologiya, : Popova, I.; Sachanski, S.; Inheritance of characters determining the yield of some crosses between two Pisum sativum subspecies. Genetics and breeding, (2): Popova, Y.; Naseldyavane na kolichestveni i kachestveni priznatsi pri krastoski mezhdu normalni i Afila sortove furazhen grah (Pisum sativum L.), Genetika i selektsiya, : Rodeva, R.; Kosturkova, G.; Georgieva, I., Mehandjiev, A.; Evaluation of pea genotypes for ascochyta blight resistance in vivo and in vitro. Beirage zur Zuchtungsforschung, (1): Rodeva, R.; Kosturkova, G.; Mehandjiev, A.; Searching for resistance to ascochyta blight in pea (Pisum sativum) by conventional and

7 biotechnological methods and approaches. Plant Science (Bg), (3): Romero, G.E.; Frey, K.J.; Inheritance of semi dwarf ness in several Wheat crosses, Crop. Sci., : Sharma, A.K.; Singh, S.P.; Sharma, M. K.; Genetic variability, heritability and character association in pea (Pisum sativum L.). Crop Res., (1): Snood, B.; Arthur, A.E.; Genetical studies of quantitative characters in peas. I A seven parent diallel cross of cultivars, Euphytica, (2): Sorphi, S.P.S.; Yadav, R.; Malik, S.; Genetic variability, correlations and path analysis for seed yield and its component characters in pea (Pisum sativum L.). Plant Arch., (2): Styanova, M.; Penchev, E.; Mihov, M.; Protein content in pea collection. Proc. Of Balkan Symposium on Vegetables and Potatoes, Novi Sad, : Voskresenskaya, G.S.; Shpota, V.I.; Transgressiya priznakov u gibridov Brassica i metodika kolichestvennogo ucheta etogo yavleniya, Doklady VASHNIL, : Received: January 17, 2014 Accepted: April 23,

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