F1-F2 hybrids from a half diallel set of crosses involving 7 early Cajanus cajan lines were studied for yield and a number of yield-related traits. Parental performance was a good predictor of breeding value for most traits in the F1, but in the F2 a significant positive association between GCA effects and the parental performance was seen only for number of days to flowering. GCA variance predominated for all traits in the F1 and F2. The results suggested exploitation of the additive genetic variation in breeding for improvement of the cro

Byth, D E

Lacy, I H de

Saxena, K B

Wallis, E S

ICRISAT Open Access Repository

GENE ACTION IN SHORT-DURATION PIGEONPEAS K.B. Saxem*, D.E. Bytb, E.S. Wallis and I.H. De Lvcy Department of Agricullurc, University of Queensland. St. LuciaJOb7, Australia ABSTRACT A halfdiallet of ,even short-duratlon pigeonpea lines was evaluated in the F1 and FZ pcnerationtl at the University of Qucensland, Rcdland Bay Farm (27 S). Australia. The PI trial was clpbce planted and thc F2 trial wan grown in normal plant dennity. The rssulta indicated tbat GCA variance pmdominatad for all characters considered. SCA variance was significant for some characters but was small compared with GCA. There atas a clear effect of method of avaluation on the expression of generic parameters among the pr0genk.s. This paps discus~s the implicationn of these results in relatlon to the accuracy and mcaningfulne~a of genetic parameters eslimated In this and other sinlilat studies and those of other workers b r  breeding stntqisr. Additloud1 bdex words : Pigsoapu, Cojonur &n, dlallel, combining rblllty. INTRODUCTION Most of the present day pigeonpea [CaJhnus oqjon (L.) Millsp.] cultivars are land race8 which are maintained by open-pollination. The national average yield of pigeonpea in India continue to be around 600 kglha and to  date, imporcment in yield through breeding has been limited (Singh, 1971). This may, to  some extent, be related to  the relatively poor genetic knowledge of Important economic characters. Information on genetic variability, gene action and breeding value for yield and yield cmpanents  is of importance to  broeders in developing strategies and methods for improving yield. Srivastava (1986), reviewing the limited genetic information gvailable in this crop, tnentioned that for yield both additive and  nonadditive genetic variances are important, but the former seems to  be more predominant. In pigeonpea the availability of genetic information is limited a8 well a s  conflicting and, therefore, the present study on gene action was undertaken. MATERIAL AND METHOD A half.diellel of seven relatively early flowering lines was evaluated in the f i  and E, geaeratians at  Redland Bay Farm of the University of Queendand, kustralh (27%3). The parental material cansistcd of seven short-duration pigeon- pea oultivare. Of these four (QPL 1 ,  2, 3, 4) were selected at the Uhivetbity of Qua~ndand; UPAS 120 and Prabhat w e n  intraduced fram India, and 3D8103 war inl;r&u*ed ftoim IITA, Nigeria. Tho 21 F l s  and F2s were evaluated separately in  ra@deaiiaad oompiete block dealgas with thrce replications. find were sow o n  -Wwbt eddm* : Pi@mpC8 Qraodhr, It!R&.4T, htrndherlz J02 324, A.P., ladla. 104 Legume Research, 12 (3) : 1989 hlXkbtr 30, 1979. h e  to limited sead qeaetitiee the FI d W  w e b  SW'J p h t d  (1 m x 1 m) ip. two mw plots $ m long. The F2 trial ffid dxmw p b ~  3 mlowwitb ~i .$ mx.?, m #wing, Tht orp4timentrl rite baa 8 humid risact;ot a m r l r o t r ~ t  r$b a &ap, t&h e l ~ y  &isel &+am-) dl. h t i l h ~  *&pi rp:bJlenf tb wtniS16.318 lq N tiad 45 kg B,Cr, pa &actme. SWY 'e ldun war Wd M a w a r y  a &void wistura s t m ~ ,  Tht or* WM p m d  f r a t  wacif f&tegWtkim and ~ B W S  damage to eoqw Mod growth. Data wew ~ r Q w . 4  on day@ to flower, pod number, wsd daer, ostdqlpod, yisld/pl$nt, and plant %eight (only in F1). 1x1 the F1 test, @burrvations wcm d o  on all the plaatn w'hercar in the F2 plots, ten competitive #ants p r  plot were sampled at rmdom, Days to flowering was recorded on a whole p i 4  basia for the F2 trial, and a bulk harvest was done for the plot yield. Statistidal trnalyeia was done on plot means acoording to method 4, model 1 of G t l n g  (1956), RESULTS AND DISCUSSION The coefficients of variation for each character was almost similar in both the generations inferring that the characters had similar precision of determination in these trials (Tabk 1). Substantial genetic variation existed for plant beight, days to flower, and yield but the genetic coafiicient of variation valuas were relatively low for seed size and wds/pod. In the PI generation siguificant differencas wm found among entries for all characters (Table 1). In the F2 tent, aigoificant differences existed among the populations only for days to flower, aced aim, seedslpod, and yieldlplot. For yieldjplant and pod number the genotypes differed sigaificantly only in the Fl, but not in the F2. It seems that the expreesion of genetic variability for these characters was influenced by the diffemces in inter-plant m p e t i t i w  in the two trials. in the F2 test, the use of higher plant density perhaps reaultcd in greater competition and thia suppressed the cxprelllsion of tba esnetic potmtw for these characters. In contrast in the rpam-planted PI trial, plants were vifiuaily non.cornpotitive which possibly resulted in the full exptessioa of growth potential, This resulted in much larger plants in some crosses, particularly tho& involving t~ later flowering parents, and thucr in tbe realization of a relatively higher varianca components (Table 1). Consequently, the expression of genetic differmm was eonfounded with the nature of the test environment in these trials. Ye- and and Allard (1961) also reported influence of environment on the expression of ganes in flax. Most of the SCA e&ots for the charactern studied in tho F] and FZ generations were hot ~IgfIifl~bIntly different from Zerod ffowever, far ; o f i ~  UpdS !@x 3D 8103, them woa a dgaificaut negative SCA effect for dayl to flow@ (FI and F2) and for phnt W&bt I F t ) ,  iadiwthlg a r i g u i h a l  o o n t r i b u t h ~ f  m- additive gehe adtion in the expmion of tho~e traits in this cmr.  NoO# &the T&e 1. Mean s q u a ~ s  and slatMics from aaalysis of combining abiility of varidns cbamters in the PI nod F. from a 7 x  7 dialfel Days to Rowr Piant Pod Number Seed Size Seeds!pod Sieldlplant Yield/ Height P w  l, 'Mean square of OCA and SCA effects respecuvely crosses involving Prabhat or QPL line, had a signiflcadt SCA effect for days to flower. This probably rehcts the relatively narrow genetio baw for genes influencing flowering itl thew parents and suggested that the prospwte for selecting rcoomblnants earlier thbn the parents are low. For seed yield, the data showed a marked difference in GCA acct between yield/plant in the F1 and yield per plot in the F2. Paronts 3D8103, UPAS 120, and Prabhat conditioned high seed yield/plane in the F1, but 3D 8103 was inferlor in GCA to the other two lines for yield/plot in the F2. Further, QPL lines had similar negative GCA effects for yield/plant in the F1, but QPL 3 and 4 had large positive QCA efP&ts for yield per plot in the F2. These difference8 reflect the aeota of differential inter-plant competition. This suggects that evaluation of mating ddgns Bhould be in cultural conditions simulating normal practice, at loaat as far as plant &nsity is concerned. There was a cloae association between the GCA effeats of the partats and their performance in the FI (Table 2); that in, parental performanw per 8e Was a good predictor of breeding value for most of the characters. Thus the earliegt flowering parents conditioned relatively early flowering, short plants, low pod number and low yield per plant, whereas tbe progenies of the later flowering 3D 8103 tended to be relatively late flowering and taller with large pod number, small sccd aize and high yield/plant in the Fl trial. On the other hand in the F2 a significant positive association between GCA effect and parental performance was observed only for days to flower, The raswlts emphasize the need to restrict parentage of mating design to a relatively narrow range of pbenology, otherwise, it may prejudice the muracy and meaningfulness of the genetic parameters. Tabla 2. GCA effects in a sctvaa-paceat diallei ia Pa and F,. -. ..- Days lo flmr Plant Pod Setd size fbcdsl Yialdl Yield1 Pafeat --*-- Height Number - -  Pod Plant Plot PI Fa l-5 I% '1 F8 F1 '1 ' ' 8  Prabhat 0.39 0.54 0.48 71.26 -0.17 -0,02 0.27 29.69 79.70 UPAS-120 9.23 8.64 20.20 61.10 0.96 0,41 0,04 30.21) 177.62 3P4103 20a42 1164 57.47 187 34 0.08 -0 49 -Q.OS 40.24 -553.89 OPL-2 -6.05 -4,96 113.82 -40.71 4 . 0 7  -0.19 -0.11 -17.34 -i3.21 QPGl -8.26 -5.96 -21.71 -103.53 -0.18 0.17 -0.0) --X.83 -2984 QPL-4 -11.68 -6.23 42.39 -78.78 0.02 0.29 -009 -#).a 179.37 QPL.3 -7.04 -6.16 -17.27 -Y6.67 4.24 -8.18 4 . 1 5  -3r.31 la.% LSDQ& 1.73 2.76 4.09 134.JI 0.13 0.22 0.14 30.41 "x 67.6 64,s 111.6 949.6 7.2 7.3 1 8  195.2 3 M , 7  r ~ a , ' j 5 ~ L  b,89*j 0.97" O.Q9** 0.64 b e  4.45 0.92** 0,81W 0,21 C 4 CaaalMion d OCX QW of a, PRO# with it# m & n  WO&UIW, K.B. Saxena er al. 107 To understand the potentidl influcnce of the parental phenology on the estimates of genetic parameters, the diallel set was reanalyzed eliminating the latest flowering parent 3D 8103 (Table 3). The elimination or 3D 8103 crosses had no influence on the significance of differences among the progenies (Table 1, 31, but SCA variance for days to flower and SCA and GCA variances for yieldlplot were not significant in 6x 6 diallel. For most of the characters, there was a substantial reduction in the size of OCA and SCA components. As in the 7 x  7 analys~s, GCA variance dominated all characten but the ratio of the GCA to SCA component was greater in 6 x  6 analysis. In the present study GCA variance predominated for all characters studied (Table 1) both in F1 and F2 generations. SCA variance was significant for some characten, but was small relative to GCA. These results are similar to those of Sharmr et al. (1972), Sharma el a/. (19731, and Dnhlyr and Btar (1977) in pigeonpea. On the other hand, Reddy et 01. (1977) and Retldy er al. (1979 a, b) reported predominance of SCA variance for all characters btudied, including days to flower, plant height and seed yield and its components. The cause of this total difference between these studies is not known, but it may be related to the genetic material used in the study. Plant height and seed size are known to be highly heritable characters in pigeonpea (Munoz and Ahrams. 1971; Khan and Rachie, 1972; Shrrma et al., 1972,' Sharma et al.,  1973), and this is in disagreement with estirrates indicating a preponderance of SCA variance for such traits (Reddy et a/.,  1977. Reddy et a1 , 1979 a, b). Estimates of the form of genetic variation have quite fundamental influence of the definition of breeding strategies and methods. On the basis of their estimates of gene action, Sharma et al. (1973) suggested the development of compostte varieties; Dahaya and Brar (1977) preferred the use of the bulk population improvement over pedigree methods, and Roddy atal. (1979 b) suggested the exploitation of non-additive gene action through hybrids. Kbrn (1973) and Frey (1975) advocated population breeding using naturalout crossing in the crop. Interestingly. the reproductive biology of pigeonpea permits both options. Varietal hybrids based on genetic male sterility and natural out crossing have exhibited considerable heterosis for seed yield (Saxeaa et al., 1986). Also, pedigree or bulk breeding methods directed towards pure lines have been used sucessfully in pigeonpeas (Gupta and Saxena, 1985). Bytb et 01. (1981) reported a modibation of floral morphology which enforces self- pollination is pigeonpea, and purposed its use in breeding scheme aimed for the production of both improved pure lines and populations. In view of the conflicting and limited literature on gene action, in pigeonpea i t  Is nat possible to define breeding strategies directed exclusively at the exploitation of additive vmus non-additive genetlc variance. However, the present results and t & o ~  ~fghmm et al. (1972), Sharma et al. (1973) and Dabiya and Brrr (1977) a~ggest hat for the improvement of pigeonpea tbe additive genetic variation should &. exploited through appropriate breeding procedw~e. rdk 3. Mean squaed and atatiatia Cmm anatysin of combining ability for various cbars&cs in the P, and F, from a 6x6 diallcl. iloera D a y r t o f l ~ t e  Phal Height Pod Number Scad Size Seedslpod Yiddlptant Yiild/plot oGcoNic m a e a t  of variation. K. B. Sarena et al .  ACKNOW1,EI)GEMENT This work was supported by tbc lnteroatiorlal Crops Rcscarch Institute for the Semi-Arid Tropics (ICRISAT). One ot [IS (KBS) was suppurled by a Cornnlon- wealth Special Research Grant.  'The tcchn~cal hclp to Mr. C. Brauos IS  gratefully acknowledged. R IiFERENCES Bytb, D.E.; Walllu, E.S. and Saxena, K.R. (1981) Adaptation and breeding strategies for pigeonpea. Proc. Intl. Workshup on pigconpee. In~ernatinnal Crops Research lnstitrlle for the Semi-Arid Trop~cs (ICRISAT), Hyderabad, India. Dec. 15-19, 1980. Vol. 1 : 450-465. Dahlya, B.8. and Brar, J.S. (1977) Diallel analysis of gcnetlc variat~on in ptgeonpea (Cojnnus c&n). EXPI. Agric.. 13 : 193-200. Frey, K.J. (1975) Breeding concepts aod techniques for aelf pollinated crops. Proc. Inti. Workshop on Grain Lrgumes, ICRISAT, Hyderabad, India. Jan. 13-16. PP. 257-278. GrUtlng, 0.  (1956) Coocept of general and specific combining ability in rclat~on to diilllel croasing systems. Aust J ,  Biol. Sci ,9 : 463-493. Gupta, S.C. sad Snxena, K B. (1985) Breeding for high yield and disease resistance in early maturing pigeonpeas. Paper presented at AICPIP Kharif Pulses Workshop 16-19 May 1985- TNAU. Coirnbatore. Ind~a. Kban. T.N. and Rachle, K 0. (1972) Preliminary evaluation and utilization of pigeonpca germplasm in Uganda. East Afr. Apn'c. & For, J. 38( 1) : 78-82. Khan, T.N. (1973) A new approach to the breeding of pigeonpeas (Lbjanus cajan Millsp.) : Formation of composites. Euphytica, 22 : 373-377. Munoz, A.A. and Abrema, R. (1971) lnhcr~tance of some quantitative characters in pigeonpeas [Cru'anus cojan (L.) Millsp 1. J.  A ~ r i c .  Otriv. Puerto Rlco, 55(1) : 23-43. Reddy, R.P.; Azeem, M.A. and Rao. N.G.P. (1'177) Brecdiog suoerior pigeonpeas. 3rd Inter- national Congress of SABRAO, Canberra 811-816. Reddy, H.P.; Rae, K.V. and Rae, N.G.P. (1979a) Mcterosls and combining ability in pigeon- pea. Ind. J .  Genet., 39 : 240-246. Reddy, R.P.; Azeem, M.A.; Itso, K.V. and Rao. N.G.P. (1979b) Combin~ng ability and index selection in F2 penoralion of pigeonpea crosoea. Ind. I. Genet., 39 : 247-254. Saxens. K.B.; Faris, D G.; Reddy, L.J.; Sharmr, D.; Reddy. B.V.S.; Cupta, S.C, and Green, J.M. (1986) Prospsctr for hybrid pigeonpea. In Now Frontiers in Breeding Rewarches. Proc. Pifth Intl. Congress. SABRAO PP. 378-388. Sbarma, D.: Slngh, L.; Baghel, S,S, and Sharma, H.K. (1972) Gcnetic analysis of seed size in pigeonpea (Cajanus cajan). J .  Genet. Cytol., 14 : 545-548. Sbarma. H,K.; Siagh, L. and Sharma, D. (1973) Combining ability in diallel crosses of pipeon- pea. Ind J. Agrl. Scl., 43 : 25-29. Srlvantava, H K. (1986) pigeonpea Genetics and Broedinfi. In Pigeonpea Genetics and Improvement, E!d. S.B.S, Tikka Sharda hblishing Academy. pp. 33-55. SIngb, L. (1971) Resultj of the All India Coordinated Varietal Trials of orhar 1971-71, Paper presented at the Fifth All India Research Workers' Workshop, Indian Agricultural Research Institute, New Delhi. September, 1971 (Mimeographed). Yermnos, D.M. rod Allard, RW. (1961) The detection of epistatic gene action in Bax. Crop Sci.,l : 307-310. 

Gene action in short-duration pigeonpeas

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Gene action in short-duration pigeonpeas

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