Gene Symbols in Pigeonpea

Pigeonpea (Cajanus cajan (L) Millsp.), also called as ‘red gram’ in India, ‘Nandolo’ in Malawi, ‘Gandul ‘ in Puerto Rico, ‘Mu Dou’ in China and ‘Quinchocho’ in Venezuela, is an important pulse crop of the semi-arid tropics regions of the Indian subcontinent, Africa, and the Caribbean. A large genetic variation exists within this species for different quantitative and qualitative traits (Remanandan, 1990; Saxena and Sharma, 1990). A number of genetic studies have been conducted in the past and gene symbols assigned. In this paper an attempt has been made to compile information on gene symbols for a number of traits. The studies conducted by different authors have been grouped together appropriately and we have not made any attempt to equate them or to reinterpret the published data. A compilation of originally assigned gene symbols in pigeonpea and in some interspecific crosses, involving pigeonpea and its wild relatives, is respectively summarized in Table 1 and 2.

This compilation is the first such attempt in pigeonpea and the information will be useful in assigning gene symbols in future. Also, it will avoid duplications of the gene symbols. In most cases the information has been obtained directly from the published articles and attempts have been made to reproduce them in the original form. However, the authors could not acquire two research papers (marked with *) and the information has been cited using other related publications. The authors would appreciate comments from readers regarding any errors of content or omission. In this review an attempt has also been made to provide information about genetic variation considered for study, the details about the genetic control of various phenotype(s) is not discussed for the sake of brevity and for such details the readers are advised to refer the original articles.

Table 1. Summary of gene symbols in pigeonpeas

Variation studied F1 phenotype F2 ratio Gene symbol(s) Reference(s) Remarks
Flower colour
Yellow, white Yellow 49:15 W1, W2, Iy Patil and D’Cruz, 1962 W1 and W2 duplicate genes produce white flowers, Iy inhibits and produces yellow flowers.
Creamy white with red veins on back of standard,yellow-red veins on back of standard Yellow with red veins on back of standard 3:1 Y fl Patil et al., 1972  
Yellow, creamy white Yellow 207:49 Yvsda1, Yvsda2, Yvsdb1, Yvsdb2 Jambhale et al., 1978  
Yellow, yellow with red veins Yellow with red veins 45:19 Rvdsa, Rvdsb,Rvdsc Chopde et al., 1979  
White, yellow Yellow 49:15 Wfll, Wfl2, I-Wfl Kolhe and Nayeem, 1977  
Colour of ventral surface of standard petal
Yellow, pale yellow Yellow 3:1 Yvs D’Cruz and Deokar, 1970a  
Orange yellow with diffused purple base, yellow Orange yellow with diffused purple base 9:7 Oyvsa, Oyvsb Deokar and D’Cruz, 1972a Oyvsa pleiotropic for colour of dorsal surface of standard and purple spotting on white seed
Yellow, lemon yellow Yellow 13:3 Llt, I-Llt Deokar and D’Cruz, 1971a I-Llt common to colour of dorsal surface of standard and to colour of veins on dorsal surface.Also pleiotropic action on pod colour (duplicate action with gene Blp of pod Colour)
Yellow, pale yellow Yellow 3:1 Yvs D’Cruz et al., 1971c  
Yellow, lemon yellow Yellow 162:94 Yvsa,Yvsb2, Yvsc2, Yvsd2 Deokar et al., 1972c The subscript of 2 has been to three out of 4 genes because these three have duplicate complementary action, any two of the three are effective.
Yellow, orange yellow Orange yellow 3:1 Oyrs Chaudhari and Thombre, 1983  
Red, yellow Red 3:1 Pds Narkhede et al., 1980  
Cream, white Yellow 9:7 Yvsa, Yvsb Ghatge and Kolhe, 1985  
Colour of dorsal surface of standard petal
Orange yellow with purple veins and purple patch at the base, yellow Orange yellow with purple veins and purple patch at the base 3:1 Oyvsa Deokar and D’Cruz, 1972a Common with one of the two genes for colour of ventral surface of standard
Yellow, light purple with purple base Yellow 13:3 Llt, I-Llt Deokar and D’Cruz, 1971a  
Yellow with deep red veins and base diffused red, yellow with light red veins Yellow with deep red veins, and base diffused red 3:1 Rvbds Shinde et al., 1972  
Orange yellow, yellow Orange yellow 45:19 Oydsa, Oydsb1, Oydsb2 D’Cruz et al., 1971b  
Yellow with purple veins, yellow, orange yellow Yellow with purple veins, orange yellow 9:7 Rvdsa RvdsbOydsa, Oydsb D’Cruz et al., 1973  
Yellow red vein, yellow Yellow red vein 183:73 Rvds, I-Rvds, A-I-Rvds1, A-I-Rvds2 Marekar and Chopde, 1985  
Yellow, yellow with purple veins Yellow with purple veins 3:1 Rvds D’Cruz et al., 1974  
Purple, diffused with deep purple veins Yellow with faint purple veins 1:2:1 Pds Narkhede et al., 1980  
Colour of vein of dorsal surface of standard petal
Purple, yellow Purple 39:25 Llt, I-Llt, A-I-Llt Deokar and D’Cruz, 1971a  
Dark red, Purple red Dark red 21:43 Drv, I-Drv Chaudhari and Thombre, 1975  
Yellow, red Red veins 9:7 Rvdssa, Rvdssb Chaudhari and Thombre, 1977  
Yellow with purple vein
Yellow with self coloured veins 		Yellow purple veins 15:1 Pvds1, Pvds2 Deokar et al., 1971b  
Yellow with purple veins, yellow Yellow with purple veins 3:1 Rvds D’Cruz et al., 1974  
Yellow, red Red 9:7 Rdvdsa, Rdvdsb Kolhe et al., 1972  
Yellow, red Red 9:7 Rdvdsa Rdvdsb Patil, 1970  
Red, yellow Yellow 13:3 Rvds,I-Rvds Ghatge and Kolhe, 1985  
Calyx color
Pigmented, yellow Pigmented 9:7 A, B Singh and Srivastava, 1981  
Partially cleistogamous flower
Normal, partially cleistogamous Normal 3:1 Pct Saxena et al., 1992  
Normal, closed Normal 15:1, 3:1 Clf 1, Clf 2 Mehetre et al., 1992 Mutant form governed by duplicate genes
Response to photoperiod
Sensitive, insensitive Sensitive 3:1 PS3,PS2, PS1 Saxena, 1981 PS3, >PS2, >PS1. Therefore, 3:1 every time
Flowering habit
Early, late Early 45:19 Efla, Ef1b, Eflb2 Marekar, 1982  
Early, late Early 45:19 Efla, Eflb2,Eflc2 Marekar and Chopde, 1985  
Male sterility
Fertile, sterile Fertile 3:1 ms2 Saxena et al., 1983  
Fertile, sterile Fertile 3:1 ms1 Reddy et al., 1978  
Fertile, sterile Fertile 15:1 Stfl1, Stfl2 Mehetre et al., 1989 Duplicate genes
Pod colour
Maroon blotched, green Maroon blotched 15:1 Gppd1, Gppd2 D’Cruz and Deokar, 1970a  
Maroon blotched + green with blackish purple streaks,maroon blotched + green with blackish purple shades Maroon blotched with blackish purple all over pod 15:1 Blp, I-Llt Deokar and D’Cruz, 1971a Unripe pod colour
Maroon blotched red grained, maroon blotched round leaf Maroon blotched 195:61 BIp1,BIp2,I-BIp,A-I-BIp Deokar et al., 1972b Unripe pod colour
Purple, green with purple shade Purple 9:3:4 Gpstpd Gpshpd Deokar et al., 1971b  
Green with purple streaks, green Green with purple streaks 3:1 3:1 Gppd D’Cruz et al., 1971b
D’Cruz et al., 1971c		  
Dark purple, purple streaks Dark purple 39:25 Ppd, I-Ppd, A-I-Ppd D’Cruz et al., 1974
Purple with green streaks, green with Purple streaks Purple with green streaks 117:139 PGPDa, PGPDb, I-PGPDab, A-I-PGPDab Deokar et al., 1972c  
Green with black colour diffused,green with black color in streaks Green with black color diffused 3:1 Bldpd Shinde et al., 1972
Green with purple streaks, green with black streaks Green with black streaks 3:1 Blpd D’Cruz et al., 1970b  
Greenish black, maroon blotched Greenish black 3:1 Blp Patil and D’Cruz, 1965 Unripe
Greenish black, maroon blotched Greenish black 3:1 Blpd Patil, 1970 Unripe
Green, black streaked Black streaked 45:19 Gbpda, Gbpdb1, Gbpdb2 Jambhale et al., 1978 Duplicate complementary
Plain, maroon Plain 3:1 Bpd Nayeem, 1978  
Plain, maroon Plain 3:1 Ppd Kolhe and Nayeem, 1977  
Pod development
Normal, open carpel Normal 3:1 Cd1 Saxena et al., 1988a  
Seed colour
Reddish brown, white Reddish brown 9:7 Brsda, Brsdb D’Cruz and Deokar, 1970a  
Blackish purple, white Chocolate 3:6:3:1:2:1 Oyvsa, Brsd Deokar and D’Cruz, 1972a  
Reddish brown, white Reddish brown 9:7 Llt, Brsd Deokar and D’Cruz, 1971a  
Reddish brown, white Reddish brown 3:1 Rsd Deokar et al., 1972b  
Brown, white Brown 3:1 Brsd Deokar et al., 1971b  
Brown, white Brown 3:1 Brsd D’Cruz et al., 1974  
Blackish purple, white Blackish purple lighter than parent 9:3:3:1 Brsd Wpsd D’Cruz et al., 1971b
Brown, white, blackish purple Brown Blackish purple 9:7 9:3:3:1 Brsd, Wpsd Brsda, Brsdb D’Cruz et al., 1973  
Reddish brown, white Reddish brown 63:1 Brsd1, Brsd2, Brsd3 Patil et al., 1972  
Red, white Red 9:7 Rsda, Rsdb Marekar and Chopde, 1985  
Black, white, brown Brown 39:13:9:3 Brsd1 , Brsd2 , Brsd3 Singh and Srivastava, 1981  
Brown, white Brown 39:25 Brsd, BrsdI, BrsdIA Patil, 1970  
Brown, purple Purple 3:1 Plsd Chaudhari and Thombre, 1975  
Pinkish white, brown Brown 9:7 Brsda, Brsdb Chaudhari and Thombre, 1977  
Brown, fawn Light brown 9:3:4 Brsd, Fsd Ghatge and Kolhe, 1985  
Seed size
Bold, small Bold 27:37 Bsda, Bsdb, Bsdc Marekar, 1982  
Bold, small Bold 129:127 Bsd, I-Bsd, A-I-Bsda A-I-Bsdb Marekar and Chopde, 1985  
Growth habit
Determinate, indeterminate Indeterminate 13:3 Id, D Waldia and Singh, 1987a Id had an inhibitory effect on the gene for determinate habit, D
Determinate, indeterminate, semi-determinate Determinate, indeterminateor semi-determinate 3:13:112:3:1 Dt1Dt2s Gupta and Kapoor, 1991 two epistatic genes
Plant type
Erect, creeping Erect 3:1 Egr Deokar and D'Cruz, 1971a  
Creeping, prostrate Erect with spreading branches 45:9:10 Cgra, Cgrb1, Cgrb2 Deokar et al., 1971b
Erect, prostrate Erect 54:10 Egra2, Egrb2, Egrc2 D'Cruz et al., 1974 The subscript 2 indicates that any two of the three are necessary for production of erect which is dominant.
Erect, creeping Erect 13:3 Cgr I-Cgr Shinde et al., 1972  
Erect, creeping Erect 13:3 Cgr, I Patil and D'Cruz, 1965  
Erect, creeping Erect 13:3 Cgr, I Patil, 1970  
Erect, creeping Erect 3:1 Egr Chaudhari and Thombre, 1975  
Erect, semi-erect Erect 54:10 Egha1, Egha2, Egha3 Chaudhari and Thombre, 1977  
Plant height
Tall, dwarf Tall 3:1 Tsta D’Cruz et al., 1971a  
Tall, dwarf Tall 49:15 Tht, I-Tht2I-Tht2x Marekar, 1982  
Tall, dwarf Tall 165:91 Tht, I-Thta, I-Thtb, A-I-Tht Marekar and Chopde, 1985 One basic, two inhibitory complementary and one anti-inhibitory gene
Tall, dwarf Tall 3:1 d Sen et al., 1966  
Tall, dwarf Tall 3:1 t3, t3h Saxena et al., 1989  
Tall, dwarf Tall 15:1 t1, t2 Waldia and Singh, 1987b  
Tall, dwarf Tall 3:1 t4, t5 Gupta et al., 1992  
Stem colour
Purple, green Purple 3:1 Pst D’Cruz and Deokar, 1970a  
Purple, green Purple 45:19 Psta ,Pstb, Pstc Deokar and D’Cruz, 1972a  
Purple, green Purple 3:1 Pst D’Cruz et al., 1971a  
Purple, green Purple 3:1 Pst D’Cruz et al., 1971b  
Purple, green Purple 3:1 Pst D’Cruz et al., 1974  
Purple, green Purple 3:1 Pst D’Cruz et al., 1971c  
Purple, green Purple 45:19 Psta1, Pstb1 Nayeem, 1978  
Purple, green Purple 9:7 Psta, Pstb Kolhe and Nayeem, 1977  
Purple, green Purple 3:1 Pst Narkhede et al., 1980  
Stem surface
Corky, smooth Smooth 13:3 sm, Ck Saxena et al., 1988b Dominant form of sm masks the expression of Ck allele, resulting in smooth surface
Straight, wavy Straight 45:19 Strsta, Strstb1, Strstb2 Marekar, 1982 One complementary and two duplicate complementary genes
Branching habit
Spreading, erect Spreading 3:1 Sbr D’Cruz and Deokar, 1970a  
Spreading, erect Spreading 3:1 Sbr Deokar and D’Cruz, 1972a  
Spreading, erect Erect with spreading branches 54:10 Sbra2 Sbrb2 Sbrc2 D’Cruz et al., 1971c  
Spreading, erect Spreading 9:7 SBRa, SBRb Deokar et al., 1972c  
Spreading, erect Spreading 3:1 Sbr D’Cruz et al., 1971b  
Spreading, erect Spreading 3:1 Sbr Kolhe et al., 1972  
Spreading, erect Erect 111:145 Sbt, A-I-sbra, A-I-sbrb, A-I-sbrc   Marekar and Chopde, 1985  
Close, semi-spreading Close 63:193 Clbra, Clbrb, I-clbrc, I-clbrd Marekar, 1978  
Spreading, erect Spreading 9:7 Sbra, Sbrb D’Cruz et al., 1970b  
Spreading, erect Spreading 9:7 Sbra, Sbrb Nayeem, 1978  
Semi-erect, spreading Spreading 243:13 Sbra, Sbrb,Sbrc, Sbrd Chaudhari and Thombre, 1983  
Spreading, erect Spreading 3:1 Sbr Narkhede et al., 1980 and Ghatge and Kolhe, 1985  
Spreading, semi-spreading Semi-spreading 27:37 Ssbra, Ssbrb, Ssbrc Chopde et al., 1979  
Close, erect Close 21:43 Clbr, I-Clbra, I-Clbrb Marekar, 1982  
Leaflet shape
Lanceolate, obcordate Lanceolate 3:1 Llt D’Cruz and Deokar, 1970a Pleiotropy with wing shape, wing position, keel shape, habit of keel
Lanceolate, obcordate Lanceolate 3:1 Llt Deokar and D’Cruz, 1972a Pleiotropic effect with wing shape, wing position, keel shape, habit of keel
Lanceolate, round Lanceolate 3:1 Llt Deokar and D’Cruz, 1971a Common to the two genes governing colour characters of standard petal.At the same time it showed complementary action with the gene for seed colour.
Lanceolate, obcordate Lanceolate 3:1 Llt D’Cruz et al., 1971a Pleiotropic effect with wing shape, wing position, keel shape, habit of keel
Lanceolate, round Lanceolate 3:1 Llt Deokar et al., 1972b Linkage with gene for seed coat colour
Lanceolate, obcordate Lanceolate 3:1 Llt D’Cruz et al., 1971b Linkage with genes for stem colour and leaf thickness
Lanceolate, obcordate, round Round, lanceolate indifferentcrosses 3:1
3:1		 Llt, llt, lltr D’Cruz et al., 1973  
Lanceolate, obovate Lanceolate 3:1 Llt Patil and D’Cruz, 1965 Linked with genes for growth habit and unripe pod colour.First linkage group reported.
Lanceolate, obovate Lanceolate 3:1 Llt Patil, 1970  
Lanceolate, obcordate Lanceolate 3:1 Llt Marekar, 1978 Linked with one of the two complementary genes of branching habit
Lanceolate, round Lanceolate 39:25 Llt, I-Llt, A-I-Llt Chaudhari and Thombre, 1977  
Lanceolate (pointed apex), obcordate (depressed apex) Oval (blunt apex) 9:3:4 Llt, Obclt Patil and Deokar, 1980 One of the two genes shows supplementary gene action.
Round, obcordate connate Obovate (oblong obovate) 27:9:9:9:3:3:4 Llt, Rlt, Clt Ghatge and Kolhe, 1984  
Obovate, lanceolate Lanceolate 3:1 Llt Nayeem, 1978  
Lanceolate, obcordate Lanceolate 3:1 Llt Chopde et al, 1979  
Obcordate, lanceolate Oblong 27:21:16 Llt, Oblta, Obltb Jambhale et al., 1978 One basic and two complementary genes
Leaflet number
Trifoliate, multifoliate Trifoliate 183:73 Tf, I-Tf, A1-I-Tf, A2-I-Tf Patil et al, 1972 A1 and A2 duplicate genes having anti-inhibitory effects
Trifoliate, multifoliate Trifoliate 189:67 Tf, Tfa1, Tfb2, Tfc3 Marekar, 1982  
Trifoliate, multifoliate Trifoliate 45:19 Tfa, Tfb1, Tfc2 Marekar and Chopde, 1985  
Leaf thickness
Thick, thin Thick 3:1 Dg lt D’Cruz et al., 1971c Same gene for leaf colour
Thin, thick Thin 3:1 Tn lt D’Cruz et al., 1971b Linked with genes for leaflet shape and stem colour
Leaflet base
Broad, narrow broad 9:7 Bdlba, Bdlbb Kolhe et al., 1972 One of the genes along with the gene for leaf apex (Llt) shows duplicate gene action for the production of petiolule.
Leaflet apex
Obtuse, acute, retuse obtuse 117:75:64 Alta, Olta, I-Olta, A-I-Olta D’Cruz et al. 1971c  
Notchless, notched Notchless 3:1 Llt Kolhe et al., 1972 Pleiotropic effect on keel shape, keel habit and type of inflorescence
Round, pointed Pointed 3:1 Llt Nayeem, 1978  
Pointed, depressed Blunt 9:3:4 Llt, Obclt Patil and Deokar, 1980  
Petiolule
Present, absent Present 15:1 Bdlb, Llt Kolhe et al., 1972  
Leaf colour
Light green, dark green Light green 3:1 Dg lt D’Cruz et al., 1971c Same gene for leaf thickness
Light green, dark green Light green 3:1 Lglt Ghatge and Kolhe, 1985  
Petiole length
Long, short Long 3:1 Lpt D’Cruz and Deokar, 1970a  
Long, short Long 3:1 Lsta Deokar and D’Cruz, 1972a Represents one of the two genes governing stipel length
Long, short Long 15:1 Lpt1 Lpt2 D’Cruz et al., 1971a  
Stipel length
Long, short Long 3:1 Lst D’Cruz and Deokar, 1970a Closely linked with genes for stem colour and petiole length
Long, short Long 9:7 Lsta, Lstb Deokar and D’Cruz, 1972a  
Mutant gigas leaf
Normal, gigas Normal 3:1 Nh, nh Pokle, 1976 Pleiotropic action
Disease resistance
Alternaria susceptible, resistant Susceptible 3:1 al1 Singh et al., 1988  
Alternaria susceptible, resistant Susceptible 3:1 abr1 Sharma et al., 1987  
Sterility mosaic susceptible, resistant Susceptible 43:21 13:3 Sv1,Sv2 sv3,sv4 Singh et al., 1983 Sv1 and Sv2 showed duplicate dominant epistasis and sv3 sv4 showed duplicate recessive epistasis for resistance.
Phytopthora susceptible, resesistant Resistant 3:1 Pd1 Sharma et al., 1982  
Wilt resisistant, susceptible Susceptible 3:1 pwr1 Jain and Reddy, 1995  


Character Variation studied F1 phenotype F2 ratio Gene symbol(s) Reference(s) Remarks
Strophiole Present, absent Present 13:3 NS, SDI Reddy et al., 1981
Strophiole Present, absent Present 15:1 SS1, SS2 Pundir and Singh, 1985 Duplicate gene action
Seed mottling Present, absent Present 9:7 Msda, Msdb Reddy et al., 1981    
Twining nature Twining,non-twining Non-twining 13:3 I, T Pundir and Singh, 1985    
Pod surface Hairy, non-hairy Hairy 3:1 Hp Pundir and Singh, 1985    
Plant type Erect, spreading Intermediate 1: 1: 14 Eg1 Eg2 Pundir and Singh, 1985 2 genes partial dominance
Seed colour Black, orange Grey 1:2:1 Osc Pundir and Singh, 1985 Partially dominant
Leaflet shape Lanceolate, obovate Intermediate 1:2:1 L1 Pundir and Singh, 1985 Partially dominant

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Compiled by: Dr.K.B. Saxena, Dr. Rajini Raina and Mr.R.V. Kumar

For more information contact:
Dr.K.B.Saxena
ICRISAT-Patancheru
Andhra Pradesh
India
Email:k.saxena@cgiar.org

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