2) Gene symbols and information on male sterility

T. K.INOSHITA

Professor Kmeritus. Hokkaido University. Sapporo. 060 Japan

Male sterility is an important character for hybrid seed production on a large scale. Information on genes responsible for various types of male sterility in rice is summarized as shown in Tables, 1, 2 and 3, which were partly transposed from the data of rice genes mentioned in Rice Genetics Newsletter Vol. 12 (Kinoshita 1995).

1. Cytoplasmic male sterility (CMS)

CMS was first reported by Katsuo and Mizushima (1958). They introduced nuclear genome of Fujisaka 5 into the cytoplasm of 'Chinese wild rice' (Oryza f. spontanea=0. rufipogon) by successive backcrosses. In hybrid rice breeding 'Chinsurah boro' cyto-

Table 1. Male sterile cytoplasms

Cytoplasm

Name

Maintainer

Restorer

Reference

[cms-bo]

Chinsurah boro II cytoplasm

Taichung 65

Rfl-a Rfl-a

Shinjyo 1969 1975,

etc.

Kadowaki et al. 1988, 1990

[cms-ld]

Lead rice cytoplasm

Fujisaka 5

Rf2 Rf2

Watanabe 1971

[cms-TA]

TA820 (Tadukan) cytoplasm

Norm 8

Kitamura 1962a, 1962b

[cms-CW]

Chinese wild rice cytoplasm

Fujisaka 5

Katsuo & Mizushima

1958

[cms-WA]

Wild abortive cytoplasm

IR24 etc.

Rf3 Rf3

Lin & Yuan 1980.

Rf4 Rf4

Kakowaki et al. 1988,

Virmarn et al. 1989

[cms-HL]

Red awned wild cytoplasm

IR54753A etc.

Lin & Yuan 1980

[cms-ak]

Akebono cytoplasm

Lien-Tong-Tao

Rfak Rfak

Yabuno 1977,

([cms-jp])

Sakamoto et al. 1990

[cms-ARC]*

ARC13829-16 cytoplasm

IR54755

IR42 etc.

Virmani et al. 1989

[cms-GAM]*

Gambiaca cytoplasm

Chao Yang 1

IR58 etc.

Lin & Yuan 1980,

etc.

Kakowaki et al. 1988,

Virmani et al. 1989

[cms-sp] *

MS577A cytoplasm

IR42 etc.

Kakowaki et al. 1988,

Virmani et al. 1989

[cms-UR89]*

UR89F cytoplasm

Taichung 65

Rfl-b Rfl-b

Kakowaki et al. 1988,

Shinjyo 1990

[cms-UR102]*

UR102F cytoplasm

Taichung 65

Rfl-c Rfl-c

Kakowaki et al. 1988,

Shiniyo 1990

[cms-UR104]*

UR104F cytoplasm

Taichung 65

Rfl-d Rfl-d

Kakowaki et al. 1988,

Shinjyo 1990

[cms-UR106]*

UR106F cytoplasm

Taichung 65

Rfl-e Rfl-e

Kakowaki et al. 1988,

Shinjyo 1990

[cms- UR27]*

UR27F cytoplasm

Taichung 65

Shinjyo 1990

[cms-54257]*

54257 cytoplasm

Ling et al. 1989

[cms-Khiaboro] *

Khiaboro cytoplasm

Akibare

Nagamine et al. 1995

[cms-IR66707A]*

Oryza perennis Ace 104823

IR64

Dalmacio et al. 1992, 1995

cytoplasm

*Cytoplasm symbols are not registered in RGC.

Rice Genetics Newsletter Vol. 14

Table 2. Fertility restoring genes

Gene

Explanation (Name/Cytoplasm/Origin)

Locus

Reference

Rfl

Pollen fertility restoration-1

/[cms-bo]/Chinsurah boro II/Rfl-a,

Rfl-b, Rfl-c, Rfl-d (multiple alleles)

Chr.l0:12

7.5cM to XNpb291, 3.7cM to OSRRf,

1.5% to fL601

Close to fL60lB

Shinjyo 1975,1990, Sato el at. 1985, Vimiani & Shinjyo 1988, Fukuta rial. 1992, YueiaJ. 1995, Afaigielal. 1996, Yokomla et al. 1996

Ichikawa n al. 1997

Rf2

Pollen fertility restoration-2

/[cms-ld]/Fukuyama

chr.2:132

Watanabe 1971,

Shinjyo & Sato 1994

Rf3* (R2, Rf2)

 

 

Rf4* (Rl, Rfl, RfWAl)

Fertility restoration-3/[cms-WA]

/IR24 etc.

 

Fertility restoralion-4/

[cms-WA] /IR24 etc.

chr.l:

1.9cMtoRG532

 

chr.7:

near XNpb379

Rfl mil Rf4:

Lu & Zhang 1986, Bharaj et al. 1991,1995,Teng& Shen 1994, Chang a al. 1994, 1997, Zhang & Lu 1996

RfWA2

Weaker fertility restoration/

[cms-WA]/IR36

chr.l0;

triplo-10

Bharaj et al. 1995

Rf2, 3,4, 5

Fertility restoration (QTL)/

[cms-Wa]DH from ZYQ/JX

chr.2:

RZ404C-RG241B chr3:

RG694-RG413

chr 5:

C22-RG449D

chr.5:

RG435-RG172A

Zhu et al. 1996

Rf5(t)" (Rf(t))

Fertility restoration-5/[cms-WA]

/Fen. revertant from 1132A (WA-tpe)

chr.l:

5.3cm to OPB07-640

Shen et al. 1993a, 1996a 1996b

Rfa. b, c

Pollen fertility restoration/[cms-bo]

/H-406/complernentary

Maekawa 1982

Rja, b', c', d'

Pollen fertility restoraton/[cms-bo]

/H-103/complementary

Maekawa 1982

Rfak (Rfjp)

Pollen fertility restoration-ak/[cms-ak] /Akebono

Yabuno 1977

lfr

Induction of fertility restoration

/[cms-bo]/partially sterile mutant from Taichung 65CMS

Sano & Eiguchi 1991,

Sano et al. 1992

* Gene symbols are not registered in RGC.

plasm or BT cytoplasm designated as {cms-bo] is extensively used for japonica hybrid rice development. The mode of inheritance of fertility restoring genes and molecular nature of mtDNA have been studied as a basis for breeding (Shinjyo 1975: Kadowaki et al. 1986, 1988, 1990-, Shikanai et al. 1989; Yamato et al. 1992 etc.). On the other hand, the cytoplasm of a male sterile line, 'wild abortive' designated as [cms-WA] is also successfully used for indica hybrid rice development (Lin and Yuan 1980). There are several cyto-plasmic sources as mentioned by Virmani and Shinjyo (1988). Virmani et al. (1989) demonstrated the cytoplasmic diversity among five cytoplasms, cms-bo, cms-WA, cms-GAM, cms-ARC and cms-sp depending on the spikelet fertilities of the progenies from test-crosses with a set of 28 cultivars. As to the utilization of wild Oryza species, Shinjyo (1990) classified at least four kinds of new cytoplasms derived from UR-lines belonging to 0. rufipogon depending on the interaction with different genotypes of Rf1 alleles. Ling et al. (1989) also found a CMS line, 54257 among somaclonal variants regenerated from anther culture and the mutant CMS was controlled by a cytoplasmic factor designated as

Report of Committee on Gene Symbolization

Table 3. Genetic male sterility

Gene

Name/Origin/Mutagen Locus

Reference

Ms1 (sf)

male sterile-l/Fukukame/spontaneous chr.6:27

Hara 1946

ms2

male sterile-2/md-strain/spontaneous

Group A (ms2-ms6),

ms3

male sterile-3/Bufumochi/spontaneous

Shibuya 1973

ms4

male sterile-4/Fujiminori/spontaneous

ms5

male sterile-5/Otori/spontaneous

ms6

male sterile-6/Bufumochi/spontaneous

ms7

male sterile-7/K-l:Koshihikari/EI chr.3:80

Group B (ms -ms6);

ms8

male sterile-8/K-2:Koshihikari/EI chr.7:20% to rfs

Ko & Yamagata 1987, 1989

ms9

male sterile-9a/RTI-3aT65/spontaneous chr.6:100

Sato & Shinjyo 1991

ms9

male sterile-9b/E-l:Etsunan 77/EI

ms10

male sterile-10/T-l:Toyonishiki/EI chr.9:5% to Dn1

msll

male sterile-ll/T-2:Toyonishiki/EI

msl2

male sterile-12/T-3:Toyoi1ishiki/EI

msl3

male sterile-13/S-32:Sasanishiki/EI

msl4

male sterile-14/S-40:Sasanishiki/7-ray chr.5:11% to nl1

ms15

male sterile-15/S-55:Sasanishiki/EI

msl6

male sterile-16/S-59:Sasanishiki/EI

msl7

male sterile-17/S-81:Sasanishiki/7-ray chr.2:35% to gh2

ms18*

male sterile- 18/MS27: Nihonmasari/EI

Group C (msl8~ms45);

msl9*

male sterile-19/MS28:Nihonmasari/ γ-ray

Fujimaki et al. 1977,

ms20*

male sterile-20a/MS15:Nihonmasari/EI

Hiraiwa & Tanaka 1980,

ms20*

male sterile-20b/MS29:Nihonmasari/ γ-ray

Fujimaki & Hiraiwa

ms2l*-ms23*

male sterile 21-23/MS:Nihonmasari/γ-ray

1986, Tamaru 1994

ms24*

male sterile-24a/MS4:Nihonmasari/ γ-ray

ms24*

male sterile-24b/MS9:Nihonmasari/ γ-ray

ms25*-ms30*

male sterile 25-30/MS5-10:Nihonmasari/γ-ray

ms31 * ~ms45*

male sterile 31-45/MSII-26:Nihonmasari/EI

ms46*

male sterile-46/M201/streptomycin

Group D (ms46~ms63);

ms47*, ms48*

male sterile-47, 48/MIOI/γ-ray

Trees & Rutger 1978,

ms49*

male sterile-49/M201/EMS

Mese et al. 1984, Hu &

ms50*

male sterile-50a/Calady/spontaneous

Rutger 1992

ms50*

male sterile-50b/Earirose/spontaneous

ms51*

male sterile-51/M201/EMS

ms52*

male sterile-52/MIOI/γ-ray

ms52*

male sterile-53a/MIOI/γ-ray

ms53*

male sterile-53b/M201/EMS

ms54*~ms56*

male sterile 54-56/M201/EMS

ms57*

male sterile-57/MIOI/γ-ray

ms58*

male sterile-58/M201/EMS

ms59*

male sterile-59a/MIOI/γ-ray

ms59*

male sterile-59b/M201/EMS

ms60*

male sterile-60a/MIOI/γ-ray

rns60*

male sterile-60b/M201/EMS

ms60*

male sterile-60c/Calrose 76

/anther culture

ms61*

male sterile-61/M201/EI

ms62*

male sterile-62/MIOI/7-ray

ms63*

male sterile-63/Caloro

/spontaneous

msIR36*

male sterile (ms-IR36)/IR36/EI

Group E (mslR36 ~

msIR36*

male sterile (5495ms)/Line5495

msm77(t));

/spontaneous

Singh & Ikehashi 1981,

msIR36*

male sterile (5683ms)/Line5683

Suh et at. 1989, 1991

/spontaneous

mslR36*

male sterile (Milyang 54ms)

/Milyang 54/spontaneous

 

Rice Genetics Newsletter Vol. 14

(Continued)

Gene

Name/Origin/Mutagen

Locus

Reference

msm55(t) *

 

msm67(t)*

msm77(t)*

male sterile (Milyang 55ms)

/Milyang- 55/spontaneous

male sterile (Milyang 67ms)

/Mi]yang 67/spontaneous

male sterile (Milyang 77ms)

/Milyang 77/spontaneous

chr.l:31

 

chr.l:15% to mp1

ms1(t)*

 

ms2(t)*'

male sterile/Co40XVaigai F1

/anther culture

male sterile/Co40XVaigai F1

/anther culture

Group F (ms1(t).

ms2(t);

Kaul 1986

oms

open hull male sterile/spontaneous

Takeda 1987

tmsl

thermosensitive male sterility-I

/5460:IR54/spontaneous

chr.8: Sun et al. 1989. 6.7cMtoTGMSI,2 Yang et al. 1992 (OPB-19) Wang et al. 1995. 1996

tms2

thermosensitive male sterility-2

/Reimei/ γ-ray

Maruyama et al. 1991

tms3(t)*

thermosensitive male sterility-3

/IR32364/ γ-ray

chr.6:

7.7cM to

OPAC3-640

Borkakati & Virmani

1993, Subudhi et al.

1995, 1996, 1997

thermosensitive male sterility

/26 Zhaizao S/ γ-ray

Shen et al. 1993b

pms1* (pgms, ms1P)

 

 

pms2* (ms2P)

IP) photoperiod-sensitive male sterility-I /Nongken 58s/spontaneous

 

photoperiod-sensitive male sterility-2

/Nongken 58s/spontaneous

chr.7:

4.3cM to RG477

 

chr.8:

near RG191

Group G (pms1, pms2);

Zhu & Yu 1989,

Wang et al. 1991, Shao

&Tang 1992, 1995

Zhang et al. 1993

Pms(t)*

photoperiod-sensitive male sterility

/M201/EMS

Oard et al. 1991,

Oard & Hu 1995

Allelism tests were carried out only within each group.

*Gene symbols are not registered in RGC.

[cms-54257]. Following this, Dalmacio et al. (1992, 1995) produced a CMS line, IR66707A as a new source in the combination between the cytoplasm from (O. perennis Ace 104823 and the nuclear genome of IR64. Nagamine et al. (1995) also selected a new cytoplasmic source from boro varieties introduced from Bangladesh. The cytoplasm was different from [cms-boro] using test-crosses with several restorers and molecular structure of mtDNA, and was named as [cms-Khiaboro]. These new CMS systems are useful to enlarge the cytoplasmic diversity in hybrid rice breeding.

Many kinds of somatic and cytoplasmic hybrids were produced by protoplast fusion using Ory'w species (Kyozuka et al. 1989). Transfer of [cms-bo] cytoplasm was achieved by donor-recipient method (Aviv et al. 1984) in japonica varieties. It is effectively used not only to shorten the time for breeding of CMS lines, but also to elucidate the genetic nature of CMS. Later it was demonstrated that a new chimeric gene, rpsi is also responsible for CMS in mtDNA besides atp6 (Kinoshita and Mori 1996).

Report of Committee on Gene Symbolization 17

It is generally accepted that Rf followed by a numeral is used to designate the fertility restoring genes. Hitherto Rfl and Rf2 located on chromosomes 10 and 2 respectively are used for the restorer genes for {cms-bo} and {cms-ld} (Table 2). It is indicated that two dominant restorer genes have additive effects for the restoration of fertility in [cm^-W^]. The two genes are located on chromosomes I and 7 respectively (Zhang and Lu 1996; Zhang et al. 1994, 1997). These genes were named as Rf3 and Rf4 by Zhang et al. ( 1997) replacing Rf2(R2) and Rf] (/?/) which were formerly used in several papers (Bharaj et al. 1991.1995: Teng and Shen 1994; Zhang et al. 1994). Teng and Shen ( 1994) showed that there is no evidence for interaction and pleiotropic effects between the fertility restoring genes for {cms-bo} and [cm^-W4]. In addition, a male fertile revertant was induced with Co"" gamma ray treatment from a CMS line, 11-32A having [c»M-W4] and the mutant acted as a fertility restoring gene (tentatively named as Rf5(t)) and is different from Rf3 and Rf4 present in IR24 and Minghui 63 (Shen et al. 1993a, 1996a, b). Fine mapping of these genes is progressing by using RFLP markers around OPB07-640 (chromosome 1) which are closely linked with the gene (Shen et al. 1996a). DNA markers closely linked to /?/-genes provide useful tools in marker-aided selection for the sterile maintainer and fertility restoring genes (Akagi et al. 1996).

Induction of partial male fertility in {cms-bo} Taichung 65 was controlled by the gene designated lfr (Sano and Eiguchi 1991: Sano et al. 1992). II. Genetic male sterility (GMS)

Usually single recessive genes condition male sterility and number over 70 in rice. As shown in Table 3, a numeral attached to ms were assigned without the coordinated allelism tests by the respective research group. In a few cases, duplicate recessive genes were reported, while no dominant genes have been reported. Type and stage of pollen abortion are various among the mutants (Hu and Rutger 1992; Tamaru 1994).

Environmentally conditioned male sterility (EGMS) can be efficiently used for hybrid seed production and 'two lines method' (EGMS and pollen parent) is gradually replacing 'three lines method' (CMS, maintainer and restorer). Sometimes EGMS are divided into thermosensitive MS (TGMS) and photoperiod-sensitive MS (PGMS or PSMS) depending upon their mode of response.

In the case of TGMS, male sterility is conditioned under high temperature but returns to normal under low temperature. Three single recessive genes, tmsl, tms2 and tms3(t) have been reported (Sun et al. 1989; Yang et al. 1992; Maruyama et al. 1991; Subudhi et al. 1995). Both tms2 and tms3(l) were induced with gamma irradiations, while tmsl occurred as a spontaneous mutant. The TGMS line containing tms3(t) shows complete male sterility at day and night temperatures of 32° and 24°C, but is partially fertile at 27° and 21 °C or 24°C and 18°C in the phytotron (Borkakati and Virmani 1993). In RFLP analysis, tmsl and tms3(t) were located on chromosome 8 and 6 respectively (Wang et al. 1995, 1996:Subudhicf<3/. 1995, 1996, 1997). ApromisingTGMSmutantnamedas26 Zhaizao S was induced by gamma-rays and the mutant showed low fertility especially at 30°C though there is no inheritance data (Shen et al. 1993b). In the case of PGMS, a spontaneous mutant was found in the field of Nongken 58.

Rice Genetics Newsletter Vol. 14

The mutant exhibited various types of pollen abortion under the restricted condition of long day (14hr/day) (Wang et al. 1991). Genetic analysis indicated that at least two recessive genes, pins I and pms2 are responsible for the trait and are located on chromosomes 7 and 3 respectively (Shao and Tang 1992; Zhang et al. 1993). Main effects of pmsl were about three times larger in the average than that of pms2 showing a considerable interaction between these two loci (Zhang et al. 1993). There is a possibility that one of the two sterility inducing genes from Nong-Ken 58S (NK58s) is allelic to one of the fertility restoring genes for [cw.s'-lVA] (Shao and Tang 1995). Similar PGMS mutants were found in progenies of the ethyl methanesulfonate-treated materials of the rice cultivar M-201 (Oard et al. 1991). Pollen fertility increased 3 to 44-fold when the mutant plants were ratooned and grown in a growth chamber with a 12hr daylength. PGMS was governed by one to three recessive genes. From the experiments with red or far-red light treatments, it was found that phytochrome is involved in regulation of pollen fertility (Oard and Hu 1995).

A spontaneous mutant named as open hull male sterile was controlled by a single recessive gene, oms. The pleiotropic effects of mutant gene resulted in lower seed fertility, smaller caryopsis, darker leaf color and good ratooning (Takeda 1987).

In order to promote gene identification and allelism tests, free exchange of the germplasm of male sterile mutants among different research groups is urgently needed.

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Rice Genetics Newsletter Vol. 14

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