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CROP BREEDING, GENETICS & CYTOLOGY

S^h and S_c—Two Complementary Dominant Genes that Control Self-Compatibility in Buckwheat

^a Kade Research Ltd, Research Centre, Unit 100-101 Route 100, Morden, MB, Canada R6M 1Y5
^b Dep. of Plant Science, Univ. of Manitoba, Winnipeg, MB, Canada R3T 2N2

^* Corresponding author (Rachael_Scarth{at}umanitoba.ca)

	ABSTRACT

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION REFERENCES

 
Fagopyrum homotropicum Ohnishi, a wild diploid (2n = 2x = 16) species with self-compatibility expressed by homostylic flowers, has been used for improving cultivated buckwheat, F. esculentum Moench, a self-incompatible diploid (2n = 2x = 16) species with heterostylic pin and thrum flowers. Four crosses were made between F. homotropicum and F. esculentum pin flowers, assisted by ovule rescue in vitro, to study the inheritance and interaction of the two breeding systems in the genus Fagopyrum. The presence of homostylic or pin flowers was used to determine the expression of self-compatibility or self-incompatibility, respectively. The segregation ratios of the F₂ progeny derived from F₁ single plants, the BC₁F₁ generation and the F₃ progeny derived from homostylic plants were used to study the inheritance of self-compatibility. Five F₂ populations fit a one-gene 3:1 segregation ratio and did not fit a 9:7 ratio, while the other three F₂ populations fit a two-gene 9:7 ratio and did not fit a 3:1 ratio. The BC₁F₁ and F₃ progeny segregation confirmed these observations. These results support a two-gene model with three alleles at the first locus S and two alleles at the second locus S_c. The proposed model has S for self-incompatible thrum, S^h for self-compatible homostyly, and s for self-incompatible pin, with the intrallelic interaction S > S^h > s at the first locus and S_c for homostyly and s_c for pin (S_c > s_c) at the second locus. The two complementary dominant genes S^h and S_c control self-compatibility (homostyly) in F. homotropicum. The one gene or two gene segregation patterns are the result of interspecific crosses with different F. esculentum genotypes.

	INTRODUCTION

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION REFERENCES

 
THE GENUS Fagopyrum is composed of 18 species (Ohsako et al., 2001) and contains both diploid (2n = 2x = 16) and tetraploid (2n = 4x = 32) forms, with two breeding systems, self-incompatibility and self-compatibility. The predominant system in Fagopyrum, self-incompatibility, was first discovered by Darwin (1877) and belongs to the sporophytic system (Dahlgren, 1922) expressed in the pin and thrum flower type. The pin flower has long styles with short stamens, whereas the thrum flower has the opposite arrangement with short styles and long stamens. The two flower types are produced on separate plants. Fertilization can only occur between pin and thrum flowers, and pollination is accomplished by insects and wind. The prevention of selfing occurs as the result of the inability of selfed pollen to promote seed set, and not due to the architecture of the flower (Nettancourt, 1977). The inhibition of pollen tubes in an incompatible style can occur one to several hours after pollen germination (Nettancourt, 1977) and the expression of self-incompatibility is so strong that almost all of the pollen tubes failed to reach the base of the style (Adachi, 1990). Self-compatibility is expressed by homostylic flowers in Fagopyrum, with styles and stamens of the same height. It is believed that homostyly in this genus was derived from the breakdown of heterostyly as a general trend in evolution (Yasui et al., 1998).

Common buckwheat, the cultivated species, is a self-incompatible diploid species. It has been widely accepted that self-incompatibility in F. esculentum is controlled by one gene (Althausen, 1908; Dahlgren, 1922; Eghis, 1925; Garber and Quisenberry, 1927; Saknarov, 1946). The genetic model represents the thrum genotype as Ss and ss for pin flowers. A 1:1 ratio is produced in the progeny from crosses between thrum and pin flowers. A supergene complex has been proposed by Sharma and Boyes (1961), consisting of five subgenes corresponding to stylar incompatibility, pollen incompatibility, style length, pollen size, and stamen height. Fesenko (1985)(1986) has suggested a similar model with three subgenes controlling style length, stamen length, and pollen size. However, this model has not been confirmed in view of the report by Sharma and Boyes (1961) in which an irradiation treatment produced a thrum plant with branches bearing both pin and homostylic flowers, inferring that one of subgenes had been mutated. There is the possibility that other genes in the irradiated buckwheat were affected by the irradiation treatment (Nettancourt, 1977).

Self-incompatibility in F. esculentum, expressed in the heterostylic flowers, has been identified as one of major causes related to yield instability in buckwheat (Ruszkowski, 1990; Campbell, 1997). The self-compatible wild species F. homotropicum is the most closely related species to F. esculentum in the Fagropyrum genus (Yasui and Ohnishi, 1998) and is one possible source of the self-compatibility character for interspecific introgression into F. esculentum.

Fagopyrum homotropicum has both diploid and tetraploid forms, collected by Ohnishi (1995) in southwestern China. This species has several desirable characters in addition to its self-compatibility, including high seed set and frost tolerance, and has been hybridized with F. esculentum, with the objective of creating a new self-pollinated buckwheat (Campbell, 1995; Wang and Campbell, 1998; Woo et al., 1999). It is important to buckwheat improvement programs that the inheritance and the interaction of the two breeding systems in the interspecific crosses is understood. Since the two breeding systems exist in the two species, segregation analysis based on the interspecific hybrids would be the first step to approach the problem. The Mendelian segregation ratio is frequently aberrant in the progeny of an interspecific hybrid between a cultivated species and a wild relative (Zamir and Tadmor, 1986). However, this problem could be reduced in the interspecific hybrid between F. esculentum and F. homotropicum as the two species are closely related (Yasui and Ohnishi, 1998) and the fertile interspecific hybrids in the previous reports (Campbell, 1995; Wang and Campbell, 1998; and Woo et al., 1999) were produced due to normal cell division at meiosis.

Based on a cross between F. homotropicum diploid and F. esculentum thrum, Woo et al. (1999) proposed that self-compatibility in F. homotropicum is controlled by the same S gene locus as in F. esculentum and designated S^h as the self-compatibility S allele. The relationship between the self-compatibility and self-incompatibility alleles is described as S > S^h > s with S the self-incompatible (thrum) allele dominant to both S^h for self-compatible (homostyly) and to s, the self-incompatible (pin) allele. The limitation to this study was the small population sizes, as each F₂ population contained only 10 to 14 plants, and the occurrence of one pin plant out of 11 plants in the BC₁F₁ progeny of F₁ thrum plants crossed with pin plants of F. esculentum, which could not be explained by the model.

Fesenko et al. (1998) obtained three types of flowers in a BC₁F₁ progeny of randomly mated F₁ thrum plants with F. esculentum pin. These results suggest that the homostyly of F. homotropicum is not controlled by the same alleles as the heterostyly in F. esculentum, but they did not propose a different model from Woo et al. (1999). In addition, using the self-incompatible pin type of F. esculentum in interspecific crosses is better than using the self-incompatible thrum type, as the thrum type carries the alleles of both pin and self-compatible homostyly. Use of the only the pin type plants in crosses with homostylic plants eliminates the thrum phenotype and allows the selection of homostyly from pin.

There was no genetic study conducted previously based on the cross between F. esculentum pin and F. homotropicum homostyly. Furthermore, new collections of F. homotropicum are now available that exhibit genetic diversity for morphology and allozyme patterns (Ohnishi and Asano, 1999). The interspecific hybrids derived from these new collections would not only be valuable sources of increased genetic variability for breeding programs, but could also help in the understanding of the inheritance of desirable traits such as self-compatibility.

The objectives of this study were: (i) to study the inheritance of self-compatibility in F. homotropicum through hybridization between F. homotropicum diploid and F. esculentum pin type using new accessions of F. homotropicum; and (ii) to study the genetic relationship between self-compatibility in F. homotropicum and self-incompatibility in F. esculentum. A better understanding of the genetic relationship is important for developing successful breeding strategies for buckwheat.

	MATERIALS AND METHODS

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION REFERENCES

 
The two diploid (2n = 2x = 16) accessions of F. homotropicum, K980854 and K980855, were new collections and distinct from the accessions used in the previous studies (Fesenko et al., 1998; Woo et al., 1999). The three diploid (2n = 2x = 16) F. esculentum elite lines were BM940364, BM94999.1, and X98088. The homostylic flower of the two F. homotropicum accessions was homozygous as self-incompatible heterostylic flowers (pin or thrum) have never been found in the population. The F. esculentum population is heterogeneous with a 1:1 ratio of pin and thrum plants, but there were no self-pollinated homostylic flowers found in the population. The pin type of F. esculentum was used for the crosses. Fagopyrum homotropicum was used as the female parent in three of four crosses (K980854 x BM94999.1, K980854 x BM940364, and K980855 x BM940364), and as the male in one cross (X98088 x K980854) (Table 1). The self-compatible homostylic flowers of F. homotropicum were emasculated one day before crossing. No emasculation was required when a self-incompatible F. esculentum pin plant was the female parent. Pollination was performed in the morning between 9:00 and 11:30 AM and the crossed flowers were bagged for two days following pollination. In this study, the crosses were made in only one direction as there was no maternal effect on flower morphology including style length in our previous hybridization between F. esculentum and F. homotropicum (Wang et al., 2002).

The ovule cultures were maintained at 22°C under continuous light (40 µmol m^–2 s^–1). The embryos emerged from the ovules in approximately 1 to 2 wk. Some embryos formed normal plantlets which were transferred into half-strength MS medium for further growth. Other embryos produced plantlets with poorly developed shoots and roots and these plantlets were transferred into modified MS medium, as described by Samimy et al. (1996), for shoot induction. When the shoots grew to approximately 2 cm in height, plantlets were transferred onto half-strength MS medium for rooting. Fully developed plantlets were transplanted into a soil mixture with a ratio of 2:2:1 soil/perlite/peat moss in 12.7-cm (5-inch) pots (one plant per pot). These plants were covered by an inverted 10.2-cm (4-inch) pot for one day to maintain humidity during plant hardening.

Population Development and Analysis
F₁ hybrid plants were backcrossed with F. esculentum pin plants in pairwise crosses between single plants in each cross. Although a number of backcrosses were attempted for all crosses, only two backcrosses used for analysis in this study, because seed set was low. The tiny buckwheat flowers make emasculation and pollination laborious and production of large amounts of seed difficult. In addition, the interspecific barrier was still present in the first backcross. The two backcrosses which produced more than 40 seeds were used in segregation analysis. Eight F₂ populations derived from selfing single F₁ plants were produced with a population size of 80 to 117 plants for each population. Due to insufficient seed set in the F₂ single plants and the limited greenhouse space, only three of the eight F₂ populations were used for the F₃ progeny. These three populations were chosen because they represented the two types of segregation patterns in the F_2. The populations were scored for homostylic and pin flowers to determine the segregation of self-compatibility and self-incompatibility, respectively. Chi-square analysis was used to test the goodness of fit to the expected gene segregation ratios.

All plants were grown in a greenhouse which was maintained at a minimum temperature of 22°C with natural light supplemented by high pressure sodium lamps to give a 16-h day and 8-h night photoperiod.

	RESULTS AND DISCUSSION

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION REFERENCES

 
Segregation of F₁, F₂, and BC₁F₁
Fifty-two F₁ hybrids were obtained from the four crosses through ovule rescue in vitro (Table 1). All F₁ plants were self-compatible with homostylic flowers, indicating that self-compatible homostyly is dominant to self-incompatible heterostylic pin.

Of the eight F₂ populations, five populations fit a 3:1 homostyly/pin ratio and did not fit a 9:7 ratio, while the remaining three populations fit a 9:7 ratio but did not fit a 3:1 ratio (Table 2). The three F₂ populations with a 3:1 ratio also fit a 13:3 ratio indicating a epistatic interaction, but the other two populations with a 3:1 ratio did not fit a 13:3 ratio (² = 6.5 and 10.76).

The population sizes have been increased in this study with 80 to 117 plants for each F₂ population and approximately 50 plants in the two backcrosses (Table 2) compared with the previous study (Woo et al., 1999) where only 10 to 14 plants in each F₂ population and 11 plants in one backcross population were used for analysis.

F₃ Progeny Testing
Fourteen F₃ lines derived from homostylic plants in three F₂ populations were used for progeny testing (Table 3). In the two F₂ families which segregated 3:1, the F₃ ratios were 1:0 and 3:1 homostyly to pin. Three segregation ratios, 1:0, 3:1, and 9:7, were produced in the F₃ progeny from the F₂ family which segregated in a 9:7 ratio (Table 3).

We propose a two-gene model with three alleles at the first locus as described by Woo et al. (1999). The homostylic self-compatible allele was designated as S^h in the one-gene model proposed by Woo et al. (1999), where the relationships between the alleles which produce the three types of flowers, were described as S > S^h > s, that is, the thrum self-incompatible S allele was dominant to the homostylic self-compatible S^h allele and S^h dominant to the pin self-incompatible s allele. The second locus has two alleles, S_c a self-compatibility allele dominant to s_c, a self-incompatibility allele. The two alleles S^h and S_c confer the self-compatible (homostyly) phenotype and s and s_c represent the two alleles for the self-incompatible (pin) genotypes.

The proposed genotype for F. homotropicum is S^hS^hS_cS_c as F. homotropicum is pure-breeding for homostyly and heterozygosity at either loci would result in the production of homozygous recessive genotypes and the expression of a pin phenotype. Fagopyrum esculentum pin and thrum plants, on the other hand, have more than one possible genotype. At the first locus, the proposed genotype for pin is homozygous recessive ss and for thrum is Ss, resulting in a 1:1 ratio between the two types of flowers in the progeny with no homostyly phenotypes produced in the species. The pin and thrum phenotypes may be homozygous or heterozygous at the second locus. Therefore, the proposed genotypes for F. esculentum pin plants are ssS_cS_c, ssS_cs_c, or sss_cs_c and for F. esculentum thrum plants are SsS_cS_c, SsS_cs_c, or Sss_cs_c.

Using the proposed model to predict the genotypes of the three flower types in the two parental species, it is possible to explain the observed segregation ratios in the F₁, F₂, F₃, and BC₁F₁ produced in the crosses between the two species (Tables 2 and 3), including the crosses between F. homotropicum and F. esculentum pin (Fig. 1) and between F. homotropicum and F. esculentum thrum (Fig. 2). The genotype S^hsS_cS_c in the F₁ would result in a 3:1 ratio in the F₂, while the genotype S^hs S_cs_c in the F₁ would produce a 9:7 ratio in the F₂. The F₃ progeny segregation ratios of 1:0 and 3:1 supported the occurrence of the two possible F₂ genotypes S^hS^hS_cS_c and S^hsS_cS_c. The three segregation ratios in the F₃ from the 9:7 segregating F₂ population indicated that the genotypes of the F₂ homostylic plants were S^hS^hS_cS_c, S^hsS_cS_c, and S^hsS_cs_c.

View larger version (14K): [in this window] [in a new window]   Fig. 1. The possible segregation patterns in the F1, F2, F3, and BC1F1 generations of crosses between Fagropyrum homotropicum homostyly (h) and F. esculentum pin (p) using the two-gene model with the following gene notation: two alleles at the first locus, Sh homostyly and s pin; two alleles at the second locus, Sc homostyly and sc pin. The proposed genotype for F. homotropicum is ShShScSc. The proposed genotypes for F. esculentum pin plants are (1) ssScSc, (2) ssscsc, and (3) ssScsc.

View larger version (15K): [in this window] [in a new window]   Fig. 2. The possible segregation patterns in the F1, F2, F3, and BC1F1 of crosses between Fagropyrum homotropicum homostyly (h) and F. esculentum thrum (t) using the following gene notation: three alleles at the first locus, S thrum, Sh homostyly, and s pin; two alleles at the second locus, Sc homostyly and sc pin. The proposed genotype for F. homotropicum is ShShScSc. The proposed genotypes for F. esculentum thrum plants are (1) SsScSc, (2) Ssscsc, and (3) SsScsc.

The proposed two-gene model therefore explains the controversial results in the previous reports (Fesenko et al., 1998; Woo et al., 1999). More support for this model has been provided by observations in our ongoing breeding program. With further backcrossing and crossing among selected progeny, all four genotypes, S^hS^hS_cS_c, S^hsS_cS_c, S^hS^hS_cs_c, and S^hsS_cs_c, for homostyly and five genotypes, S^hS^hs_cs_c, S^hss_cs_c, ssS_cS_c and ssS_cs_c, and sss_cs_c, for pin were produced. If all the possible genotypes for pin are crossed with all the genotypes for homostyly, the F₁ progeny should segregate in either a 1:0 (ss x S^hS^h) or a 1:1 (ss x S^hs) in the one-gene model, but five possible segregation ratios, 1:0, 1:1, 3:1, 1:3, and 3:5, could be produced in the two-gene model (Table 4). Even though it is not easy to separate the ratio of 3:5 from the ratio of 1:1, the ratios 3:1 and 1:3 can be easily separated from the 1:1 ratio statistically to differentiate between the two models. The ratios 3:1, 1:3, and 3:5 in the F₁ progeny from the crosses between pin and homostyly are the best support for the existence of the second gene S_c or at least that two genes control the trait. In total, five F. homotropicum accessions have been used for interspecific hybridization in our breeding program, including one accession used in the previous report (Woo et al., 1999). All five segregation patterns have been observed in the backcrosses between the hybrids and F. esculentum pin or intercrosses among the progeny of pin and homostylic plants (data not shown). To verify the proposed model, more powerful evidence such as molecular analysis would be required.

Application to buckwheat breeding
Introgression of self-compatibility from F. homotropicum to F. esculentum involves crosses between the homostylic and heterostylic species. In the present study, only heterostylic pin flowers were used in the interspecific crosses. Because of the differences between pin and thrum in morphology and genetics, utilizing the pin plant has advantages over thrum plants in a crop improvement program. Thrum flowers are more difficult to cross than pin flowers as the styles of thrum are imbedded among the stamens. It is also more difficult to distinguish homostyly from thrum in the small buckwheat flowers (3- to 4-mm diameter). In addition, interspecific crosses between thrum and homostylic plants could produce progeny with all three types of flowers (Fig. 2). The use of only pin type plants in crosses with homostylic plants eliminates the thrum phenotype and facilitates the selection of homostyly from pin as the style of the pin flower is conspicuous above the stamens.

	ACKNOWLEDGMENTS

 
The authors would like to thank Dr. Y. Yasui for his advice in the gene notation.

Received for publication November 3, 2003.

	REFERENCES

TOP ABSTRACT INTRODUCTION MATERIALS AND METHODS RESULTS AND DISCUSSION REFERENCES

 

• Adachi, T. 1990. How to combine the reproductive system with biotechnology in order to overcome the breeding barrier in buckwheat. Fagopyrum 10:7–11.
• Aii, J., M. Nagano, S.H. Woo, C. Campbell, and T. Adachi. 1999. Development of the SCAR markers linked to the S^h gene in buckwheat. Fagopyrum 16:19–22.
• Althausen, L. 1908. Zur Frage über die Vererbung der langgriffeligen und kurzgriffeligen Blutenform beim Buchweizen und zur Methodik der Veredelung dieser Pflanze. Russ. J. Exptl. Landwirtsch. 9:568.
• Campbell, G.C. 1995. Inter-specific hybridization in the Genus Fagopyrum. p. 255–263. In T. Matano and A. Ujihara (ed.) Proc. 6th Int. Symp. Buckwheat, Shishu, Japan. 24–29 Aug. 1995.
• Campbell, G.C. 1997. Buckwheat Fagopyrum esculentum Moench. Promoting the conservation and use of underutilized and neglected crops. 19. Inst. Plant Genet. Crop Plant Res. Gatersleben/IPGRI, Rome.
• Dahlgren, K.V.O. 1922. Vererbung der Heterostylie bei Fagopyrum (nebst einigen Notizen über Pulmonaria). Hereditas 3:91–99.
• Darwin, C. 1877. The different forms of flowers on plants of the same species. John Murray, London.
• Eghis, S.A. 1925. Experiments on the drawing up of a method of buckwheat breeding. Bull. Appl. Bot. Genet. Plant Breed. 14(1):235–251.
• Fesenko, N.N. 1985. Self-incompatible homostylous form of buckwheat with short floral organs. (In Russian.) Geneticheskie osnovy selektsii i semenovodstva grechikhi 1985:61–65. (PBA, 57:7038, 1987).
• Fesenko, N.N. 1986. The occurrence of subgene G in buckwheat outside the heterostyly locus. (In Russian.) Tsitogenet. i podkhody k izuch. Biosistem. Dokl. MOIP 1984. Obshch. Biol.:57–58. (PBA, 57:10837, 1987).
• Fesenko, N.N., A.N. Fesenko, and O. Ohnishi. 1998. Some genetic peculiarities of reproductive system of wild relatives of common buckwheat Fagopyrum esculentum Moench. p. VI:32–35. In C. Campbell and R. Przybylski (ed.) Proc. 7th Int. Symp. Buckwheat, Winnipeg, Canada. 12–14 Aug. 1998.
• Garber, R.J., and K.S. Quisenberry. 1927. The inheritance of length of style in buckwheat. J. Agric. Res. 34:181–183.
• Murashige, T., and F. Skoog. 1962. A revised medium for rapid growth and bioassays with tobacco tissue cultures. Physiol. Plant. 15:473–497.[CrossRef]
• Nettancourt, D. de. 1977. Incompatibility in Angiosperms. Springer-Verlag, Berlin.
• Ohsako, T., S. Fukuoka, H.P. Bimb, K. Baniya, Y. Yasui, and O. Ohnishi. 2001. Phylogenetic analysis of the genus Fagopyrum (Polygonaceae), including the Nepali species F. megacarpum, based on nucleotide sequence of the rbcL-accD region in chloroplast DNA. Fagopyrum 18:9–14.
• Ohnishi, O. 1995. Discovery of new Fagopyrum species and its implication for the studies of evolution of Fagopyrum and of the origin of cultivated buckwheat. p. 175–190. In T. Matano and A. Ujihara (ed.) Proc. 6th Int. Symp. Buckwheat, Shinshu, Japan. 24–29 Aug. 1995.
• Ohnishi, O., and N. Asano. 1999. Genetic diversity of Fagopyrum homotropicum, a wild species related to common buckwheat. Genet. Resour. Crop Evol. 46:389–398.[CrossRef]
• Ruszkowski, M. 1990. The phenomenon of dominance, competition and overcompensation in buckwheat productivity. Fagopyrum 10:5–6.
• Saknarov, V.V. 1946. Inheritance of the characters of heterostyly and the system of breeding in autotetraploid buckwheat. Rep. Acad. Sci. USSR 52:259–262.
• Samimy, C., T. Bjorkman, D. Siritunga, and L. Blanchard. 1996. Overcoming the barrier to interspecific hybridization of Fagopyrum esculentum with Fagopyrum tataricum. Euphytica 91:323–330.[CrossRef]
• Sharma, K.D., and J.W. Boyes. 1961. Modified incompatibility of buckwheat following irradiation. Can. J. Bot. 39:1241–1246.
• Wang, Y.J., and C. Campbell. 1998. Interspecific hybridization in buckwheat among Fagopyrum esculentum, F. homotropicum, and F. tataricum. p. I:1–13. In C. Campbell and R. Przybylski (ed.) Proc. 7th Int. Symp. Buckwheat, Winnipeg, Canada. 12–14 Aug. 1998.
• Wang, Y.J., Scarth, R., and Campbell, C. 2002. Comparison between diploid and tetraploid of Fagopyrum homotropicum in intraspecific and interspecific crossability and cytological characters. Fagopyrum 19:23–30.
• Woo, S.H., T. Adachi, S.K. Jong, and C.G. Campbell. 1999. Inheritance of self-compatibility and flower morphology in an inter-specific buckwheat hybrid. Can. J. Plant Sci. 79:483–490.
• Yasui, Y., and O. Ohnishi. 1998. Phylogenetic relationships among Fagopyrum species revealed by the nucleotide sequences of the ITS region of the nuclear rRNA gene. Genes Genet. Syst. 73:201–210.[Medline]
• Yasui, Y., T. Ohsako, and O. Ohnishi. 1998. Evolutionary processes of Fagopyrum inferred from the molecular phylogenetic analyses. p. VI 50–60. In C. Campbell and R. Przybylski (ed.) Proc. 7th Int. Symp. Buckwheat, Winnipeg, Canada. 12–14 Aug. 1998.
• Zamir, D., and Y. Tadmor. 1986. Unequal segregation of nuclear genes in plants. Bot. Gaz. (Chicago) 147:335–358.

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Crop Science 2005 45: vii. [Full Text]  

This Article Abstract Figures Only Full Text (PDF) Free Alert me when this article is cited Alert me if a correction is posted Services Related articles in Crop Science Similar articles in this journal Similar articles in Web of Science Alert me to new issues of the journal Download to citation manager Citing Articles Citing Articles via Web of Science (1) Citing Articles via Google Scholar Google Scholar Articles by Wang, Y. Articles by Campbell, C. Search for Related Content PubMed Articles by Wang, Y. Articles by Campbell, C. Agricola Articles by Wang, Y. Articles by Campbell, C. Related Collections Other Grain Crops Crop Genetics