Comparative-Genetic Analysis – a Base for Wheat Taxonomy Revision

Czech J. Genet. Plant Breed., 41, 2005 (Special Issue)

N. P. G��

Institute of Cytology and Genetics of Siberian Branch of Russian Acadeny of Sciences, Novosibirsk 630090, Russia, e-mail: [email protected]

Abstract: Comparative-genetic analysis performed at the same time on cultivated wheat species and their wild relatives permits the definition of an introgressive hybridization strategy and reconstruction of the origin of the taxonomy important genes. The analysis also allows clarification of the origin of wheats and their differentiation into species, thereby proving a basis for a successful revision of Triticum L. genus system.

Keywords: Triticum genus; comparative-genetic analysis; taxonomy

The genus Triticum L. has a diphyletic origin (D��- Random amplified polymorphic DNA analysis �� & K�� 1979). Its includes di- (2n = 14), was used to assess the phylogenetic relationships tetra- (2n = 28) and hexaploid (2n = 42) species, among these five morphological groups of hexa- with the phylogeny of most of them more or less ploid wheat. These results are in agreement with clarified (T�� 1968). The main result of the those based on morphological classification (C�� wheat domestication process was the reconstruc- et al. 2000). As to di- and tetraploid wheat species, tion of rachis and glumes, which has converted no such genes controlling taxonomically important fragile spikes of the wild species into non-fragile, traits has been as yet identified and, accordingly naked grain cultivated species. These traits under- taxonomists usually use a species “radical’, in lie L�� (1753) classification of Triticum. His Vavilov’s terms, i.e. use a trait set with so far un- hexaploid wheat classification was exceptional in known genetic determination. One taxonomically dividing such wheats into species based only on important gene P is found only in T. polonicum. differences in awned-awnedlessness, but also in The poorly studied genetics of di- and tetraploid spring vs. winter growth habit. Later researchers wheats is the main weakness in M�� (1989) have demonstrated that the traits underlying the genetic classification of Triticum. Despite this the classification of Linnaeus and his followers are idea of developing a genetic classification of the controlled by a small number of oligogenes that genus Triticum has not been abandoned. The goal lead to formation of their hexaploid wheat species. of the current study was to disclose and identify There are four such taxonomically important genes genes providing taxonomic clues for the classifica- and, on their basis, hexaploid wheat genotypes may tion of Triticum species and to produce a revised be listed as (S�� & R�� 1961): classification.

T. spelta L. – qq cc SS vv T. macha Dek. et Men. – qq CC SS vv MATERIALS AND METHODS T. vavilovii Jakubz. – qq cc SS VV T. compactum Host – QQ CC SS vv Accessions of di-, tetra- and hexaploid wheats T. aestivum L. – QQ cc SS vv from the Institute of Cytology and Genetics (No- T. sphaerococcum Persiv. – QQ CC ss vv. vosibirsk, Russia), the Vavilov Institute of Plant

52 Proc. 5th International Triticeae Symposium, Prague, June 6–10, 2005 Czech J. Genet. Plant Breed., 41, 2005 (Special Issue)

Industry (St-Petersburg, Russian), the Small Grains controlled by differebt way – a pair complementary Collection (Aberdeen, USA) and the Germplasm recessive genes, sc1 and sc2, in tetraploid wheats Plant Institute of Kyoto University (Japan) collec- (G�� 1997). tions were used in the present investigation. Compact spike is also the trait that distinguishes The segregation ratio for each cross was deter- T. compactum Host from other hexaploid wheat spe- mined and compared with the expected segregation cies (D�� & K�� 1979). It is controlled ratio using the chi-square test. by a single dominant gene, C, located on chromosome 2D. This gene is not present in the di- and RESULTS AND DISCUSSION tetraploid wheat genomes. The gene that controls compact spike in hexaploid wheats T. sphaero- The number of genes that control morphological coccum and T. antiquorum is not allelic to the gene traits, which are simultaneously localized in three C of T. compactum (Table 1). We identify it as C2. wheat genomes, is small (G�� 2002). There The origin of the dominant genes C and C2 in hexa- are no complete genetic collections of the species ploid wheats is unknown because compact spike for Triticum and Aegilops L. Nevertheless, genetic accessions of Ae. squarrosa, the D genome donor collections provide real opportunities for compara- of hexaploid wheats, have not been discovered tive-genetic investigations in the genus Triticum. (G�� et al. 2002). The C gene may be pres- They are also used for phylogenetic studies. The ent in one of the subspecies of T. macha. drawback of wheat phylogenetic schemes currently Elongated glume. Spike and flower glumes are proposed consists in the absence of data on the very long in Triticum polonicum L. (about 30 mm, character (type) of genetic control of the morpho- on the average); in durum wheats they are about logical traits that are identical in their phenotypic 10 mm-long. It is a frequently studied trait because manifestation. These external morphological traits it is correlated with the valuable agronomic trait are very often arbitrary. ‘long grains’. In T. polonicum, the long glumes are Extremely dense or compact spike. Naked grain controlled by a single dominant gene (Table 2). This and compact spike are the main traits used to dis- dominant gene P is localized on chromosome 7A. tinguish T. sinskajae A. Filat. et Kurk. from T. mono- Other tetraploid wheat species T. ipahanicum Heslot coccum L. The every traits is under the control of possesses a non-allelic gene P2. Long glumes also independent single recessive genes (K�� occur in hexaploid wheat T. petropavlovskyi Udacz. et 2005). As that time the compact spike character is Migusch., but it is more weakly manifested than in

Table 1. Allelic test of genes controlling compact spike in T.compactum, T.sphaerococcum and T.antiquorum

Segregation into spike shape χ2 value for ratio* Cross combinations in F2 generation Compact Normal 3:1 15:1 Vakka T.compactum × K-20900 T.aestivum 47 19 0,51 57,22 Vakka T.compactum × K-56397 T.antiquorum 79 31 0,59 90,30 K-23790 T.sphaerococcum × CI 3090 T.compactum 105 17 7,97 1,86

*Values for signiﬁcance of P = 0.005 is 3.84

Table 2. F2 segregation for glume shape in T.polonicum

2 Segregation into glume shape in F2 generation χ value for ratio* Cross combinations long glume short glume 3:1 15:1 Gaza × T. polonicum 189 60 0,11 135,35 BS2E × K-39297 T. polonicum 63 30 2,61 107,36

*Values for signiﬁcance of P = 0.005 is 3.84

Proc. 5th International Triticeae Symposium, Prague, June 6–10, 2005 53 Czech J. Genet. Plant Breed., 41, 2005 (Special Issue)

Table 3. F2 segregation for spelta in hexaploid wheat

2 Segregation into spike shape in F2 generation χ value for ratio* Cross combinations Spelta Norma 3:1 15:1 Triple Dirk D1 × K-43376 T. petropavlovskyi 48 24 2.67 90.13 Triple Dirk B1 × K-1731 T. spelta 59 28 2.40 90.86 Masljaninskaya1 × K-1728 T. spelta 122 44 0.20 116.24 3:1 13:3 K-152 × T. spelta 129 28 4.30 0.09

*Values for signiﬁcance of P = 0.005 is 3.84; 1T. aestivum; XW – Xinjiang Wheat

T. polonicum. Long glume was observed in offspring T�� (2001) demonstrated that the wild of a F1BC1 K-44126 T. petropavlovskyi × K-20538 tetraploid species Triticum dicoccoides Körn. has T. spelta cross. The glume was long in 25 plants a dominant gene controlling spelt characters in and short in 33, thereby proving that long glume is this species. indeed under the control of single dominant gene. Spelt accessions have not been described for W�� and I�� (2002) have demonstrated diploid wheats. However, Ae. speltoides Tausch, the that genes controlling long glumes are allelic in B (G) genome donor of polyploid wheat, possesses T. petropavlovskyi and T. polonicum. Triticum pet- the spelt-gene. However, its inheritance cannot be ropavlovskyi shows no interspecific polymorphism studied, because non-spelt accessions are absent for grain storage proteins (W�� et al. 2004). in the species. Thus, it appears very likely that T. petropavlovskyi is a man-made synthetic wheat. CONCLUSION Awnlessness. No awnless accessions are known for diploid Triticum urartu Thum. ex Gandil., T. mo- Wheat classification reflects the presence of three nococcum, or T. boeoticum Boiss. (D�� & K�- ploidy levels in the genus. This fact was revealed at th �� 1979; G�� 2002). Triticum sinskajae the beginning of the 20 century. Nevertheless, divi- is the sole diploid wheat species characterized by sion of wheat species by their ploidy did not always awnless spikes; the trait is controlled by a recessive underlie the assignment of the species to sections gene (K�� 2005). However, awnlessness is (see, for example, D�� & K�� 1979). Our under the control of a dominant gene in the diploid idea was to identify wheat species according to the species Aegilops longissima Schweinf. et Muschl. of subordination principle inferred from their compara- sect. Sitopsis (G�� 2002). tive value and genetic control pa�ern. A single or a There are no awnless accessions in tetraploid spe- few taxonomically important traits were assigned cies of non-hybrid origin (D�� & K�� domination in the classification. Taken together, the 1979; G�� 2002). results allowed to us assign the 29 wheat species to A new dominant gene Hd2 that controls awnless- 5 sections (G�� 2002 with additions): ness has been detected in common wheat of Asian I. Sectio Monococcon Dum. includes 4 diploid origin (G�� 2002). (2n = 14) species, T. urartu Thum. ex Gandil., Spelt. Despite frequent studies, spelt has not T. boeoticum Boiss., T. monococcum L. are hulled been accurately defined. In together spelt form of and T. sinskajae A. Filat. et Kurk. is naked; ear, the other correlated traits are inherited as a II. Sectio Dicoccoides Flaksb. includes 10 tet- monogenic dominant addition characters, i.e. brittle raploid (2n = 28) species, T. dicoccoides (Körn. rachis and hulled glumes. The results for studies ex Aschers. et Graebn.) Schweinf., T. dicoccum on the spelt traits in a few wheat species are shown (Schrank) Schuebl., T. karamyschevii Nevski, in Table 3. The trait is controlled by non-allelic T. ispahanicum Heslot are hulled and T. turgidum genes in T. petropavlovskyi and T. spelta. L., T. durum Desf., T. turanicum Jakubz., T. po- Spelt accessions have not been described in lonicum L., T. aethiopicum Jakubz., T. carthlicum tetraploid wheats (M�� 1986). However, Nevski are naked;

54 Proc. 5th International Triticeae Symposium, Prague, June 6–10, 2005 Czech J. Genet. Plant Breed., 41, 2005 (Special Issue)

III. Sectio Triticum includes 7 hexaploid (2n = 42) G�� N.P. (1997): Comparative-genetic analysis species, T. macha Dekapr. et Menabde, T. spelta of common wheat tetraforms with eliminated D ge- L., T. vavilovii (Thum.) Jakubz. and man-made nome. Russian Journal of Genetics, 33: 549–552. synthetic T. dimococcum Schieman et Staudt are G�� N.P. (2002): Comparative genetics of Wheats hulled and T. compactum Host, T. aestivum L., and their Related Species. Siberian University Press, T. sphaerococcum Perciv. are naked; Novosibirsk. (In Russian with English Summary) IV. Sectio Timopheevii A. Filat. et Dorof. includes G�� N.P., K�� E.Y�., K�� T. 2 tetraploid (2n = 28) species, T. araraticum Jakubz. (2002): Inheritance of dense spike in diploid wheat and T. timopheevii Zhuk. and hexaploid (2n = 42) and Aegilops squarrosa. Hereditas, 137: 96–100. species, T. zhukovskyi Menabde et Erizjan. All K�� E.Ja. (2005): Genetics of diploid wheats: are hulled; I. Triticum sinskajae. In: Proc. 8th Genetic-Breeding IV. Sectio Compositum N.Gontsch. includes tet- School. So RASKhN, Novosibirsk (In press). (In raploid (2n = 28) species, T. erebuni Gandil. (syn. Russian) T. palmovae G. Ivanov), hexaploid (2n = 42) species, L�� C. (1753): Species plantarum, exhibentes plan- T. kiharae Dorof. et Migusch., 2 octoploid (2n = tas rite cognitas, ad genera relatas, cum differentiis 56) species, T. soveticum Zhebrak, T. flaksbergeri specificus, nominibus trivialibus, synonymis selectis, Navr. and decaploid (2n = 70) species, T. borisii locis natalibus, secundum systema sexuale digestas. Zhebrak. T.1.: Impensis Laurentii Salvii, Holmiae. 560 s. It is hoped that our revision proposed within the M�� J. (1989): Genus Triticum and its systematics. framework of classical taxonomic traditions has In: S�� V.K. (ed.): Vavilov’s Legacy in Modern fewer drawbacks than the previous classifications. Biology. Nauka, Moscow, 170–185. (In Russian) It is also hoped that the proposed revision would M�� М. (1986): The vulgare super gene, Q: its promote conservation of the man-made wheat universality in durum wheat and its phenotypic ef- synthetics in the world gene banks. Currently, fects in tetraploid and hexaploid wheats. Canadian the proposed classification is being checked by Journal of Genetics and Cytology, 28: 30–41. molecular genetic analysis. S�� M.S., R�� M.V.P. (1961): Macro-mutations and sub-specific differentiation in Triticum. Wheat Acknowledgements. We are grateful to Dr. H. B�� Information Service, 13: 9–11. (the National Small Grains Collection, Aberdeen, USA), T�� H. (2001): Production of near-isogenic lines Dr. T. K��a (Graduate School of Agriculture of Kyoto and marked monosomic lines in common wheat Univ., Kyoto, Japan), Drs. R.A. U�� and N.A. A�� (Triticum aestivum) cv. Chinese Spring. Journal of (N.I. Vavilov All-Russian Institute of Plant Industry, St-Pe- Heredity, 93: 254–259. tersburg, Russia) for supplying seeds of wheat species and T�� K. (1968): Origin and phylogenetic differ- a few synthetic amphiploids. The research was partially entiation of common wheat revealed by compara- rd financed on the Subprogram II of Program basic research tive gene analysis. In: Proc. 3 Int. Wheat Genet. N25 of the Russian Academy of Sciences “The Origin and Symp., 71–85. Evolution of Biosphere”. W�� N., I�� I. (2002): The inheritance and chromosomal location of a gene for long glume phe- References notype in Triticum petropavlovskyi Udacz. et Migusch. Journal of Genetics and Breeding, 57: 221–227. C�� W., S�� G., H�� P., C�� R.N. (2000): Phy- W�� N., B�� S.V., G�� N.P. (2004): logenetic relationships of five morphological groups Inheritance and chromosomal location of the gene of hexaploid wheat (Triticum aestivum L. em Thell.) for long glume phenotype found in Portuguese land- based on RAPD analysis. Genome, 43: 724–727. races of hexaploid wheat, “Arrancada”. Journal of D�� V.F., K�� O.N. (eds) (1979): Wheat. Genetics and Breeding, 58: 273–278. Kolos, Leningrad. (In Russian)

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