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D Genome Donors for Aegilops Crassa (Ddmcrmcr, Ddd2d2mcrmcr) and Ac

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D Genome Donors for Aegilops Crassa (Ddmcrmcr, Ddd2d2mcrmcr) and Ac Jpn. J. Genet. (1982) 57, pp. 349-360 D genome donors for Aegilops crassa (DDMCrMCr, DDD2D2McrMcr) and Ac. vavilovii (DDMCrMCrSPSP)deduced from esterase analysis by isoelectric focusing1) BY Yasuo NAKAI Laboratory of Genetics, Faculty of Agriculture, Kyoto University, Kyoto 606 (Received February 15, 1982) ABSTRACT Putative D genome donors to Aegilops crassa (2n = 4x = 28, DDMCrMCrand 2n=6x=42, DDD2D2MCrMcr)and Ae. vavilovii (2n=6x=42, DDMCrMCrSPSP) were studied for esterase isozyme similarities. Esterase isozymes examined by gel isoelectrof ocusing reveal no inter- or intraspecific variation in Ae. crassa and Ae, vavilovii. The putative parents, Ae. squarrosa (2n=2x=14, DD) and Ae, comosa (2n=2x=14, MM), each exhibit three isozyme phenotypes. The natural tetraploid form of Ae. crassa has an isozyme pattern correspond- ing to a mixture of esterases from type 2 squarrosa and type 1 comosa. We conclude from this that the D genome donor to Ae. crassa 4x is type 2 squar- rosa. The hexaploid form of Ae. crassa possesses the same D genome as the tetraploid form, suggesting a duplication of the type 2 squarrosa D genome. Moreover, the D genome of the hexaploid form is the same as the D genome of Triticum aestivum cv. Chinese Spring (bread wheat, 2n=6x=42, AABBDD). It was also found that the putative D genome donor to Ae. vavilovii corres- ponds to type 2 squarrosa. 1. INTRODUCTION Aegilops crassa Boiss. is both tetraploid (2n = 28, genome constitution DDM~rM~r)and hexaploid (2n=42, DDD2D2MCrMCr) This species is assumed to have originated as hybrids between the two diploid species Ae. squarrosa L. (2n =14, DD) and Ae, comosa Sibth et Sm. (2n =14, MM), or Ae, uniaristata Vis. (2n =14, MUMU)(Kihara 1954, 1963). The tetraploid plants are widely distributed from Central Asia to the Near East through Western Asia, while the hexaploid plants are distributed in Tadjik, Uzbek, and Turkmenian SSR of USSR (Zhukovskyi 1928; Jaaska 1981) and the northern province of Afgha- nistan (Kihara et al. 1965) . Drs. H. Kihara and K. Yamashita collected hexa- ploid forms in Pul-i-Khumri, Maimana, and Laman, Afghanistan, in an expedi- tion to the Karakoram and Hindukush in 1955, but they did not find hexaploids in the southern part of Afghanistan. Ae. vavilovii (Zhuk.) Chen. is a hexaploid. This species formerly was clas- 1) Contribution from the Laboratory of Genetics, Faculty of Agriculture, Kyoto University, Kyoto No. 440. 350 Y. NAKAI Fig. 1. Spikes of Ae. crassa 4x (1), Ac. crassa 6x (2), and Ac. vavilovii (3). sified as a member of the Ae. crassa complex (Ae, crassa Boiss. var. palestiana Eig), but recently it has been reclassified as Ae. vavilovii (Zhuk.) Chen., and separated from the crassa complex, since its geographical distribution and cytogenetic characteristics differ from those of the hexaploid Ae. crassa of Afghanistan (Chennaveeraiah 1966). Kihara (1963) has suggested that Ae. vavilovii has a genome constitution of DDM~rM~rS1S1(DDMCrMCrSPSP, after Kihara and Tanaka 1970). The plants are distributed intermittently from Northern Iraq to Jordan through Syria. The putative parents, Ae. squarrosa and Ae, comosa, or Ae. uniaristata, are not found in these areas, but another putative parent, Ae. longissima (2n =14, S1S1),is found in the Palestinian area, that overlaps Ae. vavilovii. The morphological characteristics of the tetraploid and hexaploid forms of Ae. crassa, and Ae. vavilovii are recognizably unique for each species (Fig. 1). The apical upper margin of the empty glume of Ae. crassa 4x has been sug- gested to have its origin in the M genome, while the truncate upper margin of the empty glume can be traced to the D genome of Ae. squarrosa (Kihara 1954; Kihara et al. 1965). Both species, Ae. crassa and Ae. vavilovii, have the D genome commonly found in Ae. cylindrica Host (2n=28, CCDD) and Triticum aestivum L. (2n=42, AABBDD). However, zymogram patterns of Esterase isozymes, Ae. crassa, Ae. vavilovii 351 esterase isozymes among the Ae. squarrosa lines show that T. aestivum (bread wheat) has the D genome found in type 2 squarrosa (Nakai 1979), and that Ae. cylindrica has the D genome found in type 3 squarrosa (Nakai 1981). In the present study, strains of Ae. crassa and Ae. vavilovii from different localities were investigated for esterase isozymes. Based on the results, pos- sible donors of the D genomes and other genomes to Ae, crassa and Ae. vavilovii are discussed. 2. MATERIALSAND METHODS Materials: The strains of Aegilops crassa Boiss. of the tetraploid form (2n=28, genome constitution DDMCrMCr),that of the hexaploid form (2n=42, DDD2D2McrM~r),and Ae. vavilovii (Zhuk.) Chen. (2n=42, DDMcrM~rSPSP)used in the present study are shown in Table 1. All of these strains were kindly provided by Dr. M. Tanaka, the Plant Germplasm Institute of Kyoto Univer- sity, Kyoto. For comparison, strains of their putative parental species, Ae. squarrosa L. (2n=14, DD), Ae. comosa Sibth et Sm. (2n=14, MM), Ae. heldrei- chii Holzm. (2n=14, MM), Ae. uniaristata Vis. (2n=14, M°MU), Ae. longissima Schw. et Musch (2n=14, S1S1), Ae, sharonensis Eig (2n=14, S'S'), Ae. searsii Kislev et Feld. (2n =14, SSSS),Ae. bicornis (Forsk.) Jaub. et Sp. (2n =14, S"S"), were also used. Some of the strains were provided by Dr. Tanaka, and five strains of the M genome group were provided by Dr. J. G. Waines, from the University of California, Riverside, California, USA. The seeds of Ae. longis- sima and Ae, searsii were a gift from Dr. M. Feldman of the Weizman Insti- Table 1. Number of strains of Aegilops crassa (4x and 6x) and Ae. vavilovii collected from different countries 352 Y. NAKAI Fig. 2. Isoelectric focusing of esterase isozymes of Ae. crassa and Ae. vavilovii: (1) Ac. crassa var. macrothera 4x (KUSE 2316), (2) Ae. crassa var, glumiaristata 6x (KUSE 2309), and (3) Ae. vavilovii (BEM 2435). tote of Science in Rehovot, Israel. Ae. searsii is a new species, morpholo- gically similar to Ae. longissima. Methods: For esterase isozyme analysis, the disc (5WX 80h mm) and vertical (160WX 120h X 2d mm) isoelectric focusing method were used. Seeds were placed in a germination chamber for 24 hr at 23° C. Enzyme extracts were prepared by homogenizing the germinated seeds in a glass mortar in 1 ml (per one grain about 15-20mg) of 0.05M cold potassium phosphate buffer (p117.0). The extracts were centrifuged at about 20,000 X g for 20 min at 0°C. The supernatant was placed on a polyacrylamide gel containing a carrier ampholite (KLB producter AB) with a pH range of 6.0 to 8.0. The anode vessel on the top and the cathode vessel on the bottom were filled with 0.02 M hydrochloric acid and 0.02 M ethylenediamine, respectively. The electric current was stabilized at 250 V and was maintained for 3 hr. Gels were removed from the tubes or the glass board, and stained with 0.2/ Fast Blue RR Solt and 0.02% a-naphthyl acetate (w/v) in a phosphate buffer (pH 7.0,1/15 M) for 20-30 min. The homo- logy of the esterase isozymes was determined by using a mixture (1:1 by weight) of esterase extracts. Marker proteins were also used for estimation of pH values at different points on the gels. Esterase isozymes, Ae. crassa, Ae. vavilovii 353 Fig. 3. Photograph of esterase zymograms and a schematic drawing of the isozymes for Ae. squarrosa: (1) type 1 (Ae, squarrosa var. strangulata, KUSE 2135), (2) type 2 (Ae, squarrosa var, typica No. 1), and (3) type 3 (Ae. squarrosa var. typica No. 2). 3. RESULTS Zymogram phenotypes of Ae. crassa and Ae. vavilovii (1) Ae. crassa 4x: All 23 strains show identical zymogram phenotypes which consist of four major (3, 5, 6 and 8B), two moderate (2 and 4), and two minor (1 and 17) isozymes (Fig. 2-1). The isoelectric points of isozymes 4 and 5 are close. (2) Ae. crassa 6x: All 12 strains show identical zymogram phenotypes which include the tetraploid phenotype (Fig. 2-2). (3) Ae, vavilovii: All three strains show zymogram phenotypes identical with those of the hexaploid Ae. crassa. A minor difference is seen in the intensity of isozymes 4 and 5 between Ae. vavilovii and Ae. crassa 6x: The intensity of isozymes 4 and 5 of Ae. vavilovii is higher than that of the corresponding isozymes of Ae. crassa 6x (Fig. 2-3). Zymogram phenotypes of the putative parental species (1) Ae. squarrosa: The isozyme patterns of each of the three zymogram phenotypes (types 1, 2 and 3, Nakai 1979, 1981, Fig. 3) show that type 1 has three major isozymes, 3, 5 and 6 (Fig. 3-1), and that type 2 has, in addition to these, one extra major isozyme, 8B (Fig. 3-2), and that type 3 has four major 354 Y. NAKAI Fig. 4. Photograph of esterase zymograms and a schematic drawing of the isozymes for Ac. comosa; (1) type 1 (Ac. comosa var. typica; Greece), (2) type 2 (Ac. comosa var, typica; Greece), and (3) type 3 (Ac. comosa var. typica; Turkey). 6, 8',10 and 12), three moderate (4, 5 and 6'), and two minor (1 and 3) isozymes (Fig. 3-3). Type 1 is found only in one strain. Factorial analysis of these three zymogram phenotypes reveal that one pair of allelic genes is responsible for the difference between each pair of types. (2) Ae. comosa, Ae. heldreichii and Ae. uniaristata: As shown in Fig. 4 and Table 2, three zymogram phenotypes are found in the samples of the Ae.
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