ITEMS FROM THE UKRAINE

YURJEV PLANT PRODUCTION INSTITUTE

National Centre for Plant Genetic Resources of Ukraine, Moskovs'kiy pr., 142, Kharkiv, 61060, Ukraine.

Using synthetic hexaploids with the ABD genomes in crosses with bread wheat.

N.P. Novoseltseva.

The genetic diversity of Ae. tauschii in combination with various wheat genotypes is of scientific and practical interest. The SH wheats created by hybridization with different cultivars and lines of tetraploid wheat with Ae. tauschii are being used even more often. Crossing these synthetic wheats with bread wheat may successfully transfer genes responsible for economically valuable characters.

In 1995, CIMMYT kindly gave 99 SH lines to the National Centre of Plant Genetic Resources of Ukraine. Seeds of 24 of the lines were viable. The SHs are of spring growth habit. For morphological traits, the lines are typical speltoid, with an ABD-genome structure. Field evaluation of the lines was according to the recommendations of the VIR. Resistance to disease was made on a 1-9 scale, where the 9 meant complete immunity and 1 indicated complete susceptibility. The bread wheat cultivar Kharkivs'ka 6 served as the standard check.

The lines were studied between 1996-99. Environmental conditions were different for each year. In droughty years (1996, 1998, and 1999), plants were strongly effected. Increased rainfall in 1997 favored development of fungal diseases, and resistant lines were identified. The following lines were resistant (IT = 7-9) to powdery mildew, Septoria, and leaf rust: D67.2/P66.270//Ae. tauschii (217), DVERD 2/Ae. tauschii (221), DOY 1/Ae. tauschii (515), D67.2/P66.270//Ae. tauschii (257), and Green/Ae. tauschii (458); to leaf rust and Septoria, DOY 1/Ae. tauschii (515); and to powdery mildew and Septoria, Arlin/Ae. tauschii (283). On an individual disease basis, we identified lines with resistance to powdery mildew (CETA/Ae. tauschii (174) and CETA/Ae. tauschii (1024)); leaf rust (DVERD 2/Ae. tauschii (1027) and CETA/Ae. tauschii (1027)); Septoria (68.112/WARD//Ae. tauschii (369), 168.111/RGB-U//WARD/3/Ae. tauschii, and 68.111/RGBU//WARD RESEL/3/STIL/4/).

Concurrently, the check Kharkivs'ka 6 had ITs of 3 to powdery mildew and 5 to leaf rust and Septoria. In the severe drought of 1999, the 1,000-kernel weight of Kharkivs'ka 6 was 39.1 g. Among the SHs, only one line (DVERD 2/Ae. tauschii (1027) exceeded the standard with a 1,000-kernel weight of 45.4 g. Spike lengths were from 3.5 to 10.5 cm (average 7.0 cm). Accordingly, the number of spikelets/spike was from 8 to 13 (average 10.5), grain weight/spike ranged between 0.19-1.12 g (average 1.0 g), and grain number/spike was 4-28 (average 16).

With the purpose of using the genetic potential of the SHs, we included them in crosses with Kharkivs'ka 6 in 1996. Seed set was high, 95-97 %. In 1997, 30 spontaneous hybrids with bread wheat were identified for some SHs. In 1999, 690 F3 artificial and 160 spontaneous hybrids were grown and analyzed for a complex of characters. As a whole, they were grouped in 72 morphological types; six were semi-speltoid, and the others were typical of bread wheat.

Spike length of the hybrids ranked from 4.2-13.0 cm (average 7.0 cm) and spikelet number/spike from 10-21, exceeding the respective parameters of both the Shs and Kharkivs'ka 6. Grain weight/spike ranged from 0.10 to 2.18 g (average 1.0 g), and the number of grains/spike was 4-60 (average 32). Grain number /spike of a majority of the hybrids was higher than that of the SHs, and seed set, as a rule, was in the 1st and 2nd florets of the spikelets in the upper and lower parts of an spike. In the SHs, the seed set was in three florets of the spikelet and was distributed more or less evenly along the length of the spike.

For 1,000-kernel weight among the hybrids, the line 262-9 (a descendant of D67.2/P66.270//Ae. tauschii (217) stood out at 51.3 g; eight samples exceeded the check but not significantly, and others were either equal to or less than the check. All hybrid samples were equal to Kharkivs'ka 6 for grain filling.

One of the main wheat attributes and the purpose of our work is grain quality. Analyses of the protein contents in the grains of F3 plants and parentals have shown that this value is higher than that of the standard Kharkivs'ka 6 both in the SHs (16.2-23.2 %) and in the majority of hybrids (15.1 %). Some outstanding plants were identified with high sedimentation values. In the SH parental lines, sedimentation values were between 38 and 55 mm; Kharkivs'ka 6 had a value of 61 mm. The hybrids ranged from 52 to 85 mm; they appear to be transgressive for this attribute. Samples 244-5 (from the SH 68.112/WARD//Ae. tauschii (369) and 251-1, 253, and 254-1 (from D67.2/P66.270//Ae. tauschii (217) with sedimentation parameters from 80 to 85 mm especially stood out. The first two typical bread wheats, whereas the last two were intermediate between bread wheat and spelta. Other samples were equal to the standard for this parameter. The best hybrid for practically all parameters was from the cross of D67.2/P66.270//Ae. tauschii 217 and Kharkivs'ka 6, numbering 60 plants. Among the hybrids, we found samples with thick straw walls or half executed straw (241-1, 241-2, and 241-3 from CROC 1/Ae. tauschii 517//Kharkivs'ka 6; 243-2, 255-2, and 255-3 from D67.2/P66.270//Ae. tauschii 217/3/Kharkivs'ka 6; and 248-1 and 248-2 from Arlin/Ae. tauschii 283/3/Kharkivs'ka 6); executed lower internodes (248-5 from Arlin/Ae. tauschii 283//Kharkivs'ka 6), and in some cases wholly executed straw (255-1 from D67.2/P66.270//Ae. tauschii (217)/3/Kharkivs'ka 6). These are valuable traits that determine resistance to the Cephus pest, which has become harmful in the eastern Ukraine in last 2 years.

Our work with the hybrids will be continued in 2000.

The author acknowledges with thanks the scientists of CIMMYT, which created the described SHs and kindly gave them to us, and also for the help of researchers E.M. Dolgova, L.V. Rogulina, and R.G. Parkhomenko, scientists of Institute for Plant Production V.Ja. Yurjev.

The evaluation of the ecological adaptability of spring bread wheat.

O.V. Golik, I.A. Panchenko, and V.P. Kolomatskaya.

Creating varieties with a wide ecological adaptability is one of the most important aims in plant breeding. However, selection in the early phases of the breeding process is not greatly effective mainly because of a lack of traits. A few cultivars sown over a wide area over several decades, Senatore Capelli, Saratovskaya 29, and Kharkovskaya, may
provide evaluation of ecological adaptability based on grain yield in contrasting years or on varietal testing in geographical areas with the different soil-climatic conditions. Quantitative assessment of a variety's ecological adaptability is based on the coefficient of variation and regression on indices of the ecological conditions (Eberhart and Russel 1966; Litun 1979). All these parameters are from a one-dimensional analysis.

A systemic approach using multidimensional space metrics is necessary for the integral evaluation of biological traits and evaluation of the adaptive potential of a variety. Picking traits is important for interpretation of the initial testing.

We analyzed the bread spring wheats Kharkovskaya 93, Kharkovskaya 6, Kharkovskaya 18, Kharkovskaya 26, Kharkovskaya 28, Kharkovskaya 30, Kollektivnaya 5, and Saratovskaya 29 and the promising lines 91583 and 91-689 in between 1995 and 1999 under the climactic conditions of eastern Ukraine (Kharkov). The 17 traits analyzed were crop capacity, cup weight, hardness, raw-gluten content in the flour, gluten quality, gluten type, elasticity, extensibility and balance of dough, flour strength, bread volume from 100 g of flour, common bread rating, common bread-making rating, protein content, common plot rating, 1,000-kernel weight, and grain yield. The humidity varied over the year; 1995 and 1996 were dry; 1997 was extremely damp; and 1998 and 1999 had severe droughts. The plot size was 10 m2. The varieties and lines were sown in experimental field after peas.

Seven genetic mechanisms were determined by means of factor analysis. These factors stipulated the dynamics of the variability (Table 1). The highest value for each factor indicated the major trait or group of traits that control the genetic mechanism. The first factor included traits for baking, including bread volume from 100 g of flour, common bread rating, and common bread-making rating; elasticity and balance of dough. The second factor included traits of dough quality. The third factor included crop capacity and cup weight. Grain yield was the fourth factor; protein content was the fifth factor; extensibility of dough was the sixth factor; and gluten quality was the seventh factor.

Coefficients of ecological plasticity were calculated for each factor. This coefficient is the regression of the factor value on the index of the ecological conditions. The analyzed varieties and lines were sorted into three clusters by means of cluster analysis (Fig. 1). These clusters have different reactions for the ecological conditions. Cluster 1 contains the varieties Kharkovskaya 28, Kollektivnaya 5; cluster 2 contains Kharkovskaya 93, Kharkovskaya 26, Kharkovskaya 30, and lines 91-583, 91-689; and cluster 3 contains varieties Kharkovskaya 6, Kharkovskaya 18, and Saratovskaya 29.

All tested forms had similar reactions to the environment conditions for the majority of traits, except for dough quality. Kharkovskaya 6, Kharkovskaya 18, Kharkovskaya 28, Kollektivnaya 5, and Saratovskaya 29 from the first and third clusters had specific reactions to the environment, which were displayed in dough quality. Evaluation of the initial data revealed reductions of elasticity, extensibility, and balance of dough, indicating dry and very dry years. Qualitative and quantitative indicators showed that varieties Kharkovskaya 93, Kharkovskaya 26, and Kharkovskaya 30 and lines 91-583 and 91-689 from the second cluster were most adapted to the environment of the region by.

Publications.

• Eberhart SA and Russel WA. 1966. Stability parameters for combining varieties. Crop Sci 6(1):36-40.
• Litun PP. 1979. Interaction genotype - environment in the genetic and plant breeding investigations and ways of it study. Problems of selection and evaluation of plant breeding material. Kiev:Science Idea:63-93 (in Russian).

Results CIMMYT durum wheat trial in East Forest-Steppe condition of Ukraine.

O.Yu. Leonov and N.K. Il'chenko.

The National Centre for Plant Genetic Resources of Ukraine annually evaluates durum wheat cultivars and breeding lines from the CIMMYT nurseries. About 800 such samples were introduced in 1995-99. The best of these were studied for 3 or more years by the Group for Plant Genetic Resources of Wheat after preliminary evaluation in a quarantine nursery.

Leaf disease (S. tritici, powdery mildew, and leaf rust) and lodging resistance (scored twice during the vegetative season), spike density/unit area, and grain plumpness were determined and rated on a scale of 1-9 (1 being the worst score and 9 being the best). Duragion of the two vegetative periods, plant height, grain yield/m2, and 1,000-kernel weight also were determined for each. Resistance, protein contents, and yield components were determined selectively.

Meteorological conditions differed in each of the trial years (Table 2). Average daily temperature and precipitation in 1996 vegetative period were near normal. The year 1997 was characterized by late sowing and an abundance of precipitation. High temperatures during the heading period with lack of rain were present in 1998. The 1999 conditions included an elongated period from germination to tillering and extremely high temperatures from heading to maturity.

CIMMYT cultivars and breeding lines differed highly from those registered in the Ukraine. A cluster analysis was done on a sample of 36 cultivars with averages for 11 variables for the 1996-99 season. The minimum Euclidean distance between Kharkivska 37 (the local check) and the CIMMYT sample (Bartramia 1) was 8.82, and the maximum distance of 6.98 was between two CIMMYT samples.

On average, CIMMYT samples were 5 days longer than Kharkivska 37 for the germination-heading period and 2 day shorter for heading-maturity. In the hot years (1998 and, especially, 1999), differences were not so large. Resistance to S. tritici was 1-2 marks less than the check. The maximum pathogen development was highest in 1997, a moist year. Powdery mildew resistance was slightly less on the check, and the CIMMYT material was more susceptible. In the hot and dry years of 1998 and, especially, 1999, the CIMMYT and Ukrainian cultivars had equal degrees of resistance or susceptibility. CIMMYT durum wheat samples were more resistant to leaf rust than most of the Ukrainian and Russian lines, but no significant epidemic of leaf rust occurred during these years. All Mexican cultivars were shorter than the check, on average 46 cm, and resistant to lodging. CIMMYT samples had smaller and less plump grains, but protein contents were higher by 1.7 %. The average grain yield of trial samples was significantly less than that of the check (135 g/m2 and 286 g, respectively). Correlation analysis of the data for all the years showed that varieties with taller plants and greater spike density yielded better. High-yielding varieties had bigger and plumper grain. In most years, cultivars with a shorter period from germination to heading were preferable. Only in the hot weather during the heading period in 1998 did plants with a shorter germination to heading period not perform well.

Some cultivars and breeding lines had greater yields than the CIMMYT checks. Mexicali 75, Altar 84, and Aconchi 89 also had other valuable qualities (Table 3).

Comb Duck 5, May Fowl 2, Om Rabi 5, and Bartramia 1 headed earlier than the other samples and had a longer heading-maturity period. Lavandera 1, Elwag 2, Scooper 1, and Daption 1 were most resistant to S. tritici. May Fowl 2, Yuan Yang 5, Chaika 1, Frailecillo 2, CD98295, and Podiceps 9 were less susceptible to powdery mildew. Altar 84-Alto 1, Ghoo 1, Frailecillo 2, Scooper 1 were resistant to head and loose smuts. The Mexican cultivars did not lodge as much in the Kharkiv conditions, so taller samples were preferable (correllation between plant height and grain yield r = 0.60). Lotus 1 and Cascadena 1 yielded more than Aconchi 89 when they were of the same plant height. May Cock 2, Comb Duck 5, Altar 84-Alto 1, CD72648, Ghoo 1, Dabchick 1, Scooper 1, Lotus 1, and Cascadena 1 had the highest spike density/m2 among the CIMMYT samples. Wagtail, Lavandera 1, SDW2184, Silver 2, and Cascadena 1 had the highest grain weight from the main spike. Jabiru 4, May Cock 2, Bleater 9, Burhinus, and Bartramia 1 had big, plump grains. Aconchi 89, Frailecillo 2, and Scooper 1 contained protein levels above 17 %.

Factor analysis by the maximum likelihood factors method with varimax normalized rotation identified three common factors: productivity (grain yield, spikes density, 1,000-kernel weight, grain plumpness, and plant height variables in descending factor loadings order); vegetation period (days from germination to heading and from heading to maturity); and resistance to powdery mildew (first and second marks). Jabiru 4, May Cock 2, Lotus 1, and Cascadena 1 had the maximum first-factor scores after Kharkivska 37.

Parentage analysis (according to information from the Genetic Resources Information Package for Wheat - Version 1 (GRIP I) and The International Wheat Information System (IWIS), Version 1) show that a majority of the lines have Jori or its descendants Ruff, Flamingo, Yavaros, Sterna, or Altar in their pedigrees. Some new lines with Plata, Silver, Altar, Shag, Lotus, and Rascon in their pedigrees were high yielding in the dry climate of 1999, but further trials are necessary.

Efficiency of chemical treatment of winter wheat seeds under water deficit.

Yu.G. Krasilovets, V.V. Sotnikov, and A.E. Litvinov.

The experimental farm of the Yurjev Plant Production Institute is situated in the southern suburb on the left edge of the forest-steppe region in the Ukraine. In 1998-99, we studied the efficacy of treating winter wheat seed with systemic chemicals using Vitavax 200 FF, 34 % s.t.c. (at the rate of 2.5 l/t); Baytan-Universal, 19.5 % s.p. (at 2 kg/t); Raksil, 2 % s.p. (at 1.5 kg/t); and Sumi-8, 2 % s.p. (at 1.5 kg/t). Water consumption was 10 l/t of seed. The conditions for a pathogen development were natural, lower than the economic threshold. The degrees of plant damage by Helminthosporium-Fusarium root rots were 16.3-21.4 % at spring tillering and -5.8-11.3 % at the waxy-ripe stage. Smut diseases were absent. Tiller damage by stem pests was 9.2-12.7 %. Damage by mandibulate insects on the spikes and stems was 2.4-13.2 %. Chemical treatment was not applied to the control. Field experiments were made under dry conditions. The soil was a common chernosem with a humus layer content of 5.9-6.1 %.

Cultivation of winter wheat is common for areas with unstable water provision. Following a crop of black fallow, the winter bread wheat Kharkiv'ska 96 was sown. The plot size was 26.46 m2, with three replications. The first of the two sowing dates was during the second half of the optimum sowing time and the second planting was at the end. Seeding rate was 4.3 x 10^7^ seed/ha for the first sowing date and 4.9 x 10^7^ seed/ha for the second sowing. Seed was planted at two depths; the optimum of 5-6 cm and 7-8 cm, a depth recommended under dry conditions. The crop overwintered satisfactorily.

The yield level was limited by extremes in climactic conditions. A drought, high air and soil temperatures, and lack of effective rainfall accompanied the periods for sowing and plant growth and development. Hydrothermal coefficient indices during the months of wheat cultivation in relation to the average were 37 % during the month before sowing, 20 % during the month of sowing, 44 % for the month before harvest, and 33 % during harvest. For the period from the start of shooting to waxy-ripeness, rainfall was 95 mm. During the entire spring-summer vegetative period, it was 114 mm or 68 % that of the average yearly rainfall.

The results indicate that under a water deficit during sowing, germination was 81.4 % for untreated seeds (control) sown at the optimal date (12 September, 1998) and at the optimal depth. When sown at the 7-8 cm depth, germination fell to 57.7 %. Germination was much lower, 52.6 % and 40.0 % at optimal and lower depths, respectively, when seeds were treated with Baytan-Universal. In all the experiments, total and productive tillering was close to or above that of the control.

An improvement in soil-water consumption 10 days after the second sowing date (21 September, 1998) produced an increase in germination of the untreated seeds up to 95.8 % in the control and 68.0 % for the deeper planting. Under these conditions, small reductions occurred in germination of seeds with chemical treatment and sowing at the 5-6 cm soil depth and a considerable decrease (up to 45.5 %) occurred with the use of Baytan-Universal at the same depth and in sowings at the 7-8 cm soil depth.

The highest protection against Helminthosporium-Fusarium root rots at the spring tillering was found in seed treated with Baytan-Universal (53.0-77.9 %). Protection ranged from 32.2 to 54.1 % depending on the sowing date and depth of planting in the experiments with Vitavax 200 FF, Sumi-8, and Raksil.

Under a water deficit during sowing and spring-summer vegetative periods, a considerable reduction in grain yield (by 520 kg/ha in comparison with the control) was found only when seed was treated with Baytan-Universal and sown during the second half of the optimal period at a 5-6 cm soil depth (see Table 4). In the rest of the trials, soil planting depth and seed treatment had no effect on yield at either sowing date.

A history of the ancient and modern Ukrainian wheat cultivars used in breeding of the Krasnodar winter wheat cultivars and an analysis of the structure their high-molecular-weight glutenins.

S.V. Rabinovich, O.Yu. Leonov, I.A. Panchenko, R.G. Parkhomenko, Z.V. Usova, and A.A. Kushchenko.

Sources of information on the breeding history of the Krasnodar winter wheat cultivars were breeders' publications (Luk'yanenko 1973; Puchkov et al. 1987, 1996, 1997; Nabokov 1987; Bespalova et al. 1996a, 1996b, 1998; Timopheev 1996; Timopheev et al. 1999; Lysak et al. 1998a, 1998b; Gritsay et al. 1999); personal communication with Luk'yanenko and Yu.M. Puchkov and other breeders; and visits to experimental fields between 1967-99. Pedigrees of the Krasnodar cultivars and their ancestors from Ukraine, Russia, France, Serbia, Bulgaria, Canada, the U.S.A., Mexico, and Australia are from Luk'yane-nko (1973); Rabinovich (1972); Dorofeev et al. (1976); Martynov et al. (1990); Pushkov et al. (1996); Zeven at al. (1976, 1991); and some others. The HMW-glutenin composition of the Krasnodar and other Russian and Ukrainian varieties were deduced according to data from the Grain Quality Department at our institute and partially published in Rabinovich et al. (1997, 1998) and also in Morgunov et al. (1990, 1992) and Sobko (2000). Data for cultivars from other countries are according to Branland et al. (1985); Vapa et al. (1988, 1995); Lukow et al. (1989); Graybosch (1992); CIMMYT (1997); Bushuk (1998); and Ivanov et al. (1998). The gliadin composition of some of the Krasnodar cultivars are in accordance with Timopheev (1996), Timopheev et al. (1999), and Lysak et al. (1998b).

According to Payne et al. (1997), an index of quality (quality score (QS)) is defined for each subunit or subunit group for the HMW glutenins. In this report, high positive correlations were detected between the presence of subunits 1 or 2* (QS = 3) at locus Glu-A1; 7+8 (QS = 3), 7+9 (QS = 2) and/or 13+16 (QS = 3), 17+18 (QS = 3) at locus Glu-B1; and 5+10 (QS = 4) in locus Glu-D1 and high baking properties (Lukow et al. 1989, Bushuk 1998, and others). Thus, QS = 10 in wheats with alleles 1 or 2 *, 7+8 or 17+18 or 13+16, and 5+10; QS = 9 in wheats with 1 or 2 *, 7+9, 5+10; QS = 8 in wheats with N, 7+8, or 17+18 or 13+16, 5+10; QA = 7 in wheat with 1 or 2 *, 6+8 or 7 or 20, 5+10; N, 7+9, 5+10; or 1 or 2, 7+9, 2+12; QS = 6 in wheats with N, 6+8 or 7 or 20, 5+10; or 1 or 2 *, 6+8 or 7 or 20, 2+12; QS = 3-5 in wheats with N, 7+9, 2+12; and QS = 1 in cultivars having 2 or 3 alleles.

Winter wheat breeding at Krasnodar began in 1920 by V.S. Pustovoit (academician and future expert in sunflower breeding) and in 1930 by P.P. Luk'yanenko (Luk'yanenko 1973, Bespalova et al. 1996b). Ukrainian cultivars or its descendants were used for breeding. The Canadian cultivar Marquis (QS = 9, 1 7+9 5+10) and the Australian cultivar Florence were descendants of Red Fife (QS = 9, 2* 7+9 5+10), which was introduced into Canada from Galicia (now the Lviv region, Ukraine). The Canadian cultivar Kitchener (QS = 10, 1 7+8 5+10) and U.S. cultivar Thatcher (QS = 9, 2* 7+9 5+10) were descendants of Marquis. The winter wheat Krymka was taken to the U.S. in 1884 under the name Turkey (QS = 8.5, 1/2 7+8/7+9 2+12/5+10) and used in the breeding of Oro and Kanred (QS = 9). The highly winterhardy wheats Odes'ka (OD) 3 (QS = 9.5) and OD 16 (QS = 9) (see pedigree in Fig. 1) were used in breeding Krasnodar cultivars that were released in the 1960s-70s. The Saratov cultivar Hostianum (HST) 237 (QS = 6.5) , a local variety from Kharkiv through Saratovskaya 3, was the source of high winter hardiness for Krasnodarskaya (KRD) 39 released in the 1970s. Krasnodar cultivars from the 1980s-90s were bred from the OD 16-derived OD 51 (QS = 9.5), OD 66 (QS = 9.5), and Obrij (QS = 9.5) from 1969, 1979, and 1983, respectively. The Ukrainian cultivars Myronivs'ka (MYR) 264, MYR 808, and MYR yuvileina (all with QS = 9) also were used successfully in Krasnodar breeding.

Winter wheat breeding in the 1930s-50s was directed by P.P. Luk'yanenko and was aimed at selection of soft winter wheats that were productive and adapted to the central and southern Kuban (Puchkov et al. 1996). The first crossings of high quality, Canadian, spring wheats Marquis and Kitchener with the winter-type, Krasnodar cultivar Ferrugineum 13 (a selection from a local variety) were made by V.S. Pustovoit in 1925. The winter wheats Pervenets and Krasnodarka were selected from these crosses by P.P. Luk'yanenko and released in the late 1930s. Later, the high quality, Ukrainian wheat Ukrainka (susceptible to brown rust , QS = 9, 1 7+9 5+10) was crossed with Marquis (QS = 9). From this cross, Novoukrainka 83 (QS = 9, 1 7+9 5+10) and Novoukrainka 84 (a selection from Novoukrainka 83) were selected and released in 1945 and 1953, respectively. These cultivars were strong (high quality) and resistant to leaf rust.

Skorospelka (SKRP) 1 and SKRP 3 (both QS = 6, N/2* 7+9 2+12) were selected from crosses made by Luk'yanenko in the 1930s between the U.S. line 266287 (Kanred (QS = 9, 2* 7+9 5+10)/Fulcaster (QS = 9, 1 7+8 2+12/5+10)) and the Argentinian spring wheat cultivar Klein 33. Skorospelka 1 and SKRP 3 were released in 1952 and 1955, respectively. An unreleased variety SKRP 2 also was obtained later from this cross. The more productive and earlier maturing SKRP 3B had better bread-making quality (QS = 7, 1/2*.7+9.2+12).

In 1944, Bezostaya (BEZ) 4 (released in 1955) was selected from a cross of the Ukrainian cultivar Lutescens (LUT) 17 (local variety/Ukrainka) with an SKRP 2 elite plant. Repeated, individual selection from BEZ 4 by P.P. Luk'yanenko identified one elite plant from 600, which formed the basis for the new, high-quality cultivar BEZ 1, the most widely spread Ukrainian cultivar in the world. Subunit 2* in BEZ 1 was inherited from SKRP 2, the alleles 5+10 were from Ukrainka or, probably, Krymka through Kanred, and alleles for subunits 7+9 were present in all the BEZ 1 parents. The high-quality alleles of the Ukrainian wheat HMW glutenins were derived from BEZ 1 and its derivatives and are found in a majority of cultivars in many countries in the world. BEZ 1 (QS = 9, 2* 7+9 5+10) was released in 1959 and is still planted in Russia, Ukraine, and some countries of eastern Europe. BEZ 4 and BEZ 1 descendants are the Ukrainian cultivars Krymka and Ukrainka and their offspring.

The most remarkable results in the world of wheat breeding with T1BL·1RS translocation came from the work of P.P. Luk'janenko, which began in the 1950s. In the early 1970s, Aurora (AUR) (QS = 8, N /2* 7+9 5+10), Kavkaz (KVK) (QS = 8, 2*/N 7+9 5+10), and BEZ 2 (all bred from of Neuzuct (pedigree DEU/BEZ 4//BEZ 1, QS = 9); Skorospelka (SKR) 35; and Predgornaja (PRG) 2 (also with the German line Neuzucht in their pedigrees) all were developed under his leadership. The cultivars AUR and KVK were grown widely in most of the former USSR and in many countries of Europe and Asia in the 1970s and 1980s. In 1973, under cultivation on over 2 million hectares in the USSR, they proved susceptible to a new biotype of leaf rust, race 77. Subsequently, the area sown to these cultivars was reduced. Cultivars developed from AUR and KVK were not released in the USSR, with the exception of the Ukrainian cultivars OD 66 and five others and the Belorussian cultivars Nadzeja and Garmoniiya, which were released in late 1970s to the mid-1990s (Rabinovich 1998).

The first Krasnodar derivatives of AUR, KVK, and BEZ 2 after 1973 were Soratsnitsa, Ejka, Yugtina, Delta, Kroshka, Gorlitsa, Ekho, and Kupava, which were released in the Russian Federation between 1993-99. In the 1970s and 1990s, AUR, KAV, SKR 35, BEZ 2, and PRG 2 were used in breeding in a number of countries in Europe, Asia, North and South America, and Oceania. Some are still under production.

Winter-hardy wheat cultivars. Winter-hardy and drought-hardy, winter wheat cultivars bred for steppe regions of the northern Caucasus began in 1947 after the organization of what is now the Northern Kuban branch of the Krasnodar Agricultural Research Institute in the settlement of Leningradskaya (Puchkov et al. 1996). Odesa cultivars that were winter hardy enough for Krasnodar conditions with a qualitative HMW-glutenin composition were used widely in Krasnodar breeding for winter hardiness (see Fig. 2). Krasnodarskaya (KRD) 6 (QS = 8, pedigree: OD 3 (QS = 9.5)/SKRP 3 (QS = 6), N/2* 7+9 5+10) was the first cultivar released by this program in 1967. Derivatives Severokubanka (QS = 6) and KRD 70 (QS = 7) were later important in winter wheat breeding and production. KRD 46 (pedigree: BEZ 1/OD 16//BEZ 1, QS = 9, 2* 7+9 5+10) had a moderate level of winter hardiness and was released in 1976. In the 1980s and 1990s, derivatives of KRD 46, Zamena (QS = 8), Umanka (QS = 9, MYR 808 and MYR Yuvileina in pedigree), and Demetra (QS = 10, MYR 808 in pedigree) were created for special ecological niches and supplemented the widely adapted cultivars (Bespalova et al. 1996b).

Olimpiya 2 (QS = 10, 2* 7+8 5+10), a descendant of the Ukrainian cultivars OD 51 (QS = 9.5) and MYR 808 (QS = 9), is characterized by high winter and frost hardiness, potential productivity up to 9.5 t/ha, and a high intensity of spring growth. Olimpiya 2 is the most winter hardly cultivar among those for late sowing. A new cultivar with a short vernalization period, Delta, is a descendant from Olimpiya 2. Soratnitsa (QS = 9, 1 7+9 5+10, pedigree OD 66 (QS = 9.5)/Partizanka YUG (QS = 9)) is a significant achievement in breeding for high yield, stable productivity, and complex hardiness to abiotic factors and main pathogens (Puchkov et al. 1996). Another new cultivar, Lira Krasnodara (QS = 9, 1 7+9 5+10), was obtained from crossing Soratnitsa with the spring wheat Budimir (QS = 8, 2* 7+8/17+18 2+12). Released in the 1990s, Yuna (QS = 10), Nika Kubani (QS = 9), Rufa (QS = 9.5), and Leda (QS = 9.5) are all descendants of the Ukrainian cultivar Obrij.

To increase winter hardiness in the Krasnodar wheats, Luk'yanenko (1973) selected BEZ 1 for crossing with two Saratov cultivars Lutescens (LUT) 329 (created in 1929 by selection from Polish variety Sandomirka) and Saratovskaya 3 (synonym LUT 319, created in 1955 by crossing HST 237 with wheat-rye hybrid 434-134) and a large number winter hardy Saratov cultivars. Severo-kubanskaya 43 was created from the first cross but was not released. Krasnodarskaya 39 (pedigree: Saratovskaya 3/BEZ 1, QS = 7, N 7+9 5+10) was released in 1973 and was grown (maximum 0.9 x 107 ha) not only in the Krasnodar region, but also in the eastern Ukraine and Volga regions. Krasnodarskaya 39 is still planted in Russia. The cultivar Pavlovka (PVL) (QS = 7.5, 2*/N 7+8/7+9 5+10/2+12) was created from a selection from KRD 39 in 1980. In the 1990s, KRD 39 descendants KRD 90 (QS = 9.5, 2* 7+8/7+9 5+10) and Zimorodok (QS = 8, 2*/N 7+8 2+12/5+10) were released. Quite probably, the HMW-glutenin subunit composition was inherited from the Ukrainian local variety Kharkivs'ka through HST 237. A HMW-glutenin analysis in the Grain Quality Department at our Institute showed a subunit composition of N 7+8/7+9 5+10/2+12 in HST 237 in seed received in 1996 from Saratov. New cultivars from PVL include Russa (QS = 9, 2* 7+9 5+10) and Lepta (a selection from Russa) (Lysak et al. 1998a).

On the basis of KRD 39, a new breeding program was established for producing productive, leaf rust-resistant cultivars with the winter hardiness level of KRD 39 (Puchkov et al. 1996). Severokubanka (QS = 6, N 7+9 5+10/2+12) was selected from KRD 39 (QS = 7) and KRD 6 (QS = 8) crosses and released in 1980. Severokubanka inherited the low-quality alleles N and 2+12 and 7+9 from Russian (HST 237 and SKRP 2) and Ukrainian wheats (Krymka, Kharkivs'ka, and OD 3). Severokubanka is one of a few cultivars with a combination of winter hardiness, drought hardiness, resistance to leaf and strip rusts, and high productivity and was used widely in breeding and production.

Prikubanka and Olimpiya (QS = 9, 2* 7+9 5+10) were created with the use of Severokubanka and other high-quality cultivars (MYR 808, BEZ 1, and KRD 57 (QS = 9.5, 2* 7+8/7+9 5+10). Released in the 1990s, Sfera, Otrada, and Zimorodok (QS = 8-9.5) were KRD 57 derivatives or selected from the line KRD 57-324 (Fig. 3). Winter hardy Ukrainian wheats Kharkivska, OD 16, MYR 808, and Russian Albidum 114 (QS = 10, 1 7+8 5+10) from the Volga region also were in Zimorodok's pedigree.

Winter-hardy and early maturing cultivars. Krasnodar breeders (Nabokov 1987; Puchkov et al. 1997) created some winter wheat cultivars with these combined characteristics. The best from this plan, from our point of view, is KRD 70 (released in 1990, QS = 9, 1/2* 7+9 5+10) (see Fig. 3). The Krasnodar line LUT 319 was obtained by crossing a northern Caucasian wheat Osetinskaya 3, the Ukrainian winter-hardy Ferrugineum 1239 (QS = 8, 1 7+8 2+12), and a Krasnodar early maturing hybrid F2 (pedigree: Novoukrainka 83 (QS = 9)/SKRP 1 (QS = 6)). KRD 70, a Ukrainian wheat (spring Red Fife and winter Krymka and Ukrainka) descendant, is the first cultivar released that was created from the highly winter hardy, but very late-maturing Kharkiv breeding cultivar Ferrugineu 1239 in almost 70 years after its release.

Early maturing wheats of this group include Ejka (QS = 7, pedigree: Donskaya Polukarlikovaya (DPK)/Daritsa, N/2* 7+9 2+12 5+10) and Yugtina (QS = 8.5, pedigree: Zagore BLG/DPK (QS = 9.5)//KRD 57-273 (selected from KRD 57 (QS = 9.5)/Daritsa (QS = 8))), which were released in 1994. Winter and frost hardiness was inherited from MYR 808 (through DPK) for these wheats and from the local variety Kharkivs'ka (through HST 237, KRD 39, and KRD 57) for Yugtina. Early maturity has also come from the Bulgarian wheat Rusalka through DPK and early maturing and frost resistance from Daritsa. The Mexican wheat Siete Cerros (QS = 8) also has Ukrainian varieties in its maternal pedigree (through Turkey and Thatcher) and in its paternal pedigree Donskaya Ostistaya (QS = 9) and the winter-hardy Ukrainian cultivars OD 3 and OD 16. Although not released but of breeding interest, the two sister lines Zimdar (QS = 9) and Volgodar, produced in 1988 and 1990, respectively, inherited early maturity from the Bulgarian cultivar Rubin and high frost resistance from Kharkivs'ka (through KRD 39, which is used two times in their pedigree).

The hybrid combination Erythrospermum 3217h6 was obtained from crossing very early types (Genrumil and Korea/Yugtina) followed by repeated selections in the later generations. A transgressive form (Puchkov et al. 1997) that successfully combined very early maturity with high frost resistance and productivity level and resistance to leaf rust and Septoria was Yugtina (QS = 8.5, 1/2* 6+8/7+9 5+10). Uskoryanka (QS = 6, N 7+8 2+12) also was selected from this combination with the best Glu-1B allele from a Korean cultivar.

Semidwarf cultivars for fallow and irrigation. Breeding of semidwarf cultivars for fallow and irrigation was begun under the guidance of P.P. Luk'yanenko in 1965. Various height reduction sources were involved in the breeding process, such as the Japanese cultivar Norin 10 and it derivatives, Tom Thumb from Tibet, Olesen's Dwarf from Zimbabwe, and other dwarf mutants (Bespalova et al. 1996). However, the best result in winter wheat semidwarf varieties breeding was obtained using the induced mutant BEZ 1 (2* 7+9 5+10)-Karlik 1 (QS = 8.5, 1 7+9/7+8 2+12/5+10) in crossings with widely adapted BEZ 1 derivatives. Several semidwarf varieties were obtained (see Fig. 3) including Polukarlikovaya (PK) 49 (QS = 9, pedigree: MYR Yuvileina, UKR (QS = 9)/Karlik 1 (QS = 8.5)//Rannyaya 12 (QS = 8)); Krinitsa (QS = 9, pedigree: PK 49/Rostovchanka (QS = 9)); Spartanka (QS = 9, sibs Krinitsa/PVL (QS = 7.5)); Birlik (QS = 9, pedigree: PK 49/Yubilei, BLG); Skifyanka (QS = 9, selection from Spartanka), Nika Kubani (QS = 9, pedigree: Obrij, UKR (QS = 9.5)/Krinitsa), and Novokubanka (QS = 9.5, pedigree: Okhtyrchanka, UKR (QS = 9)/PK 49). Spartanka, released in 1986, and Skifyanka, released in 1992, were the most widely spread of these, grown in southern Russia, Ukraine, Transcaucasia, and central Asian countries on up to 2 x 107 ha. In the pedigrees of the above-mentioned cultivars are the high-quality Ukrainian wheats MYR Yuvileina (QS = 9) in PK 49; MYR 264 (QS = 9) in Rostovcanka (QS = 9) in Krinitsa and its derivatives; local variety Kharkivs'ka through SAR 3 and KRD 39 in PVL and its derivatives Spartanka, Skifyanka, and its offspring Obrij in Nika Kubani (QS = 9).

Widely grown since 1992 in southern Russia, Ukraine, and central Asia, the winter wheat Yuna (QS = 10) is an offspring of Ukrainian wheats Obrijand MYR 808 through the Yugoslavian cultivar Novosadska Rana 2. Yuna is interesting also because of its fitness to alternate plant-growing culture. Released between 1995-99, Zhirovka (QS = 9), Kroshka (QS = 9), and Pobeda 50 (QS = 9.5) are widely spread in the Krasnodar region. Kroshka is a MYR 808 descendant. Ukrainian wheats derivatives were used to create Zhirovka (PK 49, Rostovchanka, and PVL), Kroshka (Spartanka), and Pobeda 50 (Skifyanka).

A spontaneous hybrid was selected in a seed-production crop of KRD 39 (N 7+9 5+10) in 1974. The wheat Pavlovka (2*/N 7+8/7+9 5+10/2+12), released in 1982, was selected from this seed. The cultivar was named after Pavel Luk'yanenko. The high productivity of the new cultivar is due to a high degree of tillering. Pavlovka is an early maturing cultivar (4-5 days earlier BEZ 1), short, and resistant to leaf rust and has a much higher protein content. More than 10 varieties were created using Pavlovka and the local variety Kharkivs'ka. In 1988-92, Spartanka, Skifyanka, and Istok (QS = 9.5, with OD 3 and OD 16 in the pedigree) were obtained. Istok is interesting as initial material for breeding for resistance to drought and grain shedding. All three cultivars are grown not only in Russia, but also in the steppe zone of the Ukraine. Kroshka (QS = 9), a PVL derivative, is one of the most productive semidwarf wheats. Using PVL, the cultivars Goritsa (QS = 9) and Ekho (QS = 9) also were created for the mountain and foothill agriculture zones.

Cultivars with a short vernalization period. Connected with changes in the economic situation in the countries of the former USSR in the 1990s, a need developed for cultivars with low agronomical-input levels and intermediate-type development that were suitable for both autumn and early spring sowing in the so-called 'February-March window' and for a renewal of winter wheat crops after unfavorable winter conditions (Bespalova et al. 1998).

Created by P.P. Luk'yanenko and released in 1952, the winter wheat SKRP 1 did not require vernalization under spring sowing (Bespalova et al. 1996a). Such forms (SKRP 1 among them) originated more often in winter-spring hybrid combinations. These types also are possible in hybrid combinations resulting from crosses with two winter and a spring wheat in the pedigree. Widely spread in 1990s, Yuna, with a short (20-25 days) vernalization period was the most suitable for sowing between January through the first 2 weeks of March and would yield high in such conditions. The spring wheats Siete Cerros 66 and Red River 68 (USA, QS = 10, through Obrij) were involved in the pedigree of Yuna. The ancestors of these cultivars were Ukrainian winter wheats Krymka (Turkey) and the spring wheats Thatcher and Florence (AUS) (old Ukrainian spring wheat local variety and Red Fife descendants). Yuna (QS = 10, 1/2* 7+8 5+10) can be referred to as a conditional, intermediate-type wheat. For breeding such cultivars in Krasnodar, 8-10 new winter wheat varieties and lines with the appropriate pedigree were sown at 2-3 days intervals between 5 March and 31 March. The wheat Ofeliya (QS = 9.5, 1/2* 7+8/7+9 5+10), a Spartanka sister line (see Fig. 2) is similar to Yuna in morphological and biological characteristics and is recommended as a more winter-hardy alternative to Yuna.

The new cultivar Delta is a highly productive winter crop and has good reaction to mineral fertilizers. Delta has a short (25-30 days) vernalization period and can be sown in the spring, but not later than 5 March. Delta was created by crossing Olimpiya 2 with Albidum 4431h86-4 (Russian wheats Karlik 1 (QS = 8.5), Kavkaz, Severodonskaya (a MYR 808 derivative), Bulgarian Rubin, and Yugoslavian Partizanka (QS = 8)). Even though all of the ancestors are of high-quality glutenin composition, Delta (QS = 8, 2* 20 5+10) has the Glu-B1 low-quality subunit 20 (QS = 1), the origin of which is unknown. Another new cultivar, Yara (QS = 9.5, 2* 13+16/7+9 5+10, pedigree: INIA 66, MEX (QS = 1, 1 13+16 5+10)/Thatcher descendant with Lr19//Skifyanka (QS = 7)) is characterized by frost resistance at the level of Rannyaya 12 (pedigree: BEZ 4 /SKRP 3B) and will head after spring sowing during March (Bespalova et al. 1998). This wheat can be sown up to 10 days later in the spring than Yuna, i.e., up to the middle of March, and still yield between 3.0 and 6.0 t/ha.

In 1979, during seed production of PVL, a spontaneous hybrid with early maturity (3-4 days earlier PVL because of shorter ripening period), was selected and the medium-height, winter-hardy line PVL 102 was released. The PVL gliadin profile (421112) differs from that of PVL 102 (711121) for four of the six loci. Using this line, the Krasnodar cultivars KRD 57 (QS = 9.5) and Daritsa, and the Bulgarian wheat Pliska (QS = 7), a short, early maturing cultivar Otrada (QS = 9) was obtained. In 1987, PVL 102-87 was selected from PVL 102. PVL 102-87 headed 6-8 days earlier than PVL and for 10-12 days earlier than BEZ 1. This line is characterized by intensive growth in the early spring and was registered in Russian state as Russa and is present in the State Register of Breeding Achievements Permitted to Utilization in Russia.

Russa was sown around a collection nursery during a period when many early matured China samples were under study. Quite probably, very early Russa hybrids emerged spontaneously, and the initial plant of the new variety Lepta was selected in 1993. Lepta headed 3-4 days earlier and had straw 5-7 cm shorter than Russa, with winter hardiness at the BEZ 1 level (Lysak et al. 1998a, 1998b). Russa and Lepta belong to the group of wheat with a double-growth habit. A short vernalization period (20-25 days) enables Russa to be sown until 5 March and Lepta until 10 March (Bespalova et al. 1998).

Repeated selection from crosses made in 1993 between Russa and the above-mentioned very early hybrid combination Erythrospermum 3217h6 (Genrumil, Korea/Yugtina) has allowed selection of a transgressive segregant of an early maturing plant that heads 6-7 days earlier than Russa and has more intensive spring growth. This form was named Samska.

Cultivars for the foothills and mountain regions. Cultivars created for this zone are distinguished by use of ecologically and genetically remote forms in hybridization. Excellent quality, strong, and valuable wheats with a little lower winter hardiness dominate.

The first wheat created from this program was the cultivar Rannyaya 12 (QS = 8, 1 7+9 5+10/2+12), obtained from the cross 'BEZ 4/SKRP 3' and released in 1967. Nearly 20 years later (Fig. 4), the wheat Istok (QS = 9.5, pedigree: PVL/Donskaya Ostistaya) and Kolos 80 (QS = 9) were released. The Ukrainian roots of the parental forms of these cultivars already have been mentioned (see Fig. 2). In the 1980s to early 1990, eight cultivars obtained from the geographically and genetic remote crossings with excellent grain quality were planted in state trials. Ukrainian wheat descendants Daritsa (QS = 8, 1 7+9 5+10/2+12) and Labinka (QS = 9, 1/2* 7+9 5+10), both from the cross 'Siete Cerros 66, MEX/Kavkaz//Donskaya Ostistaya'; Massiv (QS = 9, 2* 7+9 5+10) and Tokmachanka (QS = 9, 1/2* 7+9 5+10), both from the cross 'Kavkaz/Atlas 66, USA (QS = 8)//PK 49; and Dialog (QS = 9, 1 7+9 5+10) and Rada (QS = 9, 1/2* 7+9 5+10) were among them. Daritsa and Massiv were used in the breeding of new cultivars (Bespalova, Puchkov, Kolesnikov et al. 1996b).

The high-quality cultivars Rufa and Leda (both QS = 9.5) were obtained in the mid-1990s using high-quality (strong) Ukrainian wheats MYR 264, MYR 808, MYR Yuvileina, and Obrij. These wheats are for the forest-steppe ecological area and are recommended for cultivation on medium-fertile soils. Crosses of AVR and a line created in France, Mexico 50/B 21 (Versallies is a sister line of Courtot (QS = 8, 2* 7+8 2+12)) the exotic wheat Kniiskh 60 was obtained. Krymka and derivative Oro, U.S., Red Fife and offspring Thatcher and Florence all have Kniiskh 60 in their pedigrees. Results of crossing PVL with Kniiskh 60 and the other Krasnodar varieties Gorlitsa (QS = 9) and Ekho (QS = 9) were obtained and released in 1996 and 1997, respectively, for growing in the mountain and foothill zone. These cultivars have narrow vertical leaves characteristic of wheats of the steppe ecological type, but differ by having a high water-holding capacity, evidence of their drought hardiness.

Kupava (QS = 7, 2* 7+9 2+12), released in 1998, and the new variety Naslednitsa (QS = 9, 2* 7+9 5+10) belong to the valuable wheat quality class. A sister line of Kupava (QS = 7), Goryanka (N 7+8 2+12) has QS of only 6. Kupava, with a genetic potential of producing 810 t/ha, was obtained by crossing two selection lines 9021-10 (pedigree: Rusalka, BGR/Severodonskaya (QS = 9)/3/12549, BGR/Rostovchanka (QS = 9)//Dniprovs'ka 846, UKR/KVK/4/Zlatna Dolina, YUG/KRD 46 (QS = 9)) and 3161A29-2901 (pedigree: KVK/Atlas 66//PK 49). IRussian, Ukrainian, Bulgarian, Yugoslavian, and USA cultivars were used in the breeding of Goryanka. The Russian cultivars Rostovchanka, Severodonskaya, and KRD 46 are direct derivatives of the Ukrainian wheats MYR 264, MYR 808, and OD 16. Vadimovka (QS = 9, 1 7+9 5+10) was selected from the cross 'Massiv/4/KVK/Atlas 66//MYR Yuvileina/Karlik 1/5/MYR 808/4/Biserka, YUG (QS = 8)//BEZ 2/MYR 808/3/Partizanka, YUG (QS =9)'. Vadimovka is high-yielding variety in Kabardino-Balkaria and is recommended for release in 2000.

Winter wheat cultivars with the wheat-rye T1BL·1RS translocation-hexaploid triticale derivatives. Substitutions or translocations in wheat-rye crossings are extremely rare, whereas a 'triticale bridge' can use the valuable T1BL·1RS translocation (Timopheev 1996; Timopheev et al. 1999). Between 1974-95, more than 900 wheat-triticale populations were investigated for use in breeding winter wheat. All Krasnodar winter wheat varieties from the 1980-90s have hexaploid triticale in their pedigree (Fig. 5). Panatseya, Polovchanka, Knyazhna (probably including the sib line Krasota), and Bystritsa are nearly homogeneous in glutenin structure 1(or 1/2*) 7+9 5+10 and have a QS = 9-9.5. The same glutenin structure rarely occurs in other varieties with T1BL·1RS.

Panatseya and 425 ( selected from cross 'AD 206, UKR/Rubin, BGR, F1//AD 170 T1'), Polovchanka, Knyazhna, and probably Krasota have the gliadin composition 631321. Varieties obtained from other crosses with the same glutenin structure, Bystritsa (431311) and Masha (124232), have different gliadin structures. Masha (Kniiskh 22) does not have Gli-1B3, evidence of the lack of the T1BL·1RS translocation.

Polovchanka, Knyazhna, and Krasota have Gli-1B3 from the triticales AD 206 and AD 170T1. These wheats have a good root system, a high productivity potential, a positive response to high levels of agrotechnics, and stable grain yields after late forecrops, on rice-irrigation fields, or on solonetzic soils. The cultivars are heat, air, and soil-drought hardy and resistant to rust, powdery mildew, Septoria, root rot, and FHB. The T1BL·1RS translocation derived from hexaploid triticale differs qualitatively from that of KVK (Gli 432116) (Timopheevi at al. 1999). The T1BL·1RS translocation in KVK is from the German line Riebezel 47-51 (Pedigree: Criewener 194 /Petkus rye) through the German variety Neuzucht (Rabinovich 1998). The two differ is in a set of adaptive genes necessary for wheat plants in the varied soil-climatic conditions of the northern Caucasus zones.

Of the semidwarf cultivars for fallow and irrigation released in the 1990s, the high-quality winter wheat Khazarka (QS = 9, 2* 7+9 5+10 and 537312) was created from the cross 'KVK/Atlas 66//Odes'ka Napivkarlykova, UKR (QS = 9)/3/PK 49/Rostovchanka//Pavlovka' (see Fig. 3). The last three cultivars are Ukrainian wheat offspring. The presence of Gli-1B3 and T1BL·1RS (inherited from Kavkaz) in Khazarka demonstrates that this block is not an insurmountable obstacle for of creation high-quality wheats.

After Bezostaya 1, the dominant subunit is 2* in Glu-A1 in the Krasnodar cultivars (37.5 % of 70, see Table 5). However, in the group of wheats for the foothills and, especially those that are triticale descendants, the dominant subunit is 1. The Glu-B1 subunit 7+9 dominates, but in crosses using the Ukrainian cultivar Red River, subunit 7+8 appeared. The Glu-D1 subunit 5+10 dominates.

Table 5. Composition of HMW-glutenin subunits in different groups of winter wheat cultivars of P.P. Luk'janenko Krasnodar Agricultural Research Institute. The groups of Karasnodar wheat cultivars include A, medium-height, widely grown in many countries and bred in 1945-72) by under P.P. Luk'janenko; B, winter-hardy wheats bred in 1967-99); C, semidwarf wheats for fallow and irrigation bred in 1979-98; D, wheats for foothills and mountain regions bred in 1967-98); and E, derivatives of winter triticale, 2n=42, bred in 1986-99.

Glutenin subunit and quality score	Number and percent (%) of cultivars for all HMW-glutenin subunits in the different groups of Krasnodar wheats.
	A		B		C		D		E		All
	No.	%	No.	%	No.	%	No.	%	No.	%	No.	%
Glu-A1
1 (3)	1.5	25	5.5	21	2.5	17	8.5	49	4.5	90	22.5	32
2* (3)	3.0	50	14.5	56	12.0	80	7.5	42	0.5	10	37.5	54
N (1)	1.5	25	6.0	23	0.5	3	2.0	10	--	--	10.0	14
Glu-B1
7 + 8 (3)	--	--	4.5	17	3.0	20	3.5	19	0.5	10	11.5	16
13 +16 (3)	--	--	--	--	0.5	3	1.0	6	--	--	1.5	2
17 +18 (3)	--	--	--	--	--	--	--	--	--	--	--	--
7 + 9 (2)	6.0	100	20.0	77	11.5	77	13.5	75	4.5	90	55.5	79
6 + 8 (1)	--	--	0.5	2	--	--	--	--	--	--	0.5	1
20 (1)	--	--	1.0	4	--	--	--	--	--	--	1.0	2
Glu-D1
5 +10 (4)	4.0	67	22.5	87	14.0	93	14.0	78	5.0	100	59.5	85
2 +12 (2)	2.0	33	3.5	13	1.0	7	4.0	22	--	--	10.5	15
Numbers of cultivars by group.
	6		26		15		18		5		70
Quality score average by group.
	7.8		8.4		9.0		8.5		9.2		8.6

Among 17 Krasnodar cultivars bred in 1945-99, the average QS = 8.6. Fourteen Krasnodar cultivars released between 1945 and 1980 have an average QS = 7.8; 16 released between 1981 and 1990 have QS = 8.8; and 40 released between 1991 and 1999 have a QS = 8.7.

In the HMW-glutenin composition of six medium-height, widely spread, wheat cultivars bred by P.P. Luk'janenko in 1945-72 (Group A), 4.5 cultivars (75 %) have the high-quality subunits 1 (5.5, 25 %) and 2* (3, 50 %) in Glu-A1; subunits 7+9 are present in all cultivars in Glu-B1; and four had 5+10 (67 %) in Glu-D1. The average of QS for the Group A wheats is 8.7, 76 % have a QS = 9-10 and 14 % have a QS = 7-8 (Tables 5 and 6).

Of the 26 winte-rhardy wheats bred from 1967-99 (Group B), 20 (77 %) have high-quality subunits 1 (5.5, 21 %) and 2* (14.5, 56 %) in Glu-A1; 24 wheats (94 %) with Glu-B1 have 7+8 (4.5, 17 %) or 7+9 (20, 77 %); and 5+10 in Glu-D1 in 22.5 (87 %). Among the wheats of Group B, the average QS = 8.4, 58 % have QS = 9-10, and 38 % have 7-8.

In the HMW-glutenin composition of 15 semidwarf wheats for fallow and irrigated conditions bred in 1979-98 (Group C), 14.5 cultivars (97 %) have high-quality subunits 1 (2.5, 17 %) and 2* (12, 80 %) in Glu-A1; all cultivars have Glu-B1 high-quality subunits 7+8 (3, 20 %), 13+16 (0.5, 3 %), and 7+9 (11.5, 77 %); and 5+10 (14, 87 %) in Glu-D1 are present. The average QS of Group C wheat is 9.0, 87 % have a QS = 9-10, and 13 % are 7-8.

Among wheats bred between 1967-98, 18 wheats for the foothills and mountain regions comprise Group D. Sixteen (97 %) have high-quality subunits 1 (8.5, 49 %) and 2* (7.5, 42 %) in Glu-A1; all cultivars have 7+8 (3.5, 19 %), 13+16 (1, 6 %), or 7+9 (13.5, 75 %) in Glu-B1; and 5+10 (14, 78 %) in Glu-D1. Of the Group D wheats, the average QS = 8.5; 72 % have a QS = 9-10, and 17 % have a QS = 7-8.

In the HMW-glutenin composition of five wheats that are derivatives of winter triticale bred between 1986-99 (Group E), all cultivars have the Glu-A1 high-quality subunits 1 (4.5, 90 %) and 2* (0.5, 10 %); 7+8 (4.5, 90 %) or 7+9 (0.5, 10 %) at Glu-B1; and 100 % have 5+10 in Glu-D1. The average of QS for the Group E wheats is 9.2, with 100 % of samples having a QS = 9-10.

A cluster analyses indicated that the winter-hardy and semidwarf wheats were most closely linked (Fig. 6). Medium-height, widely spread cultivars, bred in 1945-72 and wheats for the foothill and mountain regions were included in the second group. The wheat cultivars that were Triticale derivatives were the most remote from other groups.

References.

• Bespalova LA, Kolesnikov FA, Puchkov YuM, Timopheev VB, Kostin VV, Zhogin AF, and Fomenko NP. 1996a. Alternative type development wheat breeding singularities in Krasnodar conditions. In: Scientific works, 95th Academician P.P. Luk'yanenko anniversary publication. Krasnodar, pp. 65-71. (in Russian).
• Bespalova LA, Puchkov YuM, Kolesnikov FA, Timopheev VB, Kostin VV, Zhogin AF, and Fomenko NP. 1996b. A role of a selection material created by Academician P.P. Luk'yanenko in development wheat breeding in Krasnodar Agricultural Research Institute. In: Scientific works, 95th Academician P.P. Luk'yanenko anniversary publication. Krasnodar, pp. 55-64. (in Russian).
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• Rabinovich SV. 1972. Modern wheat varieties and their pedigree. Urozhay, Kyiv. 328 pp. (in Russian)
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• Rabinovich SV, Panchenko IA, Parchomenko RG, and Bondarenko VN. 1998. HMW glutenin subunit composition of spring bread wheats grown in the Ukraine and Russian Federation between 1995-97 and its connection with pedigrees. Ann Wheat Newslet 44:236-251.
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• Timopheev VB, Dudka LF, and Kovtunenko VYa. 1999. Effectiveness of triticale-wheat crossings and selection in bread winter wheat breeding. Scientific basis for stabilization of plant production: Reports theses of international conference devoted to 90-years from V.Ya. Yur'ev Institute for Plant Production founding. Kharkiv. pp. 98-100 (in Russian).
• Vapa LJ and Savic M. 1988. HMW glutenin subunit composition of wheat varieties bred in Yugoslavia. In: Proc 7th Internat Wheat Genet Symp (Miller TE and Koebner RMD eds). Institute of Plant Science Research, Cambridge, UK. 2:1019-1023.
• Vapa LJ and Dencic S. 1995. Genetic variation in Glu-1 loci and breadmaking quality in wheat. Cereal Res Commun 23:161-166.
• Zeven AC and Reiner L. 1991 . Genealogies of 3200 wheat varieties. Institute of Plant Breeding I.V.P., Agricultural University, Wageningen, the Netherlands, and Crop Husbandry Technical University. 79 pp.
• Zeven AC and Zeven-Hissink NCh. 1976. Genealogies of 14000 wheat varieties. Institute of Plant Breeding I.V.P., Agricultural University, Wageningen, the Netherlands, and Crop Husbandry Technical University. 117 pp.
• 5th Facultative and Winter Wheat Observation Nursery 1995-1996: Appendix 1. Prepared and distributed by the National Wheat Improvement Program, Turkey, CIMMYT, ICARDA, and Oregon State University. p. 1-4.

The history of ancient and modern Ukrainian wheat cultivars used in breeding spring wheat cultivars of the U.S., Mexico, and western and eastern Europe and an analysis of their HMW-glutenin structure.

S.V. Rabinovich, O.Ju. Leonov O.Ju, R.J. Pena *, G. Fedak **, O. Lukow ***, V.M. Bondarenko, and A.A. Kushchenko.

* CIMMYT, Lisboa 27, Colonia Juares, Apdo. Postal 6-641, 06600 Mexico, D.F., Mexico and Agriculture and Agri-Food Canada, ** Central Experimental Farm, Bldg 50, Ottawa, Ontario K1A 0C6 and *** Cereal Research Centre 195 Dafoe Road Winnipeg, Manitoba R3T 2M9, Canada.

Information was gleaned from 55 publications, published between 1970-99, on the HMW-glutenin subunit structure of 226 spring wheats bred between 1919 and 1995 in the U.S., Mexico, and 10 western and two eastern European countries. The HMW-glutenin structure in the genealogy of many cultivars from these different countries is given according to the publications of Branlard and LeBlanc 1985; Lawrence 1986; Ng et al. 1988; Khan et al. 1989; Lukow et al. 1989; Morgunov et al. 1990; Graybosch 1992; Lookhart et al. 1993; Parchomenko et al. 1996; CIMMYT 1997; Rabinovich et al. 1997, 1998; Redaelli et al. 1997; Bushuk 1998; and Fedak et al. 1998.

The genealogy of wheats from many countries of the world is given according to Zeven et al. 1976, 1991; family trees and release dates are according to Almer et al. 1998; Anonymous 1973, 1996a, 1996b, 1997; Borojevich 1970; CIMMYT 1995; Dorofeev et al. 1976; Hanisova et al. 1998; Kohli 1986; Kraic et al. 1996; Martynov et al. 1990; Rabinovich 1972; Sontang et al. 1986; Uhlen 1990; and Villereal and Rajaram 1988.

The majority of North American and European cultivars with high quality inherited their glutenin subunits from the ancient Ukrainian spring wheat Red Fife directly or from its derivatives, sometimes after only a few generations. Red Fife, from Galicia (now the Lviv region of the Ukraine) was brought through Poland and Germany to Canada in 1842, and, from there, later into the U.S. Among Red Fife (2* 7+9 5+10, quality score (QS) = 9) derivatives are the Canadian spring wheats Marquis (Hard Red Calcutta, IND/Red Fife); Garnet (Preston (QS = 9, Ladoga, RUS/Red Fife) and Onega, RUS in the pedigree) (N 7+9 5+10, QS =7); and some others that played an important role in improving the quality of European winter wheats. The role of the U.S. spring wheat Thatcher (Jumillo, ITA, T. durum/Marquis// Kanred, winter (QS = 9, a selection from Krymka, UKR/Marquis)) (2* 7+9 5+10, QS = 9) created in 1934 is very important in grain quality improvement not only for European wheats, but also on other continents where this cultivar was used in breeding. An important role in quality improvement of some European wheats has been played by U.S. winter wheats Oro and Kanred (an offspring of the Ukrainian variety Krymka that was sent into U.S. in 1884 under the name of Turkey).

Almost all the CIMMYT semidwarf wheats and the hundreds of derivatives worldwide have inherited dwarfness from the U.S. hybrid (Norin 10, JPN (1 7+8 2+12)/Brevor, USA (2* 7+9 2+12)), with Ukrainian wheats in their pedigree including Krymka, Oro, and the Australian spring wheats Florence (a direct descendant of Red Fife) and Federation (a derivative of Red Fife through the U.S. cultivar Improved Fife). In the genealogy of some European wheats, the French spring wheat Noe (a selection from an old Ukrainian local variety from the Odesa region named Synya (Didus 1967)) also was used.

The analysis of HMW-glutenin subunits of spring wheats indicates that among the 122 wheats from the U.S. and Mexico, subunits 2* (64 %) and 1 (28 %) predominate at the Glu-A1 locus, 7+9 (44 %); 17+18 (23 %), and 7+8 (18 %) predominate at Glu-B1; and 5+10 (73 %) is the main subunit at Glu-D1. Among 104 wheats from 12 western and eastern European countries, subunits 1 (47 %) and 2* (33 %) predominate at locus Glu-A1; 7+9 (47 %) and 7+8 (21 %) at predominate at Glu-B1; and 5+10 (69 %) at Glu-D1. However, among wheats from seven countries (mainly Sweden and Germany), subunit 14+15 (14 %) occurs at locus Glu-B1, which is absent in Northern America cultivars. Subunit 17+18 is present in Northern America cultivars (23 % total and 32% in Mexico) more often than in European wheats (3 %).

In the HMW-glutenin composition of 63 U.S. wheat cultivars released between 1927 and 1985, 53.5 (90 %) had high-quality subunits 1 (9, 15 %) or 2* (44.5, 75 %) at Glu-A1; 54.5 (93 %) had 7+8 (15.5, 26 %), 13+16 (1, < 1 %), 17+18 (8, 14 %), or 7+9 (30.5, 52 %) at Glu-B1; and 47 (80 %) had 5+10 at Glu-D1 (see Table 7) (Ng et al. 1988; Khan et al. 1989; Lukow et al. 1989; Graybosch 1992; Lookhart et al. 1993; Redaelli et al. 1997; Fedak et al. 1998).

Among U.S. cultivars, 76 % of samples have a QS = 9-10 and 14 % have 7-8. The average QS of these cultivars is 8.7 (Table 8); however, HRSWs have a QS = 9.1 and SWWs have a QS = 5.5. The HRSWs from Montana have a high QS of 9.4 , those from South Dakota = 9.0; those from Minnesota = 8.9; and a group of cultivars created by corporations in California, Colorado, Nebraska, and some other states have the highest QS = 9.7. CIMMYT germ plasm with the high-quality subunits 17+18 and 7+8 predominates as the initial material for the breeding of these cultivars. The average QS of North Dakota wheats is 8.5, because of the low QS of the old cultivar Hope (6.5) and is 7 in two newer lines. Hope, released in 1927, was created by crossing a high-quality Canadian cultivar Marquis with a stem rust-resistant T. dicoccum line from Russia. Hope has stable resistance to the pathogen, but is low in grain quality.

Thus, most of the U.S. HRSWs have a high index for the glutenin complex (Table 9) that was inherited from high-quality spring cultivars Marquis, Thatcher, Apex, Reward, Canus, and other Red Fife offspring and the winter wheats Kanred, Oro, and other offspring of Krymka from Canada and the U.S.

The SWSWs from Idaho and Washington bred in the 1979s-80s have low quality scores, 5.0 and 5.6, respectively, even though the high-quality wheats Marquis, Thatcher, Florence, Federation, Krymka and descendants, and Oro are in their pedigrees. High quality is difficult to combine with bunt resistance, which is needed in this growing region.

Among 63 wheats from Mexico bred between 1950-95, 59 (94 %) have high-quality subunits 1 (26, 41 %) and 2* (33, 53 %) at Glu-A1; 61 (97 %) have 7+8 (6.5, 10 %), 13+16 and 13+19 (11, 18 %), 17+18 (20, 32 %), and 7+9 (23.5, 37 %) at Glu-B1; and 42.5 (68 %) have 5+10 at Glu-D1. The average QS for Mexican wheats is 8.8; 64 % of the samples have a QS = 9-10 and 37 % have 7-8. (Table 7 and Table 8) (Ng et al. 1988; Payne et al. 1988; Lukow et al. 1989; Graybosch 1992; Randal et al. 1993; Silvera et al. 1993; Tahir et al. 1995; CIMMYT 1996b, 1996c, 1997; Rabinovich et al. 1998).

The average QS for 34 Mexico wheats bred between 1950-1970 is 9.1. Of 18 others and 10 CIMMYT lines selected in 1981-95, the quality scores were 8.5 and 8.4, respectively. Thus, a dominance of high-quality subunits 1 and 2* at the Glu-A1 locus varied little over the years. At the Glu-D1 locus, the presence of high-quality subunits 5+10 (QS = 4) is near 65 % for older cultivars and 70 % for cultivars and lines created after 1981. At the same time, the high-quality subunits 17+18, 13+16, and 7+8 (QS = 3) at Glu-B1 in 28 cultivars and breeder lines created after 1981 have decreased from a high of 82% to 34 % for 34 cultivars released in 1950s-1970s.

In our opinion, subunit 17+18 arose from Australia. The Australian cultivars Gabo and Timstein (probably sister lines) were used widely in the CIMMYT breeding programs. Timstein was used in the 1950s in the Mexican wheats Yaqui 53 and Yaqui 54, and Gabo 55 and Gabo 56 were created from Gabo. Since that time, the use of Mexican wheats as the genetic basis for crosses has resulted in the wide spread of subunit 17+18 in wheats of many countries of the world.

According to data of Macindoe et al. (1968), Gabo has the pedigree 'Bobin/Gaza (from Palestine) T. durum//Bobin sel.' The glutenin structure of Gabo according to Lawrence et al. (1980) is 2* 17+18 2+12. However, Wrigley et al. (1977) state that it is necessary to be extremely cautious about the Gabo family tree. Research on the gliadin structure of Gabo and its ancestors from 25 years earlier indicated that Bobin (Thew (a derivative of Fife, UKR)/Steinwedel, AFR) actually could be the Australian cultivar Gular (Wagga 13/Marshall's No. 3), and the Timstein and Lee pedigrees quite possibly used Gular instead of Bobin.

Rabinovich et al. (2000) made analyses of more than 140 cultivars bred in 26 countries and found that Mexican wheats were the genetic basis for the wide use of subunit 17+18 in wheat breeding from Great Britain to Australia, including the U.S., and a majority of South America, Asia, and Africa but saw limited use in the Ukraine and Russia.

Between 1969 and 2000, the largest number of the Mexican wheats with subunit 17+18 (26 in nine countries of the world) were created using Sonora 64 (a derivative of Yaqui 54), 10 cultivars in five countries were created using II 8156 (a descendant of Gabo 55), 11 cultivars in seven countries were created with Siete Cerros 66 (a selection from II 8156), 17 wheats in six countries were created with the involvement of Ciano 67 (a derivative of Sonora 64), and 20 cultivars in eight countries with Bluebird (II 8156, Sonora 64, and Ciano 67 in the pedigree). With Yecora 70 (a selection from Bluebird), Bluejay (derivative of Siete Cerros and Paloma), and Pavon 76 (descendant of Ciano 67 S and Bluebird), four or five cultivars were created in three to four countries.

Lawrence (1986), Lukow et al. (1989), Redaeli et al. (1991), Graybosch (1991), Lookhart et al. (1993), Randal et al. (1993), Cornish et al. (1995), Tahir et al. (1995), Jackson et al. (1996), and CIMMYT (1996a) contain information on 60 wheat cultivars from 12 countries released between 1930-90 and verify that subunit 13+16 at locus Glu-B1 is of Brazilian origin. The Brazilian cultivar Fronteira (Polissu (a selection from LV Guapole)/Alvredo Chaves 8 (a selection from LV)) (2* 13+16 2+12) and the derivative Frontana probably inherited the 2* subunit from old local Brazilian wheats and spread it through Surpreza (sib of Fronteira), Supremo 211 (U.S.), and Lerma Rojo (MEX) to the Mexican cultivar Lerma Rojo 64. Lerma Rojo 64 and derivatives INIA 66, Norteno 67, and Jupateco 73 are the genetic sources of subunit 13+16 for several cultivars from Mexico, Pakistan, South Africa, and Australia.

A marked drop in the average QS from 9.1, for Mexican cultivars released in the 1950s-70s, to 8.4-8.5, for cultivars and lines bred mainly in the 1980s-90s, is connected with a modification in the qualitative structure of the wheats used in crosses in the different decades. Thus, in the pedigrees of Mexican wheats from the 1950s-70s, the same parents (offspring of Red Fife and Krymka) were used as in U.S. wheat breeding programs (see footnote of Table 9) and their Mexican derivatives. In the pedigrees of cultivars and lines bred in the 1980s-90s, together with the earlier bred wheats, a series (see entry numbers 26-30, 32, 33,44, 45, and 53 in the footnote of Table 10) of mainly low- and medium-quality French winter wheats with QS = 4-6 (derivatives of the spring wheat Noe and the Ukrainian wheat Synya offspring with unknown HMW glutenins) were included. Among the French winter wheats are Iga Blondeau (QS = 5) and derivative Coutishes (Red Fife in pedigree). Teeter and the Romanian Fundulea 29 were used in breeding the Mexican wheat Tatbird, Riley 67 (U.S., a descendant Red Fife) was used in Batera Benhur, and seven HRWWs and SRWWs with QS = 6-10 (see numbers 52, 54-58, and 60 in footnote of Table 10) from the states of NE, KS, OR, and IN and the Russian wheat Bezostaya 1 were used in Oringa. Many of these are offspring of Ukrainian wheats.

The Mexican wheat Alondra, the derivative Curinda 87, and some other wheats with T1BL·1RS (Rabinovich 1998) inherited the wheat-rye translocation from the German winter wheat Weique (Criewener 192/Petkus rye). Veery (QS = 9), obtained by selection from Ures 81 (QS = 8), Genaro 81, Glenson 81, Seri 82 (QS = 9), and derivatives Bacanora 88 (QS = 9), Cumpas 88, Mochis 88 (QS = 7), Arivechi 92, and probably Choix 95 inherited T1BL·1RS from the German line Neuzucht (Weique synonym or sister line) through the Russian wheat Kavkaz. Bobwhite, Teeter, Bagula, and derivatives Angostura 88 and, probably, Catbird have inherited this translocation (the last four mentioned wheats through Fundulea 29) from Neuzucht through the Russian wheat Avrora (sib of Kavkaz).

Of eight wheats bred in Sweden in the 1970s-90s, five (63 %) have high-quality subunits 1 (25 %) and 2* (38 %) and three (37 %) have subunit 21* (QS unknown) at Glu-A1; all have subunits 14+15 (63 %), 17+18 (12 %), or 7+9 (25 %) at Glu-B1; and only three (38%) have subunit 5+10 at Glu-D1. The average QS of samples from Sweden is 8.8; 64 % have a QS = 9-10 and 37 % have 7-8 (see Tables 7 and 8) (Payne et al. 1983; Lindahl et al. 1988; Johanson et al. 1993; Kazman et al. 1996; Groeger et al. 1997). At the Glu-B1 locus in Swedish wheats, subunit 14+15 predominates . This subunit may have a northern origin.

Many of the spring wheats from Sweden (Karn I, Karn II, Svenno, and Pompe) are Red Fife offspring. The cultivar Kadet is widely spread among these and has a good HMW-glutenin structure (1 7+9 5+10, QS = 9). In the Kadet family tree are the Red Fife offspring Ring (SWE), Kolibri (GER), Noe, Sappo, Nemares, and Trol (see Table 11).

Among 18 wheats from Finland bred between 1919-81, 75 % have high quality subunits 1 (33%) and 2* (42%) at Glu-A1; 94 % have 7+8 (44 %) and 7+9 (50 %) at Glu-B1; and 78 % have 5+10 at Glu-D1. The average QS for cultivars from Finland is 8.4; 56 % have a QS = 9-10 and 33 % have a QS = 7-8 (Tables 7 and 8) (Sontang et al. 1986; Jackson et al. 1996).

Ruskea, Pika, and Pika II, bred at the beginning of 20th century and with an average QS = 8.5, have the high-quality subunit 7+8 at Glu-B1 and the low-quality subunit 2+12 at Glu-D1 (Table 11). Improving grain quality in Finland and other countires of northern and western Europe was by the use of high-quality foreign germ plasm. The 1930s cultivars Sopu (Marquis/Ruskea (QS = 8)) and Hopea (Marquis/Pika) inherited a QS = 9 from Marquis. Later cultivars (upto the 1980s) Kimmo, Touko, Kinru, Terra, Ruso, Veka, Taava, Tahti, Ulla, and Luja had the good HMW-glutenin structure from Red Fife offspring Sopu, Hopea (FIN), Karn I, Svenno, and Reward (CAN, through Ruso); and Kinru, Tahti, and Ulla through the Australian cultivar Aurore.

All 15, analyzed Norwegian wheat varieties have high-quality subunits 1 (7 % of samples) and 2* (93 %) at Glu-A1; 7+8 (50 %), 13+16 (7 %), and 7+9 (36 %) at Glu-B1; and 5+10 (50 %) at Glu-D1. Forty-seven percent of the Norwegian cultivars have a QS = 9-10 and 53 % have 7-8. The average QS of Norwegian wheats is 8.5 (Tables 7 and 8) (Uhlen 1990; Jackson et al. 1996).

Wheats created in the first half of the century in Norway had low-quality subunit 2+12 and a QS = 7-8. Around the 1950s, Norrona and Nora (both with QS = 10) inherited the high-quality subunits 5+10 from Red Fife through Marquis and Sopu and 2* and 7+8 from the Norwegian wheat From II. Norrona has the high quality of Rollo (QS = 10). Rollo and the German wheat Els (a Red Fife descendant of the Canadian Garnet and the Ukrainian wheat Synja, a derivative of Noe) are Runar (QS = 10) ancestors. Els, the Swedish Karn II, and McIntosh from Canada (through the Finish cultivar Tammi) are Reno (QS = 9) ancestors. The Bastian family tree together with the Norwegian, U.S., Mexican, Argentinian, and Japanese wheats (many of which are derivatives of Ukrainian varieties) are included.

British, Swedish, German, Dutch, and Indian wheats were included in the genealogy of four wheats from Great Britain with a QS = 7-10 (Cooke 1995; Kazman et al. 1996). Chablis and Shiraz were created with the participation of the Dutch cultivar Jerico (the Ukrainian winter wheat Myronivs'ka 808 in pedigree). A number of Ukrainian wheats and descendants are ancestors of modern English cultivars.

Among 10 French cultivars released from 1951-84, four have the high-quality subunits 1 (20 %) and 2* (20 %) at Glu-A1; all have 7+8 (25 %), 14+15 (10 %), 17+18 (10 %), and 7+9 (55 %) at Glu-B1; and only 50 % have 5+10 at the Glu-D1 locus. Among French cultivars, 30 % of samples have QS = 9-10 and 30 % have 7-8. The average QS of French wheats (7.0) is the lower than that of other spring wheats of Europe (Tables 7 and 8) (Branland and LeBlanc 1985; Payne et al. 1988; Jackson et al. 1996).

Many cultivars represented in Table 11 are Red Fife derivatives (Marquis, Thatcher, Garnet, Els, White Fife, Preston, Florence, and Federation) or Noe (Synya derivative) offspring. The U.S. winter wheat Oro was used in creating of two French spring wheats. Among the French cultivars, Florence-Aurore is of the greatest interest. Both parents were selected in Australia from Red Fife through Jacinth (FRA), Improved Fife, and Saskatchewan Fife (CAN) from Eden (AUS). Florence-Aurore was cultivated in France, many African countries including in Tunisia and Algeria and probably Spain under the name Ariana 8. Two new spring wheats in the Ukraine were created using Ariana 8.

Of German wheats created from 1959-95 25, 84 % have the high-quality subunits 1 (80 %) and 2* (4 %) at Glu-A1; 80 % have 7+8 (7 %), 14+15 (16 %), and 7+9 (60 %) at Glu-B1; and 88 % have 5+10 at Glu-D1. The average QS for German wheats is 8.2; 52 % have a QS = 9-10 and 48 % have 7-8 (Tables 7 and 8) (Ng et al. 1988; Margiotta et al. 1993; Kazman et al. 1996; Groeger et al. 1997).

Of the nine German cultivars with known family trees (Table 11), two are Red Fife offspring through Marquis, six through Garnet, and one (Mephisto) through both cultivars. The French cultivar Noe (a Synya derivative) was used to breed seven German cultivars. In Ralle, Red Fife, Noe, the winter wheat Ukrainka (UKR), and Kanred were used through the Austrian winter wheat Perlo and Bezostaya 1.

Most widely used in German breeding programs was Opal (twice a descendant of Garnet) (QS = 9). Opal and derivatives Laval 19 (QS = 9), Dundas (QS = 9), Vernon (QS = 9), and Messier (QS = 10) were cultivated in the Canadian provinces of Quebec and Prince Edward Island for many years. The spring wheats Beloruskaya 12 and Beloruskaya 80 (both Opal offspring) were widely grown in Belarus, Ukraine, and Russia.

In the 1990s, Erwin (QS = 8) had the most genetically diverse family tree of four Austrian spring wheat cultivars with known HMW-glutenin structure and QS = 8-10 (Tables 7 and 8) (Groeger et al. 1997) . The maternal parent is the German cultivar Mephisto (an offspring of Red Fife from Marquis, Garnet, Newthatch (U.S.), and Krymka. The widely grown Ukrainian winter wheat Myronivska 808 was used in breeding the Dutch cultivar Minaret.

Subunit 14+15 is found in the HMW-glutenin structure of the Swiss wheat Lona (QS = 7) (Kazman et al 1996; Groeger et al. 1997) of unknown pedigree and Axona (QS = 9) from the Netherlands (Cooke 1995). Axona is an offspring of Red Fife through Garnet, Koga II, and Maris Dove from Great Britain. Subunit 14+15 often occurs in Sedish cultivars but less often in German wheats.

The HMW-glutenin composition of eight Czech wheat cultivars released in 1975-97, seven had the high-quality subunit 1 at Glu-A1; 7+8 (19 %), 7+9 (69 %), or were unknown (12 %) at Glu-B1, and six had 5+10 at Glu-D1 (Cerny et al. 1992; Kraic et al. 1996; Almer 1998; Hanisova et al. 1998). Among the Czech cultivars, 74 % have a QS = 9-10 and 13 % have 7-8, with an average of 8.5 (Tables 7 and 8).

Foreign wheats are frequently in many Czech cultivar pedigrees (Table 11), among them Nadadores 63 and Siete Cerros 66 (offspring of Red Fife through Marquis, Thatcher, Florence, and Federation); Krymka (through Oro and Kanred); Koga I and Koga II (descendants of Red Fife through Preston and Garnet); Kolibri (a derivative of Red Fife from Garnet and Els); and Synya (from Noe) and MYR 808. The new Czech cultivar Aranka (QS = 9) inherited subunit 14+15 at Glu-B1 from ancestors of the German cultivar Planet (QS = 8).

Among four the Yugoslavian cultivars with known HMW glutenins structures (Table 7 and Table 8), the QS varies from 4 to 10 (Vapa et al. 1988). Jarka is the lowest (QS = 4, from Red Fife, Thatcher, Florence, Federation, Krymka, Siete Cerros 66, Ukrainka, Bezostaya 1, and the Serbian wheat NS 975). Planinska, with a QS = 10, inherited subunits 17+18 and 7+8 in Glu-B1 from the Mexican wheats.

The most interesting cultivar of the five Spanish wheats is Ariana (a sib of the French wheat Florence-Aurore, derived from Red Fife) (Payne et al. 1988; Silvera et al. 1993).

A cluster analysis of the frequency of HMW-glutenin subunits in wheats from these different countries segregates European countries into two clusters; one with wheats from GBR, AUT, CZE, SPA, YUG, SWE, and DEU and a second with wheats from FIN, NOR, and FRA. The U.S. wheats and especially the Mexican wheats, strongly differ from those from Europe (Fig. 7) . Possibly, the difference is connected with the spread of the 17+18 subunit at the Glu-B1 locus in Mexican and U.S. cultivars. This subunit is present less often in cultivars from other countries.

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