ISSN 0095-4527, Cytology and Genetics, 2009, Vol. 43, No. 3, pp. 169–176. © Allerton Press, Inc., 2009.Original Russian Text © I.I. Motsnyi, V.I. Fayt, E.M. Blagodarova, 2009, published in Tsitologiya i Genetika, 2009, Vol. 43, No. 3, pp. 26–35.

169

INTRODUCTION

The present gene pool of bread wheat is deficient ingenes, providing its resistance to unfavorable abioticand biotic conditions [1]. The most important problemof wheat cytogenetics is the development of introgres-sive bread wheat forms, containing different amountsof foreign genetic material. Some species and even gen-era allied to wheat can be used as the source of suchmaterial [2]. The aim of our study was to determinekaryotypes and to study morphological and agronomi-cal characteristics, including resistance to diseases, intwo introgressive lines of winter bread wheat obtainedby an intergeneric hybridization.

MATERIALS AND METHODS

The objects of our study were two sister F

12

lines ofwinter bread wheat, Hostianum 273/97 (H273) andHostianum 274/97 (H274), derived by the hybridiza-tion of octoploid triticale AD825 (bread wheat Hos-tianum 237 rye VSHI) with durum wheat (cv. Cherno-mor). The lines are constant concerning the complex ofmorphological characteristics and represent the off-spring of the individual selection for winter hardinessand frost resistance, performed among the progeny ofone F

5

plant [3]. In the case of a wide-row sowing, linesH273 and H274 were notable for big ears, even andsometimes strong stems, and a high grain mass perplant.

In the period from 2002 to 2005, the genomic, cyto-logical, biochemical, and phytopathological analysis ofthe lines H273 and H274 was carried out. Within theframework of a genomic analysis, we studied highchromosome associations in their F

1

hybrids with breadand durum wheat and winter rye (cv. Kharkovskaya60). In the case of a cytological analysis, we studied the

level of conjugation of metaphase chromosomes in tenand nine plants from the lines H273 and H274, respec-tively. The line of cv. Odesskaya 267 (Od267), derivedin 2001 from the progeny of one typical isolated plant,was used as a control and also as a pollinator for thegenomic analysis. Since we did not reveal any differ-ences among the control Od267 plants concerning thefrequencies of meiotic associations, the data were inte-grated. The F

1

hybrids between (1) Od267 and ryeKharkovskaya 60 (one plant) and (2) Chinese Spring(CS) and rye Kharkovskaya 60 (nine plants) were usedas a control for wheat–rye polyhaploids. All materialsused in the studies and hybridization were from sam-ples taken from the collection of the Department ofGenetics of the Plant Breeding and Genetics Institute.

The study of somatic chromosomes was carried outin root tip cells, which were pretreated with a saturatedmonobromenaphthalene solution, fixed in glacial aceticacid, subjected to hot hydrolysis with 1 N HCl, andstained with Schiff reagent according to the Feulgentechnique [4]. The meiosis study was performed usinga material fixed in Caroy’s fluid (6 : 3 : 1). Anthers weretreated with 4% iron alum and stained with 2% aceto-carmine solution. For each sample, we examined 8–11 plants and at least 30 clear plates per plant at themetaphase I stage (MI). The number of chromosomeassociation sites (CAS) per one maternal pollen cell(MPC) was determined, considering that each junctionof two chromosome arms corresponds to one CAS.The conjugation level index

ϕ

was calculated after theFisher transformation:

where

C

is the ratio of the empirical CAS to the maxi-mum CAS possible for the given cell [5]. This index issuitable for the comparison of data obtained for plants

ϕ 2 C,arcsin=

Identification and Characteristics of the 1R (1B)Substitution Lines of Bread Wheat

I. I. Motsnyi, V. I. Fayt, and E. M. Blagodarova

Plant Breeding and Genetics Institute, National Center of Seed-Growing and Cultivar Investigations,Odessa, 270036 Ukraine

e-mail: motsnyyii@gmail.com; fayt@paco.net; blagodarova@rambler.ru

Received May 15, 2008

Abstract

—The 1R (1B) chromosome substitution is identified in two introgression bread wheat lines, derivedfrom a distant hybridization between octoploid triticale and durum wheat. The substitution is marked by theoriginal alleles of the secaline-coding loci

Sec1

and

Sec3

. The lines, demonstrating a rather low level of conju-gation, high winter hardiness and frost-resistance, and high crop capacity in both favorable and unfavorableyears, can be differentiated on the basis of the indicated traits.

DOI:

10.3103/S0095452709030050

170

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MOTSNYI et al.

with differing numbers of chromosomes, since it repre-sents an inversely weighted value. The distribution ofthis index is close to normal, and this index can be cor-rectly treated using parametric methods [6].

The electrophoretic separation of gliadins and glu-tenins was performed in an acidic polyacrylamide gelwith acetic acid–glycine buffer in accordance with theoriginal technique, developed in the Department ofGenetic Basis of Breeding of the Plant Breeding andGenetics Institute (PBGI) [7]. Genetic formulas for thegluten protein loci were determined using the cataloguewritten by F.A. Popereli.

Agronomic characteristics were studied under natu-ral conditions on the experimental field of the Depart-ment of Genetics of PBGI. For a close sowing of lineson the plots (the registration area was 3 m

2

), we used anSSFK-7 tractor seeder, whose seeding rate was500 seeds/m

2

. The following cultivars were used ascontrols: highly productive cv. Albatros odesskiy (Alb,a national standard of Ukraine); cv. Odesskaya 267(Od267, the most common cultivar); two cultivars,Odesskaya 16 (Od16) and Obriy, demonstrating highand low winter hardiness, respectively; and also one ofthe parental forms, cv. Hostianum 237 (H237). Thefrost resistance was assessed during the germ stage at

−

11

°

C [8] and for tillered plants (25–35 plants of eachgenotype) [9], selected in the field in the first ten daysof December (–13

°

C), second ten days of January(

−

14

°

C), and the first ten days of March (–11

°

C). Thewinter hardiness was assessed under field conditions(we registered the number of plants at the end of Octo-ber and the number of overwintered plants in spring).During the freezing of tillered plants, we registered thenumber of tillering shoots (NBS) for each plant beforethe freezing and the number of shoots grown after thefreezing (NSG). During vegetation and harvesting, weexamined ten plants of each sample for the followingparameters: epycotil length (EL)—before wintering;number of additional tillering shoots (ATS) and the dateof earing—in spring; plant height (PH), average eargrain mass (EGM), grain productivity of the plot (GP),and the number of productive stems per 1 m

2

(NPS)—in summer. The phytopathological assessment of thematerial was carried out under field conditions (natural

infection background) and in the case of an artificialinfection with leaf and stem rust. The level of plantresistance was determined in accordance with the inten-sity of infection using an integrated nine-point scale[10].

The uniformity of the groups of performed cytolog-ical studies and the reliability of differences betweenmean values and fractions were determined using a95% confidence interval

±

t

0.05

, indicated in alltables of this article; the standard error of the proportionwas determined using a formula for alternative variabil-ity [6]. The statistical treatment of the data of fieldexperiments was carried out using a variance analysis.

RESULTS AND DISCUSSION

The study of somatic chromosomes revealed signif-icant differences between the lines concerning theiraneuploidy levels. Lines H273 and H274 contained2.7

±

3.7% and 24.7

±

11.0% aneuploids, respectively,and both euploid and aneuploid plants of these linescontained only two explicit satellite chromosomes intheir karyotypes (Fig. 1a). The sampling study of theMI stage in these plants and in their F

1

hybrids (H273

×

H274) revealed a frequently occurring high chromo-

some association, , in all 42-chromosome plants P

1

,P

2

, and F

1

and, therefore, the homogeneity and identityof the genomic composition of both lines. The aneup-

loids of the line H273 had 2

n

= 41 and + 1

II

in theform of a high association. Similar karyotypes wererevealed for most of the aneuploids from the line H274.In addition, we found one 43-chromosome plant and one

plant with 2

n

= 41 and a high association + 3

I

(436–2/04). A possible reason for the differences in the ane-uploidy level between the lines is the fact that, duringthe line H273 reproduction, we twice (2001 and 2002)used the progeny of only one plant, whereas in the caseof the line H274 we did not perform such selection.

Meiotic observations in F

1

hybrids (H273 or H274

×

Od267) at the MI stage revealed that, irrespective of aline, for all plants studied, 2

n

= 42 and the high associ-

x Sx

213II

203II

193II

(a) (b) (c)

Fig. 1.

(a) Somatic chromosomes of the euploid plant from the line H273 and (b) chromosome associations at the MI meiotic stagein the F

1

hybrid between H273 and Od267, 20

II

+ 2

I

; (c) segment of the AII meiotic stage for the H273

×

Od267 F

1

hybrid with thedelay of a satellite chromosome. Arrows indicate satellite chromosomes.

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IDENTIFICATION AND CHARACTERISTICS 171

ation was + + 2

I

(Table 1), except for one cell,

where it was + 2

I

. At the same time, univalent chro-mosomes did not conjugate between themselves in all322 cells studied, which confirms the presence of awheat–rye substitution in these lines. In several cases,we observed the presence of a satellite for one of theunivalents (Fig. 1b). Obviously, this univalent was rep-resented by a missing chromosome (1B or 6B), inher-ited through a paternal gamete. Since satellite chromo-somes usually do not differ from other chromosomes atthe MI stage, then, according to Sears’ recommenda-tion [11], the presence of a satellite for one of the uni-valent chromosomes was confirmed at the AII stage,when this chromosome several times was delayed at theequator (Fig. 1c). Besides the substitution, a structuralchromosome reconstruction possibly occurs in the linesH273 and H274 and complicates the conjugation ofhomologous chromosomes in the hybrids betweenthese lines and wheat (Table 1); a practically total lackof multivalents in the lines and F

1

hybrids indicates theabsence of a reciprocal translocation affecting homeo-logic wheat chromosomes.

In the case of the crossing of the lines with rye, allexamined F

1

hybrids had 28 chromosomes. However, atthe MI phase, we observed a significant variation in themeiotic association frequencies and the total level of con-jugation (from 0.98

±

0.23 to 3.75

±

0.49 CAS/MPC),depending on the plant studied. This fact can be explainedby the genetic heterogeneity of rye concerning suppres-sors of a diploidizing system of wheat [12], which is con-firmed by the CAS/MPC ratio variation in the control

193II 10

II

203II

(0.21

±

0.15 in the hybrid Od267

×

rye and from 0.07

±

0.06 to 1.41

±

0.32 in the hybrid CS

ψ

rye). As a result, ahigh diversity of high chromosome association classeswas observed (Table 1). Nevertheless, we observed theformation of one or, sometimes, two ring bivalentspractically in all F

1

hybrids, though with varying fre-quencies (from 0.08

±

0.07 to 0.73

±

0.19 at MPC). Theonly exceptions were two descendants of the plant 436–

2/04 (karyotype + 3

I

), derived from the fusion of apaternal rye gamete with an ovule containing the fullset of wheat chromosomes, though in these hybrids one

of every 118 cells had the + 26

I

karyotype (the aver-age number of ring bivalents for a MPC was 0.01

±

0.02). In the control, we observed random ringbivalents only in the CS

×

rye combination and only inthe plants with a high level of conjugation, which didnot differ significantly in these characteristics. The fre-quency of these ring bivalents was very low (0.012

±

0.009 on average for an MPC) and corresponded to theearlier observed in CS

×

rye F

1

hybrids [13]. Therefore,the conclusion about the presence of a wheat–rye sub-stitution in these lines was confirmed. One should notethat, in spite of the presence of two homologous ryechromosomes in the wheat–rye F

1

hybrids, the fre-quency of a ring bivalent formation was low (dependingon the plant, it varied from 7.8

±

6.7% to 63.6

±

14.5%of MPCs). The frequency of microsporocytes withoutany bivalents varied from 1.6

±

3.1% to 37.1

±

12.3%,i.e., the rye chromosome obtained from triticale weaklyconjugates with the homologous chromosome of therye cv. Kharkovskaya 60. Possibly, it was caused by

193II

13II

Table 1.

High chromosome associations in F

1

hybrids, derived from the crossing of H273 and H274 lines with bread wheat,rye, and durum wheat

Line Hybrids with bread wheat Hybrids with rye Hybrids with durum wheat

H273 + + 2

I

(4; 98) + + 24

I

(2; 127) + + 8

I

(1; 72)

+ + 22

I

(1; 66) + 8

I

(1; 72)

+ 9

I

(1; 72)

H274 + + 2

I

(5; 160) + 20

I

(2; 118)

+ + 2

I

(1; 64) + + 22

I

(1; 64)

+ + 18

I

(2; 128)

+ + 16

I

(1; 60)

+ + 22

I

(1; 64)

+ + 18

I

(1; 44)

Note: The lower indices

3

and

0

designate ring and rod bivalents, respectively. The number of examined plants and MPCs is shown in

parentheses. Two hybrids were obtained in the progeny of the plant 436-2/04 (karyotype + 3

I

).

193II 10

II 13I 10

II 123II 10

II

13I 20

II 133II

133II

193II 10

II 40II

183II 20

II 13II 20

II

13II 40

II

13II 50

II

23II 10

II

23II 30

II

193II

172

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MOTSNYI et al.

some modifications of the rye chromosome in the triti-cale karyotype [14] or the negative influence of wheatchromosomes [15].

Three examined F1 plants (H273 × T. durum) had2n = 34 and 2n = 35 and the high associations 13II + 8I

and + 9I, respectively (Table 1). So, it is possibleto conclude that (1) the rye chromosome replaces oneof the chromosomes of the AB genome; (2) this chro-mosome (or one of the D-genome chromosomes) iseliminated in aneuploids; and (3) possible structural

133II

reconstruction, complicating the conjugation of homol-ogous chromosomes in wheat F1 hybrids, affects the Dgenome.

The electrophoretic analysis of storage proteinsrevealed the presence of secalines, rye glutene proteins,in the studied material. Among the alcohol-soluble pro-teins (gliadins), we identified the block of componentsregulated by the Sec1 locus, being a genetic marker forthe short arm of the 1RS rye chromosome [16]. Theallele of this locus, detected in H273 and H274, is iden-tical to that of triticale AD825 (Fig. 2) and differs fromany of the typical wheat homeoalleles and also from theprolamin Gld 1B3 block (according to Popereli’s clas-sification), which marks a 1BL.1RS translocation, typi-cal of the cvs. Aurora and Kavkaz [17]. The electro-phoregrams of gliadins from triticale AD825, lineH274, and also introgressive lines and well-known cul-tivars containing the alleles of the Gld 1B locus (partic-ularly, the Gld 1B3 allele), typical of winter breadwheat, are shown in Fig. 2. The new block is similar tothe Gld 1A17 allele (according to the Popereli cata-logue, [18]), which can be detected in cultivars with the1AL.1Rs translocation from the cv. Amigo [19]. How-ever, in the studied material, this new block is encodedby the 1B chromosome, since other gliadin-coding loci1A, 1D, 6A, 6B, 6D of the chromosomes of lines H273and H274 contain blocks typical of wheat, particularlythe Gld 1A2 allele of the Gld 1A2 locus (Table 2),whereas the wheat components of the Gld 1B locuswere not observed.

In the spectrum of glutenins, alkali-soluble high-molecular-weight proteins, we revealed componentsregulated by the secaline locus Sec3. This locus marksthe long arm of the 1R chromosome [16] and replacesthe corresponding homeologic locus of the 1B chromo-some in the studied lines (Fig. 3), which is evidence ofthe substitution of both arms in the 1B chromosome.The presence of some components unusual for wheat intriticale (which also has some wheat components) andin the line H274 is shown in Fig. 3; at the same time, thetypical Glt 1B1 (Glu–B1 7 + 8) or Glt 1B2 (Glu-B1 7 + 9)

1 2 3 4 5 6 7 8 9 10 11 12

Fig. 2. Electrophoregrams of wheat storage proteins (glia-dins): (1) AD825; (2) cv. Chernomor; (3) E200/97–2-B;(4) E217/97-B; (5) H242/97–2-B; (6) H274/97. Standards:(7) cv. Albatros odesskiy; (8) cv. Selyanka; (9) cv. Ku-yalnik; (10) cv. Glenlea; (11) cv. Veselka; (12) cv. PerlynaLisostepu. Arrows indicate the components of rye Sec-1locus alleles: three components (lanes 1 and 6) represent theallele from rye cv. VSHI and four components (lanes 3, 5,11, and 12) represent the allele from wheat cv. Aurora.

Table 2. Genetic formulas of the alleles of glutene protein loci in introgressive lines, their initial forms, and standard cultivars

MaterialGliadins (Gld)

Glutenins (Glt)

LMW HMW

1A 1B 1D 6A 6B 6D 2–1A 1A 1B 1A 1B 1D

H237 2 2 1 1 + 4 2 4 1 2 2 2 2 1 + 2*

AD825 2 + N*** 2 + S1** 4 1 2 4 1 2 2 + S1 1 + 2 2 + S3** 2

cv. Chernomor 6 + 3 2 – 3 4 – 3 6 2 2 2 –

H273 2 S1 4 1 1 1 1 2 S1 2 S3 1

H274 2 S1 4 1 2 1 1 2 S1 2 S3 1 + 2

Albatros odesskiy 4 1 4 4 2 2 3 4 1 2 2 1

Note: LMW, low-molecular-weight subunits; HMW, high-molecular-weight subunits; * 1 + 2, heterozygote; ** S1, Sec1 block; S3, Sec3block; *** N, unknown protein.

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IDENTIFICATION AND CHARACTERISTICS 173

blocks are absent. The fact that the substitution does notaffect the 1A chromosome is confirmed by the presenceof a typical wheat allele Glt 1A2 in the lines H273 andH274. The presence of typical blocks in the loci Gld 1Dand Glt 1D indicates that there are no reconstructions inthe 1D chromosome in the lines studied.

Thus, we can consider that the presence of a 1R(1B)substitution in the studied lines, when the 1R rye chro-mosome (cv. VSHI), transferred via triticale AD825,replaces the 1B wheat chromosome, is confirmed. Theloss of the satellite of the 1R chromosome in a wheatkaryotype, including the 1R chromosome of triticale[14], was registered by many researchers and can beexplained by the shortening of a satellite thread.

In general, both lines had a reliably lower level ofconjugation (P = 0.05) than the control (cv. Od267).The only exception was one 41-chromosome plantfrom the line H274, in which we revealed an unexpect-edly high level of conjugation of 20 homologous chro-mosome pairs. The reduction of the level of conjugationoccurs mainly via a desynapsis, which increases thenumber of rod bivalents and univalents (Table 3). Onaverage, only 13.1 ± 3.2% (from 3.1 to 26.4%, depend-ing on the plant) of 436 examined microsporocytes

from euploid plants had ; 22.9 ± 4.0% (from 5.6 to46.9%) of cells had univalents, and most of the uni-valents (except those which are obligatory in aneup-loids) had a clear desynaptic origin, i.e., sharp edgesand symmetric arrangement. According to [15], thereduction of the level of conjugation in substitutedwheat lines is typical mainly of the rye chromosomepair. It is possible that the aneuploid plant with a highlevel of conjugation (Table 3) has a rye chromosome asthe univalent, whereas other chromosomes conjugate inthe ordinary way. The formation of univalents at the MIstage results in the aneuploidy and, according to thedata from Table 3, is caused by both aneuploid and eup-loid plants.

The study of agronomic and morphological quanti-tative traits showed that both lines H273 and H274 hada shortened epicotyl comparing to the old winter wheatcultivars (initial H237 form and the best winter-hardycultivar Od16); its length practically corresponded tothe level of the modern frost-resistant cultivars, Alba-tros odesskiy and Od267. This fact provides the lower-ing of the tillering node by 10–15 mm compared to thesame value in old cultivars. According to [20], geno-types with a shortened epicotyl have a better resistanceto unfavorable wintering conditions; at the same time,such genotypes have the same or even less physiologi-cal frost resistance. In our study, this fact influenced theincrease in the regrowth intensity after the spring frosts(Table 4); however, the lines did not show any addi-tional tillering in spring. Concerning such parametersas DE, PH, EGM, NPS, and crop capacity in favorableyears (2002 and 2005), the lines were similar to themodern cultivars, though their crop capacity was lessthan that of the standard cultivar (Table 4). In the unfa-

213II

vorable year (the severe winter of 2002/2003 and thesevere drought in spring of 2003), the lines were similarto the old cultivar Od16 (this cultivar is the most resis-tant to abiotic factors), surpassed H237 in crop capac-ity, but were inferior to the modern highly adaptive cul-tivar Od267. At the same time, the line H273, demon-strating a rather low EGM value, tended to a highercrop capacity in unfavorable years owing to its NPSlevel; the line H274 demonstrated the same tendencyfor favorable years. Also, both lines H273 and H274tended to higher tillering in autumn and spring, respec-tively. In general, the level of tillering of the lines wasrather weak, whereas winter-hardy cultivars H237,Od16, and Od267 tended to a strong autumn tillering.

Concerning the frost and winter resistance, sampleswere differentiated in the following way (Table 5). Aswas expected, cv. Obriy always was the least frost andwinter resistant, and, on the contrary, cv. Od16 had thehighest resistance to these two factors. Both studiedlines had a high frost resistance, surpassing that of theline H237; however, they differed from one anotherdepending on the time and method of freezing. Thefrost resistance of H274 and H273 germs was equal tothat of cvs. Albatros odesskiy and Od16, respectively.This fact corresponds to the published data [3],obtained by a similar way at T = –10°C and T = –12°C.

1 2 3 4 5 6 7 8 9 10 11 12

Fig. 3. Electrophoregrams of wheat storage proteins (glu-tenins): (1) AD825; (2) cv. Chernomor; (3) E200/97–2-B;(4) E217/97-B; (5) H242/97–2-B; (6) H274/97. Standards:(7) cv. Albatros odesskiy; (8) cv. Selyanka; (9) cv. Ku-yalnik; (10) cv. Glenlea; (11) cv. Veselka; (12) cv. PerlynaLisostepu. Arrows indicate high-molecular-weight compo-nents of the rye Sec-3 locus of cv. VSHI (lanes 1 and 6).

174

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MOTSNYI et al.

During the freezing of tillered plants at the beginningand in the middle of winter, the frost resistance of H273was about the same as that of Od16 and surpassed thefrost resistance of H274, which, in turn, was similar tothat of H237. At the end of winter, the frost-resistanceof both lines was about the same as that of Od267,being less than that of Od16, but more than that ofH237. Weather conditions during the winter of2002/2003 were very good for a winter hardiness

assessment (Table 5). The line H274, having a lessphysiological frost resistance, overwintered better thanthe line H273; its winter hardiness was practically thesame as that of Od16, whereas the winter hardiness ofH273 was the same as that of the initial form. Duringthe years favorable for wintering, the variety of linesand cultivars concerning their frost resistance andpotential winter hardiness had little influence on theresults of the overwintering owing to the insufficient

Table 3. Chromosome conjugation type at the MI stage of meiosis

Line

Number of examined

2n

Average number per cell ± t0.05

plants cells III

III CAS* ϕrod ring total

Od267 8 315 42 0.3 ± 0.1 1.7 ± 0.2 19.1 ± 0.2 20.8 ± 0.1 0 40.0 ± 0.2 2.759 ± 0.025

(0–4) (0–7) (13–21) (19–21) (33–42) (2.179–3.142)

H273 8 236 42 0.6 ± 0.1 2.1 ± 0.2 18.6 ± 0.2 20.7 ± 0.1 0.004 ± 0.008 39.3 ± 0.3 2.692 ± 0.032

(0–6) (0–7) (12–21) (18–21) (0–1) (31–42) (2.067–3.142)

2 64 41 1.2 ± 0.2 2.2 ± 0.3 17.7 ± 0.4 19.9 ± 0.1 0 37.6 ± 0.4 2.684 ± 0.047

(1–5) (0–6) (14–20) (18–20) (34–40) (2.346–3.142)

H274 6 200 42 0.5 ± 0.1 2.6 ± 0.3 18.1 ± 0.3 20.8 ± 0.1 0 38.9 ± 0.3 2.644 ± 0.017

(0–4) (0–9) (12–21) (19–21) (33–42) (2.179–3.142)

1h 35 41 1.1 ± 0.1 1.1 ± 0.5 18.8 ± 0.5 20.0 ± 0.1 0 38.8 ± 0.5 2.888 ± 0.085

(1–3) (0–6) (14–20) (19–20) (34–40) (2.346–3.142)

11 32 41 1.3 ± 0.2 3.0 ± 0.7 16.8 ± 0.8 19.8 ± 0.2 0.06 ± 0.09 36.7 ± 0.7 2.595 ± 0.078

(1–3) (0–8) (12–20) (18–20) (0–1) (32–40) (2.214–3.142)

1* 32 41 3.5 ± 0.3 2.1 ± 0.5 16.5 ± 0.5 18.6 ± 0.2 0.06 ± 0.09 35.3 ± 0.5 2.627 ± 0.064

(3–5) (0–4) (13–19) (17–19) (0–1) (32–38) (2.324–3.142)

Note: CASes are chromosome association sites, ϕ is the ratio between the number of revealed CASes and the maximum possible CASnumber, transformed for the purpose of its normalization [6]. The subscripts indicate the level of conjugation: high (h) or low (l).

* The plant 436-2/04 with a high association + 3I. The variation range is shown in parentheses.

x Sx

193II

Table 4. Quantitative traits of introgressive lines, parental forms, and standard cultivars

Genotype EL, mm ATS DE

(May)PH, cm

EGM, g CC, kg/m2 NPS, pieces/m2 NBS, pieces

U F U F U F 1 2 3

H237 28.1 7.0 25 125 0.52 0.59 0.13 0.30 352 448 3.3 2.9 3.2

H273 16.6 4.0 18 88 0.42 1.38 0.18 0.49 408 499 2.5 2.9 2.4

H274 18.2 5.7 17 87 0.76 0.97 0.16 0.52 312 534 1.8 2.7 2.9

Obriy 22.2 11.1 15 85 0.76 1.11 0.21 0.48 28 423 2.3 2.6 2.1

Od16 24.5 5.6 23 123 0.56 0.99 0.18 0.42 408 517 3.4 3.2 2.7

Albatros odesskiy 20.4 6.5 16 84 0.80 1.52 0.09 0.55 148 479 2.8 2.6 2.2

Od267 16.1 8.3 19 98 0.79 1.10 0.22 0.51 472 422 3.2 2.7 2.5

LSD0.05 7.3 4.5 3 12 – 0.50 – 0.20 – 158 1.5 0.7 1.6

Note: EL, epicotyl length; ATS, additional tillering in spring; DE, date of earing; PH, plant height; EGM, average ear grain mass; CC, cropcapacity; NPS, number of productive stems; NBS, number of tillering shoots (1—summer, 2—winter, 3—spring); U, data for theunfavorable year (2003); F, averaged data for two favorable years (2002 and 2005).

CYTOLOGY AND GENETICS Vol. 43 No. 3 2009

IDENTIFICATION AND CHARACTERISTICS 175

stress load, though the ranks of the lines remainedunchanged. There are some data [21] about the success-ful use of triticale AD825 for substantially increasingthe frost resistance and winter hardiness of winterwheat; however, this publication does not contain anyinformation on the cytogenetic or genomic analysis ofsamples. At the same time, it is difficult to say anythingabout the role of the revealed 1R (1B) substitution insuch a high manifestation of this trait.

Concerning phytopathological diseases, in certainyears, the studied lines were resistant to powdery mil-dew and moderately susceptible to stem rust; possibly,this was due to variations in the race composition andintensity of the natural infection background. We areinclined to connect this fact with the presence of a ryechromosome, since the interaction of wheat and ryechromosomes often entails resistance to powdery mil-dew and stem rust [10]. It seems that the aforemen-tioned chromosome does not contain any other efficientgenes of resistance to the studied diseases, which isconfirmed by a high susceptibility of the lines (Table 5).In the case of an artificial infection with Fusariumgraminearum under laboratory conditions, lines H273and H274 also remained highly susceptible: the level ofinfection was 83 and 84%, the germination energy wasreduced by 34 and 38%, and the germination wasreduced by 73 and 80%, respectively (the same param-eters for the cv. Albatros odesskiy were 35, 23, and25%).

In spite of a very high resistance to unfavorable win-tering factors, good ripening rate, evenness, short-stalk-ness, and good crop capacity, the possibility of the prac-tical use of the lines studied is under question. On theone hand, the aneuploidy of the studied material alwaysnegatively influences its fertility and stability; on theother hand, the presence of rye proteins (Sec1 and Sec3loci) significantly lowers the quality of flour produced

from grain with such genotype [16]. Probably, suchlines can serve as an initial material for the selection offorage cultivars. In any case, it seems advisable to makea selection of euploid plants in the line H274 for its sta-bilization.

CONCLUSIONS

The 1R chromosome of rye (cv. VSHI), transferredto the lines H273/97 and H274/97 via triticale AD825,replaces a wheat 1B chromosome. This chromosomepoorly conjugates with a homologous rye chromosomefrom cv. Kharkovskaya 60 in the rye–wheat F1 hybridsat the MI stage of meiosis. The lines have a low level ofconjugation and form aneuploids. The percentage ofaneuploidy in the line H274/97 is significantly higherthan in the line H273/97. The presence of the 1R chro-mosome in the karyotype is marked by original secalinecomponents of Sec1 (gliadin spectrum) and Sec3 (glu-tenin spectrum) blocks. The lines are characterized bya high rate of ripening, short-stalkness, high frost resis-tance and winter hardiness, and good crop capacity.The line H273/97 is more frost resistant, whereas theline H274/97 demonstrates a higher winter hardiness.In certain years, the lines were resistant to powdery mil-dew and moderately susceptible to stem rust, but werestrongly infected with leaf rust, winter wheat leafblotch, barley yellow dwarf virus, and seedling blight.

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H237 27.2 11.1 25 85 87 53 59 96 4–6* 1–5 2 2–5 2–4

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H274 20.5 38.1 63 86 88 78 71 100 4–8 1–5 3–4 3–6 3–5

Obriy 38.3 31.7 9 77 92 35 4 98 3–4 1–5 2–8 2–6 2–5

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