INSTITUTE OF EXPERIMENTAL BIOLOGY
EE 3051 Harku, Harjumaa, Estonia.
Behavior of meiosis in monosomic F1 hybrids.
T. Enno.
Plants of the common wheat mutant 146-155,
induced in the cultivar Norroena
after mutagenic treatment with N-methyl-N-nitrosourea,
were crossed with the tetraploid wheat species T. timopheevii,
a source of disease resistance. The introgressed line 146-155-T
with resistance to leaf rust and powdery mildew was selected in
the progeny of an interspecific hybrid (Enno and Peusha 1992;
Peusha and Enno 1996). Aneuploid analysis using the set of 21
monosomic lines of wheat cultivar Chinese Spring was made and
the genes conferring resistance to diseases in line 146-155-T
were located on chromosomes 1B and 6B.
Chromosome pairing in PMCs at the MI was analyzed
in monosomic F1 hybrids. Every PMC was scored for
univalents, bivalents, and multivalents. Cytological analysis
showed that some chromosomes of 146-155-T affected the
pairing at MI. Chromosomes 3A, 6A, and 2B decreased pairing,
but chromosomes 6B, 1D, and 4D increased pairing at MI.
Structural differences between the chromosomes of
Chinese Spring and the chromosomes of 146-155-T were
supported by the data of the chromosome associations and multivalent
formations in monosomic F1 hybrids.
Some hybrids with trivalent configurations, with
or without the presence of univalents, were observed at MI among
the 21 monosomic crosses. The critical monosomic hybrids on chromosomes
3A, 4A, 6B, and 7D had trivalent configurations without univalents,
and, consequently, were involved in chromosome translocations.
On the basis of the frequency of trivalent associations, we believe
that the introgressed line 146-155-T has the reciprocal
translocations 3A/4A and 6B/7D in relation to Chinese Spring.
A comparatively low frequency of trivalent configurations
in PMCs indicated that rather small segments of chromosomes were
involved in translocations. Transfer of alien chromatin from
T. timopheevii into the genome of 146-155-T
was analyzed by fluorescent cytological analysis using in situ
DNA hybridization (Enno et al. 1995; Jarve et al. 1996).
References.
Enno T and Peusha H. 1992. Introgression of genes
for rust resistance from Triticum timopheevii to common
wheat. Vortrage fuer
Pflanzenzuchtung. 24:197-199.
Enno T, Peusha H, Jarve K, Timofeyeva L, Tsimbalova
E, and Priilinn O. 1995. Introduction of alien genetic variation
by means of interspecific hybridization. EWAC Newsletter. pp.
65-67.
Jarve K, Peusha H, Tohver M, Tamm S, Timofeyeva L,
Tsimbalova E, Priilinn O, and Enno T. 1996. Alien chromatin
detection and disease resistance gene identification in introgressive
lines of common wheat. Ann Wheat Newslet 42:79-80.
Peusha H, Enno T, and Priilinn O. 1996. Genetic
analysis of disease resistance in wheat hybrids, derivatives of
Triticum timopheevii and T. militinae. Acta Agronomica
Hungarica 44(3):237-244.
Screening for powdery mildew resistance in common wheat
cultivars and breeding lines.
H. Peusha, A. Ingver
(Joegeva Plant Breeding Institute, Estonia), and O. Priilinn.
The main objectives of wheat breeding in Estonia
are grain yield, protein content, protein quality, and resistance
to diseases. Powdery mildew is the most serious of the wheat
diseases. The use of genetic resistance to control the powdery
mildew is of major importance throughout the world, and the production
of genetically resistant cultivars should remain an important
breeding strategy.
Alleles of the powdery mildew-resistance genes have
been identified at 20 loci, located on different chromosomes (Zeller
et al. 1993a). However, in the commercial wheat cultivars grown
in western and central Europe, only nine major resistance genes
are utilized in breeding for resistance: Pm1, Pm2,
Pm3c, Pm3d, Pm4b, Pm5, Pm6,
Pm8, and Pm9. Wheat cultivars grown in Europe usually
possess one or a combination of these genes (Zeller et al. 1993b).
There is little information on the occurrence of
genes for resistance to diseases in wheat cultivars and breeding
lines of Scandinavian origin growing in Estonia. In our earlier
investigations of wheat cultivars grown in Finland, genes controlling
resistance to powdery mildew were identified (Peusha et al. 1996).
The gene Pm4b was found to be the only resistance gene
in commercial cultivars grown in Finland. The spring wheat cultivar
Tapio, has Pm3d, but is no longer grown commercially.
The Finnish landrace NGB 43 has the Pm6 gene and may
serve as potential genetic resource for breeding resistant wheat
cultivars.
The wheat collection from the Joegeva
Institute of Plant Breeding was screened in 1996. Evaluations
for powdery mildew resistance were done on segments of primary
leaves of host seedlings grown in a phytotron. The leaf segments
were placed in petri dishes with 6 g/l agar and 35 mg/l benzimidazole
(Lutz et al. 1995). The differential E. graminis tritici
isolates used were collected from different parts of Europe (Felsenstein
et al. 1991). The major resistance genes found in the cultivars
were postulated from comparisons of the response patterns of differential
wheat cultivars and isogenic lines (given in Table 1) and application
of the gene-for-gene concept of Flor (1956). Table
1 presents the different types of interactions between cultivars
and lines of wheat with known major genes for resistance to powdery
mildew and 11 pathogen isolates. Host-plant reaction was classified
as follows: R = resistant, S = susceptible, and I = intermediate.
Compound ratings-R,I
and I,S-- indicate
both reaction types. The rating S/I means that in the plant population,
both susceptible and intermediate responses were observed.
The response patterns of common wheat cultivars and
breeding lines were differentiated from the wheat lines possessing
documented resistance genes (Table 2).
Finnish wheat cultivars Luja, Laari, Manu, Ulla,
Mahti, and Heta were tested for powdery mildew resistance after
inoculation with 11 differential E. graminis tritici isolates.
The cultivar Luja had a susceptible reaction to all test isolates.
This cultivar lacks the major resistance genes listed in Table
1. The cultivar Laari had resistance similar to that observed
in wheats with gene combination Pm3a + Pm5 + Pm?.
Manu wheat had the pattern of Pm4b resistance. Manu apparently
received Pm4b from Runar, which was developed from a derivative
of the German wheat breeding strain possessing Pm4b. The
cultivar Ulla showed resistance to powdery mildew isolates typical
of wheats with gene Pm1. Mahti wheat was resistant to
isolates 10 and 14, a reaction that was typical for wheats with
the Pm5 gene. Breeding line Cebeco 1036, which
was in the pedigree of cultivar Mahti (Table 3), possessed the
resistance gene Pm8 (Krivtchenko 1988). However, the response
pattern of Mahti did not show the presence of Pm8 gene.
The cultivar Heta had resistance to seven powdery mildew isolates,
and exhibited intermediate reaction after inoculation with the
5 and 17 isolates. We failed to identify resistance genes of
Heta using available test-isolates (Peusha et al. 1996).
Table 1. Reaction of cultivars and differential lines with genes for resistance to powdery mildew inoculated with 11 isolates of Erysiphe graminis f. sp. tritici.
| Cultivar / line | Resistance gene(s) | Isolates of E. graminis tritici | ||||||||||
|---|---|---|---|---|---|---|---|---|---|---|---|---|
| 2 | 5 | 6 | 9 | 10 | 12 | 13 | 14 | 15 | 16 | 17 | ||
| Axminister/CCR*8 1 | Pm1 | R | S | R | I,S | R | S | S | S | R | S | S |
| Ulka/CCR*8 1 | Pm2 | S | R | R | S | R | S | S | S | R | S | S |
| Asosan/CCR*8 1 | Pm3a | R | S | R | R | R | S | R | R | S | S | I |
| Chul/CCR*8 1 | Pm3b | R | S | S | R | R | R | R | R | S | R | I,S |
| Sonora/CCR*8 1 | Pm3c | R | S | S | I | R | S | R | I,S | S | S | S |
| Kolibri | Pm3d | S | S | S | R | S | R | S | R | R | S | R |
| W150 | Pm3e | S | I,S | I,S | I | R | I,S | R | R,I | S | S | S |
| M.A./CCR*8 1 | Pm3f | R | S | S | S | R | S | R | I,S | S | S | S |
| Khapli/CCR*8 1 | Pm4a | S | R | S | R | I | R | S | S | I | S | I |
| Armada | Pm4b | S | R | S | R | R | R | S | S | R | S | S |
| Hope | Pm5 | S | S | S | S | R | S | S | R | S | S | S |
| TP 114/STK*2 2 | Pm6 | S | R,I | R,I | R | R,I | S | R,I | R,I | R,I | I | S |
| Disponent | Pm8 | R | S | S | R | S | R | S | S | S | S | R |
| Normandie | Pm1+2+9 | R | R | R | R | R | S | S | S | R | S | S |
| BRG 3N 3 | Pm16 | R | R | R | R | R | R | R | R | R | R | R |
| Amigo | Pm17 | I | I | I,S | I | I | I | R | S | I | R | R,I |
R = resistant, S = susceptible, I = intermediate.
1 Eight backcrosses to Chancellor.
2 Two
backcrosses to Starke.
3 BRG 3N /76-F2-205-derivate
of T. turgidum var. dicoccoides.
The Swedish cultivars Satu, Tjalve, Polkka, Fagott,
Dragon, and Dacke were screened for powdery mildew resistance.
Satu exhibited resistance to isolates 5, 6, 9, 15, and 17; susceptibility
to isolates 10, 12, 13, 14, and 16; and an intermediate reaction
to isolate 2. However, origin of disease resistance in Satu remains
unknown. The cultivar Tjalve was resistant to all test-isolates,
except isolate 2. The resistance gene in Tjalve was not identified,
and the origin of its resistance also remains unknown. Polkka
wheat exhibited a response pattern typical of wheats with the
gene combination Pm2 + Pm3c + Pm?. Fagott
wheat showed the pattern of gene combination Pm3d +
Pm5 + Pm? The breeding line CI 12632, which was in pedigree
of Fagott, possessed the resistance genes Pm2 and Pm6.
The resistance in Fagott most likely derived Pm2 and Pm6
from this breeding line, although disease reaction after inoculation
did not support this assumption. The cultivar Dragon was resistant
to all 11 differential isolates. The wheats Sicco and Sappo with
resistance genes Pm5 and Pm2 + Pm4b, respectively,
are in the pedigree of Dragon wheat. Dragon wheat may possibly
possess some combination of resistance genes, or only one unknown
gene. The cultivar Dacke also exhibited the same response pattern,
and the gene(s) for resistance remains unknown.
Table 2. Reaction of wheat cultivars/lines from Finland, Sweden, and Norway inoculated with 11 isolates of Erysiphe graminis.
| Cultivar / line | Resistance gene(s) Pm | Isolates of E. graminis tritici | ||||||||||
|---|---|---|---|---|---|---|---|---|---|---|---|---|
| 2 | 5 | 6 | 9 | 10 | 12 | 13 | 14 | 15 | 16 | 17 | ||
| Luja | - | S | S | S | S | S | S | S | S | S | S | S |
| Laari | 3a + 5 + ? | R | S | R | R | R | S | R | R | S | S | R |
| Manu | 4b | S | R | S | R | R | R | S | S | I | S | S |
| Ulla | 1 | R | S | R | I,R | R | S | S | S | R,I | S | S |
| Mahti | 5 | S | I,R | S | S | R | S | S | R | S | S | S |
| Heta | ? | R | I | R | R | R | S | R | R | R | S | I |
| Satu | ? | I | R | R | R | S | S | S | S | R | S | R |
| Tjalve | ? | S | R | R | R | R | R | R | R | R | R | R |
| Polkka | 2 + 3c + ? | R | R | R | R | R | S | R | S | S | S | R,I |
| Fagott | 3d + 5 + ? | S | S | R | R | R | R | S | R | R | R | R |
| Dragon | ? | R | R | R | R | R | R | R | R | R | R | R |
| Dacke | ? | R | R | R | R | R | R | R | R | R | R | R |
| Reno | 4b | S | R | S | R | R | R | S | S | I,R | S | S,I |
| Runar | 4b | S | R | S | R | R | R | S | S | R | S | S |
| Bastian | ? | R | R | R | R | R | R | R | R | I | S | R |
| Hja 24201 | 2 + 4a + ? | S | R | R | R | R | R | R,I | R | S/I | S | R |
| Hja 24471 | ? | R | R | R | R | R | R | R | R | R | R | R |
| Hja 24472 | ? | R | R | R | R | R | R | R | R | R | R | R |
| Bor/Hja 25077 | ? | S/R | S/I | I,S | I,S | R | S | S/R | I,S | S | S | I |
| Bor/Hja 25084 | ? | R | S | S | R | R | S | S | I,R | S | S | S |
| Bor/Hja 25092 | 2 | S | R | R | S | R | S | S | S | S | S | S |
| Bor/Hja 25115 | 6 + ? | R | R | R | R | R | S | R | R | S | S | R |
| Bor/Hja 25145 | 5 | S | S | S | S | R | S | S | R | S | S | S |
| Bor/Hja 25191 | (1 + 2 + 9) + 3d | R | R | R | R | R | R | R | R | R,I | S | R |
| Bor/Hja 25203 | 3a + ? | R,I | R | R | R | R | S | R | R | S | S | R |
| qBor/Hja 25229 | 3a + 8 + ? | R | R | R | R | R | R | R | R | S,I | S,I | R |
R = resistant, S = susceptible, I = intermediate
Evaluation of three Norwegian cultivars showed that
Reno and Runar have resistance gene Pm4b. These cultivars
apparently received gene Pm4b from the cultivar Els (Paderina
et al. 1995). Bastian wheat was susceptible to test-isolate
16, and exhibited intermediate reaction to isolate 15. The resistance
gene in this cultivar remains unknown.
Screening Finnish breeding lines for powdery mildew
resistance revealed two lines, Hja 24471 and Hja 24472, which
were resistant to all eleven test-isolates. We assumed that
the cultivar WW 21220 was the source of unidentified resistance
gene in these lines, because it is present in the pedigrees of
both lines. The other nine Finnish lines possessed the following
resistance genes or combination of genes: Pm2; Pm5;
Pm6 + Pm?; Pm2 + Pm4a + Pm?;
Pm (1 + 2 + 9) + Pm3d; Pm3a
+ Pm?; and Pm3a + Pm8 + Pm?.
The present study concluded that Scandinavian wheat
cultivars and breeding lines possess mainly the same powdery mildew-resistance
genes as the cultivars of western Europe. Only the Finnish
cultivar Laari and the two breeding lines Bor/Hja 25203 and Bor/Hja
25229 have the resistance gene Pm3a in combination with
an as yet unidentified Pm gene, which probably is not present
in the wheat cultivars produced in Europe.
Table 3. Genealogies of wheat cultivars and lines.
| Cultivar/line | Pedigree |
|---|---|
| Luja | Svenno // Hopea / Tammi |
| Laari | Villa glori / Touko |
| Manu | Ruso / Runar |
| Ulla | Hankijan Ulla // Tammi / Ta4431 |
| Mahti | Cebeco 1036 / Hja 20519 |
| Heta | Hja a 1105 / Hja a 1099 |
| Satu | Snabbe / Drabant / T106 / Snabbe |
| Tjalve | T9111 / 449-73 / 15432 |
| Polkka | (SV 70415 / Snabbe // Norroena / Kaern 2 /3/ Snabbe) |
| Fagott | SW 79452 / C1 12632-Prins 6 // Drabant |
| Dragon | WW 24380 / Sicco / W12502 /3/ Sappo |
| Dacke | ? |
| Reno | Els // Tammi / Kaern 2 |
| Runar | Els / Rollo |
| Bastian | Bajo66 / Runar /4/ Jaktana / Norin 10 / Brevor /3/ Mogystad /5/ Rollo / Magnif /4/ Sonora / TZPP / Nainari /3/ Mogystad |
| Hja 24201 | Hja 2216 / WW19018 |
| Hja 24471 | WW 21220 / Hja 22058 |
| Hja 24472 | WW 21220 / Hja 21600 |
| Bor/Hja 25077 | WW 24042 / Luja |
| Bor/Hja 25084 | WW 24042 / Luja |
| Bor/Hja 25092 | WW 24042 / Hja 234117 |
| Bor/Hja 25115 | WW 24042 / Polkka |
| Bor/Hja 25145 | Matador / Luja |
| Bor/Hja 25191 | Polkka / Hja 23145 |
| Bor/Hja 25203 | Polkka / Luja |
| Bor/Hja 25229 | WW25187 / Hja 23171 |
References.
Flor HH. 1956. The complementary genetic systems
in flax and flax rust. Adv Genet 8:29-54.
Felsenstein FG, Limpert E, and Fischbeck G. 1991.
Wheat mildew populations in the FRG and neighbouring regions
-
some aspects of their change. Integrated control of cereal mildews:
virulence patterns and their change (Jorrgenses
JH ed). Riso Natl Lab, Roskilde, Denmark:1-7.
Krivtchenko VI. 1988. Catalogue of World Wheat
collection. Cereal cultivars with known genes of resistance to
fungal diseases. VIR, Leningrad. p. 79.
Lutz J, Limpert E, Bartos P, and Zeller FJ. 1992.
Identification of powdery mildew resistance genes in common wheat
(Triticum aestivum L.). I. Czechoslovakian cultivars.
Plant Breed 108:33-39.
Paderina EV, Hsam SLK, and Zeller FJ. 1995. Identification
of powdery mildew resistance genes in common wheat (Triticum
aestivum L. em Thell.). VII. Cultivars grown in Western
Siberia. Hereditas 123:103-107.
Peusha H, Hsam SLK, Enno T, and Zeller FJ. 1996.
Identification of powdery mildew resistance genes in common wheat
(Triticum aestivum L. em Thell.). VIII. Cultivars and
advanced breeding lines grown in Finland. Hereditas 124:91-93.
Zeller FJ, Lutz J, and Stephan U. 1993a. Chromosome
location of genes for resistance to powdery mildew in common wheat
(Triticum aestivum L.). I. Mlk and further alleles
at the Pm3 locus. Euphytica 68:223-229.
Zeller FJ, Stephan U, and Lutz J. 1993b. Present
status of wheat powdery mildew resistance genetics. Proc 8th
Inter Wheat Genet Symp (Li ZS and Xin ZY eds), Beijing, China.
pp. 929-931.
Utilization of wheat storage proteins for identification
of chromosome translocations.
M. Tohver.
Genes controlling the synthesis of wheat storage
proteins are localized on the short arms of the group 1 and 6
chromosomes (gliadins, LMW glutenins), and on the long arms of
the group-1 chromosomes (HMW glutenins) (Shepherd 1968; Payne
et al. 1980).
In Volume 42 of the AWN, we described translocations
of the Gli-1 and Gli-2 loci in the introgressed
line 146-155-T (a derivative of T. timopheevii).
These translocations were detected by A-PAGE. The Gli-B2
locus translocation on chromosome 6Bin the hybrid line is from
T. timopheevii (Jarve et al. 1996).
SDS-PAGE subsequently was used for identification
of translocations at loci Glu-A1, Glu-B1,
and Glu-D1 located on the long arms of chromosomes
1A, 1B, and 1D, respectively. The HMW-glutenin subunits are encoded
by genes at these loci. Total wheat protein of wheat mutant 146-155,
the introgressed line 146-155-T, and T. timopheevii
was extracted from one kernel and fractionated by SDS-PAGE
using 10 % polyacrylamide gels. Cultivars Chinese Spring, Dragon,
and Dacke were used as standards.
The HMW-glutenin subunit compositions of 146-155
and 146-155-T were identical and differed from T.
timopheevii subunits. We concluded that there were no translocations
of the loci coding for HMW glutenins.
References.
Jarve K, Peusha H, Tohver M, Tamm S, Timofeyeva L,
Tsimbalova E, Priilinn O, and Enno T. 1996. Alien chromatin
detection and disease resistance gene identification in introgressive
lines of common wheat. Ann Wheat Newslet 42:79-80.
Payne PI, Law CN, and Mudd EE. 1980. Control by
homoeologous group 1 chromosomes of the high-molecular-weight
subunits of glutenin, a major protein of wheat endosperm. Theor
Appl Genet 58:113-120.
Shepherd KW. 1968. Chromosomal control of endosperm
proteins in wheat and rye. In: Proc 3rd Inter Wheat Genetic
Symp (Findlay KW and Shepherd KW eds). Austral Acad Sci, Canberra.
pp. 86-96.
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