doi: 10.15389/agrobiology.2012.5.70eng

УДК 633.521:631.524.86:577.2


S.N. Kutuzova

The resistance to diseases may be a more important trait than high productivity in variety, as a damage caused by the pathogen sometimes becomes the limiting factor in cultivation. The first domestic varieties of fibre flax relatively resistant to rust have a polygenic resistance and maintain this property for a long time. An extension of such resistant varieties permitted to reduce the loss of crop and to terminate the lesion by 1998. From 1990, the varieties with short-lived oligogenic resistance are grown, and they currently occupy more than 80 % of acreage. That can be dangerous for flax-growing, if to use the effective resistance genes out of control. For creating varieties with the most reliable protection from the pathogen, secured by effective dominant genes against the background of polygenic resistance, the donor varieties, combining both types of resistance were isolated in N.I. Vavilov Institute of Plant Industry (VIR), and now they may be proposed for use in breeding programs.

Keywords: flax, varieties, genes of resistance to rust, oligogenic resistance, polygenic resistance, donors of resistance.


The resistance to diseases is often more important trait of a plant variety than productivity, because a damage caused by the pathogen can be a limiting factor in cultivation. Plant immunity to infectious diseases was in focus of the first significant scientific work of N.I. Vavilov (1), and he had been studying this issue until the last days. N.I. Vavilov investigated a large number of cereals and other crops in terms of their resistance to pathogens, which data he used to formulate theoretical fundamentals of plant immunity to parasitic organisms (2). Later it became a basis of Vavilov’s theory of plant immunity.
Flax rust is a disease of flax caused by the fungus Melampsora lini (Pers.) Lev. It is widespread over the world and regularly affects flax fields at various degrees up to epiphytoties, which causes a substantial damage (3-6). The most promising way of crop protection is the establishment of resistant varieties protected by one or more dominant resistance genes providing complete immunity to particular race of the pathogen, though not excluding sensitivity to the others. Today there are 34 R-genes with various efficiency described in different regions of the world (7, 8). The weak point of such resistance is the probability that pathogen overcomes it by changing the racial composition of population, which really happened many times in a history of flax growing (6, 9-11). This reduces  the reserve pool of effective R-genes; for example, in South America, the immunity to flax rust by 1975 had been controlled by only two genes - P4 and M6 (11). Solving the problem of immunity in flax through the more prolonged polygenic (horizontal) resistance, as it was done in other crops, particularly cereals, hasn’t been reported in the world scientific literature.
In Russia, flax rust has been known since 1885; the most severe expansion of this disease occurred in the early XX century (12), and later it was described as widespread over the whole acreage. Flax rust epiphytoties were recorded in 1924-1927 (13), 1934-1935 (14), 1959-1960 (15), and in 1966-1967 (16). In 1977, flax rust caused damage in Kirov and Perm regions, all the territory of Ukraine, Central Russia and Siberia (17). In the period from 1977 to 1980 the rust disease of flax was recorded in Volga-Vyatka region and Ukraine, where a warm wet weather persisted in late summer (17). In following years, the disease annually occurred in some areas at somewhat smaller degree, without epiphytoties (18, 19). By 1984, the disease affected only 1-2% crop. Since 1998 flax rust hasn’t been longer observed (20).
The crop damage from flax rust was reducing proportionately to the involvement in large-scale use of rust-resistant varieties. In the mid-1960s, medium-susceptible and resistant flax varieties were grown in 15-18% acreage and the disease affected 85% of the crop (20). In this period the loss in seed production from flax rust ranged from 92-94% up to 100%. The yield of flax fiber decreased by 2-3 times, its quality – in 3-9 grades (21). Then, by 1984, resistant varieties occupied 94% of the area, and 19 of 24 zoned varieties were relatively immune to flax rust, so the disease affected only 2,1% of the crop. The exception was the Tomsk region, where flax rust damaged 85,2% area of the susceptible variety Tomsky 10. In 1998, 19 flax varieties of the 31 included in the National Register of Breeding Achievements were medium-resistant, 10 - highly resistant, so resistant varieties were grown in 95,2% area (20). According to the report of 2010 (22), highly resistant varieties are grown in more than 80% acreage. However, the world history evidences that disregarding even a terminated disease may be erroneous and quite dangerous.
The nature of rust resistance in domestic flax varieties was investigated using multi-level evaluation (23); it was revealed that relatively resistant varieties grown in the 1960-1980s were protected by a polygenic (horizontal) resistance to flax rust, while none of them had somehow effective dominant genes (vertical resistance). This fact was revealed in low infectious background with 50 clones of the pathogen (24). Such resistance was manifested by longer incubation period, smaller size and number of pustules, smaller yield and lower viability of spores along with earlier transition to teleutospore stage while the reduce in number of uredo-generations. All of these features suppress propagation of the pathogen and reduce probability of epiphytoties. Such type of resistance is controlled by a great number of minor genes individually weakly manifested and enabled in different periods of plant development. This type of resistance is more effective the greater is epiphytotic severity and expansion, and it is almost independent on racial composition of the pathogen’s population. In cv Orshansky 2 and its derivative VNIIL 11 polygenic resistance is complemented by age-dependant immunity – after flowering the infestation of plants by uredospores doesn’t occur, which also reduces the risk of epiphytoties. Total effects of all available types of resistance provided quite high horizontal resistance in zoned flax varieties in the 1980-ies (K-6, Orshansky 2, Vperyod, Orshansky 72, Progress, etc.) (24).
The varieties with polygenic resistance were established through the individual selection from hybrid populations against provocative and infectious backgrounds in years of natural epiphytoties. The donor of resistance was cv L-1120 zoned in 1951. This variety and its derivatives were used as a basis for development of 37 new cultivars (10 of them included in the State Register of Breeding Achievements permitted for use). These varieties are protected by genetically homogeneous rust resistance, which doesn’t exclude epiphytoties. However, as it has been shown (24, 21), the immunity of cv L-1120 is provided by polymeric genes and quite effective age-dependant resistance. These mechanisms provide the retention of relative rust resistance in flax for nearly 60 years.
Tomsky 16 is the first selection variety with effective vertical resistance zoned in 1990. By 1998 there were already 10 oligogene-protected resistant varieties among 31 included in the State Register of Breeding Achievements, or 32% of total, and it became 56% by 2010 (Table 1).

1. Distribution of zoned flax varieties (relative to a total number, %) by types of resistance to Melampsora lini (Pers.) Lev. according to the data of testing in artificial infectious background (St. Petersburg-Pushkin, 1963-2010)

Resistance type

The year of attachment to the Stage Register of Breeding Achievements




























Total number of varieties







The improvement of rust resistance in zoned flax varieties in the period from 1960-ies up to nowadays was based mainly on polymeric genes, and since 1990-ies – on dominant R-genes. In the last few years the number of medium-resistant varieties with polygenic immunity has been reducing, while the proportion of oligogene-protected ones - sharply increasing, and susceptible varieties are becoming rarer.
The dominance of varieties with oligogenic resistance can be quite risky when effective genes are used with no attention to territorial location of varieties. The international experience of using R-genes in cereals shows that the more effective is a resistance gene, the less frequent it should be in a crop (25).
Uncontrolled inclusion in different varieties of the same gene or set of genes from a small pool of diverse R-genes will provide complete immunity, but it also can provoke the occurrence of supervirulent pathogen races and eventually – a new epiphytoty. Varieties with monogenic resistance are usually unstable and easily overcome by the pathogen. A negative experience of this sort is known from the past of domestic flax industry. By the 1935-1936 in the USSR there were established and zoned 9 varieties of flax with complete immunity to rust - 1288/12, Pryadil’schik, Svetoch, Pobeditel’, etc.; they maintained the resistance for 7-10 years (26). However, by the 1943-1945 they were attacked by flax rust even severier than older non-resistant varieties (10). The reason of this fact some authors see in a super-aggressive pathogen resulting from changes of races in the population (27, 28).
At the same time, the number and originality of genes that protect modern resistant varieties included in the State Register of Breeding Achievements are still unknown. Unfortunately, the description of variety at its attachment to the Register does not bind specifying how many and which genes or donors of genes were used at its establishment. This all means that favorable epidemiologic situation on flax rust persisting for a few last decades under successful polygenic control may be compromised. Prevention of risks requires the use of approaches providing longer effect of R-genes and varieties with stable immunity, which must be done considering both international experience of breeding for resistance and local characteristics of the pathogen.
Deceleration of pathogen’s adaptation to new resistant varieties is the purpose of special programs on establishing convergent varieties with several R-genes providing better protection than a single gene (29), and multi-line varieties consisting of lines with different R-genes. These genes located separately were found to be more effective, because such state simulates stabilizing selection (30). However, such programs are very complicated, time-consuming and expensive. Another approach is mosaic arrangement of cultivated varieties protected by different resistance genes (31). This “geographical” equivalent of growing multi-line variety is much cheaper since it doesn’t need many compulsory backcrosses and “fixing” a breeder on one recurrent variety for years (30). Because the major task of breeding for immunity is not a complete suppression of disease but prevention of epiphytoties, the actual practice is aimed at the development of component varieties based on the principal of heterogeneous lines and compositions of lines where one third has a specific resistance. It is believed that the mix of three or four lines is better and more reliable for a long-term resistance (32).
The most efficient and secure method of breeding for disease resistance is establishing the varieties with both oligogenic and polygenic resistance, which can be done by introducing the maximum number of minor genes along with major genes.  The variety immune to field races of the pathogen and simultaneously resistant to all its races is considered as the best for prevention of epiphytoties. Though this is a difficult task since it’s impossible to assess polygenic resistance of such variety (33).
Practical studies of flax rust have shown that its local populations in our country have consistently low virulence (21, 34, 35). The pathogen reproduces itself only through annual sexual process, after which pathogenicity of the population weakens due to heterozygosity of races. During the summer, reproduction of uredospores is influenced by stabilizing selection, which also eliminates the races with a broad range of virulence (34). Despite of it, the facts suggest probability of new virulent races aggressive to previously resistant varieties (35, 36), so no this risk is still actual.
Breeding flax for rust resistance by decelerating the pathogen’s adaptation to a new variety can be successful when it involves a large pool of effective genes, the source of which is the global gene pool of flax maintained in N.I. Vavilov All-Russia Research and Development Institute of Plant Industry (VIR) (37). This collection was started in 1922 by N.I. Vavilov; he investigated flax in more than 20 countries over the world and collected more than 560 original samples (38). Today there are 5795 samples including 2225 samples of fiber-producing flax.
Rust resistance is a quite rare trait in fiber flax. Only 0,6% rust-resistant types – donors of original efficient R-genes - were identified during a preliminary assessment among the 1,5 thousand samples of VIR collection in 1980 and further genetic studies and screening of resistant samples (39, 40). Most of them were representatives of several indigenous “kryazh” (a local form of flax producing unique type of fiber naturally associated with specific soil, climate and quality of seeds) derived from different provinces of Russia as earliest in the collection (41). Several donors were revealed among foreign varieties of flax (42). A large number of effective R-genes were detected in oilseed flax (43); later they were identified (44).
19 donors of rust resistance with a number of commercially valuable traits were established through a series of saturating crosses of lines with highly-efficient R-genes (37, 45). 11 of these donors – descendants of a recurrent variety Orshansky 2 – carry the original R-genes effective against all modern races of the pathogen, and they express horizontal resistance which was successfully passed through three backcrosses (Table 2).

2. Donors of resistance to Melampsora lini (Pers.) Lev. established on the base of recurrent variety Orshansky 2 (line 3-3) (45)

№ in VIR catalogue


Maternal form

Gene or locus



F8BC3 line 1-1 of Leona (k-6297 )



VIR -2

F8BC3 line 5-1 of Bombay (k-4069)

N  or Q


VIR -3

F6BC4 line 10-1 of Yaroslavsky kryazh k-759 (k-7704)



VIR -6

F8BC6 line 3-1 of Pskovsky kryazh k-729 (k-7701) 



VIR -7

F8BC6 line 7-1 of Yaroslavsky kryazh k-744 (k-7703)



VIR -8

F9BC6 line 6-1 of Soletsky kryazh k-726 (k-7700)



VIR -9

F8BC3 line 4-1 of GDS-3 k-5375 (k-7705)



VIR -10

F7BC3 line 3-1 of Pskovsky kryazh k-733 (k -7702)

M or P


VIR -11

F8BC5 line 3-1 of Currong k-6608 (k-7706)

N, P orK


VIR -15

F9BC3 line 3-1 of Svalof k-6658 (k-7707)



VIR -16

F8BC3 line 4 of Natasja k-7213 (k-7708)


Note. VIR – N.I. Vavilov All-Russia Research and Development Institute of Plant Industry.

These donors produce a fiber of high quality, good yield of seed, lodging resistance; some of them are relatively resistant to Fusarium wilt. They slightly yield to modern varieties in fiber content, so breeding work on these samples should include hybridization with high-fiber parents (45).
Using the donors with the phenotype of cv Orshansky 2 allows a reliable prevention of epiphytoties through the introduction of effective R-genes for polygenic resistance into flax varieties. Should one R-gene lose its efficiency, the plants will express a chlorosis with very small uredopustules not leading to teleytospores, so the crop will be secure.
The variety Prizyv 81 was used as a base for establishing 8 donors that express rust resistance (Table 3) along with early maturation, high content and quality of fiber, and good productivity. These donors may carry the same or other R-genes, because each line is protected by two genes. Thus, it was found that VIR-15 and VIR-13 both developed upon a common parental line carrying two R-genes, are protected by different resistance genes. Donors VIR-16 and VIR-14 also have different R-genes.
The transfer of rust resistance genes in a new developed variety now is possible using just a single hybridization with a donor. F1 hybrids should be threshed individually by plants. Among hybrid F2 families grown in a hard infectious background the breeder should select the lines not splitting for rust resistance, which after testing the immunity of following generation can be used in any breeding programs; in this case the split off susceptible individuals is highly improbable.

3. Donors of resistance to Melampsora lini (Pers.) Lev. established on the base of recurrent variety Prizyv 81 (45)

№ in VIR catalogue


Crossing scheme

Gene or locus



F6BC3 {k-7472 × line 3-1 of Pskovsky kryazh k-729 (k-7701)} ×  line 7 of k-7472



VIR -5

F6BC3 {k-7472 × line 4-1 of GDS-3 k-5375 (k-7705)} × line 7 of k-7472



VIR -12

F9BC3 {k-7472 × line 3-1 of Pskovsky kryazh k-729 (k-7701)} ×  line 7 k-7472



VIR -13

F15BC3 {k-7472 × line 2-1-1 of Svalof 60132 k-6658 (k-7707)} × line 7 k-7472



VIR -14

F5BC3 {k-7472 × line 4-1 of Natasja k-7213 (k-7708)} ×  line 7 k-7472



VIR -17

F10BC3 {k-7472 × line 3-1 of Currong k-6608 (k-7706)} ×  line 7 k-7472



VIR -18

F10BC3 {k-7472 × line 10-1 of Yaroslavsky kryazh k-759 (k-7704)} × line 7 k-7472



VIR -19

F7BC3 {k-7472 × line 3-1 of Pskovsky kryazh k-733 (k-7702)} ×  line 7 k-7472

M og P

Note. VIR – N.I. Vavilov All-Russia Research and Development Institute of Plant Industry, k-7472 — cv Prizyv 81.

The introduction of polymeric R-genes into the genotype of a recipient variety requires several continuous backcrosses with a donor established upon cv Orshansky 2; susceptible individuals must be eliminated in every generation before the start of flowering along with selection by phenotype. After the last backcross the lines not splitting for rust resistance should be isolated, their progeny – tested for resistance (45).  
The All-Russia Research and Development Institute of Flax (VNIIL, Torzhok) is another scientific center that carries out investigations aimed at obtaining sources and donors of flax rust resistance against the infectious background. Among the world gene pool of flax there were revealed a significant number of samples with different levels of rust resistance (46). Several highly producing lines protected by one or two R-genes effective against the local population of flax rust were obtained through a series of simple and complex crosses; one of such genotypes carries two R-genes one of which presumably controls age-dependant immunity (47). The varieties of flax established in VNIIL show a number of important commercial traits, although using them in breeding work must consider that most of them are protected by the same gene inherited from cv Uspekh.
The establishment of a new variety usually takes 15-17 years. Using a donor of R-genes, which is actually semi-finished variety, can twice reduce the time needed for creation of varieties with long-lasting immunity to flax rust. VIR collection was used to develop the new high-quality source material suitable for any breeding programs of creation of new varieties with reliable long-term immunity to flax rust.
So, in 1980-1990-ies domestic varieties of flax expressed relatively stable rust resistance supported by polymeric genes. The widespread use of these varieties contributed to a gradual decline in crop damage up to its termination by 1998. However, since 1990 flax varieties with oligogenic resistance are being increasingly involved in a large-scale use. Today, olygogene-protected varieties are 56% of total, and they are grown in more than 80% acreage. In this case, uncontrolled use of resistance genes can bring a risk on a whole flax industry. To obtain new varieties with good resistant to flax rust pathogen it is recommended to use the donors of rust resistance developed in N.I. Vavilov All-Russia Research and Development Institute of Plant Industry (VIR) that combine both types of resistance.


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N.I. Vavilov Research Institute of Plant Industry, RAAS,
St. Petersburg 190000, Russia,

Received May 10, 2012