View of Initial material for solving the main problems encountered in breeding winter rye in the northwestern zone of the USSR

Maataloustieteellinen^Aikakauskirja Vol. 60: ^215—221, 1988

Initial material for solving the main problems encountered in breeding

winter

in

of

USSR

V.D. KOBYLYANSKIJ

The N.I. Vavilov All-Union

Scientific

Research Institute

of

190000 Leningrad, USSR

Abstract. The main problems encounteredinbreedingwinterryeinthe northwestern region are:winterhardiness,lodgingresistance, disease resistance and grain quality. Winterhardiness inthe northwest of the USSR depends^onthe resistance of^ryetosnowmould. Varieties from the USSR (Vyatka, Falenskaja, Udinskaja, Narymskaja23andsomeothers) and Austria (Lun- gauer Tauern,Schlager)aswell^{as many}varieties from Finland^present valuable material for breedingfor winlerhardiness. The breeding of short-strawed varieties waspromotedconsider- ably bytheuseof donors with the dominanttypeof inheritance of the short straw characteris- tic (EM-1, k-10028) and manyforms and varieties created ^on their basis. Nine commercial varieties withahigh yield potentialand winterhardinesswerebred. Short-strawed donors resistant to powdery mildew and brown rust (Imering4,Fletera2,Immunnaja5), uniqueintheir kind and veryvaluable,werebred by introgression of thegenesforresistance, which had been dis- coveredinthe populations of wild species and older varieties. Astrategyof breeding rye for disease resistance is suggested.Inbreeding to improve grain quality,a genebank ofvarieties, inwhich the stability of high grain quality isretained, is valuable. Some varieties havea pro- tein contentinthe grain of^10%,withalysine contentinthe protein of4 °/o or more(Marien- hofer, Wieselburger, Kisvardai). Four short-strawed analogues of commercial varieties (UraTskaja HI,TchishminskajaHI, KustroHI, StalrugHI) areclose to the variety Otelloas totheir baking qualities. The material presented^onthe inheritance of characteristics contrib- utes tothe selection ofdonors whennewvarieties are bred.

Index words: winter ^rye, winterhardiness, diseaseresistance, grain quality,donors

Introduction

During the last 10 years, certain progress has been observed in the breeding of winter rye in the USSR. Some varietieswerebred that

have improved winterhardiness and lodging resistance _as compared to formed varieties.

New commercialvarieties, ^such as Voskhod 1, Voskhod 2, Chulpan, Saratovskaja 5, Yaroslavna, Kharkovskaja 12 and some JOURNAL OFAGRICULTURAL SCIENCEIN FINLAND

others, ^haveayield potential upto6—B t/ha.

However, realization of the potential yieldand of grain quality in many varieties is seldom attained in actual crop production, because of unsolved problems involvingwinterhardiness, lodging resistance, disease resistance ^and resistance ^to grain germination in the ear.

These problems areof particular importance torye breeding in northwestern regions of the Europeanpart ofthis^country. The future task of breeders isto createvarieties retaining high productivity under varying climatic conditions (Kobylyanskij 1982, ^{Popov et} al. 1986, SOLODUKHINA 1986).

With the object of solving different problems of breeding, the All-Union Scientific Research Institute disposes of the gene bank of specimens in thecollection, in whichnear- ly all the world’s rye is presented. Improve- mentof the^sourcesofthe genes, which regu- late characteristics useful to breeding pro- grammes, contributesto the breeders’ possi- bilities _to improve varieties, bringing them closer to the ideal stipulated by the pro- gramme.

Material and methods

The gene of rye comprises: ^sources and donors with characteristics of high winter- hardiness ^(frost resistance, ^{snow mould} resistance), lodging resistance (short-strawed donors and dwarf donors with a dominant and recessive genetic control), donors with dis- ease resistance (powderymildew, ^{brown and}

stemrusts, snowmould), sourcesof the genes with high grain quality (high protein and ly- sine ^content,resistance of the graintogermi- nation in the ear).

Modern methods of breeding are basedon the use of multiple crossings which make it possibletocombineamaximum of necessary characteristicsand properties inone genotype.

In manycases a cycle ofcrossesis carriedout, using the method ofconvergent breeding and the method ofrecurrent orreciprocal_crosses when I—21—2 valuable characteristics ^{are re-} quired for improvement of the variety. The

latter in combination with selection is very ef- ficient in improving varietiesastothe charac- teristics ofwinterhardiness, ^{immunitytofun-} galdisease,shortness ofstrawand grain qual- ity.

When intraspecificand, particularly, inter- specific hybridization is carried out,notonly the combination^ofdesired characteristicsⁱⁿ plants is achieved, ^but also ^therearises the possibility of obtaining valuable transgressive characteristics absent from the varieties_orspe- cies to be crossed. Selection from ^hybrid populationsis^thenefficientwhen pairor mass cross-pollinations of plants are carried out, these having been singled out previously ac- cording to flowering under provocativecon- ditions (Kobylyanskij 1982).

The most widely used method of breeding winter rye in the USSR is a manifold indi- vidual-strain selection and strain-group selec- tion from cross-pollinated intervarietal hybrid populations knownasthe method of compli- cated populations (Krasnyuk 1946). During the last 10 years, creation ofvarieties, ^orsyn- thetics, has been popular, and is based on combining strains with general combining ability (GCA), the effect of populational hete- rosis being used. When creating the synthetic variety Voskhod2, Goncharenko (1979) dis- covered heterosis in the strains with the best grain yield of 6.5—33.4 ^%, 1000 grain weight 3.8 °7o,grain number in theear 3.1 */o.

Results and discussion

Advances in the breeding of winter rye are universally determined by the rich genetic potential of theinitial material, its suitability to the soil and climatic conditions of the_zone of breeding, and knowledge of the principles governing the inheritance of characteristics used for breeding work.

In Europe, winterhardiness is determined by resistance to damping off, resulting from being affected by thesnowmould pathogen, Fusariutn nivale. In winter the lowtempera- tures under the snow cover never reach the level dangerous to plant life. The search for

initial material^{to use}in breeding for resistance was conducted inclimatic zonesfavourable to the development of thesnowmould pathogen.

Sources of resistance ortolerancetobeing af- fected bysnowmouldwerediscovered among rye species and varieties, the provenance of which^areboth mountainregions of the Cau- casusand Asia and lowland northern regions of Europe and Asia.

In the populationsof ^some varieties, 10^— 20 ^% of thegenotypes are immunetothe ef- fects of F. nivale under conditions of ^severe epiphytotics. These include varieties from Siberia Udinskaja, Mnogokoloska, local rye k-7629, k-9612, k-3851, and from Austria Lungauer Tauern, Schlager. Most of^the locally bred varieties from Finland and north- ern regions of the USSRare noted for ahigh degree of tolerancetosnow mould infection.

With the object of creating resistantvarieties, the following varieties^maybe recommended:

from the USSR Vyatka, Vyatka Severnaja, Falenskaja, Narymskaja23,Kamalinskaja 13, Onika; local varieties from Austria Las-

saer, Kefermarkter; local varieties from the FRG Otterbacher, Bergroggen; varieties from Finland Onni, Ponsi, Ensi, ^Kelpo, Jussi. Of the wild species, the subspecies Kuprijanovii possesses the highest degree of resistance.

The dominance of resistance or tolerance characteristicsto snowmould contributedto theuseof backcross without interruption, the aim being to transfer resistance to modern winter rye varieties.

Specimens from the collection tolerantto snowmould infectionwereused in the breed- ing ofnew winter-hardy short-strawed^varie- ties: Korotkostebel’naja 69,Korotkostebel’na- jaTetra, Volkhova.

Resistance to lodging

A study of the gene bank of short-strawed winter rye varieties and specimens contributed to discovering the genetic differentiation of the short-strawcharacteristic,the independent genetic control and the linkage of everytype

of shortness of straw with productivity ele- ments andgrain yield. Nocorrelation^{was dis-} covered between the short-strawcharacteris- tic and winterhardiness in plants (Kobylyan-

sku 1982).

Of the four main short-strawedtypes, only two are well suited to creating the highest yielding varieties.

1. The polygenic recessive typeof shortness of straw, withintermediate inheritance^{of the} characteristic and influence of the protoplasm onthe degree of its manifestation. The^sources of thischaracteristicarethepresentWest Eu- ropean varieties and ^somevarieties from^the USSR created on the basis of varieties from Petcus, GDR: Danae, ^{Kustro, Kungs 11,} Dominant, Voskhod 1, Kombaininyaj, Tul’vi, Saratovskaja 5 and someothers. Thistypeof shortness ofstraw is widely and successfully used in ^rye breeding.

2. The monofactor dominanttypeof short- ness of straw is controlled by the HI gene (Dwarf 1). The donors of resistance are the naturalmutantEM-1, ^singlegenotypesfrom local Bulgarian rye k-10028 and the specimen from Poland k-11. Because of the rather low yield ability of the above donors, ^breeding programmes use numerous winter rye forms and varieties with high yield, createdon the basis of the mentioned donors.

The dominant shortness of straw charac- teristic^{was foundnot to}be genetically linked with othercharacteristicsreducing grain yield and decreasing the winterhardiness of plants.

The pleiotropic effect of the gene regulating the short^strawcharacteristic results in shorter internodes andstrawup to40®7o, anincrease in the bush habit of plants, in leaf and ear sizes, and in spikeletsand grain in theear. In- dependent genetic control of the characteris- ticcontributestocombine the dominant short- strawtype with othertypes of recessive short- ness of straw(Kobylyanskij 1975).

The dominanttypeof shortness ofstrawis used inrye breeding programmes by all the breeding institutions in the USSR. During a short period since 1979, nine short-strawed commercial ^{varietieswere}created: Chulpan, 217

Chulpan3,^Talovskaja 12,Khar’kovskaja 78, Orlovskaja 9,Kryzhachek, Verasen’,Korot- kosbel’naja 69,Tetra,Korotkaja. The lasttwo arecultivated inwestern Siberia.In different climatic regions of thecountry, over40 °7o of the winterrye sowingsare of the above varie- ties.

Resistance tofungal diseases

Intensification of the technology of winter rye cultivation made topical theproblem of plant protection from harmful pathogens. The harmfulness of fungal diseases increasedcon- siderably when cultivatingshort-strawed ^var- ieties. In sowing the microclimate is made favourable to the development of all the pathogens, while shortening of the shorter stem increased the role of the leaves in the storage of the products of photosynthesis, whichare necessary forgrain yieldformation.

The situationarosein which similar infection of the leaves of long-strawed and short- strawed plant varieties have a greater effect on the short-strawed varieties. In short- strawed diploid rye, powdery mildew infection of 30—60 °7o reduces grain yield by 28—

47 ®/o. Thesame analogy is also observed with brown_rust infection. Plant infection of50^— 70 ^%results in a 21 —39 ^%reduction of the yield, due to a decrease in the 1000 grain weight, the harmfulactionof the pathogen be- ginning with _a damaged leaf _area of 10 ^%.

Reduction of the yield bymorethan50 ^%fol- lows from stem rust and stemfusarial rot epiphytotics (Solodukhina 1986).

In all the world’s winter rye varieties, ^there isnovariety completely resistanttopathogens.

The search forsourcesofresistance is carried

out in populations of wild species and in old cultivated rye varieties. In populations of wild species, Secale ^montanum subspecies Kupri- janovii, collected in the region of the Cauca- sus, single_genotypesimmune topowdery mil- dew, brown leaf rustand stem rust weredis- covered. Singlegenotypes partly resistant to powdery mildew, brownrust and stem rust werefound in old fodder and grain rye varie-

ties: Griinschnitroggen (The GDR), Sangaste, Novozybkovskaja 4, Derzhavinskaja 29, Khar’kovskaja 60, and insome other related varieties. ^{Based onthe} discovered^sources of resistance, methods of selection and back- crossesgive highly productive donors of these characteristics, the object beingto use them in breeding programmes.

Resistanceto powderymildew, brown^rust and ^{stem rust was}found ^topredominateover susceptibility in all the cases. In each indi- vidual case, resistance is controlled by one dominant gene the Er and Pm 2 genesfor powdery mildew, ^the Pd, ^Pd 1 genes for brown leaf rust. In our days donors _were received combining both genes (Er, Pm 2) for resistance to powdery mildew and shortness ofstraw Imerig4; donorscombining short- ness of^{straw and} resistance ^topowdery mil- dew (Er, Pm 2) and tobrown leafrust (P) Hetera2; those combining partial resistance to powdery mildew (Er) with resistance to brown rust (Pd) and donors combining dominant shortness of straw (HI) andresist- ancetosnowmould Immunaja 5. If should be observed that, ^{in all the cases, resistance}

to leafrust hasarace-specific heterogeneous nature;here,along with necroticspots(super- sensitivity reaction), differently developed pustules of the fungus are found.

Dominant genetic control of resistance to the pathogens determines the methods of selection when backcrosses without interrup- tion and introgression of the genes for resistance in highly productive genotypes of ryeare used. The selection of resistance in the genotypes in each backcross progeny should be carriedout underartificialinfected condi- tions previous toflowering of plants.

The _strategyof breeding rye for resistance todiseases is based _ondifferences in the genes forresistance, ^eachtypebeing resistant _{to a} definite group of the pathogen ^races,and on the heterogeneity of varietal populations con- tributing _to the long-term maintenance of resistance. In the breeding programme, four more probable directions ^maybe included:

creation of varieties withone typeofrace- specificresistance;

creation of polyresistant complicated populations obtainedon the basis ofcom- bining plant varieties and strains with differenttypesof race-specificresistance;

creation of polyresistant varieties by in- trogression in their populations of the genes from differenttypesof race-specific resistance;

creation of varieties of polyresistant popu- lations combining both race-specific and race non-specific resistance (Kobylyanskij and Solodukhina ^1987).

Genetic control of resistance to diseases regulated by independent genes contributes^to combining inonepopulation notonly differ- ent types of resistance_to pathogen, butalso resistance to various harmful diseases powderymildew,^{leafrust and}stem^rust, snow mould and some others.

Grain quality

The concept of grain quality in rye com- prises the following main aspects: 1) food value (protein and essential amino acid_con- tent) and 2) technological properties (mainly baking qualities). The breeding of rye for an increased protein content in the grain ad- vancespoorly,_as the problem associated with protein has been solved in many countrieson account of other high-protein plants.

The gene bank of rye comprises a large diversity ofvarieties, forms,and species with respect to protein and essential amino acid content, particularly that of lysine. Among old and bred varieties, of special interest for breeding_are the specimens witha 14^%pro- teincontentin the grain and a morethan3 ^% lysinecontentin protein: Manychskaja, Kam- linskaja 13, Sitnikovskaja (USSR); Mestnaja k-9514 (Yugoslavia); Balbo (USA); Explorer (Canada); Tisca Kozi Feher (Hungary);

Gregerovo (Czechoslovakia). Still _a higher protein content in the grain (over 17 ^%) and of lysine in protein (over 3 ^%) is discovered in single samples of weed rye, collected in regions of the Caucasus: k-5806, k-9633, k-10552. Weed rye is easily crossed with cul- tivated rye, but is primitive as to othereco- nomically valuablecharacteristics ^itis ^not lodging resistant, and gives poor yields.

The crossing of high protein varieties with low protein types gives intermediate in- heritance of protein in the grain. Hybrids of ahigher protein_type(by 15 ^%)areobtained when the maternal components of thecross have ahigher protein content.

With the object of breeding for theutmost high lysine content, initial material is disco- vered with aprotein _content in the grain of

10^%andalysinecontentin protein over4^% (Table 1).

Among thevarieties^withahigh lysinecon- tent, specimens comprising high grain quali-

Table I. Rye varieties high in lysine content.

Variety Provenance 1000grain Protein ^% Lysine

weightg (Nx 5.7) ^{~ ~ ~}

gm/100 vo of

per g protein grain

Marienhofer Austria 38.2 10.3 0.431 4.19

Wieselburger ^» 41.4 10.2 0.427 4.20

LungauerTauern ^» 31.5 10.1 0.410 4.07

Fleischmanns Hungary 31.0 10.9 0.464 4.27

Lovaszpatonai ^» 36.0 11.3 0.464 4.11

Kisvardai ^» 41.0 10.8 0.461 4.28

Nagucenki ^» 35.7 10.6 0.437 4.13

Falenskaja USSR 28.6 10.0 0.423 4.23

Gibridnaja2 ^» 36.8 10.5 0.428 4.07

Volzhanka ^» 31.3 11.1 0.446 4.03

tytogether with large grains, this beingaposi- tive fact for^yieldincrease, have special value.

Among high lysinecontentvarieties,apartic- ularly large grain (38.2 —41.4 g) is found in Marienhofer,Wieselburger (Austria) and Kis- vardai (Hungary).

The problems of improving the baking quality of rye may be solved by breeding work when varieties resistant togerminationarein- volved in the hybridization. The Swedishvar- iety Otello is very suitable for this purpose.

Among local old rye varieties from the USSR (Yakutia, Kazakhstan), the samples k-10409 and k-8016 were found superiortothe world standard Otello as to this characteristic.

Inrecentyears, basedonshort-strawed_ana- logues of rye varieties and forms, reliable sources of resistance to grain germination in the ear were created. EM-1, the donor ^of shortness of^{straw,possessesa}high degree of resistance to germination, close to that of Otello.The value of the amylogram of EM-1 grainwas notedto increase from 50u.am. in 1976to 430 u.am. in 1986,reproduction of the seeds having been carried ^{out during 10} years under conditions ofhigh air humidity (in the surroundings of Leningrad). In this connection, it is suggested that systematical year-to-year growing of short-strawed rye forms under conditions contributing to ger- mination of the grain in theearautomatical- ly results in the elimination from the popula- tion of thegenotypes inclinedtogermination, and also results in thestorageof thegenotypes

Table2. Short-strawed analoguesof winterryevarieties best for grain quality.

Variety Amylogram Fallingnumber

x range x range

Otello 325 225—430 275 201—346

Ural’skajaHI 407 300—490 264 224—335 Chishminskaja HI 300 240—370 226 170—291

Kustro 373 290—520 234 193—261

Stalrug 347 280—405 320 280—354

resistant to germination. In short-strawed sawings of rye, a higher air humidity is re- tained as compared to that in long-strawed

ones,particularly in the sawings in the north- western region of thiscountry.

Newsources of high baking quality^{are non-} lodging short-strawedanalogues of commer- cial varieties Ural’skaja HI,Chishminskaja 3

HI,^{KustroHI,StalrungHI,}all of them being closeto the variety Otelloas totheir techno- logical properties (Table 2).

Hybrid analysis revealedadominant digenic additive genetic control of the characteristic of low alpha-amylase activity in the grain, i.e.

of resistance to germination (Muller and Peters 1979). The above determines the selection of the methodtouse for breeding rye for grain quality. The method comprises the use ofthe sources of resistance ⁱⁿ backcross crosses along with individual strain selection of the plantstobe backcrossed under provoca- tive conditions.

References

Goncharenko, A.ASelekciyaisemenovodstvo. 1979,No 1. 9—ll.

Kobylyanski,V.D. 1975.Rozh. Avtoreferat Dissertacii.

L. 57p.

1982.Rozh. Geneticheskiye^osnovyselekcii. M.:271 pp.

&Solodukhina, ^{O.V. 1987.} Strategiyaselekciirzhina

ustoichivostk boleznyam.Bull. ^VIR, No. 171.

Krasnyuk,A.A. 1946.Agrobiologiya,No. 2: 44 —51.

Muller, H. ^&Peters, C., 1979.Stand und Möglich-

keiten der Entwicklung ertragreichter Roggensorten mit verbesserer Auswuchsfestigkeitinder DDR. Archiv

fiirZiichtungsforschung.H.5. 8.9.

Popov, G.1., Vasko, V.T. ^& Pucach, N.G. Selekciya ozimoi rzhi. 1986. L.: 240.

Solodukhina, O.V. 1986.Sozdaniyedonorov ustoichi- vostikburoi rzhavchineimuchinistoirosedlyaselek- cii diploidnoirzhi. Avtoreferat dissertacii. Leningrad.

19p.

SELOSTUS

Neuvostoliiton luoteisalueensyysrukiin- jalosluksen perusongelmien ratkaisemisen lähtöaineisto

V.D. Kobylyanskij

The N.I. Vavilov All-UnionScientific

Research Institute ofPlant Industry Herzen Str. 44

190000Leningrad, USSR

Luoteisen alueen rukiinjalostuksen pääongelmat ovat talvenkestävyys,lakoontumattomuus, taudinkestävyys ja jyvänlaatu. Talvenkestävyysonriippuvainenrukiin kes- tävyydestälumihometta vastaan. Arvokasta aineistoa tal- venkestävyysjalostusta varten tarjoavat neuvostoliittolai- set lajikkeet Vjatka, Falenskaja, Udinskaja, Narymska- ja 23 jaeräät muut, itävaltalaiset Lungauer Tauern ja Schlagersamoinkuin monet suomalaiset rukiit.

Lyhytkortisen^rukiinjalostusonedistynytsuuresti,^kun risteytyksissäonollut käytettävissä lyhytkortisuuden val-

litsevana ominaisuutena sisältäviälähteitä,kuten EM-1 jak-10028. On jalostettu yhdeksän kauppalajiketta, joilla onkorkea sadontuottokyky ja hyvä talvenkestävyys. Ly- hytkortisethärmän- ja ruskeanruosteenkestävyydenläh-

teet Imering4,^Hetera2ja Immunaja5ovat erityisenar- vokkaita. Neonjalostettusiirtämällä villien lajien ja al- kukantaisten rukiinen populaatioistakestävyyden tuot- tavia geenejä introgressioiden kautta. Artikkelissa esite- täänrukiin resistenssijalostuksen strategia.

Jyvän laadun parantamisessa tarjoavat geenipankissa ylläpidetyt hyvän jyvänlaadun omaavat lajikkeetarvok- kaan perustan. Muutamien lajikkeiden, kuten Marienho- fer, Wieselburger jaKisvardai, valkuaispitoisuuson 10 prosenttia, lysiinipitoisuuden ollessa4prosenttiatai yli sen.Neljän kauppalajikkeen, Ural’skajaHl,Tchishmin- skajaHl,Kustro HljaSlälruisHl, lyhytkortisuus-ana- logienleivontalaatu onlähellä Otello-lajikkeen tasoa.

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