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BREEDING OF PERENNIAL FODDER FORMS OF MULTIFOLIATE LUPIN (LUPINUS POLYPHYLLUS LINDL.)

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BREEDING OF PERENNIAL FODDER FORMS OF MULTIFOLIATE LUPIN (LUPINUS POLYPHYLLUS LINDL.)
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    Wild and ultivated Lupins From the l ropics to the Poles Proceedings of the lOth International lupin Conference laugarvatn Iceland 19 24 June 2002  67 BREEDING OF PERENNIAL FODDER FORMS OF MULTIFOLIATE LUPIN LUPINUS POLYPHYLLUS LINDL. B.S. Kurlovich and J. Heinanen International North Express, Lepplilaaksontie 5 -1, Fin -52420 Pellosniemi, Finland. Corresponding author s e-Mail: bkurl@mbnet.fi ABSTRACT The aim of the work was to develop methods to create fodder (sweet) varieties of Lupinus polyphyllus Lindi, with stable, low levels of alkaloids. New approaches were developed with the specific intention of avoiding the rapid restoration of alkaloid synthesis in this crosspollinated lupin species. We created sweet forage varieties of multifoliate lupin with sufficiently stable and low alkaloid content. Positive results were demonstrated with the first sweet commercial cultivar Pervenec, which was listed in the State Catalogue of Breeding Achievements in Russia. The major tasks carried out in Finland since 1996 were the breeding of cultivars with stable low alkaloid content, different types of pollination, non-dehiscent pods, winter hardiness and frost tolerance, capable of producing high yields of green matter (40 -50 Mg ha- 1  and maintaining the shading of cover crops. One of our purposes of Washington lupin breeding in Finland is to create productive sweet forms with white flowers and seeds analogous to white flowered and white seeded varieties of narrow-leafed lupin L angustifolius L. in Australia. KEYWORDS Alkaloids, allelic and complementary tests, cross-pollination, breeding methods, low alkaloid (sweet) lupin INTRODUCTION Perennial multifoliate or Washington lupin L. polyphyllus Lindi.), widely distnbuted in Belarus, Finland, Russia, Poland and other countries, was introduced into Europe as an ornamental plant and for the production of green manure. Natural popula tions of his species are found at the edges of woods and fields, along roadsides, and in pruks and on the sites of former manors. It was widelypopulariz.ed by Pryanishnikov (1962) who recom mended this plant as the best source of green manure. It was also championed by G.D. Hill as a source of forage and as an ornamental plant in New Zealand (Hill, 1994; Hill and Tesfaye, 1994). These researchers demons1rated that it is necessary to apply green manure for the reclamation of soils with low organic matter content This practice not only improves soil properties but also raises the efficiency of mineral fertiliz.ers. Perennial types oflupin, particularly L. polyphyllus can be grown for fodder and seed in countries with limited agricultural resources. Examples are the northwest areas of Russia (Kurlovich et al. 1995), Finland, (Anisz.ewski, 1993) and other far northern countries. When low-alkaloid (sweet) forms of L. polyphyllus are produced, the prospect seems quite promising for the complex utilization of this perennial species of lupin for fodder production and green manure in horticulture and in another fields. The obstacles connected with the creation of fodder (sweet) varieties of this species for different countries are discussed. MATERIALS AND MEmODS Research material was obtained from the lupin collection of the NJ. Vavilov Institute of Plant Industry (VIR.), where 150 accessions of L. polyphyllus from 8 countries are stored. Many years (1973-1996) of observing the lupin plants sown in various regions of many countries (Belarus, Russia, Finland, Ukraine, and Abkhazia) and the use of special methods developed by us and the N.l Vavilov Institute, which are applicable to lupin (Kurlovich et al. 1990), have enabled us to identify new sources of L. polyphyllus with high yield and a low alkaloid content Belorusskkij 1, Borovlyanskij, Chemigovskij, Stodolishchenskij, and Dagestanskij. However, bitter (high alkaloid) plants ap peared in these accessions during our research. This may be due to the accessi.ons containing different recessive alleles. Cross pollination of these forms resulted in F 1 heterozygotes with dominant alleles at all loci ALpl 1 Lpl 2 AL pl ,thus leading to the restoration of alkaloid synthesis. To overcome the aforemen tioned problem, two approaches were developed: (1) selected low-alkaloid plants were crossed with each other and the progeny from each cross was grown separately. Populations consisting only oflow-alkaloid plants were selected as initial material, and (2) every low-alkaloid plant was crossed with an alkaloid producing fonn, which had other valuable traits, and populations with stable, low-alkaloid levels were selected in the F 2• We also studied the N-fixing ability of L. polyphyllus using new strains of bacteria developed at the Research Institute for Agricultural Microbiology (St Petersburg - Pushkin, Russia). RESULTS AND DISCUSSION To avoid the rapid restoration of alkaloid synthesis in cross pollinated species of lupin, and in particular, in multifoliate or Washington lupin L. polyphyllus Lind .), new approaches were developed on the basis of specific crossing. The principal, distinctive feature of he approaches developed is that all initial sources of low alkaloid content, whether identified or bred, are /NE. van Santen and GD. Hill (eds). Wtld and Cultivated Lupins from 1he Tropics to the Poles. Proceedings of1he 10th International Lupin Conference, Laugarvatn, Iceland, 19-24 June 2002. Jntematicmal LupinAsoociation, Canterblll) , New z.ealand. ISBN 0-86476-153-8.  68 PROCEEDINGS 1om INTERNATIONAL LUPIN CONFERENCE Table 1. Characteristics of ow alkaloid (sweet) populations of Lupinus polyphyllus Linell. created by different breeding methods averages for 198~2000 . Number and( ) ofiew. Fresh Methods of creating low alkaloid alkaloid Seed biomass Protein Fusarium ~sweet 202ulations N I?lants xield xield con1ent wilt gplanf 1 2 kgm-  DM - -- Conventional Method 2000 98 54 7.9 19.5 5 (Voronov, 1974; all low-alkaloid plants (4.9) are hybridized without preliminacy checking for genetic compatibility) First proposed method 2000 9 40 6.1 16.8 6 (Low alkaloid plants identified are (0.45) crossed with each other for checking genetic compatibility) Second proposed method 2000 6 61 8.0 20.4 0 Evecy low alkaloid plant, created by (0.3) first method, is crossed with productive high alkaloid form) Control 1000 1000 58 8 19.9 (Initial prodictive alkaloid producing (100) form) Standard error 5.3 0.9 2.5 1.1 Values within columns with different superscripts differ significantly at < 0.05. not grown together and allowed to cross-pollinate as it was done earlier Voronov, 1974). Only compatible forms are involved in hybridiz.ation, with their low alkaloid content controlled by the same genetic system. For the practical application of this approach, two methods were developed (fable . The first method of obtaining fodder sweet) forms of L po/yphyllus involved the pairwise crossing of selected low-alkaloid plants and growing each progeny group in isolation. Populations consisting only of low-alkaloid plants were selected initially for breeding Chekalin and Kurlovich, 1989). The second method consisted of crossing of low-alkaloid plants with productive high-alkaloid plants possessing other valuable characters. Populations with a stable, low alkaloid content were selected in the F 2 Kurlovich and Chekalin, 1992). By comparing the characteristics of he low-alkaloid forage lines produced by these methods we demonstrated that both are suitable for breeding low-alkaloid varieties. The drawback: of he first method is that it is rather labor intensive because it required a large number of low-alkaloid plants, many of whom exhibited negative properties such as low productivity, susceptibility to diseases, et.c. The aim of he second method was to improve the process of obtaining high-yielding forms. Populati0ns developed by the second method were higher yielding and resistant to Fusarium wilt The productivity of hese populations was practically the same as in the initial high. alkaloid forms Table . Our goal in developing these breeding approaches was to create low-alkaloid populations without narrowing the genetic variability in the process. These methods also feature the absence of complementary interactions of non-allelic genes, which would restore high alkaloid content However, in such fodder sweet) crops as yellow, white and blue lupin, as well as in fodder low alkaloid forms of multifoliate Washington lupin, bitter plants may appear from time to time as a result of everse mutations or pollination oflow-alkaloid sweet) forms with the pollen ofwild high-alkaloid plants. In all phases of seed production involving low-alkaloid forms of multi.foliate Washington lupin, it is neces sary to ensure strict spatial isolation between different samples. Furthermore, plants need to be tested for alkaloid level and high. alkaloid plants must be removed before flowering. Because this perennial lupin species attracts insect pollinators, it seems pru dent to establish permanent seed increase nurseries and to remove wild bitter lupin plants from adjacent areas. In the first year, all plants in the nursery need to be tested for the presence of alkaloids by pressing leaves and petioles against alkaloidsensitive paper. All identified bitter plants should be removed before flowering. Subsequently, such a nursery will satisfy the demand for a substantial production of see s for many years. With these methods and the new sources mentioned above it is possible to produce sweet forage forms ofmultifoliate Wash ington lupin with sufficiently stable and accep1able low alkaloid levels. The forage cultivar Truvor was developed with this method and released in the Ukraine. The first commercial cultivarreleased from our own efforts was Pervenec. t s adapted to northwestern Russia and also listed in the State Catalogue of Breeding Achievements in Russia. Our major tasks in breeding multi.foliate lupin in Finland since 1996 has been to produce types with stable low alkaloid content, different types of pollina tion either cross- or self-pollinating), non-dehiscentpods, winter  PROCEEDINGS lOTH INTERNATIONAL LUPIN CONFERENCE LITERATURE CITED 69 hardiness and frost tolerance. Improved cultivars should also have a yield potential in the order of 40 -50 Mg green matter ha- 1 and have the ability to shade out competing species. Individual group selection and step-by-step crossing were used to enable us to combine valuable characters and properties, dis- persed between many initial forms, within one population. One of our pwposes in Washington lupin breeding in Finland is to create productive sweet forms wi1h white flowers and seeds analogous to white flowered and white seeded varieties of narrow-leafed lupin in Australia. The improved cultivars result ing from 1his approach represent a new stage in breeding and will be more competitive on 1he world marlret. An important aspect of 1he successful lupin cultivation is inoculation wi1h improved strains of Bradyrhizobium sp. (Lupinus). The best results were achieved when seed was treated wi1h strain 1625 Kozhemyakov et al. 1992; Kurlovich et al. 2000). Growing lupins in areas where 1here had previously been no lupins cultivated demonstrated 1hat inoculation is absolutely essential for successful lupin cultivation. The breeding me1hods developed may also be applied to o1her aoss-pollinating lupin species such as L. arboreus Sims., L. pe1t 111lis L., L. nootkatensis Donn., L. elegans H.B.K., L. hartwegii Lindl., wi1h promising prospects for agricultural production. Ani.sz.ewski, T. 1993. Lupin: a potential crop in Finland Studies on he ecology, productivity and quality of Lupinus ssp. PhD thesis sum- mary, Joensuuu, Finland, 50p. Chekalin, N.M, and B.S. Kurlovich. 1989. The copyright certificare on he invention, SU1499737. Moscow. Hill GD 1994. The potential of perennial lupins for Grazing. pp. 435-444JN J.M Martins and ML. Beimo daCosta(eds.). Ad vances in Lupin Research. Proc. o he J I International Lupin Conference, Evora, Portugal, 18-23 April, 1993. ISA Press, Lisbon, Portugal, 1994 Hill GD and M. Tesfaye. 1994. The characterization of Russell lupins. pp. 73-76 JN J.M Martins and ML. Beimo da Costa eds.). Advances in Lupin Research. Proc. o he J I International Lupin Conference, Evora, Portugal, 18-23 April, 1993. ISAPress, Lisbon, Portugal. 1994. Kozhemyakov, AP., N.S. Ivanov and B.S. Kurlovich 1992. Efficiency of noculation of odder Lupinus polyphyllus Lindl. wi1h nodule bacteria and root diazotrops. Research Bull. VIR, 220:3-5. Kurlovich, B.S., N.S. Nazarova, VA Rybnicova, SJ. Pilipenko, L.T. Kartuz.ova, andF. T. Tarba. 1990. Study o Lupin Samples o World Collections Methods of nvestigations, in Russian). VIR, St Petersburg, ~sia 34p. Kurlovich, B.S., and N.M Chekalin. 1992. The copyright certificare on he invention, SU1767714, Moscow. Kurlovich, B.S., M.V. Petrova, AK. Stankevich, and L.T. Kartuz.ova. 1995. Lupin, Theoretical basis of plant breeding. pp. 9-122. JN B.S. Kurlovich and SJ. Rep ev eds.). The gene bank and breed ing of grain legmnes lupine, vetch, soybean, and common bean), Vol.111. St Petersburg, VIR, Russia1995. Kurlovich, B.S., L.T. Kartuz.ova, B.M Cheremisov, TA Emeljanenko, LA Tikhonovich, AP. Koz.emyakov, and SA Tchetkova. 2000. Evaluation of he biological nitrogen-fixing abil ity oflupin (Lupinus L.). Pla.nt Genetic Resources Newsletter 123:68-77. Ptyanishnikov, D.N. 1962. Lupin on seiv:ice to socialist agriculture. pp 5-10. JN NA Majsurjan (ed) Lupin, Moscow, Russia 1962. Voronov, AT. 1974. Breeding of perennial fodder lupin. pp. 243-250. IN N.V. Turbin ( ed . Breeding, seed production and growing of lupin. Orel, Rllssia. 1974. in Russian).
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