Tuesday 16 January, 2007

BREEDING OF PERENNIAL FODDER FORMS OF WASHINGTON LUPIN




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


Kurlovich, B.S. and Heinänen, J.

International North Express Co, Finland



Abstract
The aim of the work was to develop methods to create fodder (sweet) varieties of Lupinus polyphyllus Lindl, 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 varieties of multifoliate lupin with sufficiently stable and low alkaloid content. Positive results were demonstrated 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 hardiness and frost tolerance, capable of producing high yields of green matter (40 - 50 Mg ha-1) and maintaining 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 Australia.


Key words:Alkaloids, allelic and complementary tests, cross-pollination, L. polyphyllus Lindl., methods of breeding, low alkaloid (sweet) lupin.

Introduction
Perennial Washington lupin (L. polyphyllus Lindl.), widely distributed in Belarus, Finland, Russia, Poland and other countries, was introduced into Europe as an ornamental plant and for the production of green manure. Natural populations of this species are found at the edges of woods and fields, along roadsides, and in parks and on the sites of former manors. It was widely popularized by Academician Pryanishnikov (1962) who recommended this plant as the best source of green manure. It was also championed by G. Hill as a source of forage and as an ornamental plant in New Zealand (Hill, 1994; Hill and Tesfaye, 1994). These researchers demonstrated 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 fertilizers. Perennial types of lupin, 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, (Aniszewski, 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.


Material and Methods


Research material was obtained from the lupin collection of the N.I. 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.I.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, Chernigovskij, Stodolishchenskij, and Dagestanskij.

However, bitter (high alkaloid) plants appeared in these accessions during our research. This may be due to the accessions containing different recessive alleles. Cross-pollination of these forms resulted in F1 heterozygotes with dominant alleles at all loci (AL1al1 AL2al2 AL3al3),thus leading to the restoration of alkaloid synthesis. To overcome the aforementioned 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 of low-alkaloid plants were selected as initial material, and (2) every low-alkaloid plant was crossed with an alkaloid producing form, which had other valuable characters, and populations with stable, low-alkaloid levels were selected in the F2. 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 Washington lupin (L. polyphyllus Lindl.), new approaches were developed on the basis of specific crossing. The principal, distinctive feature of the approaches developed is that all initial sources of low alkaloid content, whether identified or bred, are not grown together and allowed to cross-pollinate as it was done earlier (Voronov, 1974). Only compatible forms are involved in hybridization, with their low alkaloid content controlled by the same genetic system. For the practical application of this approach, two methods were developed.

The first method of obtaining fodder (sweet) forms of L. polyphyllus 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. Populations with a stable, low alkaloid content were selected in the F2 (Kurlovich and Chekalin, 1992). By comparing the characteristics of the low-alkaloid forage lines produced by these methods we demonstrated that both are suitable for breeding low-alkaloid varieties. The drawback of the 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, etc.
The aim of the second method was to improve the process of obtaining high-yielding forms. Populations developed by the second method were higher yielding and resistant to Fusarium wilt. The productivity of these populations was practically the same as in the initial high-alkaloid forms.
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 reverse mutations or pollination of low-alkaloid (sweet) forms with the pollen of wild high-alkaloid plants. In all phases of seed production involving low-alkaloid forms of multifoliate Washington lupin, it is necessary 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 prudent 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 seeds for many years.
With these methods and the new sources mentioned above it is possible to produce sweet forage forms of multifoliate Washington lupin with sufficiently stable and acceptable low alkaloid levels. The forage cultivar Truvor was developed with this method and released in the Ukraine. The first commercial cultivar released from our own efforts was Pervenec. It is adapted to northwestern Russia and also listed in the State Catalogue of Breeding Achievements in Russia. Our major tasks in breeding multifoliate lupin in Finland since 1996 has been to produce types with stable low alkaloid content, different types of pollination (either cross- or self-pollinating), non-dehiscent pods, winter 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, dispersed between many initial forms, within one population.

One of our purposes in 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 in Australia. The improved cultivars resulting from this approach represent a new stage in breeding and will be more competitive on the world market.
An important aspect of the successful lupin cultivation is inoculation with improved strains of Bradyrhizobium sp. (Lupinus). The best results were achieved when seed was treated with strain 1625 ((Kozhemyakov et al., 1992; Kurlovich et al., 2000). Growing lupins in areas where there had previously been no lupins cultivated demonstrated that inoculation is absolutely essential for successful lupin cultivation.

The breeding methods developed may also be applied to other cross-pollinating lupin species such as L. arboreus Sims., L. perennis L., L. nootkatensis Donn., L. elegans H.B.K., L. hartwegii Lindl., with promising prospects for agricultural production.

Literature Cited

Aniszewski, T. 1993. Lupin: a potential crop in Finland. Studies on the ecology, productivity and quality of Lupinus ssp. PhD thesis summary, Joensuuu, Finland, 50p.

Chekalin, N.M., and B.S. Kurlovich. 1989. The copyright certificate on the invention, SU1499737. Moscow.

Hill, G.D. and M. Tesfaye. 1994. The characterization of Russell lupins. pp. 73-76 IN J.M. Martins and M.L. Beirão da Costa (eds.). Advances in Lupin Research. Proc. of the 7th International Lupin Conference, Evora, Portugal, 18-23 April, 1993. ISA Press, Lisbon, Portugal. 1994.

Hill, G.D. 1994. The potential of perennial lupins for Grazing. pp. 435-444 IN J.M. Martins and M.L. Beirão da Costa (eds.). Advances in Lupin Research. Proc. of the 7th International Lupin Conference, Evora, Portugal, 18-23 April, 1993. ISA Press, Lisbon, Portugal, 1994.

Kozhemyakov, A.P., N.S. Ivanov and B.S. Kurlovich 1992. Efficiency of inoculation of fodder Lupinus polyphyllus Lindl. with nodule bacteria and root diazotrops. Research Bull. VIR, 220:3-5.

Kurlovich, B.S., N.S. Nazarova, V.A. Rybnicova, S.I. Pilipenko, L.T. Kartuzova, and F. T. Tarba. 1990. Study of Lupin Samples of World Collections (Methods of investigations, in Russian). VIR, St. Petersburg, Russia, 34p.

Kurlovich, B.S., and N.M. Chekalin. 1992. The copyright certificate on the invention, SU1767714, Moscow.

Kurlovich, B.S., M.V. Petrova, A.K. Stankevich, and L.T. Kartuzova. 1995. Lupin, Theoretical basis of plant breeding. pp. 9-122. IN B.S. Kurlovich and S.I. Rep’ev (eds.). The gene bank and breeding of grain legumes (lupine, vetch, soybean, and common bean), Vol.111. St. Petersburg, VIR, Russia.1995.

Kurlovich, B.S., L.T. Kartuzova, B.M. Cheremisov, T.A. Emeljanenko, I.A. Tikhonovich, A.P. Kozemyakov, and S.A. Tchetkova. 2000. Evaluation of the biological nitrogen-fixing ability of lupin (Lupinus L.). Plant Genetic Resources Newsletter 123:68-77.

Pryanishnikov, D.N. 1962. Lupin on service to socialist agriculture. pp 5-10. IN N.A. Majsurjan (ed.) Lupin, Moscow, Russia. 1962.

Voronov, A.T. 1974. Breeding of perennial fodder lupin. pp. .243-250. IN N.V. Turbin (ed.). Breeding, seed production and growing of lupin. Orel, Russia. 1974. (in Russian).


2 comments:

Anonymous said...

Dear Professor!

Greetings from Zhitomir area of Ukraine!

We have understood from the Internet that you are in Finland now.
We are reminding you the following:
We have started to grow sweet long-term lupin in collective farm «Victory» under your management in 1991.

Now we inform you to Finland: there are more than 100 hectares of the selected by you perennial sweet lupin (cv. Truvor) in our collective farm.
We prepare silage from it; our cows willingly eat it in a fresh kind.

We are very pleased with yours sweet lupin!

Come to us again to Ukraine!

With friendly greetings,

Chairman of collective farm «Victory»

Vladimir Birychenko

Anonymous said...

Good words.