GENETICS OF LUPINS
B.S. Kurlovich
Genetics of qualitative characters
The genus Lupinus L. is not investigated fully and deeply enough from the genetic point of view (Swęcicki W and Wolko, 1992). The nature of inheritance of commercially valuable characters is most profoundly studied in three annual domesticated species (L. angustifolius, L. luteus and L. albus). We have clustered the respective data on each species separately.
Morphological characters
Vegetative organs. The variability on coloring and degree of pigmentation of cotyledons, stalks and leaves is typical for the forms of narrow-leafed lupin. Reddish color of cotyledons and young leaves which would in time turn green is controlled by the rut gene, and their yellow-green color is controlled by the vires gene (Hackbarth, 1957b). Different hues of yellow are caused by polymeric genes (vires 1, vires 2). The shades of anthocyanin on the stem and leaves are monitored by different allelic conditions of the Pur gene. Almost red color of stems and leaves in pink-flowered forms is caused by a dominant allele of the Pur gene. This pleiotropic gene provides an effect on the color of flowers (pink) and seed (almost black, with fine white points and spots). A recessive allele of this gene causes the usual coloring of vegetative organs. Leaves owe their blue-green color to the vir gene, and salmon color, identified by Hackbarth (1957b) in an induced mutant, to the slm gene. The lat gene, discovered in a mutant form of cv. Müncheberger blue, is responsible for broad leaves. This gene is dominant and pleiotropic. It also causes tallness of plants. A recessive allele of this gene provides for narrow leaves.
Flower. The tests for allelism and complementarity of characters in the accessions of L. angustifolius have resulted in revealing four basic types according to the color of the flower corolla: blue, pink, pale violet and white (Kurlovich and Stankevich, 1990,1994). Blue color is the dominant one among them, being typical for the majority of wild forms. White color of the corolla is caused by the leuc factor (Sypniewski,1925, 1930; Hachbarth, 1957b). The forms with pink flowers, which for the first time were described by Sempolowski (1901), are designated by the term ros, and those with pale violet flowers by the term viol (Sempolowski, 1901; Kajanus, 1912). Beside the basic corolla color types, different researchers have described intermediate types. Dark blue-gray color is designated by the symbol scoer (Sypmiewski, 1925, 1930; Майсурян and Атабекова, 1974), and bluish-white color by the symbol salb (Hackbarth, 1957b). With some forms, flowers have different hues in different parts of the inflorescence, or their color changes in the process of growth and development. Such forms are designated by the term discolor (Hackbarth, 1957b). The color of the corolla is inherited in result of complementary interaction of genes. The blue (wild) type is restored in the first generation when crossing the forms with different color of flowers (pink, pale violet, and white). And there is segregation into blue, pink, pale-violet and white flowers in the second generation, depending on the genotype of parental forms (Таранухо, 1980).
Pod. Wild forms of narrow-leafed lupin have dehiscent pods. This trait, useful from the biological point of view, providing for better distribution of seed and expansion of the growing area of plants, is not necessary for cultivated plants. The character of non-dehiscence of pods was for the first time identified in Australia. According to Gladstones (1967, 1970, 1977), it is regulated by two genes: tardus (tа) and lentus (lе). The tа gene would limit dehiscence at the expense of widening the pod valves by forming a solid row of sclerenchymal cells along the whole perimeter of the pod. The lе gene reduces dehiscence at the expense of structural changes in the valves. A layer of dense pigmented tissue, looking like a grid, is developed inside them to impede curving of valves. The valves of such pods acquire a bright reddish hue by the time when seed begin to mature. The le gene has (Курлович, 1986, 1988) a visible attribute (reddish pigmentation and thickening of the pod).
Seed. Lupinus angustifolius differs by a large polymorphism of seed color. The tests for allelism and complementarity of characters in L. angustifolius accessions have resulted in revealing 8 basic types of seed colors: (1) motley, gray, with unclear spotting; (2) almost black, with fine white specks and spots; (3) gray, with white spots; (4) white with sparse brown and gray spots; (5) beige (hazel), with brown spots; (6) white, dull at the scar, without a triangular spot or strip; (7) white, with sparse brown spots; and (8) sheer white, glossy. Different researchers also described the same intermediate types of coloring. The seeds can be pure white, or one-color, two-color, and even three-color (Hackbarth, 1957b). Dominant above all is the motley or gray color of seed with unclear spotting, which is controlled by the factors designated by the symbol gris. Hackbarth (1957b) described a few alleles of this gene (gris 1, gris 2) causing various shades of coloring.
Beige (hazel) seed coat is caused by gene fer (Hackbarth, 1957b). This author described the mutation with gray-light-blue color of the seed coat without a marble pattern. Such seed were also attributed to a one-color group, designated by the symbol m (Майсурян and Атабекова,1974). Different two-color and three-color patterns of seed coat are designated as mac (Hackbart, 1957b).
White seed color is caused by several factors. White-seeded forms with a triangular stain and strip near the hilum are designated by the symbol leuc (Sypniewski, 1925, 1930; Glagstones, 1970). The gene determining this character pleiotropically causes the white color of flowers and light green color of leaves. The only white-seeded form with neither triangular stain nor strip at the hilum, discovered in 1941, was designated by the symbol niv (Troll, 1941). This factor also causes bright-green coloring of vegetative organs. The gene that determines only white colored flowers and has no pleiotropic relation to the color of vegetative organs has been named alb (Hackbarth, 1957b). Presence of a figure or triangular stain near the hilum is a monomeric dominant character, while their absence is a recessive character (Майсурян and Атабековa, 1974). The seed without a figure at the hilum was found by Roemer (1924). He marked the gene causing this character by the symbol f.
According to the opinion of Taranukho (Таранухо, 1980) and Kurlovich (Курлович, 1991), seed coat color is controlled by a group of polymeric genes. The number of dominant alleles determines its intensity. White-seeded forms crossed with various genotypes can produce a hybrid progeny of different colors. It is possible to prevent this phenomenon by making controlled crossings of the combined lines and hybrids in the process of breeding a new cultivar. Seed color, pigmentation of cotyledons, leaves, stalks, inflorescences and flowers are determined by a group of coherent genes (Tаранухо, 1979, 1980). Strong correlative interdependence evolves between these characters in the process of crossing. As a result of that, new varieties and forms occur. Genetic research was also targeted to other seed properties, besides the coloring of seed coat. For instance, the character of seed coat thinness and good water permeability is caused by the gene moll, whereas large seed size is designated by the term reticulatus (Gladstones, 1970, 1977). Genetic influences on the split seed disorder in L. angustifolius also is discussed (Walton and Francis, 1975).
Steam. Shortness of plants is caused by the gene min (Hackbarth, 1957b). In cv. Nemchinovsky 846 a form was identified where the stem expands in a fasciation-prone manner, forming a swelling at the top. Flowers are arranged there in plenty, and from this bundle a compact group of pods is developed with simultaneous ripening habit. Thus, seed yield of the main truss in this form equals to the yield of a whole plant of cv. Nemchinovsky 846. This form was identified when the seed were processed by gamma-rays. This described character of breeding value was designated by the term cofertus (Клочко and Курлович, 1990).
Physiological properties
Low alkaloid content. Active genetic researching on the inheritance of alkaloid content was launched when v. Sengbusch in Germany had discovered the first low-alkaloid forms, and when the method of analyzing plants and seed for low alkaloid content had been developed and published in the USSR (Иванов et al., 1932). This character represents a biochemical mutation (Mikolajczyk,1962; Таранухо 1980; Kurlovich et al.,1995).
Syntheses of alkaloids in plants are provided by no less than three pairs of genes. Alkaloid content in a plant becomes low if even one of these pairs of genes has mutated from dominance to recessiveness (al1al1 AL2AL2 AL3AL3 ; AL1AL1 al2al2 AL3AL3 or AL1AL1 AL2AL2 al3al3 etc.). On the basis of this phenomenon, fodder (sweet) lupin cultivars were produced in Germany (Sengbusch, 1942) and in the USSR (Федотов, 1934). However, in many of them alkaloid plants gradually came back again. It happened due to the differences in recessive genes as far as many low-alkaloid forms were concerned. Cross-pollination of these forms resulted in heterozygosis in F1 with the presence of dominant alleles in all pairs of genes (AL1al1 AL2al2 AL3al3). This, in its turn, also provided for the restoration of alkaloids. Biparous segregation in F2 into alkaloid and low-alkaloid plants, in this case, takes place more often in the proportion of 9:7 (Анохина, 1970, 1975). However, restoration of a normal process of alkaloid synthesis during hybridization can be accomplished in various ways and with different speed, depending on the number of links in the chain of biosynthesis which is broken in the process of mutations at the stage of initial parental forms (Анохина, 1975).
Biosynthesis of alkaloids is renewed especially frequently and quickly when accessions of different origin are crossed. For production of stable fodder forms there are methods based on selecting plants with identical genotypes and with absence or low content of alkaloids caused by the same genes, and their involvement in the hybridization process (Анохина, 1970, 1975; Турбин and Анохина, 1974; Чекалин and Курлович, 1989; Курлович and Чекалин, 1991). It is recommended to use controlled crossing of initial sweet plants with subsequent assessment of their progeny for presence or absence of alkaloids (tests for allelism and complementarity). Three genes of low alkaloid content were described in L. angustifolius: iuk, es and dep (Hackbarth (1957b); J.S.Gladstones, 1970). The latter gene was found in a Russian cultivar bred by Fedotov (Федотов, 1934). Each gene was characterized by a certain amount of residual alkaloid content (iuk - 0,049%, es - 0,106%). However, subsequent tests for allelism and complementarity, involving diverse and variable materials, resulted in arranging the tested accessions into 7 groups according to their complementarity potential (Турбин and Анохина, 1974). It testified to the presence of no less than 6 genes responsible for alkaloid synthesis in narrow-leafed lupin.
Features of growth and development. Rapid growth speed as well as plant development to a greater height are caused by the factor proc in Polish cultivars (Sypniewski,1925, 1930; Hackbarth,1957b). The gene efl reduces sensitivity of plants to vernalization and accelerates flowering (Gladstones, 1970). The Swedish forms with gene Ku (cv. Borre) do not require vernalization at all, and are characterized by early flowering habit when sown in spring (Gladstones, 1977; Курлович, 1991б).
The forms with determinate growth processes and especially with determinate branching have become widely used in breeding programs for the development of early-maturing grain varieties of lupin (Gladstones, 1977; Mikolajczyk et al, 1984; Delane et al, 1986; Курлович, 1988a, 1991a; Kurłowicz, 1992; Römer, 1994; Kurlovich and Ivanova, 2000). Such forms are also known as “topless”, “samozakącząnce”, or epigonal types. Determinate growth, or determinacy is the limitation of plant growth possibilities, which makes the plants ready for the next developmental stage. A peculiarity of narrow-leafed lupin forms with determinate branching is that all their branches (the main as well as lateral, if they are available) end by generative organs (flowers or flower trusses).
Our researches have shown that phenotypic manifestation of determinacy of branching in narrow-leafed lupin includes the influence of the genotype, effect of the environment, and genotype-environmental interaction (Kurlovich et al., 1995; Kurlovich and Ivanova, 2000). The influence of the genotype on the character of determinate branching is controlled in the accessions Ladny (Russia) and Mut-1 (Poland) by recessive alleles of two genes (deb1 and deb2), whereas in the accession Lanedeks-1 (Belarus) by dominant alleles of two other genes (Deb3 and Deb4). The non-allelic gene interaction takes place in all the cases, its nature being influenced by the modifying effect of environment. This problem is discussed in more detail in the section «Genetics of quantitative characters».
Resistance to diseases. Resistance to anthracnose (Glomerella cingulata) is controlled by the dominant gene An in Australian cultivars (Cladstones, 1970, 1977). This gene was found in the wild Portuguese form P.I.168535. Resistance to Stemphhylium, or gray leaf spot (Stemphylium vesicarium) is caused, according to the data of the same author, by the genes with the symbol gl. This gene of stability (gl) was discovered in a spontaneous mutant of a bitter commercial cultivar of Australian origin. The second gene of resistance (gl1) was identified in the wild Portuguese form P.I.168530. Resistance to anthracnose and gray leaf spot has already been transferred into many commercial cultivars of narrow-leafed lupin from Australia, New Zealand, USA and other countries.
Yellow lupin – Lupinus luteus L.
Morphological characters
Vegetative organs. Variation in the color of cotyledons, leaves and stalks is observed in yellow lupin as well as in narrow-leafed lupin. The most widespread is the dark green (olive) color of cotyledons and leaves. It is determined by the gene oliv (Hackbarth, 1957а). The forms with the presence of anthocyanin are designated by the term anth (Hackbarth, 1957а). Meanwhile, the presence of anthocyanin in the plants of cv. Słodziak is caused by the gene pa (Майсурян, Атабековa, 1974). The light-golden color of cotyledons, leaves and pods is designated as aur. Such recessive mutation was for the first time detected in 1938 in the strain 7844, and served as a basis for breeding the world famous cv. Weiko III. Yellow-green color of leaves with various shades in the mutant forms from Sweden is determined, according to Hackbarth (1957а), by the genes chlorina (chl 1, chl 6) or chlorinus (chlor 2, chlor 4). The group of genes supervising pale light-green color of leaves without pleiotropic effect on other parts of the plant are designated by the symbols pal1, pal2, and pal3. Besides, Hackbarth (1957а) obtained various non-chlorophyllic mutants controlled by lethal genes from the groups albinus (albi 1, albi 2) and xantbus (xan 1, xan 2). Such plants, as a rule, are not viable and perish. Light-cream coloration of vegetative parts in yellow lupin depend on a single recessive gene in the homozygous state. This gene was called dilutus (Kazimierska and Kazimierski, 1995).
Flower. The tests for allelism and complementarity of characters in L. luteus accessions have resulted in identifying four basic types of corolla color: yellow, lemon-yellow, orange and whitish (Kurlovich and Stankevich, 1994). Dominant color is yellow, which is determined by the gene Flav in cv. Słodziak (Mikolajczyk, 1862). Lemon-yellow coloring of the corolla is designated as sulf in strain 8 (Hackbarth, 1957а). However, Micolajczyk (1962, 1964) marked the same colour by the symbol cit in cv. Cytrynowy. The amber-like color in the sample Rod C is controlled by the factor hel, while ginger color in cv. Guelzower St RM by the gene ruf (Micolajczyk, 1962, 1964; Майсурян and Атабековa, 1974). The color of the corolla is inherited in result of complementary interaction of genes with subsequent restoration of yellow color in F1 (Таранухо, 1980). Lupin forms with reduced flower petals were discovered in 1951 by Hackbarth (1957a).
The gene red is responsible for this trait. In 1953, induced mutants with intervals between separate flowers in their inflorescence were identified. Such forms are designated as sec.
Pod. Non-dehiscence of pods is determined by the factor inv in yellow lupin (Hackbarth, 1957а). It is provided by a continuous cord of sclerenhymal cells along the pod perimeter. Absence of pubescence on pods (hairs fall down) is caused by the gene nud (Hackbarth, 1957a). The second gene brev causes formation of the pods with short (not more than 0.4 mm) down. The gene brach promotes development of a dense truss. The pods are arranged compactly and, as a rule, are situated in the lower part of a plant (Hackbarth, 1957a).
Seed. The color of seed is studied best of all from the genetic viewpoint. It is characterized by wide variation (from black to white). Testing the characters of L. luteus accessions for allelism and complementarity resulted in identification of 9 basic types of seed coat colors: (1) white, with black dots, without arcs (dotted); (2) white, with black spots and two light-colored arcs; (3) white, with brown spots, without arcs; (4) white, with brown spots and two dark-colored arcs; (5) white, with black spots and a wide clean space around the scar; (6) brown-and-black, with light-colored arcs; (7) black, without arcs; (8) cream-colored, with white arcs; and (9) sheer white (Kurlovich and Stankevich, 1994). Intermediate variants differ among themselves in various color of spots, maculations and presence or absence of arcs. Black seed are designated by Hackbarth (1957а) as Coloratus (Col). In his opinion, this color is dominant. Non-uniform distribution of the black color on the seed with a crescent arc near the hilum acquired the name falcatus (col/falc). Uniform distribution of the black color with fine points on white background was named parvimachlatus (col/parv). Brown pigmentation of seed is determined by the gene fuscus (Swęcicki, 1988). The forms with brown seed coat occur most frequently in Israel and Palestine. White-seeded forms can emerge due to the effect of genes alb or niv (col/niv). The gene niv is also responsible for pleiotropic determination of the light-green color of vegetative organs and the keel edge. The gene alb exerts no influence on the color of other plant organs (Hackbarth,1957a; Gladstones,1970).
Crosses between white-seeded forms of various origin (genes col/niv and alb) made clear their genetic differences and non-allelism of these genes. One of the essential features of the seed in the majority of cultivated forms is the presence of thin and water-permeable seed coat. The gene controlling this property (w) is recessive. In view of this, this character can be easily stabilized (Gladstones, 1970).
Steam. The branching on the stem of yellow lupin may be monopodial and sympodial. According to Hackbarth (1957a), the gene bbr is responsible for the absence of branches at the base of the stem (sympodial branching). The form was identified in Poland by Kazimierski and Kazimierska where the stem expands in a fasciation-prone manner, forming a swelling at the top. Flowers are arranged there in plenty, and from this bundle a compact group of pods is developed with simultaneous ripening habit. This described character of breeding value was designated by the term compactus (Kurlovich et al., 1995).
Physiological properties
Low alkaloid content. Four genes were earlier described to be responsible for low alkaloid content in yellow lupin. Three of them were identified in Germany (Sengbusch, 1931, 1942; Hackbarth, 1957а) and one in Holland (Lamberts, 1955, 1958). Low alkaloid content in Strain 8 is caused by the gene dul. Strain 80 has the gene am, and Strain 102 has the gene lib. Low alkaloid content is caused by the gene V-351 in the Dutch sample (Sengbusch, 1931, 1942). The gene dul maintains the residual content of alkaloids at the level of 0,05 %, the gene am at 0,013 %, and the gene lib at 0,010 % (Maissurjan and Atabiekova, 1974). However, Kazimierski and Novacki (1971) reported that it was the gene dulcis that conditioned reduction of alkaloids in the seed of yellow lupin to 0.01%, as compared with the wild type. This gene also governs declining fertility in low-alkaloid plants and reduces their adaptive value, so low-alkaloid forms soon disappear from a mixed or hybrid population. The character of low alkaloid content is recessive, and is inherited in result of complementary interaction of genes (Анохина, 1970, 1975; Таранухо, 1980; Курлович and Чекалин, 1991).
Features of growth and development. The usual rhythm of plant growth with a prolonged (about 30 days) rosette phase is dominant over others growth rates. It is designated by the symbol Cres. A typical form with the gene Cres is St. Treb.1 (Hackbarth, 1957a; Swęcicki et al., 1989). The recessive gene celer (cres/cel) provides for shortening of the rosette phase by 12-15 days and causes pleiotropical determination of the light-green color of leaves in Strain 8 (Gladstones,1970; Hackbarth,1957a). Forms with the gene altus (cres/alt), distinguished for their tallness, overtake the forms with cres/cel in the rate of growth, and also have light-green leaves. The dominant gene Rapidus in some wild forms from Palestine also causes rapid growth in initial phases of development, although it is not connected with the color of vegetative organs (Lamberts, 1955, 1958; Hackbarth, 1957a). Besides, Micolajczyk (1962, 1964) described the recessive gene of fast growth (promtus). In cv. Express, it also exerts pleiotropic influence on the leaf color, branching character, and duration of the vegetation period. A dwarfish plant was found in F4 of a hybrid of two yellow lupin forms.
Genetic analysis demonstrated that dwarfism was preconditioned by a recessive factor, which was named nanus (Kazimierski and Kazimierska, 1976). The inheritance of growth rates has two stages, according to Taranukho (Таранухо, 1979, 1980). The character of slow growth rate dominates in the first phases until the plant comes out of the rosette. Further on, heterosis is observed, i.e. the hybrids overtake their parents in growth. Early maturity of lupin plants has discrete nature, and was reported to be a dominant character (Хотылева and Савченко, 1988).Resistance to deseases. Fusarium wilt is the most harmful and widespread disease of lupin in Russia, Belarus and the Ukraine. Fusarium-resistant plants were identified among the wild Portuguese forms after studying lupin genetic resources of various eco-geographical origin (Wuttke, 1943; Курлович et al., 1990a). The character of resistance appeared to be monogenic and dominant; it is designated by the term Fus 1 (Hackbarth, 1957a; Gladstones, 1970). However, subsequent researches (Лукашевич, 1980, 1981; Хотылева, Савченко, 1988; Kurlovich, 1990) revealed more complex nature of inheritance of this character, as the agent of this wilt (Fusarium Lk.) is represented by a large number of species, strains and races. In view of this, the nature of resistance inheritance in lupin hybrids can be either dominant or recessive, depending on the origin of initial parental forms. Besides, it can be controlled by different numbers of genes. The resistance in cvs. Cyt, Refusa Nova and Borluta is determined by recessive genes, and in cvs. Afus, Янтарь and LLU-17537 by dominant ones (Лукашевич, 1980, 1981).
Plants resistant to powdery mildew (Erysiphe communis) were also found among the wild Portuguese forms (Gladstones, 1970). Their resistance is determined by a monogenic and dominant factor, and is defined as Er (Hackbarth, 1957a).
Genes of sterility. White-leafed forms with sterile pollen were identified in 1946 in Sweden (Hackbarth, 1957a). The factor causing sterility was designated as sterile (st). These forms also had genes chl. The second gene of sterility (st2) was identified in 1952 (Tedin and Hagberg, 1952; Tedin, 1954). Forms with this gene can be traced by a number of morphological changes in the structure of the flower.
White lupin – Lupinus albus L.
Morphological characters
Vegetative organs. The color of stalks and leaves can be whitish-green, green, and dark-violet or dark brown with a shade of anthocyanin. The presence of anthocyanin is especially typical for wild forms (subsp. graecus). Green color of stalks and leaves dominates among cultivated forms (subsp. albus). However, Hackbarth (1957c) in 1960 in Germany obtained a recessive mutant with light-green vegetative organs and with decelerated growth rate, which was named pal 1. In 1951, the same author also produced a second mutant with bright leaves. This character is controlled by the gene pal 2. The color of vegetative organs is inherited in results of complementary interaction of genes. (Таранухо, 1979, 1980).
Flower. The color of the corolla is dark-blue in the wild forms (subsp. graecus), pink-blue or light-pink in subsp. thermis, and greyish, light-blue or white in subsp. albus (Kurlovich and Stankevich, 1994). When crossing different forms within subsp. albus, the greyish-light-blue color of the corolla is dominant, and the white color is recessive. Meanwhile, the dark-blue color dominates in all cases when forms from subsp. graecus are involved in hybridization. The majority of cultivated forms have a greyish-light-blue corolla, this color being caused by a dominant gene Col (Micolajczyk, 1961, 1962, 1964). A recessive allele of this gene (col) causes white coloring of the corolla, but the keel edge in this case is stained with anthocyan. Entirely white coloring of the corolla without anthocyan on the keel edge is determined by the gene alb (Gladstones, 1970). This character is completely recessive. A mutant with the light-pink corolla was found in Poland, and received the name rоseus (Stawinski, 1988). This color presents itself as recessive also in relation to the greyish-light-blue color. All flowers appeared greyish-light-blue in F1 when a light-pink form had been crossed with the white form (complementary interaction of genes), and dihybrid splitting took place in F2: 9 plants with greyish-light-blue flowers, 3 with pink ones, and 4 with white flowers.
Seed. Most of the cultivated forms of white lupin, and the forms that have turned wild, are characterized by the white color of seed coat. Dark-brown and dotted seed coat is typical for wild forms of winter mottled-seed and semi-winter mottled-seed ecotypes (subsp.graecus). The dark-brown and dotted pattern of seed coat is dominant over the white color. All white-seeded forms with permeable seed coat and non-dehiscent pods were selected by man in the age of antiquity (Gladstones, 1970, 1974). Seed shape depends on one of the three genetic factors (Micolajczyk, 1961): quadratus III (quad III), quadratus V (quad V) and phaseolicus (phas), each of the three having a series of multiple alleles. The factor quad III causes the square (angular) shape of seed, which is dominant over the factor phas, which determines the oval form of seed. The factor quad V causing the rectangular form of seed is recessive in relation to the factor phas.
Characters of the whole plant. The forms of white lupin manifest wide variability in plant height and degree of branching. The samples from the west Mediterranean area are, as a rule, taller and have a more vigorous branching habit; the forms from the east Mediterranean are less tall. The spontaneous mutation brev causing dwarfism, fast growth and early flowering was found among the east Mediterranean forms (Hackbarth, 1957c). However, these parameters are strongly influenced by the duration of the day, sowing conditions and air temperature. Early sowing reduces the height of plants and prolongs the duration of the vegetation period by several days. Similar forms were obtained during subsequent experiments as a result of artificial mutagenesis. These mutations were designated by symbols brev 1, brev 2, brev 3 (Hackbart, 1957c). A stable form with short lateral branches was produced from the strain Reuscher 27. This character is recessive, being designated by the symbol con. Environmental conditions influence the variation of this trait to a lesser degree, than the character controlled by genes brev. The recessive mutation, revealed in 1935, was described by the presence of a short radicle in young plants (Hackbarth, 1957c). Such phenomenon is especially useful when seed are germinated in the Petri dish. This character is designated by the symbol rab.
Physiological properties
Low alkaloid content. The first low-alkaloid plant was observed in Strain 19 in Germany (Gladstones, 1970; Troll, 1941, 1943). This character appeared monogenic, and was designated by the symbol mit. In subsequent researches, low-alkaloid plants were found in cvs. Kraffquell, Gela and Ultra (gene pau), while low-alkaloid plants with the non-allelic gene nut occurred in cv. Nahrquell (Hackbarth, 1957c; Gladstones, 1970). Low-alkaloid plants with the gene red were later revealed among the bitter forms of white lupin from the west Mediterranean areas, plants with the gene sua were found in a sample from Palestine, and low-alkaloid plants with the gene exi were selected from a sample from Hungary (Gladstones, 1970). Besides, three genes of low alkaloid content were described by Micolajczyk (1961): the gene tert in cv. Biały III, the gene prim in cv. Biały I and the gene q in cv. Biały V. It is still unclear whether these three genes are allelic or non-allelic between themselves and in relation to the other genes described earlier. Features of growth and development. Growth habit of the main stem is determined by the dominant gene Alt (Micolajczyk, 1961). This gene controls the growth of plants before flowering. The identified three types of alleles in this gene – Alt 1, Alt 2 and Alt 3 – differed in the effect they produced on the growth intensity and capacity of the main stem (Головченко et al., 1984). Growth intensity and capacity of lateral branches are determined by the dominant gene Long (Micolajczyk, 1961). Four dominant alleles were identified in this gene: Long 1, Long 2, Long 3 and Long 4. The corresponding recessive alleles (long 1, long 2, long 3 and long 4) reduce the vigor of lateral branching. The height of plants and their growth rate are determined by the interaction between the genes Alt and Long. The intensity of differentiation and development of flowers on the main truss is controlled by the dominant gene Flor (Головченко et al., 1984).
The recessive allele of this gene flor detains differentiation and development of flowers. The duration of the period from young growth to the beginning of flowering and its intensity is regulated by the interaction of the genes Alt and Flor. The dominant gene Fest jointly with the recessive gene long accelerate the time of flowering on lateral branches. The recessive allele of the gene fest together with dominant alleles of the gene Long provide for an opposite effect (Micolajczyk, 1961; Головченко et al., 1984).
The list of genes
It is possible to judge about degree of genetic variability of different characters at lupins on the base of materials submitted in the Table 28. This information about genes at three species of lupin is taken from different reports (Lamberts, 1955, 1958; Hackbarth, 1957a-c; Micolajczyk, 1961; Gladstones, 1970, Maissurjan and Atabiekova, 1974, Kazimierski and Kazimierska, 1976, etc.). The genes are listed in alphabetic order for each species and the characters controllable by them are specified. It is necessary to note, that lupin is investigated in the genetic attitude more poorly in comparison with peas or soya. The morphological attributes are investigated in the greater degree at lupins.
Table. The list of genes at three species of genus Lupinus L.
(Symbol- name -and phenotype)
(Lamberts, 1955, 1958;Hackbarth, 1957a-1957c; Micolajczyk, 1961; Gladstones, 1970, Kurlovich, 1995, Maissurjan and Atabiekova, 1974, Kazimierski and Kazimierska, 1976, and others).
Lupinus angustifolius L.
..... alb- albus - Only white coloring of a seed coats without the triangular stain and strip near a hilum .....
An - -Resistance to anthraknose (Glomerella cingulata)
.....
con - confertus - Fasciation of a stalk, dense arrangement of flowers and beans
.....
deb1, deb2, Deb3, Deb4 - determinate branching; deb1 and deb2 - determinate branching at accessions Ladny and Mut-1; Deb3 and Deb4 - determinate branching at accession Lanedeks-1
.....
dep - depressus - Low alkaloidness (strain 14)
.....
dis - discolor - Various coloring of flowers on a truss
.....
efl - eflorestentia - Limited requirements to vernalization, moderately early flowering at autumn sowing
.....
es - esculentus - Low alkaloidness (strain 415)
.....
f - Presence of a figure on a seed near hilum
.....
fer - ferrugineus Beige (hazel) coloring of seed coats
.....
gl, gl1 - Resistance to grey leaf spot (Stemphylium vesicarium)
.....
gris1, gris2 - griseus - Motley or grey coloring of seed coats with unclear spotting
.....
iuc - iucundus - Low alkaloidness (strain 411)
.....
Ku - Low requirement to vernalization,
early flowering
.....
lat - latifolius - Wide leaflets
.....
le - lentus - Limited shattering of beans
.....
leuc - leucospermus - Only white coloring of flowers and seed coats
.....
m - Grey - light-blue one-color coloring of seed coat without marble
.....
mac - maculatus - Two and three-color coloring of seed coats
.....
min - minor - Low growth of plants
.....
moll - mollis - The coat of seed is thin and good permeable
.....
n - Absence of figure on seed near a hilum
.....
niv - niveus - Only white coloring of flowers and seed coats, without a triangular stain and strip near a hilum
.....
proc - proceus - Tall plants, fast rates of growth
.....
Pur - Purpureus - Presence of anthocyanin on vegetative organs (reddish color of stalks and leafs)
.....
ret - reticulatus - Large seed
.....
ros roseus Pink coloring of flowers
.....
rut - rutilus - Reddish coloring of cotyledons and young leafs
.....
salm - salmonides - The color of leafs as a salmons
.....
salb - subalbidus Bluish-white color of flowers
.....
Scoer - Subcoerulius - Dark blue-grey color of flowers
.....
ta - tardus - Limited shattering of beans
.....
viol - violaceus - Violet coloring of flowers
.....
vires1, vires2 - virescens - Vellow - green coloring of cotyledons and young leafs
.....
vir - viridis - Blue-green color of leafs Lupinus luteus L. albi1, albi2 albinus Absence of chlorophill
Lupinus Luteus L.
albi1, albi2 - albinus - Abcence of chlorophil
.....
alb - albus - White colouring of a seed coat
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am - amoenus - Low alkaloidness (strain 80)
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anth - anthozyanin - Presence of anthocyanin on vegetative organs
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aur - aureus - Light with yellowness color of cotyledons, leafs and beans
.....
bbr - no basal branches - Absence of branching in bottom parts of a plant
......
brach - brachulobus - Dense arrangement of beans on trusses
.....
brev - brevis - Short hairs (pubescence) on seed
.....
chl, chl1, chl6 - chlorina - Yellow - green coloring of leafs
.....
chlor2, chlor4 - ,chlorinus - Yellow - green coloring of leafs
.....
cit - citrinus - Lemon-yellow coloring of corolla
.....
Col - Coloratus - Black coloring of a seed coat
.....
col/falc col/- falcatus - Non-uniform distribution of a black pigment and presence of arcs on seed coat
.....
col/parv col/- parvimaculatus - Uniform (points) distribution of a black pigment on a seed coat
.....
col/niv col/- niveus - White coloring of a seed coat and light coloring of vegetative organs
......
com - compactus - Fasciation of a stalk, dense arrangement of flowers and beans
.....
Cres - Crescens - Normal speed and duration of growth
.....
cres/alt cres/- altus - Fast and high growth, light-green coloring of leafs
.....
cres/cel cres/- celer - Short phase of the rosette, lihgt- green coloring of leafs
.....
dil - dilutus - Light-cream coloration of vegetative plant organs
.....
dul - dulcis- Low alkaloidness (strain 8)
.....
Er - Erysiphus Resistance to Powdery mildew
.....
Flav - Flavus - Yellow coloring of corolla
.....
Fus - Fusariosus Resistance to fusariose at wild- growing Portuguese forms
.....
- fuscus - Presence of a brown pigment on seed coat
.....
hel helvis - Amber coloring of corolla
.....
inv - invulnerabilis - Non shattering pods
.....
lib - liber - Low alkaloidness (strain 102)
.....
- nanus - Dwarf growth
.....
niv - niveus - White coloring of a seed coat
.....
oliv - olivaceus - Dark green (olive) color of cotyledons and leafs
.....
pal1, pal2, pal3 - pallidus - Light green coloring of leafs
.....
pa - paucus - Presence of anthocyanin on on vegetative organs (cv. Clodziak)
.....
- promtus - Fast growth and early flowering (cv. Express)
.....
- Rapidus - Fast growth in initial phases of development
.....
red - reduced petals - Reduced floral bracts
.....
ruf - rufus - Rufous color of a seed coat
.....
sec - secatus - Presence of intervals between flowers on a truss
.....
st - sterile - Sterility of pollen, light leafs
.....
st2 - steriller2 - Sterility of pollen
.....
sulf - sulfureus Lemon-yellow coloring of corolla
.....
V-351 - Low alkaloidness (strain V-351)
.....
w - Thin and water-permeable seed coat
.....
xan1, xan2 - xantbus Yellow plants (absence of chlorophyll)
Lupinus albus L.
alb - albiflorus - Only white coloring of corolla
.....
Alt - Altus - The control of intensity of growth of the main stalk
.....
brev, brev1, brev2, brev3 - brevis - Dwarfism, fast growth, early flowering
.....
Col - Coloratus - Greyish-light-blue coloring of corolla
.....
con - contractus S- hort lateral branches
.....
exi exiguus Low alkaloidness
.....
Fest - Festicus - Early flowering of lateral branches
.....
Flor - Floridus - Intensity of differentiation and development of flowers
.....
Long - Longus - Intensity of growth of lateral branches
.....
mit - mitis - Low alkaloidness
.....
nut - nutricius - Low alkaloidness
.....
pal1, pal2 - pallidus - Light green coloring of vegetative organs
.....
pau - pauper - Low alkaloidness
.....
phas - phaseolicus - The oval form of seed
.....
prim - primus - Low alkaloidness
......
quad III - quadratus III - The square form of seed
.....
quad V- quadratus V - The rectangular form of seed
.....
q - quintus - Low alkaloidness
......
rab - radibrevis - Short radicle
.....
red - reductus - Low alkaloidness
.....
ros - roseus - Pink coloring of corolla
......
sua - suavis Low - alkaloidness
.....
tert - tertius - Low alkaloidness
Genetics of quantitative characters
Available today are separate fragmentary works on genetics of quantitative characters at lupin (Агаркова et al., 1980, 1991; Агаркова and Пухальская, 1991; Анохина et al., 1984; Хотылева and Савченко, 1988; Huyghe, 1990; Huyghe et al., 1994; Neves-Martins, 1994; Qiu et al, 1994; Kurlovich et al., 1995). This fact is mainly connected with difficulties in studying quantitative characters, though they determine the efficiency and quality of plants. Quantitative characters are in most cases formed and undergo changes during a certain period of development, in the process of ontogenesis, depending on the time, effect and properties of the limiting factors. Quantitative characters, being the main objects of breeding, can be improved in the opinion of Kurlovich et al. (1995) and Dragavtsev (1997) by several ways: at the expense of transgressions (breeding work with self-pollinated species), by using the phenomenon of heterosis (with cross-pollinators), or by shifting a genotype to a new ecological niche (the effect of genotype-environment interaction ). In view of this, a majority of lupin species, prone to self-pollination and cross-pollination, may be improved by all three ways. Any quantitative attributes of plant efficiency are not only a product of the action of genes or chromosomes, but also a result of interaction of the limiting environmental factors with the systems of genetic complexes. The following complex phenomena frequently take place in the process of genotype-environment interaction: 1) re-determination of the spectrum of genes determining genetic variability of a character, when the limiting factor of the environment changes; 2) modular arrangement of quantitative characters – one resulting character and two components; 3) if there are two known values of three essential characteristics: a) adaptive properties of genotypes, b) definite dynamics of limiting factors, c) genetic parameters of a population, the possibility to determinate quantitatively and definitely any third unknown characteristic (Dragavtsev, 1997). The important role in the genotype-environment interaction has also adaptability of plants, which is both complexly inherited and much affected by environment (Allard, 1977).
Inheritance and variability of resistance to diseases and pests is discussed in the section «Diseases and Pests» from the point of view of different types of plant resistance. Interactions in the host-pathogen system are conditioned adaptively and regulated generically. Distinguished are main or major genes with large effect, and minor genes producing gentle effect. Inheritance of major genes is easily identified in descendants, however they more often determine particular resistance to a definite race or group of races. Phenotypic effect of minor genes depends on their quantity, which determines the level of nonspecific resistance inherited quantitatively with polymer cleavage. Some researchers have reached the conclusion that the concepts of horizontal and vertical resistance have no genetic contents, as the genes determining resistance are the same, but their manifestation varies depending on the genetic background of accessions (Чекалин et al., 1981). Their effect provides vertical resistance, if the genes are disconnected, and horizontal, if the genes are coupled in one genotype. Investigation of quantitative characters in this connection is very difficult because the conditions of external environments constantly vary. New approaches will be necessary for studying them. Geographically arranged plantings and plant genetic resources studies by uniform techniques in different conditions, recommended by N.I. Vavilov, are very effective for this purpose. It is possible to illustrate it by the case study of such character as inheritance of determinate branching in narrow-leafed lupin. This character can be considered and studied not only as a quantitative one, but also as a qualitative feature, as the degree of the development of branching varies in lupin hybrids within very wide limits, especially in different environments (Kurlovich et al., 1995, Kurlovich and Ivanova, 2000).
In the beginning of our research, we studied this character by the methods of classical genetics as a qualitative feature. However, later we became convinced that such approach was not efficient enough, for the results were variable in different eco-geographic conditions. Nonetheless, this investigation served as a basis for further enhancement of the methods used in the research as well as for certain conclusions.
The object of the studies were the accessions of narrow-leafed lupin with determinate branching: Ladny from Russia, Mut-1 from Poland and Lanedeks-1 from Belarus. Determination in these accessions had different degrees of expression. They were crossed with common branching forms without determinacy: Nemchinovsky 846 and Timir-1 from Russia, Mirela from Poland and Illyarie from Australia.
All accessions and hybrids were assessed in 1986 - 1998 on the plots of 1-2 m2 in contrasting environments in many countries: in Russia near St.Petersburg on the experimental field of Pushkin Laboratories of VIR (Leningrad Province, 580 N. Lat., soddy-podzolic soil, spring sowing); in the Ukraine near Kiev (the Ukrainian Scientific Research Institute of Arable Farming, about 500 N. Lat., non-black-soil zone, spring sowing), and in humid subtropics of Abkhazia (Gulripsh Settlement, the former Sukhumi Experimental Station of VIR, about 420 N. Lat., autumn sowing). Besides, parental forms as well as most productive, early and stable hybrids were tested in 1997-1999 in Finland on two sites with sandy soils and clay soils (spring sowing near Mikkeli, 610 N. Lat.). More details of this investigation were presented in our earlier works (Курлович, 1991a; Kurlovich et al., 1995; Kurlovich and Ivanova, 2000; Kurłowicz, 1992).
This long-term study (1986-1999) of narrow-leafed lupin accessions with determinate branching (Ladny, Mut-1, Lanedeks-1) showed that this character was the most stable in the accession Ladny, and the least stable in Lanedeks-1. Usual indeterminate plants were segregated in Lanedeks-1 under the conditions of spring sowing in Leningrad Province, Russia, in the cold year of 1993 with low temperatures, and all plants of this accession had usual indeterminate branching under the conditions of autumn sowing in Abkhazia. However, seed obtained in the years with a cold spring season, when being sown, produced plants with determinate branching. In 1998, when this accession was tested in Finland, there also was a segregation of plants with common branching. In the accession Mut-1, determinate branching was always expressed, but either on the main stem or on lateral branches, depending on environmental conditions (Kurlovich and Ivanova, 2000).
The analysis of hybrids of the first generation in 1987 in three eco-geographic locations showed that the character of determinate branching was recessive in Ladny and Mut-1 accessions and dominant in Lanedeks-1. However, already then we paid attention to the fact that after autumn sowing in Abkhazia the F1 hybrids involving Lanedeks-1 accession had incomplete domination of determinate branching, since it manifested itself not on the main stem, but on lateral branches. When the experiment was repeated in Abkhazia in 1988, the form with ordinary branching showed domination. These circumstances urged us to continue studies with a limited set of samples in 1992-1993. The analysis of the F1 hybrids produced by crosses between the forms with determinate branching and those with usual branching under the conditions of extremely cold and rainy summer of 1993 in Leningrad Province, in contract to the data of 1987, revealed the recessive character of determinate branching inheritance in the accession Lanedeks-1. F2 populations were analyzed in 1988, and when dividing them into three classes, dihybrid segregation was observed. In this case, a different non-allelic gene interaction took place: 12:3:1; 1:6:9; 4:3:9 (chi-square = 0.21 - 4.26). However, further experiments in 1989 in Abkhazia showed in a number of combinations that inheritance of this character had also monogenic nature (chi-square = 0.15-0.17).
The effect of vernalization and duration of the photoperiod on the nature of branching was additionally tested in containers during laboratory experiments (Tab. 6 and 7). The effect of vernalization was the most evident in the accession Lanedeks-1. Under the influence of vernalization it transformed into the form with usual branching, without determinacy. Besides, the accession Mut-1 under the influence of vernalization showed determinate branching not on the main stem but on lateral branches.
Mut-1 produced flowers and developed pods only under the conditions of a long day, that is to say, it showed a long-day photoperiodic reaction. The rest of the accessions were early ripening under both photoperiodic regimes. However, under the conditions of a short day, plants of these accessions were more dwarfish. Besides, our previous researches revealed in the studied accessions of narrow-leafed lupin considerable variation of drought resistance (Курлович and Чернышова, 1986).
The results of the described studies suggest that phenotypic manifestation of the character of determinate branching is preconditioned by the influence of the genotype, the effect of environment, as well as genotype-environment interaction. With this in view, the contribution of each of the influencing factors in different accessions may be different. The most stable character of determinate branching was observed in the accession Ladny. The effect of environmental conditions on phenotypic manifestation of this character in this accession was minimal. Besides, Ladny also showed stability of this character in laboratory experiments when the influence of vernalization and different day duration were analyzed (Kurlovich and Ivanova, 2000). In this connection, the above-mentioned accession is the most valuable for future breeding of narrow-leafed lupin.
However, environmental conditions produced an essential but not similar effect on a majority of the accessions studied. This is confirmed by the differences in the nature of inheritance of this character in different sites of study. As this takes place, the manner of segregation in F2 in different hybrids is changed. This phenomenon is associated with the change in the number of active genes, their expressiveness and their different interaction. These conclusions are supported by divergent results of investigations conducted by different authors. Thus, Debely and Derbensky (Enterprise ’’Podmoskovie’’, Russia) found reciprocal differences and incomplete domination of the determinate type in cross combinations with the form Lanedeks-1 when variability and inheritance of determinate branching in F1 were studied (Debely and Derbensky, 1988). Analysis of hybrid populations of F2 between common and determined accessions showed in the framework of this investigation that the pattern of determinate branching was associated at least with the interaction of two genes. Vigorous penetration and expression of the genes’ action was noticed in this case. The pattern of the character of determinacy, like in our studies, depended on the conditions of cultivation.
In the investigations performed by Konorev, Klochko and Anikeeva almost at the same site (Timiryazev Agricultural Academy in Moscow), populations in the second hybrid generation involving accessions Ladny and Mut-1 as parental forms segregated in a ratio of 3:1, which attested to the monogenic control of this character (Конорев, et al., 1991).
In view of the above-mentioned, our conclusions (Курлович, 1991; Kurłowicz, 1992), based on the studies of 1987–1988, that the character of determinate branching is controlled by recessive alleles of two genes (deb 1, deb 2 – determinate branching) in the accessions Ladny and Mut-1 and dominant alleles of two genes (Deb 3, Deb 4) in the accession Lanedeks-1 should be slightly clarified and explained, since our last experiments (1989 and 1993) led to different results. It is possible to expect that the phenotypic expression of the character of determinate branching will also be influenced by others genes responsible for the response to vernalization and length of the photoperiod. It was shown by physiological studies (Kurlovich and Ivanova, 2000) that the studied forms differed in their response to vernalization, length of the photoperiod, drought resistance and other properties, many of which may be pleiotropically connected with the character of determinate branching. Thus, in the majority of cases, vernalized seeds of Lanedeks-1 produced to plants with common-type branching, while those without vernalization produced plants only with determinate branching. In this connection, when crossing Lanedeks-1 belonging to the determinate branching type with common branching forms and growing F1 in cold environments, the parent with common-type branching dominated. When heats and high temperatures make the process of vernalization impossible, the parent with determinate branching dominated. In Mut-1 accession, which had shown long-day photoperiodic reaction under the influence of a short day and after vernalization, determinate branching was observed not on the main stem but on lateral branches. Ladny accession appeared to be the most thermo- and photo-neutral. The nature of branching and the nature of inheritance of this character were stable, maybe owing to this fact.
Besides the environmental factors studied (vernalization and length of photoperiods), many others may also exert influence. Herewith, each of the environmental factors has effect upon the genotype in an interaction with many others and, as a result of their interactions, the final effect is displayed.
Such explanation of the phenomena of vigorous phenotypic expression of the character of determinacy in branching complies with the standpoint of Dragavtsev’s eco-genetic model associated with changeability of genetic formulae in different environments (Драгавцев, 1993; Dragavtsev, 1997).
Thus, the eco-genetic approach in the analysis of a gene pool slightly makes it possible to comprehend the mechanisms and regulations of genotypic response to limiting environmental factors. The studied character of determinacy in favorable conditions can be determined by the genes directly controlling determinate branching (deb 1, deb 2 or Deb 3, Deb 4) on the background of a limiting factor (e.g. low temperature), as well as by the genes responsible for vernalization effect, genes controlling photoperiodic sensitivity under long-day conditions, and genes of low drought resistance in drought environments. The mentioned approach opens one of the ways to understand transgression mechanisms and to forecast their appearance in particular hybrid populations. For example, if the formation of lupin productivity in a zone of selection takes place on the background of drought, the transgression on productivity can be obtained by hybridizing drought resistant accessions (yielding maximum of biomass per an area unit in drought conditions) with accessions carrying genes securing good transfer of plastic matters from the stalk and leaves to pods (Драгавцев, 1993; Курлович, 1999). Thus, modifying effect of environments may be observed not only in the nature of interaction of the genes stipulating determinacy of branching, but also in the changes of the spectrum of operational genes, and frequently in their direction. With a change in environmental parameters, or even with repetition of the experiment in nature, it becomes impossible to predict behavior and nature of inheritance of the studied character. Assuming that the yardstick of scientific knowledge is reproducibility of the results, and when they cannot be easily repeated, as in our example, it means that many compound characters, including determinacy of branching in lupin, cannot be studied by conventional methods of genetics. A legible answer about the nature of inheritance of many plant characters, especially quantitative ones, may be found in classical genetics only under monitored particular conditions: «that would be if …».
Such explanation of the results obtained seemed also appropriate after perusing the publications (Hill et al., 1998; Tigerstedt, 1993, 1994). However, to make a final conclusion and achieve full explanation of mechanisms of interaction between different genotypes in different environments, additional studies are required. These interactions are very complex and variable in different genotypes and under different conditions.
Geographic plantings organized by N.I. Vavilov and plant genetic resources studies in different environments by uniform methods appeared extremely fruitful. Such researches made it possible to study heredity problems in plants more profoundly, and to develop practical techniques of efficient application of the accumulated data. Eco-geographic investigation helped to obtain valuable materials by hybridization between forms with different variability of characters in different conditions, which often proved the non-allelic nature of the genes that controlled them, and consequently to produce transgressive forms.
As a case study, we will discuss the method of producing transgressive forms of lupin on the basis of eco-geographical approach (Kurlovich et al., 1995; Репьев and Барулин, 1998). We have found out that each quantitative character has from two to five or more types of variability. This fact is very interesting. Earlier it was rarely taken into account in the system of genetic techniques. A definite character has changed in the process of cultivation in various environments almost identically in a majority of accessions. However, this character has a different type of variability in a small number of accessions when studied in different environments (different geographic locations, or different years of study on one site). It appeared that in the hybrid progeny this character was within the same limits as in the parental forms when they had identical types of variability of such character.
However, the hybrid progeny may comprise transgressive forms, i.e. the forms with an increased or decreased value of the character when crossing parents with different types of variability. Differentiation in the variability of characters in parental forms can be found by testing them under different conditions using, however, the same techniques. With the help of this method, it is possible to obtain valuable transgressive forms with regard to any characters, from chemical structure to disease resistance (Kurlovich et. al., 1995). The results of increasing resistance to Fusarium wilt in lupin accessions are presented as a case study (Figs. 47 and 48). Lupin cultivars and lines (547 accessions) were tested for Fusarium resistance under different environmental conditions of two regions in Russia (near Bryansk and St. Petersburg), and in the Ukraine (near Kiev) on plots with artificially infested soil. A large number of accessions of different lupin species were selected for their resistance manifested on a single plot with infested soil. On two other plots, they were susceptible. So, differences in disease susceptibility in the same accessions were found in contrasting environments. Resistant forms selected in one region were crossed with accessions that showed resistance in two other regions. As a result of hybridization, two transgressive resistant forms were obtained in F4 : from the crosses cv. Frost x cv. Apendrilon (Lupinus angustifolius L.), and line G-413 x line 85 (Lupinus luteus L.). Their resistance in all three regions appeared to be higher than in their parental forms (Figs. 47 and 48). These two transgressive forms with increased resistance to Fusarium wilt were found suitable for the breeding program on Fusarium resistance in Russia, Belarus and in the Ukraine. Interesting comments on this phenomenon can be found in the article by Ride (1992) where he summarized our knowledge of recognition and response processes in higher plants in relation to fungi. The study of higher levels of specificity (e.g. the specific incompatibility between races of pathogens and certain cultivars of their host) can initiate rapid progress in understanding the mechanism underlying successful infection (basic compatibility), and from thence the higher levels of specific resistance involved in homologous incompatibility (Ride, 1992).
Our researches have also been aimed in this direction. There is basic incompatibility between higher plants and fungi at the specific level, an incompatibility that is a normal way of interaction between these two types of organisms. This type of incompatibility is synonymous to non-host resistance. It is also sometimes termed „heterologous incompatibility” in order to distinguish it from the specific incompatibility (Ride, 1992). These results of experiments explain the reason why earlier in the breeding process transgressive forms occurred very seldom, randomly, and among prolific hybrid materials. Our approach to breeding makes the process of obtaining transgressives more controlled and effective.
Theoretical and experimental substantiation of the ways of forming genetically based inter- and intra-population heterosis in yellow lupin was given in the work of Khotyleva and Savchenko (Хотылевa and Савченко, 1988). The diversity of forms by the degree of cross-pollination was observed on abundant germplasm materials from the collection of the Vavilov Institute (VIR). It became possible to identify efficient breeding sources where cross-pollination reached the proportions adequate for practical utilization. The inter-population heterosis in the hybrids was 16.0-27.0 %. The authors consider that the phenomenon of heterosis can be practically used to increase the productivity of plants. For its disclosure, it is offered to apply the method of displacement of ranks in an affiliated generation.
In the researches of Dr. Agarkova (Агаркова et al., 1980, 1991; Агаркова and Пухальская, 1991), the trend based on the use of biological multivariate statistics methods was put forward, as the application of classical hybridological analysis for productivity characters was limited. As a result, the system of genetic control over productivity characters was established on the basis of combining ability of accessions. Prevalence of additive action of recessive genes was observed in narrow-leafed lupin on a number of pods on the central truss and lateral brunches and in the 1000 seed weight. The leading role of recessive genes was revealed in determination of the levels of heterosis in F1, and occurrence of transgressions in subsequent hybrid generations (Агаркова and Пухальская, 1991). General characteristics of new initial breeding materials and commercial cultivars should incorporate the results of assessing their ecological plasticity, the necessity of which has been shown. Modifications and genotypic variations as well as the degree of their effect on the formation of economically valuable features have been studied in 36 native and foreign Lupinus luteus varieties (Анохина et al., 1984). Investigations have been carried out for 8 characters. On the basis of the data obtained on genetic variations, the conclusion has been made that breeding is most efficient for three characters, such as plant height, leafiness and 1000 seed weight. Genetic determination of the number of pods on the central and lateral trusses in narrow-leafed lupin (cvs. Severnyij 3 and N-844 from USSR, Emir and Kazan from Poland) was investigated by Pukhalskaya (Пухальская, 1984) in diallel crosses. The results of analysis indicated that the number of pods on the central cluster and lateral brunches was controlled by different genes, and the main tendency in controlling both features was dominance and over-dominance. The number of group genes determining each of the characters was not large, being limited to one or two.
Valuable researches dedicated to genetic study of the architecture of lupin plants and relationships between different lupin organs have been carried out in France (Huyghe, 1990; Huyghe et al., 1994; Harzic et al., 1994). Phenotypic variability of different plant parameters was partly explained by the effect of different growing conditions on plant architecture. The parameters were correlated to the height of the main stem, and the number of leaves on primary branches per square meter. High variability in yield could be explained by genotypic and phenotypic variations of the radiation profiles (Harzic et al., 1994). Despite the authors’ aspiration to reveal valuable characters controllable by one or more genes, or to unite similar small characters into one, their methods can effectively be used in studying the genetics of quantitative characters.
Neves-Martins (1994) studied the genetic variability within and between accessions of different population samples of white lupin and pearl lupin (L.mutabilis). Three phenotypic types of L. albus plants, located around the Iberian areas were detected using numeral taxonomy techniques on the accession/descriptors matrix data. Relationship between morphological variation and geographical origin or selection history in Lupinus pilosus was also disused (Clements et. al., 1996). The breeders work with many different characters, some of which can be controlled by genes whose individual effects can be easily detected by segregation ratios. But other traits, so-called quantitative characters, are controlled by genes whose individual effects are so insignificant that they cannot be discerned by conventional hybridological analyses used the classical genetic. Selection within these characters produces a steady response over a period of time. The biometric basis stems from the fact that quantitative traits generally follow a continuous, normal distribution, and hence can be analyzed sometimes by routine statistic procedures (Hill et al., 1998). However, this procedure is complicated enough and poorly effective with regard to the same quantitative characters.
Genotype-environment interactions can pose major problems for a scientist and practical breeder, particularly when they are of the so-called crossover type, that is when the ranking of varieties changes from one environment to the next. Some of the univariate and multivarite techniques used in their analysis are also discussed in the book by Hill, Becker and Tigerstedt (Hill et al., 1998). Virtually new approaches are necessary for future investigation of quantitative characters in different environments, which we have attempted to reflect partially in this section.
Objective regularities in variability of characters in lupin
The genus Lupinus L. numbering some hundreds species of different age and origin is a perfect object for studying the genetic bases of evolution, plant speciation and phylogenesis of plants. It is possible to make conclusions about the tendencies in the domination of characters, their change, ways of adaptation and evolution of plants in the course of time and under the effect of limiting factors, comparing the degree of development of different characters in the species contrasting in their origin and age. Most effectively it can be carried out by using Vavilov’s law of homologous series in hereditary variation (Вавилов, 1920) and differential systematic-geographical method of plant studies (Вавилов, 1931, 1935) .
Any genera or species of plants, when studied under a wide range of geographical conditions, involving inbreeding, segregates into a wide range of hereditary forms, which is difficult to understand at first sight. Yet, in studying interspecific and intraspecific diversity one can observe a number of regularities, reveal similarities according to Vavilov’s law of homologous series in hereditary variation. Laws discovered by Vavilov can help to put in order the extensive materials representing cultivated and wild plant worlds.
Vavilov marked the unity of variability describing all the family Fabaceae(Leguminosae). He established a regularity in its differentiation into separate genera and cultivars, which takes place in a set of characters pertaining to seed, pods and flowers, together with vegetative organs. It is clear from his data that, despite the differences existing between various genera, one can observe in them the similar variability of characters mandatory for all genera within the given family. He did not produce any data specially related to the genus Lupinus L. However, subsequent researches have shown that the genus Lupinus is much more illustrative of Vavilov’s law of homologous series in hereditary variation than any other genus of the family Fabaceae (Leguminosae) (Maissurjan and Atabekova, 1974, Kurlovich et. al., 1995). It is possible to find in different lupin species not only wide diversity in many thousands of forms, but also parallel series even in pathological, mutagenic and hybridological variation.
Let us recollect only a few of the numerous facts. Introducing materials for his law of homologous series in variation, Vavilov gave much attention to the character of plant color. Various coloring of plants proved to be a hereditary character transmitted from generation to generation, so it became applicable to be used in plant systematics at a distinctive feature of separate species, varieties, subvarieties and smaller taxonomic units. Now, a correlation has been established between the color of seed, inflorescences, keel edges and vegetative bodies of lupin plants. For instance, from white seeds of L. angustifolius (from the eastern hemisphere) light-green plantlets are usually produced, which further on form light-green or normal green foliage and begin to blossom with white flowers. It is possible to obtain plants with dark-green foliage and painted flowers from pigmented seeds. In L. mutabilis (from the western Hemisphere), white-seeded forms more frequently produce also light-green plantlets which would develop green foliage and begin to blossom with white, white-and-pink or white-and-blue flowers. However, even weak pigmentation on the seeds of L. mutabilis, expressed even by a small dark stain on the hilum, results in the formation of anthocyan shoots, dark-green foliage and dark-blue or violet flowers. This is a common tendency in the variability of all lupin species, as well as other representatives of the family Fabaceae (Leguminosae). However, not only parallelism in the heredity of characters is observed, but also parallelism in rejections in the development of plants. These factors facilitate the task of making intra- and inter-specific systematization of plants (Вавилов, 1935). A definite combination of the colors of seed, vegetative bodies, inflorescences and keel edges in various lupin species from both hemispheres, as well as the complete parallelism in their age variability and in morphological and biological characters, speaks about their doubtless genetic affinity. The use of the above-mentioned natural laws enabled us to identify and describe several new forms of different lupin species, so far unknown, and to develop intraspecific classification for them. Inclusion of lupin in the list of examples confirming the law of homologous series in hereditary variation will make it much more representational. But the basic essence of this law, as far as the genus Lupinus is concerned, involves the following. Lupin species more or less closely related to each other are characterized by similar series of variation with such a regularity that, knowing a succession of varieties in one species, one can forecast the existence of similar forms in other species. The similarity is more complete in the series of variation, as the species are more closely allied in the general system.
The law of homologous series in hereditary variation gives the answer to the question what material should be looked for, while the theory of the centers of origin of cultivated plants responds to the question where it should be looked for.
With reference to Lupinus, Vavilov considered the Mediterranean region and mountain areas of Mexico, Peru and others American countries as the centers of origin for this genus (Vavilov, 1926). Generalization based on the differential systematic-geographical method of crop studies provided for more precise definition of the centers of formation and origin (diversity) of some lupin species (L. albus L., L. luteus L. and L. angustifolius L.).
According to our data (Kurlovich, 1998), the center of formation of wild white lupin (L. albus L.) and the primary center of origin (diversity) of its initial cultivated forms is the Balkans where an exceptionally wide diversity of wild and local forms as well as those that turned wild is concentrated. All three subspecies of white lupin (subsp.graecus Franko et Silva, subsp. termis Ponert., subsp. albus) occur on the Balkans, and it is mainly in this region that wild forms with dotted dark-brown seeds and dark-blue flowers are found (subsp.graecus). Additional confirmation is the Greek name of white lupin: “thermos” (hot). The centers of diversity of cultivated white lupin also include the Apennines and Egypt where cultivated forms of white lupin originated in the ancient times. Moreover, in the ancient Egypt forms with pink-and-blue or light-pink flowers were spread (subsp. termis), and on the Apennines forms with grayish-and-light-blue or white flowers were distributed (subsp. albus). Existence on the Pyrenees of two close species (L. luteus L. and L.hispanicus Boiss.et Reut.) with the same number of chromosomes (2n=52), wide diversity of wild and cultivated forms of yellow lupin (L. luteus L), and the long historical period of their growing give the reasons to suppose that the Pyrenees harbored the center of formation of wild yellow lupin forms and the center of diversity of its cultivated plants (Kurlovich, 1998). For similar reasons, we also regard the Pyrenees as the center of diversity for narrow-leafed lupin (L. angustifolius L.). As a result of further agricultural development of lupins, their area of distribution gradually increased. In this connection, secondary macro- and micro-centers of diversity of cultivated lupin forms emerged on different continents. These secondary centers correspond to the numerous geotypes and ecotypes described by us. In our opinion (Kurlovich, 1998), the present-day secondary centers of diversity of cultivated white lupin (L. albus L.) include France, Germany, Poland, Belarus, Russia and Chile. Secondary centers of diversity of blue lupin (L. angustifolius L.) are, first of all, in Australia, then in South Africa, S.-E. United States, Poland and Belarus.
To the secondary centers of diversity of yellow lupin (L. luteus L.) Germany, Poland, Belarus, Russia, and Ukraine may be attributed.
Our eco-geographical researches fully confirm the concept of Vavilov about primary localization of dominant genes, first of all, in the primary centers of origin (diversity) and localization of recessive genes at the edges of the species distribution area and in the secondary centers of diversity. The forms of lupin occurring within the primary centers more often possess dominant characters, basically typical of wild forms, i.e. strongly pigmented flowers, seed and vegetative organs, dehiscent pods, monopodial branching. Domesticated forms cultivated mainly in the secondary centers typically have larger white seed, non-dehiscent pods and limited branching (recessive attributes). Vavilov’s differential systematic-geographical method of crop studies provided a possibility to perform targeted search for valuable breeding materials in various regions and, at the same time, solve the problems of phylogenesis, taxonomy and evolution. This enabled us not only to disclose the diversity of forms, but also to identify a series of regularities in their variation depending on the degree of cultivation and on the geographic, environmental and soil conditions. The annual domesticated Mediterranean species of lupin, now widely cultivated in different countries (L. albus L., L. luteus L., L. angustifolius L. and others) belong to the group of cultivated plants which maintain close connections with their wild-growing ancestors. Plants of this type only partially and to some extent may be transitioned into domesticated condition, while in the other part they retain a structure common for the wild flora of the Mediterranean and consist of various ecotypes.
Charles Darwin (Дарвин Чарлз, 1952) came to the conclusion that "... Each species was formed in one area and was settled so far, as it was possible ". The species of lupin also occupied their potential areas, and thus adapted to new habitats as a result of their development and domestication. The Mediterranean region, as the center of formation of wild-growing lupins, was at the same time one of the first places of their domestication. White lupin was domesticated in the antiquity in Greece (Gladstones, 1974). Yellow and narrow-leafed lupins were cultivated in Spain and Portugal for green manure (Майсурян and Атабекова, 1974). Some domesticated plants in these places subsequently turned wild again for different reasons, but continued to grow on the edges of fields and roads. The publication of Klinkowski (1938, 1939) testifies to it. Domesticated lupins were distributed from the Mediterranean region into other countries, in the area of influence (Sinskaja, 1969). The areas of influence for the ancient Mediterranean were Middle and Northern Europe (Sinskaja, 1969). Lupins were first introduced in these countries, and later underwent certain changes caused by different environmental conditions. Owing to these circumstances, Mediterranean and European countries possess extremely diverse genetic resources of lupin, significant part of which have been concentrated in the collection of the Vavilov Institute (VIR) by means of plant collecting and inter-country germplasm exchange. It is possible to identify the following basic groups representing many lupin species:
Local forms cultivated in different historical periods.
Plant populations turned wild.
Breeding sources, gene donors and different mutants created and collected by man.
Modern commercial cultivars and varieties.
All these five groups are represented by an extremely rich diversity, and incorporate morphological, ecological and geographical distinctions promoted by both natural and anthropogenetic factors. Wild plants evolved in various natural conditions, therefore their characters vary depending on the region, soil and climate. Local forms cultivated in different historical periods underwent a new cultural step of evolution. Transition of a part of plants into the wild state also produced an effect on their biological features. It was also one of the stages of their evolution. Sinskaja (1969) marked in this connection that a plant population that had turned wild should be distinguished from the wild ancestors of cultivated plants in each given case. There is still a possibility for them to be returned to cultivation or be used in breeding practice as sources of valuable characters.
Our researches have shown that plant populations which have turned wild ripen more regularly, have larger seed, and are more resistant to Fusarium and other diseases as compared with the proper wild forms. Breeding sources, gene donors and different mutants were developed by man for definite breeding purposes. Modern breeding cultivars have passed one more stage of evolution. However, they also differ among themselves by their adaptation to certain environments of cultivation. Great polymorphism of morphological traits is also revealed in all species of lupin. For example, 4 main colors of the flower corolla and 8 types of seed color were identified for narrow-leafed lupin, respectively 4 and 9 color types for yellow lupin, and 5 and 6 types for white lupin. Thus, a certain interrelation of these characters was found in different forms. Such regularities were taken as a basis for the development of interspecific classifications. It is well known that the species of lupin are differentiated into regional geotypes and local ecotypes over an extensive territory (Синская, 1969; Агаев, 1987; Kurlovich, 1998). They may differ among themselves in their general structure, plant height, number of stalks and branches, leafiness, length of the period of vegetation, biochemical features, disease susceptibility, frost and drought resistance, and yield parameters. Finally, a species becomes a complex population of ecotypes, or an ecosystem. Differences between ecotypes are genetic in nature. Frequently there is complex segregation in the second generation of hybrids produced from crosses between various ecotypes of the same species, since the features of an ecotype are connected with many genes.
All geotypes and ecotypes were developed in the process of interaction between genetic recombination, natural selection and environmental effect. All the studied species of lupin displayed similar ecotypes within the limits of different geotypes with variability governed by Vavilov`s law of homologous series in variation (Vavilov, 1920) and the law of spiral series (Sinskaja 1969). Thus, different geotypes of the Mediterranean area had large-seeded and small-seeded ecotypes, and L. angustifolius L. included ecotypes with very narrow leaflets and also with broader ones. However, there are no ecotypes with absolutely similar features. Geographic regularities were identified in the variability of a number of characters and properties in similar ecotypes.
Moving from the west of the Mediterranean region to the east showed an increase in seed yield, seed weight and oil content in seed for all studied species. Besides, an increase of the length and width of leaflets was observed in narrow-leafed lupin. Accordingly, the duration of the growing period, green matter weight of plants, and protein content in mature seeds were decreasing. Moreover, we have found a negative correlation between protein and oil content. These data were sufficiently confirmed by other researchers (Mota et al. 1982; Simpson and Martins 1984; Swęcicki, 1988; Cowling, 1994). The conducted studies and generalizations helped to formulate answers to the most important questions of plant introduction and breeding, i.e. what kind of genetic diversity should be collected and where, as well as what should be utilized and for what purpose. For instance, lupin accessions of the Iberian geotype proved to be most efficient in breeding for disease resistance or increased green matter yield. Samples of the Balkan-Asian and Palestinian geotypes are more promising for breeding cultivars with higher grain yield, early maturity, larger seed, drought resistance and increased content of oil. Wild-growing forms of all lupin species undoubtedly possess numerous valuable characters, such as small seed, resistance to drought, low temperatures and diseases. However, their cultivation in Russian conditions would be accompanied by considerable difficulties, while introduction of their positive properties into breeding programs requires a lot of time. More adapted to cultivation are the forms previously cultivated but later turned wild, and local varieties from Mediterranean countries. The most efficient potential breeding sources of commercially valuable characters (resistance to anthracnose, stress resistance, high seed quality) have been identified among these accessions for the conditions of Russia.
Many commercial cultivars from the countries with well-developed lupin breeding are also extremely promising for national breeding programs. Breeding programs in lupin-growing countries utilize the most diverse breeding sources and methods mainly because they have advantageous geographical setting. The countries of the Mediterranean region, Australia, New Zealand, Chile and the U.S. have favorable conditions for acclimatization of the great diversity of Mediterranean forms, particularly from the west coastal area of the Mediterranean. Russia and other countries of the former USSR as well as Poland and Germany are situated in more northern latitudes, and therefore have limited possibilities to use wild-growing Mediterranean forms. Many of them may be cultivated only in glasshouses or artificial climate rooms, although some of them are suitable for autumn sowing in the environments of Transcaucasus. Genetic resources from the east coastal area of the Mediterranean have wider application in Russia, because they demonstrate a greater scope of modification variability, have early-ripening habit, and produce larger seed. Besides, in the past years plant breeders in Russia and Poland have also widely used the induced mutagenesis. However, the best results might be attained with an optimum combination of mutagenesis and hybridization of different forms of various eco-geographical origins. Efforts of the researchers helped to disclose very perfect and, at the same time, intricate mechanisms in lupin as well as in all other living organisms. The conventional Darwin ’s theory of evolution is unable to provide exhaustive explanations as to the prime causes of the immense diversity of species and genera in nature. It may be regarded only in a narrow sense, as microevolution of a definite species, or as adaptation of separate populations to external environments or cultivation conditions. In this case, the leading role is assigned through adaptation by natural or artificial selection to the most adjusted specimens, or most useful for man, as far as crops are concerned. Development of the methods enhancing adaptive properties is of primary importance for increasing the productivity of cultivated plants.
To achieve deeper perception of the complicated genetic mechanisms of adaptation, our experiments have enabled us to recommend the methods of studying plant diversity on the basis of the eco-genetic approach by means of testing a large number of samples having different geographic origin in contrasting environments (for example, at the experiment stations of VIR) according to a unified technique (Курлович et al., 1990). With this, the most interesting are the forms identified for their yield parameters, resistance and other properties at several sites of testing, which usually witnesses to the control of these properties by polygenic systems (horizontal polygenic resistance, adaptivity to cultivation conditions, etc.). It seems useful to involve the forms selected by such technique in hybridization between themselves in order to search for positive transgressions on the required valuable characters. Using this approach made it possible to deepen the existing concepts on the nature of inheritance of quantitative plant characters preconditioned by the definite genotype, the environment, and their complex interaction, i.e. determined by a transforming spectrum of genes.
In our experiments, the modifying effect of the environment influenced not only the mode of interaction between the genes controlling determinate branching habit in lupin, but also altered the spectrum of the acting genes and, quite frequently, the direction of their action, which needs to be taken into account in breeding practice. Our researches confirmed the thesis that new cultivars need to be bred for definite environments. But it is necessary also to use sources of initial material which showed stability after testing them in contrasting conditions of environment for making new varieties with the broad ecological plasticity.
Eco-geographical investigations make possible to deliberately separate valuable material also through hybridization of forms with a different character of trait variability, which often proves about nonallelity of controlling genes, and to obtain consequently the transgressive forms. The phenomenon of dominance of characters in the hybrids of the first generation, observed for the first time by Mendel and presented as his first law (Mendel, 1865; Мендель, 1935; Corcos and Monaghan, 1993) appeared quite complex for lupin.
When studying the character of determinate branching in narrow-leafed lupin, we managed to ascertain that the direction of dominance and the degree of its expression may vary depending on the set of genotypes included in hybridization, genetic medium in hybrid organisms, and external environments, as well as in the process of phylogenesis, as a result of accumulation of certain alleles of the genes in the genotypes. The phenomenon of dominance in genetics may be compared with the singing of vocalists in a certain duet where one of the voices very often dominates (or is better discerned) due to its power and color. However, under other circumstances or in another duet such voice may be suppressed by a more powerful one.
Eco-geographic studies of numerous species, samples, hybrids and mutants of lupin revealed a complex nature of inheritance for many characters of this valuable leguminous crop. An expedient inner structure of organs and processes, intricacy and, at the same time, perfection of adaptive and protective mechanisms led us to a conclusion about the impossibility of their emergence solely by evolution, and enabled us to admit the will of the Creator. In view of this, the message of the Bible concerning the creation may be regarded as scientifically convincing evidence. It presents the main categories of plants and animals, as well as the man himself, as multiplying in their large diversity “each according to its kind, upon the Earth” (Bible, Genesis 1:11- 24).
All knowledge of Egyptian sages could not supply Moses, writer of the Genesis, with a key to the process of creation. Moses, therefore, received the revelation for the Book of Genesis from the Lord Himself, and we should trust it without any doubt, as it is proved by a much greater number of historical documents (manuscripts) than required for scientific researches.
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