Journal of Heredity 2004:95(3):265-267
© 2004 The American Genetic Association
Brief Communication |
Inheritance of Flower Color and Spininess in Safflower (Carthamus tinctorius L.)
From the Department of Agronomy and Plant Breeding, Gorgan University of Agricultural Sciences, P.O. Box 386, Gorgan, Iran (Pahlavani), and College of Agriculture, Isfahan University of Technology, Isfahan, Iran (Mirlohi and Saeidi).
Address correspondence to M. H. Pahlavani at the address above, or e-mail: hpahlavani{at}yahoo.com.
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Abstract |
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 Top Abstract Materials and MethodsResult and DiscussionReferences |
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Safflower (Carthamus tinctorius L.) flowers are used for coloring and flavoring food and also as fresh-cut and dried flowers. The most important characteristics which contribute to the ornamental value of safflower are flower color and spinelessness. The objective of this study was to determine the inheritance mode and the number of genes controlling spininess and flower color in some Iranian genotypes of safflower. The results indicated that the existence of spines on the leaves and bracts of safflower is controlled by a single dominant gene in which the spiny phenotype was completely dominant to spineless. In some crosses, flower color was controlled by two epistatic loci each with two alleles, resulting in a ratio of 13:3 in the segregating F2 population for plants with orange and yellow flowers. Also, other mechanisms of genetic control, such as duplicate dominance and duplicate recessive types of epistasis, were observed for flower color in other crosses that led to ratios of 7:9 and 15:1 for plants with orange and yellow flowers, respectively. The results suggest that for ornamental use or in the food dying industry, genotypes with orange or yellow flowers and without spines on the leaves and bracts can be produced.
Safflower (Carthamus tinctorius L.) is an annual herbaceous plant that is adapted to hot and dry environments (Li and Mundel 1996). Traditionally safflower was grown for its flower, used in coloring and flavoring food, making dyes, and in medicine. In the last 30–40 years, however, due to an increasing demand for vegetable oil in the human diet, its production as an oilseed crop has received a great deal of attention. Plants of this crop are 30 to 150 cm tall and most genotypes have many sharp spines on the leaves and bracts. Florets at the bloom stage usually display hues of yellow, orange, red, and rarely white coloration, that change to other colors at the wilted stage (Li and Mundel 1996). Flower color is generally considered neutral for seed yield and oil production, but when safflower is grown for the florets, flower color is important. The most important characteristics in safflower that determine its ornamental value are a combination of flower color (orange and yellow) and few to no spines on the flowers and bracts.
Iran is one of the centers of safflower culture in the old world (Knowles 1969) and different local populations of this crop are found throughout the country (Zeinali 1999). Bradley et al. (1999) observed that among all evaluated genotypes, Iranian accessions with orange or yellow flowers showed promise for ornamental uses as fresh-cut and dried flowers. But all of those genotypes had spiny leaves which decrease their ornamental usefulness. Joglekar and Deshmukh (1956) reported a monogenic control of yellow versus white flower in safflower, with the latter being recessive. Claassen (1952) found that four independent major genes control flower color in safflower. Also, some investigators have found that a number of independent major genes, some with epistatic effect, control floret color in safflower (Narkhede and Deokar 1986; Rao 1943). Monogenic control was also found for spiny versus spineless, with the spiny trait being completely or partially dominant (Deshmukh 1958). Claassen (1952) has reported that spininess was also modified by an unknown number of genes. Also, it was found that spines are basically dominant over spinelessness in safflower and that four genes with epistatic effects are involved (Narkhede and Deokar 1990).
Although the mode of inheritance of flower color and spininess has been investigated, little information is available about the inheritance of these traits in Iranian genotypes. Thus the objective of this study was identification of the inheritance mode and the number of genes that control flower color and leaf spininess in some Iranian safflower genotypes.
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Materials and Methods |
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TopAbstractMaterials and MethodsResult and DiscussionReferences |
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In this study, six genotypes consisting of five lines selected from Iranian populations and Saffire, a Canadian variety, were used as parents in the crosses. The parents were spiny and spineless with different flower color (Table 1). The crosses were made in the summer of 2001 at Research Farm of Isfahan University of Technology, Isfahan, Iran, artificially by hand as described by Knowles (1980). The F2 seeds of the crosses were produced in a greenhouse by bagging F1 plants in paper bags prior to the flowering period. Parental lines, F1, and segregating F2 populations were grown in rows spaced 50 cm apart, with 10 cm spacing of plants within rows, to record the flower color and spiny type of the leaves and bracts at bloom stage on the plants in 2002. Plants with no spines on the leaves and bracts were considered as spineless or nonspiny, according to safflower descriptors (IBPGR 1983).
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Chi-square analyses were applied to test the goodness-of-fit for F2 data based on a one or two gene segregation model for flower color and leaf spininess (Steel and Torrie 1980). A test for heterogeneity of the data for F2 populations of different crosses was done and the data were combined in cases where the heterogeneity chi-square was not statistically significant.
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Result and Discussion |
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TopAbstractMaterials and MethodsResult and DiscussionReferences |
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In all crosses of spiny x spineless genotypes, the phenotypes of F1 leaves and bracts were spiny. Therefore existence of spines on the leaves was considered dominant over the spineless type. In the F2 generation, individuals were classified in two distinct groups fitting a 3:1 ratio of spiny to spineless (Table 2). Therefore it was implied that spininess was controlled by one major gene with two alleles in which Sp, giving the spiny type, was completely dominant over sp, the allele for spinelessness. The low heterogeneity chi-square (Table 2) indicated that different crosses were consistent in terms of genetic control of spininess and these results are in agreement with the findings of others (Ashri and Efron 1964; Claassen 1952). The pooled data also had a suitable fit to the 3:1 ratio. Therefore results indicate that Saffire and IUTH13 should have SpSp and other genotypes should have spsp alleles in the spininess locus.
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In crosses of orange x yellow flower genotypes, all plants in the F1 generation had orange flowers, but in the F2 generation both orange and yellow flowers with different ratios were observed (Table 3). In the P1 x P3 and P2 x P5 crosses, F2 plants fitted to the ratio of 13:3 for orange and yellow flower color, indicating two genes controlling this trait. However, in cross 3 (P2 x P3), the F2 population had a segregation of 4 orange to 12 yellow flower plants. These results led to the hypothesis that flower color in safflower was controlled by two epistatic loci, each with two alleles that can be symbolized as O for orange and Y for yellow color according to Kotecha (1979). Thus the genotype of plants with orange flowers in the F1 generation could be OoYy. In the F2 population of crosses 2 and 4, plants with orange flowers can have the genotypes of O-Y-, O-yy, and ooyy, and yellow flower plants can be of the genotypes ooYY and ooYy. This type of interaction between two loci has been defined as dominance epistasis (Kearsey and Pooni 1996). A cross of orange x orange flower genotypes (P1 x P2) resulted in orange flower plants in both the F1 and F2 generations, indicating that these parental lines (Saffire and IUTC129) had the same genotypes for flower color loci. The segregation ratio of the plants in the F2 population in cross P1 x P4 indicated that the parental line of IUTM12 should have a genotype of ooYY for flower color (Table 3). In this cross, the ooyy genotype had orange flowers and genotypes ooYY and ooYy had yellow flowers. Observations of these segregation ratios for flower color in safflower were in agreement with previous reports (Rao 1943).
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The F2 plants of cross P2 x P5 (orange x yellow), segregated into a 9:7 ratio for yellow and orange color of flower. This suggests that two loci with duplicate recessive epistasis were controlling flower color in progenies of this cross. A segregation ratio in the F2 population of the cross P2 x P6 (orange x yellow) fitted to a ratio of 15:1 for orange and yellow flower, respectively, indicating a duplicate dominance epistasis for two loci controlling flower color (Kearsey and Pooni 1996). These two types of epistasis appear to be different from that described by Ebert and Knowles (1966). This suggests that, in these crosses, the O and Y loci with other types of epistatic effects could be involved in the control of flower color in safflower.
Based on the ratio of the plants in the F2 populations, Saffire (orange), IUTC129 (orange), IUTM12 (yellow), and IUTE1449 (yellow) could have the genotypes OOyy, OOyy, ooYY, and ooYY, respectively. The results of this study show that spininess is controlled by at least one locus and flower color by at least two loci, and that each trait segregates independently. The Iranian line IUTC129, with brilliant orange flowers and without spines, should have a genotype of OOyyspsp, with good potential for hand-harvesting flowers for ornamental use or dry florets for the food industry. The mode of action for genes controlling flower color and spininess showed that for ornamental use or the food dying industry, genotypes with orange or yellow flowers and without spiny leaves and bracts can be produced.
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Footnotes |
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Corresponding Editor: Prem Jauhar
Received March 23, 2003
Accepted December 11, 2003
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References |
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TopAbstractMaterials and MethodsResult and DiscussionReferences |
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Ashri A, Efron Y, 1964. Inheritance studies with fertile interspecific hybrids of three Carthamus L. species. Crop Sci. 4:510-514.
Bradley VL, Guenthner RL, Johnson RC, Hannan RM, 1999. Evaluation of safflower germplasm for ornamental use. In: Perspectives on new crops and new uses (Janick J, ed). Alexandria, VA: ASHS Press.
Claassen CE, 1952. Inheritance of sterility, flower color, spinlessness, attached pappus and rust resistance in safflower, Carthamus tinctorius. Bulletin no. 171. Nebraska Agricultural Experiment Station.
Deshmukh NY, 1958. A note on some studies on inheritance in safflower. Nagpur Agric Coll Mag. 32:32-36.
Ebert WW, Knowles PF, 1966. Inheritance of pericarp types, sterility and dwarfness in several safflower crosses. Crop Sci. 6:579-582.
Joglekar RG, Deshmukh NY, 1956. Inheritance of florets color in safflower (Carthamus tinctorius L.). Proc Bihar Acad Agric Sci. 5:90-116.
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Knowles PF, 1969. Centers of plant diversity and conservation of crop germplasm. Safflower. Econ Bot. 23:324-329.
Knowles PF, 1980. In: Hybridization of crop plants. Edited by Fehr WR and Hadly HH. Madison, WI: Am. Soc. Agron.
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Narkhede BN, Deokar AB, 1986. Inheritance of corolla color in safflower. J Maharashtra Agric Univ. 11:278-281.
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