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Inheritance of high oleic/low ricinoleic acid content in the seed oil of castor mutant OLE-1.

THE QUALITY OF SEED OILS both for food and nonfood applications is largely determined by their fatty acid composition. The seed oil of castor normally contains about 900 g [kg.sup.-1] of ricinoleic acid (D-12-hydroxyoctadec-cis-9-enoic acid) (Brigham, 1993) and is too high for use as an edible oil but gives the oil its traditional industrial usage in manufacture of polymers, lubricants, polyurethane coatings, cosmetics, plastics, and other things (Bonjean, 1991; Brigham, 1993). However, a natural mutant of castor, OLE-1, with significantly less ricinoleic acid (C18:1-OH, about 140 g [kg.sup.-1] compared with about 900 g [kg.sup.-1] in commonly grown cultivars) and increased oleic acid (C18:1, about 780 g [kg.sup.-1] compared with about 40 g [kg.sup.-1] in standard castor bean oil) has been developed by selection from a germplasm accession with high oleic content (Rojas-Barros et al., 2004). This mutant probably has altered gene(s) encoding for the oleoyl-12-hydroxylase enzyme which catalyses the hydroxylation of oleic to ricinoleic acid (Lin et al., 1996, 1998). Since high oleic acid levels are associated with oxidative stability (Friedt, 1988), the oil of the high oleic/ low ricinoleic mutant OLE-1 could have industrial uses requiring high oxidative stability such as for biofuel, or pharmaceutical applications requiring lower ricinoleic levels than the standard castor oil. Moreover, this mutant signifies an important advance toward the development of ricinoleic acid free/high oleic acid castor oil germplasm with potential for the edible oil market.

One requisite for the commercial use of the new oil is the incorporation of the modified biosynthetic pathway into commercial cultivars with good agronomic performance, which requires a knowledge of the genetic behavior of the trait. The inheritance of high oleic/low ricinoleic content in castor remains unexplained to date. However, the genetic control of high oleic content has been found to be simply inherited in different mutants of several oilseed crops. In soybean [Glycine max (L.) Merr.] and safflower (Carthamus tinctorius L.), it is controlled by multiple recessive alleles at a single locus (Takagi and Rahman, 1996: Knowles and Hill, 1964). The content of oleic acid in the corn oil (Zea mays L.) was shown to be controlled by single major gene (Widstrom and Jellum, 1984) or by two independent genes (De la Roche et al., 1971). In peanut (Arachis hypogaea L.), Moore and Knauft (1989) found two loci, designed [Ol.sub.1] and [Ol.sub.2], controlling the high oleic/low linoleic ratio in seed oil. In sunflower (Helianthus annuus L.), the high oleic acid content was found to be controlled by a single partially dominant gene designated Ol (Fick, 1984) or a dominant gene (Urie, 1984). However, later studies demonstrated that the genetic control of the high oleic acid trait in sunflower was more complex. Urie (1985) reported the existence of reversal in dominance and modifying genes. Additionally, Miller et al. (1987) described a second locus, Ml, whose recessive alleles mlml were necessary for the expression of the high oleic acid trait, and Fernandez et al. (1999) also postulated a two-gene (Ol and Ml) model, in which the high oleic acid phenotypes are the result of the expression of the genotype ololMlMl. Finally, Fernandez-Martinez et al. (1989) identified three complementary dominant genes (Old, [Ol.sub.2], and [Ol.sub.3]) controlling the high oleic acid trait in sunflower seed oil.

The objective of this study was to determine the inheritance of the high oleic/low ricinoleic acid content of the castor mutant line OLE-1.

MATERIALS AND METHODS

Plant Materials

The castor germplasm used in this study were (i) OLE-1, a high oleic/low rinoleic mutant line developed by selection from a germplasm accession with high oleic/low ricinoleic content (Rojas-Barros et al., 2004) and (ii) the line A74/18/10, with a standard seed oil fatty acid profile (low oleic/high ricinoleic) selected from breeding material of PROTOSEMENCES Toulouse (France). The fatty acid composition of these materials is shown in Table 1.

Genetic Study

Seeds of OLE-1 and A74/18/10 were individually analyzed for fatty acid composition by the half seed method, described for castor by Rojas-Barros et al. (2004), to ensure that the plants used in the genetic study bred true for seed oil fatty acid composition. A distal portion of the seed was removed with a scalpel and used to determine the fatty acid composition of seed lipids by gas-liquid chromatography (GLC). The remaining portion of the seed containing the embryo, with a known fatty acid profile was used for planting. As the trait high oleic/low ricinoleic is associated with very poor germination (Rojas-Barros et al., 2004), mature embryos of half seeds with this trait, were rescued by in vitro culture with Knudson C Modified Orchid Medium (Knudson, 1946). Plants of OLE-1 were reciprocallv crossed with plants of A74/18/10 in the greenhouse in 1999. In all cases, paper bags were placed over racemes to prevent cross-pollination with external pollen. Crossing was done by emasculating immature flower buds in the raceme of the female parent followed by immediate pollination of their stigmas with fresh pollen from open flowers of the male parent.

[F.sub.1] half seeds from reciprocal crosses as well as seeds from the parents were analyzed for fatty acid composition from plants grown in the greenhouse in 2000. [F.sub.1] plants from reciprocal crosses were self-pollinated to obtain [F.sub.2] seeds and also backcrossed to both parents. Reciprocal crosses were repeated again to obtain [F.sub.1] seeds grown under the same environment as the [F.sub.2] and B[C.sub.1][F.sub.1] seeds. An evaluation of the fatty acid composition of [F.sub.1] plants was made by averaging the GLC analyses of the [F.sub.2] seeds from each individual [F.sub.1] plant. Fatty acid composition was determined on a total of 999 individual [F.sub.2] seeds, 272 B[C.sub.1][F.sub.1] to A74/18/10 seeds, and 217)4 B[C.sub.1][F.sub.1] to OLE-1 seeds.

A total of 21 [F.sub.2] half-seeds representing all the classes for oleic acid concentration detected in this generation were selected, germinated, and grown in a field screenhouse in 2001 to obtain the [F.sub.3] generation. The study of this generation was performed through the analysis of 100 to 200 [F.sub.3] seeds from each segregating [F.sub.2] plant and about 20 to 100 seeds for each nonsegregating [F.sub.2] plant.

Statistical Analyses

Mean oleic and ricinoleic acid content was calculated for the parental lines, the [F.sub.1], and [F.sub.2] generations and compared by the LSD test. Since the results did not reveal important maternal effects for oleic and ricinoleic acid content, the fatty acid composition of segregating generations was analyzed on single seeds. The oleic acid content of B[C.sub.1][F.sub.1], [F.sub.2] and [F.sub.3] seeds was assigned to phenotypic classes on the basis of the appearance of discontinuities in the frequency distribution and the values found in the parentals grown under the same environmental conditions. The proportion of seeds observed in each phenotype class was compared with those expected on the basis of appropriate genetic hypotheses. The goodness of fit to tested ratios was measured by the [chi square] statistic. Heterogeneity [chi square] for families within a cross was nonsignificant so that data for families for the same cross were pooled for analysis.

Fatty Acid Analyses

The fatty acid composition of the seed oil was determined by simultaneous oil extraction and methyl esterification following the procedure described by Rojas-Barros et al. (2004) then analyzed by GLC on a PerkinEhner Autosystem gas-liquid chromatograph (PerkinElmer Corporation, Norwalk, CT) equipped with a flame ionization detector and with a 2-m-long column packed with 3% SP-2310/2% SP-2300 on Chromosorb WAW (Supelco Inc., Bellefonte, PA). The injector, and flame ionization detector were held at 275 and 250[degrees]C, respectively. The gas chromatograph was programmed for an initial oven temperature of 190[degrees]C maintained for 10 mm. followed by an increase of 5[degrees]C [m.sup.-1] up to 225[degrees]C, holding for 7 min.

RESULTS AND DISCUSSION

The average oleic and ricinoleic acid content of the mutant line OLE-1 were 20-fold higher and five-fold lower, respectively, than those of the standard low oleic/ high ricinoleic line A74/18/10 (Table 1). The oleic and ricinoleic acid content in reciprocal [F.sub.1] seeds differed significantly indicating the presence of partial maternal effects for these traits (Table 1). However, these differences were much smaller than those between the [F.sub.1] and each of the parents indicating that the genetic control of the contents of these fatty acids was mainly embryonic with a minor maternal effect. The differences between reciprocal [F.sub.1] seeds were not observed between [F.sub.1] plants ([F.sub.2] seeds averaged) (Table 2) revealing an absence of cytoplasmic effects for both fatty acids. Since no significant cytoplasmic effects could be detected the data from reciprocal [F.sub.2] seeds in Fig. 1 and [F.sub.1], [F.sub.2] and B[C.sub.1][F.sub.1] seeds in Fig. 2 were combined. Similar results, embryonic control with a partial maternal effects of low magnitude in some crosses and the absence of cytoplasmic effects, have been reported for oleic acid content in safflower (Knowles and Hill, 1964), sunflower (Urie, 1984, 1985: Fick, 1984: Miller et al., 1987: Fernandez-Martinez et al., 1989), rapeseed (Schierholt et al., 2001), and soybean (Takagi and Rahman, 1996: Rahman et al., 1994). There are no previous studies on maternal and cytoplasmic effects on the content of ricinoleic concentration.

[FIGURES 1-2 OMITTED]

The average oleic-acid content of [F.sub.1] seeds from reciprocal crosses (52 g [kg.sup.-1]) was very close to that of the standard low line A74/18/10 (Table 1) and much lower than the midparent value (387 g [kg.sup.-1]) indicating almost complete dominance of the standard low over the increased oleic acid content. This result is in agreement with those obtained in safflower (Knowles and Hill, 1964). However, in sunflower several authors reported dominance of high oleic over low oleic acid content (Miller et al., 1987; Fernandez-Martinez et al., 1989). The average ricinoleic-acid content in [F.sub.1] generation (855 g [kg.sup.-1]) was similar to that of the standard low oleic/high ricinoleic line A74/18/10 (Table 1) and higher than the mid-parent value (526 g [kg.sup.-1]) suggesting dominance of standard over reduced ricinoleic acid concentration.

The analysis of fatty acid composition of individual [F.sub.2] seeds from reciprocal crosses between OLE-1 and A74/18/10, showed a considerable variation for oleic and ricinoleic acids (Fig. 1), which were strongly and negatively correlated (r = -0.99, P = 0.001). There was no substantial variation for the other fatty acids. This indicated that the relative proportions of oleic and ricinoleic acids are under the control of one genetic system. The oleic acid values of the [F.sub.2] seeds from OLE-1 x A74/18/10 and A74/18/10 x OLE-1 crosses revealed a clear bimodal distribution (Fig. 1 and 2). The first class with seeds having < 110 g [kg.sup.-1] of oleic acid was assigned to the combined category "low-intermediate" with an oleic acid range between 19 and 110 g [kg.sup.-1] and the second class to the "high" category (oleic acid > 650 g [kg.sup.-1]). The observed data satisfactorily fit a phenotypic ratio 13 low-intermediate: 3 high for these classes (Table 3). This segregation suggests that the high oleic/low ricinoleic content is determined by an interaction between a recessive allele at one locus and a dominant allele at a second locus (dominant and recessive epistasis). These alleles were designated ol and Ml respectively, following the symbols previously assigned to genes controlling oleic acid content in sunflower (Miller et al., 1987). The proposed genotypes for the high oleic-low ricinoleic acid mutant line OLE-1 was ololMlMl and for the low oleic-high ricinoleic acid line A74/18/10, OlOlmlml. With this genetic model the genotypes with high levels of oleic acid would be homozygous for the recessive allele ol. The olol is recessively epistatic to Ml locus and genotypes with these alleles (ololMl_) would produce high oleic/low ricinoleic whereas the olomlml genotypes would produce low oleic/high ricinoleic. The [F.sub.2] phenotypic expression of these genotypes would be in the ratio 13 low-intermediate (including genotypes Ol_Ml_, Ol_mlml and ololmlml): 3 high (genotype ololMl_).

According to the proposed genetic model, a ratio of 1 low-intermediate (including genotypes OlolMlMl and OlolMlml): 1 high (including genotypes ololMlMl and ololMlml) was to be expected in the backcross to the high oleic plant OLE-1, whereas all the individuals were expected to be in the low-intermediate class in the backcross to the low oleic parent A74/18/10. The data observed (Fig. 2) fitted satisfactorily the theoretical ratios (Table 3) supporting the proposed model.

As a further confirmation of the genetic model proposed a progeny test was conducted on crosses by analyzing the [F.sub.3] from each of 21 [F.sub.2] plants. These plants were selected on the basis of the oleic acid content of the corresponding [F.sub.2] half-seeds, which covered the whole range of oleic levels observed in the [F.sub.2] population. The [F.sub.3] seeds derived from [F.sub.2] half-seed plants with low and intermediate oleic values (23-128 g [kg.sup.-1]) showed three different patterns (Table 4). Four of them bred true for values of this fatty acid below 50 g [kg.sup.-1]. The genotypes of these plants were identified as homozygous either for the ml or the Ol allele (genotypes _ _mlml or OlOl_ _). Five segregated for high (>700 g [kg.sup.-1]) oleic acid values with a 3:1 (low-intermediate:high) ratio and nine segregated with a 13:3 (low-intermediate:high) ratio. The segregation 3:1 (one locus) would correspond to a genotype OlolMlMl and the segregation 13:3 (two loci) would be expected for the progeny of the [F.sub.2] genotype OlolMlml. In contrast, all the [F.sub.3] progenies derived from [F.sub.2] half-seeds plants with oleic acid content higher than 700 g [kg.sup.-1] showed no segregation for this fatty acid, the oleic acid content of all the [F.sub.3] seeds being higher than 700 g [kg.sup.-1]. The genotypes of these plants were identified as homozygous for ol and Ml allele (genotype ololMlMl). However, according to the genetic model proposed, [F.sub.2] half-seeds plants with high oleic acid values (>700 g [kg.sup.-1]) hwterozygous for the Ml gene (genotype ololMlml) would be expected to segregate 1 low-intermediate (genotype ololmlml): 3 high (genotypes ololMlMl and ololMlml). The absence of this segregation in the progeny of high oleic/low ricinoleic [F.sub.2] half-seeds plants may be attributed to the fact that only three [F.sub.3] families from high oleic [F.sub.2] half seeds were evaluated due to the poor germination of [F.sub.2] half seeds with this phenotype (Rojas-Barros et al., 2004) and apparently none of them had the ololMlml genotype.

No previous studies have been reported on inheritance of altered oleic acid content in castor. However, similar genetic systems with two genes and epistatic interaction have been proposed for the control of oleic acid content in sunflower (Miller et al., 1987; Fernandez et al., 1999). These studies concluded that the desaturation of oleic to linoleic acid was controlled by a major gene which action was modified by a second gene. Marker analyses supported that the oleoyl-PC-desaturase locus (OLD7) was altered in the high oleic sunflower mutant (Perez-Vich et al., 2002) and other molecular studies (Lacombe et al., 2001) identified another locus, present only in some genotypes, that suppress the effect of the OLD7 locus on the high oleic trait. In the present study, the recessive gene ol present in the high oleic/low ricinoleic castor mutant OLE-1 could affect the action of the oleoyl-12-hydroxylase enzyme preventing the hydroxylation of oleic acid to synthesize ricinoleic acid.

Recessive alleles at the Ml locus would suppress the effect of the ol allele on the oleic/ricinoleic trait. Because of the low number of genes involved in the genetic control of the high oleic/low ricinoleic trait in the mutant OLE-1 a successful transfer of this trait into breeding lines could be performed in a few generations. Furthermore, despite a minor partial maternal effect for oleic acid concentration, the trait appears to be primarily under embryogenic control, which suggests that selection for the high oleic/low ricinoleic trait increased oleic acid (decreased ricinoleic) can be efficiently conducted at the single-seed level by means of the half-seeds technique. The use of this technique will accelerate breeding efforts for this trait.

In conclusion, the information provided by this study clarify the genetic control of the high oleic/low ricinoleic content in the castor mutant OLE-1 and establish the basis for effective breeding strategies for the development of hybrid cultivars with these characteristics.
Table 1. Mean fatty acid content and their standard deviations of the
oil for the castor line A74/18/10, mutant OLE-1, and their reciprocal
[F.sub.1] seeds, based on analysis of half-seeds
from plants grown in Cordoba (Spain) during 1999.

Germplasm                       n (p) ([double dagger])

A74/18/10                               109 (5)
[F.sub.1] (A74/18/10 x OLE-1)            98 (2)
[F.sub.1] (OLE-1 x A74/18/10)            49 (1)
OLE-1                                   206 (2)

                                    Fatty acid (g [kg.sup.-1])
                                            ([dagger])

Germplasm                            C16:0             C18:0

A74/18/10                       12 [+ or -] 2 b   15 [+ or -] 2 b
                                  ([section])
[F.sub.1] (A74/18/10 x OLE-1)   12 [+ or-] 1 b    18 [+ or -] 4 a
[F.sub.1] (OLE-1 x A74/18/10)   10 [+ or-] 3 c    14 [+ or -] 1 b
OLE-1                           17 [+ or-] 3 a    10 [+ or -] 1 c

                                     Fatty acid (g [kg.sup.-1])
                                              ([dagger])

Germplasm                             C18:1              C18:2

A74/18/10                       33 [+ or -] 6 c     57 [+ or -] 6 a
[F.sub.1] (A74/18/10 x OLE-1)   36 [+ or -] 7 c     49 [+ or -] 3 b
[F.sub.1] (OLE-1 x A74/18/10)   68 [+ or -] 28 b    55 [+ or -] 7 a
OLE-1                           742 [+ or -] 23 a   32 [+ or -] 5 c

                                    Fatty acid (g [kg.sup.-1])
                                            ([dagger])

Germplasm                            C18:3            C18:1-OH

A74/18/10                       5 [+ or -] 1 b    870 [+ or -] 10 a
[F.sub.1] (A74/18/10 x OLE-1)   6 [+ or -] 1 a    870 [+ or -] 9 a
[F.sub.1] (OLE-1 x A74/18/10)   4 [+ or -] 1 bc   840 [+ or -] 25 b
OLE-1                           4 [+ or -] 2 c    183 [+ or -] 21 c

([dagger]) C16:0 = palmitic acid, C18:0 = stearic acid, C18:1 = oleic
acid, C18:2 = linoleic acid, C18:3 = linolenic acid, C18:1-OH =
ricinoleic acid.

([double dagger]) Number of half-seed (number of single-plants)
analysed.

([section]) Mean values of parents and reciprocal crosses for each
fatty acid that have the same letter are not significantly different
(LSD, p = 0.05).

Table 2. Oleic acid content of [F.sub.1] half-seeds and mean and range
of oleic and ricinoleic acid content of the oil of [F.sub.2] seeds
from individual [F.sub.1] plants of reciprocal crosses between castor
lines OLE-1 and A74/18/10 grown in Cordoba (Spain) during 2000.

[F.sub.1] plant ([F.sub.2]          C18:1 content in
averaged seeds) or parent seed    [F.sub.1] half-seeds   n ([dagger])

                                               g [kg.sup.-1]

A74/18/10                                                     63
OLE-1                                                        100
[F.sub.1]-1 (A74/18/10 X OLE-1)            45                150
[F.sub.1]-2 (A74/18/10 X OLE-1)            25                149
[F.sub.1]-3 (A74/18/10 X OLE-1)            29                100
[F.sub.1]-4 (OLE-1 X A74/18/10)           109                150
[F.sub.1]-5 (OLE-1 X A74/18/10)           110                150
[F.sub.1]-6 (OLE-1 X A74/18/10)            19                150
[F.sub.1]-7 (OLE-1 X A74/18/10)            25                150

                                  C18:1 content in [F.sub.2] seeds

[F.sub.1] plant ([F.sub.2]
averaged seeds) or parent seed             Mean              Range

                                                g [kg.sup.-1]

A74/18/10                         31                        19-46
OLE-1                             751                       685-795
[F.sub.1]-1 (A74/18/10 X OLE-1)   199 a ([double dagger])   21-798
[F.sub.1]-2 (A74/18/10 X OLE-1)   210 a                     19-835
[F.sub.1]-3 (A74/18/10 X OLE-1)   197 a                     24-787
[F.sub.1]-4 (OLE-1 X A74/18/10)   146 a                     19-802
[F.sub.1]-5 (OLE-1 X A74/18/10)   174 a                     17-833
[F.sub.1]-6 (OLE-1 X A74/18/10)   186 a                     20-792
[F.sub.1]-7 (OLE-1 X A74/18/10)   173 a                     31-812

                                  C18:1-OH content
                                    in [F.sub.2]
                                       seeds

[F.sub.1] plant ([F.sub.2]
averaged seeds) or parent seed     Mean     Range

                                    g [kg.sup.-1]

A74/18/10                         872      843-893
OLE-1                             176      151-239
[F.sub.1]-1 (A74/18/10 X OLE-1)   710 a    117-936
[F.sub.1]-2 (A74/18/10 X OLE-1)   697 a    86-887
[F.sub.1]-3 (A74/18/10 X OLE-1)   709 a    132-888
[F.sub.1]-4 (OLE-1 X A74/18/10)   769 a    115-902
[F.sub.1]-5 (OLE-1 X A74/18/10)   741 a    103-899
[F.sub.1]-6 (OLE-1 X A74/18/10)   729 a    141-892
[F.sub.1]-7 (OLE-1 X A74/18/10)   734 ab   117-879

([dagger]) Number of analysed half-seeds; C18:1 = oleic acid;
C18:1-OH = ricinoleic acid.

([double dagger]) Mean values for each fatty acid of the reciprocal in
[F.sub.2] families that have the same letter are not significantly
different (LSD test, p = 0.05).

Table 3. Number of seeds having different oleic acid
(C18:1) content and Chi-square analyses in the [F.sub.2]
and B[C.sub.1][F.sub.1] seeds from crosses between the
standard castor line A74/18/10 and the mutant line OLE-1.

                                                    No. of seeds with
                                                    C18:1 content

                                                        Low-
                                                    intermediate
                                                      (<110 g
Generation                                          [kg.sup.-1])

[F.sub.2] (A74/18/10xOLE-1)                             118
[F.sub.2] (A74/18/10xOLE-1)                             115
[F.sub.2] (A74/18/10xOLE-1)                              78
[F.sub.2] (OLE-1xA74/18/10)                             128
[F.sub.2] (OLE-1xA74/18/10)                             122
[F.sub.2] (OLE-1xA74/18/10)                             121
[F.sub.2] (OLE-1xA74/18/10)                             125
Pooled                                                  807
Heterogeneity
B[C.sub.1][F.sub.1] ((A74/18/10xOLE-1)xA74/18/10)        74
B[C.sub.1][F.sub.1] ((A74/18/10xOLE-1)xA74/18/10)        82
B[C.sub.1][F.sub.1] ((OLE-1xA74/18/10xA74/18/10)         66
B[C.sub.1][F.sub.1] ((OLE-1xA74/18/10xA74/18/10)         50
B[C.sub.1][F.sub.1] ((A74/18/10xOLE-1)xOLE-1)            25
B[C.sub.1][F.sub.1] ((OLE-1xA74/18/10)xOLE-1)            60
B[C.sub.1][F.sub.1] ((OLE-1xA74/18/10)xOLE-1)            28
Pooled                                                  113
Heterogeneity

                                                    No. of seeds with
                                                    C18:1 content

                                                        High
                                                      (>650 g
Generation                                          [kg.sup.-1])

[F.sub.2] (A74/18/10xOLE-1)                              32
[F.sub.2] (A74/18/10xOLE-1)                              34
[F.sub.2] (A74/18/10xOLE-1)                              22
[F.sub.2] (OLE-1xA74/18/10)                              22
[F.sub.2] (OLE-1xA74/18/10)                              28
[F.sub.2] (OLE-1xA74/18/10)                              29
[F.sub.2] (OLE-1xA74/18/10)                              25
Pooled                                                  192
Heterogeneity
B[C.sub.1][F.sub.1] ((A74/18/10xOLE-1)xA74/18/10)
B[C.sub.1][F.sub.1] ((A74/18/10xOLE-1)xA74/18/10)
B[C.sub.1][F.sub.1] ((OLE-1xA74/18/10xA74/18/10)
B[C.sub.1][F.sub.1] ((OLE-1xA74/18/10xA74/18/10)
B[C.sub.1][F.sub.1] ((A74/18/10xOLE-1)xOLE-1)            23
B[C.sub.1][F.sub.1] ((OLE-1xA74/18/10)xOLE-1)            43
B[C.sub.1][F.sub.1] ((OLE-1xA74/18/10)xOLE-1)            25
Pooled                                                   91
Heterogeneity

                                                      No. of seeds
                                                    with C18:1 content

                                                     [chi square]
Generation                                          (p) ([dagger])

[F.sub.2] (A74/18/10xOLE-1)                          0.60 (0.44)
[F.sub.2] (A74/18/10xOLE-1)                          1.41 (0.24)
[F.sub.2] (A74/18/10xOLE-1)                          0.61 (0.43)
[F.sub.2] (OLE-1xA74/18/10)                          1.99 (0.16)
[F.sub.2] (OLE-1xA74/18/10)                          0.00 (0.98)
[F.sub.2] (OLE-1xA74/18/10)                          0.03 (0.86)
[F.sub.2] (OLE-1xA74/18/10)                          0.46 (0.50)
Pooled                                               0.14 (0.70)
Heterogeneity                                        4.96 (0.55)
B[C.sub.1][F.sub.1] ((A74/18/10xOLE-1)xA74/18/10)
B[C.sub.1][F.sub.1] ((A74/18/10xOLE-1)xA74/18/10)
B[C.sub.1][F.sub.1] ((OLE-1xA74/18/10xA74/18/10)
B[C.sub.1][F.sub.1] ((OLE-1xA74/18/10xA74/18/10)
B[C.sub.1][F.sub.1] ((A74/18/10xOLE-1)xOLE-1)        0.08 (0.77)
B[C.sub.1][F.sub.1] ((OLE-1xA74/18/10)xOLE-1)        2.88 (0.09)
B[C.sub.1][F.sub.1] ((OLE-1xA74/18/10)xOLE-1)        0.17 (0.68)
Pooled                                               2.40 (0.12)
Heterogeneity                                        0.73 (0.69)

([dagger]) Ratios tested: [F.sub.2] generation = 13:3 and
B[C.sub.1][F.sub.1] to OLE-1 generation = 1:1, and p is
the probability level for significance.

Table 4. Number of [F.sub.3] castor seeds having a different oleic
acid (C18:1) content in the analysis of 21 [F.sub.3] families from the
cross A74/ 18/10 x OLE-1 and Chi-square ([chi square]) analyses.

                     No. [F.sub.3] seeds in C18:1 classes

                                C18:1 of
                   C18:1 in    [F.sub.3]
                   [F.sub.2]     family         Low-
[F.sub.3] family   half-seed   ([dagger])   intermediate

                        g [kg.sup.-1]

[F.sub.3]-36           23         196           122
[F.sub.3]-73           33          39            20
[F.sub.3]-48           37         178           121
[F.sub.3]-678          39          43            20
[F.sub.3]-669          39         210           112
[F.sub.3]-675          39         269            78
[F.sub.3]-83           44          43            20
[F.sub.3]-63           48         147           119
[F.sub.3]-72           51          40            20
[F.sub.3]-664          57         233           176
[F.sub.3]-707          78         223           125
[F.sub.3]-690          87         171           128
[F.sub.3]-698          89         200           117
[F.sub.3]-674         104         192           118
[F.sub.3]-75          104         217           113
[F.sub.3]-706         105         181           168
[F.sub.3]-686         109         191           120
[F.sub.3]-114         128         190           118
[F.sub.3]-684         788         789
[F.sub.3]-99          791         832
[F.sub.3]-702         815         751

                          [chi square]   [chi square]
                           (p) 1-gene    (p) 2-genes
[F.sub.3] family   High       3:1            13:3

[F.sub.3]-36        33                   0.60 (0.44)
[F.sub.3]-73
[F.sub.3]-48        29                   0.03 (0.86)
[F.sub.3]-678
[F.sub.3]-669       36    0.04 (0.84)
[F.sub.3]-675       35    1.89 (0.17)
[F.sub.3]-83
[F.sub.3]-63        23                   0.67 (0.41)
[F.sub.3]-72
[F.sub.3]-664       61    0.07 (0.79)
[F.sub.3]-707       42    0.00 (0.96)
[F.sub.3]-690       28                   0.07 (0.80)
[F.sub.3]-698       32                   0.66 (0.41)
[F.sub.3]-674       32                   0.60 (0.44)
[F.sub.3]-75        37    0.01 (0.92)
[F.sub.3]-706       43                   0.35 (0.55)
[F.sub.3]-686       32                   0.48 (0.49)
[F.sub.3]-114       30                   0.21 (0.65)
[F.sub.3]-684      149
[F.sub.3]-99
[F.sub.3]-702

([dagger]) Mean value of analysed [F.sub.3] seeds.


Abbreviations: GLC, gas-liquid Chromatography

ACKNOWLEDGMENTS

This work was supported by the European Commission under the project No AIR3-CT93-2324.

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P. Rojas-Barros, A. de Haro, and J.M. Fernandez-Martinez, Instituto de Agricultura Sostenible (CSIC), Apartado 4084, E-14080 Cordoba, Spain. Received 16 Apr. 2004. * Corresponding author (cs9femaj@ uco.es).
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Title Annotation:Crop Breeding, Genetics & Cytology
Author:Rojas-Barros, Pilar; de Haro, Antonio; Fernandez-Martinez, Jose Maria
Publication:Crop Science
Geographic Code:1USA
Date:Jan 1, 2005
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