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Translocation breakpoints in soybean classical genetic linkage groups 6 and 8.

RECIPROCAL TRANSLOCATIONS, also known as chromosomal interchanges, are a type of chromosomal aberration arising from the exchange of broken segments of nonhomologous chromosomes. Thus, an individual with a reciprocal translocation contains a segment of one LG attached to another LG, the breakpoint being the point of attachment of the broken segments. Heterozygous translocations express partial pollen and ovule sterility (40 to 60% sterility), and this character defines the translocation breakpoint which would show linkage with mutants of the two different LGs. Marker loci close to the translocation breakpoint will also show linkage with each other, and where multiple markers are involved, the translocation-marker loci linkage relationships can furnish information needed to determine the orientation of linkage maps. However, accurate determination of the translocation breakpoint, possible with the use of multiple markers, is critical for map orientation. Reciprocal translocations are therefore useful for the location of genes to CLGs, for testing the independence of LGs, and for map orientation. The isolation and identification of a complete tester set of translocations will facilitate classical and molecular genetic linkage mapping in soybean. Sterility due to gene mutation can be complete male-sterility as in the case of [Ms.sub.1] and [Ms.sub.6], or complete male-and female-sterility as for Sts, thus distinguishing this character from the partial male-and female-sterility character in heterozygous translocations. The flower color locus, WI, (CLG 8) has been assigned to Molecular Linkage Group F (MLG F) (Cregan et al., 1999), but CLG 6 has not yet been associated with a MLG.

Earlier studies (Palmer and Kilen, 1987) have resulted in the isolation of six reciprocal translocations (Table 1). Cytogenetic studies have shown that the translocations KS172-11-3, KS175-7-3, and Clark T/T share a common chromosome that is different from the one shared by KS171-31-2, PI 189866, and L75-0283-4 (Sadanaga and Newhouse, 1982; Sellner, 1990; Mahama et al., 1999). Sadanaga and Grindeland (1984) and Sacks and Sadanaga (1984) used the KS172-11-3 translocation in linkage studies and reported a gene order of [W.sub.1], Wm, breakpoint, [Ms.sub.1]. Using the Clark T/T translocation, Palmer (1985), and Palmer and Kaul (1983) reported a gene order of [W.sub.1], Wm, [Ms.sub.1], breakpoint. In a separate study with CLG 6 loci, Palmer (1985) reported a gene order of [Y.sub.11], [Df.sub.2], breakpoint. The T locus (CLG 1) was reported as independent of the breakpoints in KS172-11-3, and Clark T/T (Palmer, 1976; Sadanaga and Grindeland, 1984). Mapping studies with multiple marker loci composed of CLG 6 and CLG 8 showed that these two LGs are the same LG (Mahama et al., 2002).

Since a common chromosome is involved in an interchange in three soybean translocations, it would be important to know the locations of the breakpoints to determine the similarities of the translocated segments. The objectives of our study were (i) to determine the location of translocation breakpoints by testing linkage with loci of CLG 6 ([Df.sub.2] and [Y.sub.11]), CLG 8 ([Adh.sub.1], [Ms.sub.1], [Ms.sub.6], [St.sub.5], [W.sub.1], and [Y.sub.23]), and other CLGs, and (ii) to confirm the orientation of these nine marker loci.

MATERIALS AND METHODS

Genetic marker types with loci in both coupling and repulsion phase linkage and six translocation lines were used in mapping studies (Tables 1, 2). All loci express complete dominance except the [Y.sub.11] locus, which expresses incomplete dominance, and plants with the genotype [y.sub.11][y.sub.11] are seedling lethal, dying shortly after germination. Plants with the genotype [Y.sub.11][y.sub.11] are viable and have greenish yellow leaves. CS is an unknown male-sterile, female-sterile mutant that has not been tested for allelism with the known male-sterile, female-sterile mutants. It is a sterile recessive gene, and dominant genotypes are fertile. Segregating populations were developed from cross-pollinations among different soybean genotypes (Walker et al., 1979). All the crosses were made in the field at the Brunet Farm near Ames, IA (Table 3). The soil was a Clarion-Nicollet Loam soil type (fine-loamy, mixed, superactive, mesic Typic Hapludolls and fine-loamy, mixed, superactive, mesic Aquic Hapludolls).

Plants of the translocation lines were used as pollen parents in cross-pollinations with the genetic marker types except for plants carrying the [st.sub.5] allele, in which case they were used as female parents. Plants of the genotype [st.sub.5][st.sub.5] are both male and female sterile. Thus, fertile plants of genotype [St.sub.5][st.sub.5], identified by progeny testing, were used as pollen parents. Following the procedures described by Mahama et al. (2002), segregation data for the different characters used as marker traits were collected from [F.sub.2] populations and [F.sub.2:3] families grown at the Brunet Farm. The [F.sub.2] populations were generated from multiple [F.sub.1] crosses. Because of poor survival, small plants, and/or small population sizes, only [F.sub.2:3] data were collected for some crosses. Because of the amount of work, cost and time involved, only [F.sub.2] populations were classified for the [Adh.sub.1] locus. Late maturity and extreme lodging made it difficult to classify certain crosses in the [F.sub.2:3] generation without serious errors. [W.sub.1] and Wm are so closely linked that we chose to classify only [W.sub.1] in the [F.sub.2:3] generation. Plants homozygous for the nontranslocated and translocated chromosomes are fertile, and under normal conditions of growth will produce mostly three-seeded pods. In contrast, plants heterozygous for the translocated chromosomes are partial male and female sterile and will produce mostly one-and two-seeded pods. Classification for partial sterility from translocation heterozygosity was done by pollen staining with iodine potassium iodide solution (Jensen, 1962) and/or visual observation of seed set at maturity (Mahama et al., 2002).

Data analyses were according to Hanson and Kramer (1950), Joachim (1947), and Kramer et al. (1954) for calculating chi-square values, and linkage intensity values involving translocations, from [F.sub.2], and [F.sub.2:3] genetic data. Appropriate coefficients for the different sources of data were entered into a spreadsheet and used to calculate chi-square values to test the independence of loci pairs, as deviations from 50% recombination, and the corresponding recombination values as percentage recombination. Data from [F.sub.2] and [F.sub.2:3] families were analyzed separately. The recessive class was not used in the calculations of [F.sub.2:3] data.

RESULTS AND DISCUSSION

KS172-11-3 and Clark T/T have been used in many linkage studies, but this is the first report on the use of KS175-7-3 in linkage studies. The marker loci [Y.sub.11] and [Df.sub.2] of CLG 6, [Ms.sub.1], [W.sub.1], [St.sub.5], and [Y.sub.23] of CLG 8, and T of CLG 1 segregated independently of the breakpoints in KS171-31-2, L75-0283-4, and PI 189866, indicating that the translocation in these three genotypes involves a different chromosome (Tables 4, 5). [F.sub.2] and [F.sub.2:3] data showed that [Df.sub.2], [Y.sub.11], [Ms.sub.1], and [W.sub.1] were linked to the breakpoints in KS172-11-3, KS175-7-3, and Clark T/T (Fig. 1, 2, and 3, respectively) with the breakpoint between [Df.sub.2], and [Y.sub.11]. The recombination values calculated from [F.sub.2] and [F.sub.2:3] data differed greatly in some cases. This is because of the fact that the chance of error is greater in the [F.sub.2] generation than the [F.sub.2:3] generation because phenotypic evaluation is based on single plants in the [F.sub.2] compared with 25 to 50 plants in the [F.sub.2:3] generation. As expected, the T locus assorted independently of the breakpoints in KS172-11-3, KS175-7-3, and Clark T/T.

Our data also revealed that [Dt.sub.1], Ep, [Fr.sub.1], G, I, [K.sub.2], Pb, [Ms.sub.3], [M.sub.4], [Rj.sub.1], [St.sub.2], [St.sub.3], [St.sub.4], [Y.sub.9], [Y.sub.10], [Y.sub.12], and [Y.sub.18] were independent of the breakpoint in Clark T/T (Tables 4, 5). This lack of linkage, together with independent assortment between [Ms.sub.1] and Ep, and [W.sub.1] and Ep (Mahama et al., 2002) are in agreement with the placement of these loci in different LGs than CLG 6 and CLG 8. The Ep locus was independent of L75-0283-4, while the breakpoint in PI 189866 was linked to the CS malesterile, female-sterile mutant (R = 24.7 [+ or -] 5.3) (Table 5). Additional linkage data are needed to confirm the observed linkage and to assign the CS mutant to a LG.

On the basis of the observation of chromosome number and constitution during meiosis, Sacks and Sadanaga (1984) reported linkage between [W.sub.1] and the breakpoint (2.0 [+ or -] 1.2) and [Ms.sub.1] and the breakpoint (18.4 [+ or -] 3.3), while Sadanaga and Grindeland (1984) reported linkage between the breakpoint and [W.sub.1] (1.9 [+ or -] 0.8) in KS172-11-3, and independent assortment with T and [Y.sub.10]. Both studies placed the breakpoint distal to [W.sub.1], and Sacks and Sadanaga (1984) placed the breakpoint between [W.sub.1] and [Ms.sub.1].

Using standard linkage estimation methods, Palmer (1985) reported linkage between the breakpoint in Clark T/T and [Y.sub.11] (18.4 [+ or -] 0.4) and [Df.sub.2] (5.8 [+ or -] 0.2), and also between the breakpoint and [Ms[+ or -]1] (8.5 [+ or -] 1.5), Wm (36.8 [+ or -] 1.6), and [W.sub.1] (39.5 [+ or -] 2.0), thus placing the breakpoint between [Ms.sub.1] and [Df.sub.2]. The values reported in our study for CLG 8 are in agreement with those reported by Palmer and Kaul (1983) and Palmer (1985), but different from Sadanaga and Grindeland (1984) and Sacks and Sadanaga (1984). These differences may be reflective of the different methods ([F.sub.2] and [F.sub.2:3] segregating data vs. microscope observations) used to obtain linkage data and the different generations of self-pollination that occurred before the translocations were used in the different studies. Whereas Sadanaga and Grindeland (1984) and Sacks and Sadanaga (1984) used the translocations shortly after their isolation following mutagenesis, five generations of self-pollination occurred before they were used in our study. Hence, their recombination values would have been subject more to any unknown instabilities or biases due to differential gametic or genotypic viabilities.

The placement of marker loci in relationship with the translocation breakpoints is based on the recombination values calculated. Linkage maps for KS172-11-3 (Fig. 1), KS175-7-3 (Fig. 2), and Clark T/T (Fig. 3) were developed from our results. Our data placed the breakpoint in all three translocations between [Y.sub.11] and [Ms.sub.1]. This placement was possible because genotypes carrying both loci were used to generate segregating populations in this study. The orientation of marker loci [Df.sub.2], [Y.sub.11], [Ms.sub.1], [W.sub.1], Wm, [Ms.sub.6], [Y.sub.23], [St.sub.5], and [Adh.sub.1] is similar to the reports of Palmer and Hedges (1993) ([Ms.sub.1], [W.sub.1], Wm, [Ms.sub.6], [Y.sub.23], [St.sub.5], [Adh.sub.1]) and Mahama et al. (2002) (YN, [Df.sub.2] , [Ms.sub.1], Wm, [W.sub.1], [Ms.sub.6], [Y.sub.23], [St.sub.5], [Adh.sub.1]). Mahama et al. (2002) reported linkage between loci of CLG 6 and CLG 8 and indicated that they are the same LG. The linkage observed between the breakpoints and [Y.sub.11] and [Df.sub.2] of CLG 6, and loci of CLG 8 ([Ms.sub.1], [W.sub.1]) serves as further evidence that the two are the same LG. On the basis of cytogenetic data, KS172-11-3, KS175-7-3, and Clark T/T were reported to share one common interchange chromosome, and KS171-31-2, PI 189866, and L75 0283-4 shared a different common chromosome (Sadanaga and Newhouse, 1982; Sellner, 1990; Mahama et al., 1999). The linkage of loci of CLG 6 and CLG 8 with the breakpoints in KS172-11-3, KS175-7-3, and Clark T/T, and the independent assortment of these loci with the breakpoints in KS171-31-2, PI 189866, and L75-0283-4 observed in this study confirm that the two groups of translocations carry different translocated chromosomes.

[FIGURES 1-3 OMITTED]

CONCLUSIONS

Recombination values indicated that the loci T, [Y.sub.9], [Y.sub.10], and [Y.sub.18], previously identified as unlinked to CLG 8 loci, as well as [Dt.sub.1], Ep, [Fr.sub.1], G, I, [K.sub.2], Pb, [Ms.sub.3], [M.sub.4] , [Rj.sub.1], [St.sub.2], [St.sub.3], [St.sub.4] and [y.sub.12], were independent of the breakpoints in chromosome translocations KS172-11-3, KS175-7-3, and Clark T/T. All three translocations showed linkage with [Y.sub.11] and [Df.sub.2], and [Ms.sub.1], and [W.sub.1], of CLG 8, with the breakpoint between [Y.sub.11] and [Ms.sub.1], while the breakpoints in KS171-31-2, L75-0283-4, and P1189866 were independent of these loci. These data confirm that KS172-11-3, KS175-7-3, and Clark T/T have a common interchange zhromosome, and that KS171-31-2, L75-0283-4, and PI 189866 do not share this common interchange chromosome. Our data are in agreement with the previous report (Mahama et al., 2002), and we propose that CLG 6 and CLG 8 be combined into one LG designated CLG 8 while dropping CLG 6.

The map orientation, once CLG 6 and CLG 8 are combined, is, in general, consistent with reported maps, and our data place [Df.sub.2] and [Adh.sub.1] at the opposite ends of the chromosome segment. We report here for the first time linkage between the CS male-sterile, female-sterile mutant and the breakpoint in PI 189866. The data reported in this study will contribute toward the assignment of genes to LGs and reassignment of LGs, the isolation of a tester set of translocations, and will enhance classical and molecular genetic linkage mapping.

Abbreviations: LG, Linkage Group; CLG, Classical Linkage Group; MLG, Molecular Linkage Group.
Table 1. Origin of six translocation lines in soybean.

Translocation    Origin

Clark T/T        Near-isogenic Clark with translocation from PI
                   101404B (Glycine soja Siebold & Zucc.) from
                   northeastern China.
KS172-11-3       Translocation from an irradiated population of
                   'Hodgson', selected by K. Sadanaga.
KS175-7-3        Translocation from an irradiated population of
                   'Steele', selected by K. Sadanaga.
KS171-31-2       Translocation from an irradiated population of
                   'Hodgson', selected by K. Sadanaga.
L75-0283-4       Spontaneous translocation in an F4 progeny row of
                   'Beeson' x 'Amsoy 71' cross, found by B.G. Palmer
                   in Illinois.
PI 189866        Glycinegracilis Skvorlzov introduction from
                   northeastern China identified by X. Delannay and
                   B.G. Palmer.

Table 2. Phenotypic description of alleles of soybean genes studied.

             Classical
              Linkage
Gene           Group             Phenotype                Reference

T                1        Tawny pubescence          Piper and Morse
                                                      (1910), Woodworth
                                                      (1921)
t                1        Gray pubescence
[Y.sub.12]       1        Normal green leaves       Weiss (1970)
[y.sub.12]       1        Whitish primary leaves,
                            yellowish-green
                            leaves
G                3        Green seed coat           Terao (1918),
                                                      Takahashi and
                                                      Fukuyama (1919),
                                                      Nagai (1921),
                                                      Woodworth (1921)
g                3        Yellow seed coat
D[t.sub.1]       5        Indeterminate stem        Woodworth (1932,
                                                      1933), Bernard
                                                      (1972)
d[t.sub.1]       5        Determinate stem
D[f.sub.2]       6        Normal plant              Porter and Weiss
                                                      (1948), symbol by
                                                      Byth and Weber
                                                      (1969)
d[f.sub.2]       6        Dwarf plant
[Y.sub.11]       6        Normal (heterozygote      Weber and Weiss
                            has greenish-yellow       (1959)
                            leaves)
[y.sub.11]       6        Lethal yellow plant
I                7        Light hilum color         Nagai (1921), Nagai
                                                      and Saito (1923),
                                                      Owen (1928),
                                                      Woodworth (1932,
                                                      1933), Mahmud and
                                                      Probst (1953)
i                7        Self dark seed coat
Ad[h.sub.1]      8        ADH band 1 present        Gorman and Kiang
                                                      (1978), Kiang and
                                                      Gorman (1983)
ad[h.sub.1]      8        ADH band 1 absent
M[s.sub.1]       8        Fertile                   Brim and Young
                                                      (1971),
                                                      Boerma and Cooper
                                                      (1978), Palmer et
                                                      al. (1978)
m[s.sub.1]       8        Male sterile
M[s.sub.6]       8        Fertile                   Skorupska and Palmer
                                                      (1989)
m[s.sub.6]       8        Male sterile
S[t.sub.5]       8        Fertile                   Palmer and Kaul
                                                      (1983)
s[t.sub.5]       8        Desynaptic sterile
[W.sub.1]        8        Purple flower             Takahashi and
                                                      Fukuyama
                                                      (1919), Woodworth
                                                      (1923)
[w.sub.1]        8        White flower
Wm               8        Purple flower             Buzzell et al.
                                                      (1977)
wm               8        Magenta flower
[Y.sub.23]       8        Normal leaves             Palmer et al. (1990)
[y.sub.23]       8        Leaves yellowish-white
                            and necrotic
R[j.sub.1]       11       Nodulating                Williams and Lynch
                                                      (1954) (as no);
                                                      symbol by
                                                      Caldwell (1966)
r[j.sub.1]       11       Nonnodulating
Ep               12       High seed coat            Buzzell and Buttery
                            peroxidase activity       (1969)
ep               12       Low seed coat
                            peroxidase activity
F[r.sub.1]       12       Roots fluorescent in UV   Delannay and Palmer
                            light                     (1982b)
f[r.sub.1]       12       Nonfluorescent
Pb               14       Sharp hair tip            Ting (1946)
pb               14       Blunt hair tip
[Y.sub.9]        14       Normal                    Probst (1950)
[Y.sub.9]        14       Bright greenish-yellow
                            seedlings
[K.sub.2]        --
             ([dagger])   Yellow seed coat          Rode and Bernard
                                                      (1975)
[k.sub.2]        --       Tan saddle on seed coat   Palmer (1984)
M[s.sub.3]       --       Fertile                   Palmer et al. (1980)
m[s.sub.3]       --       Male sterile
M[s.sub.4]       --       Fertile                   Delannay and Palmer
                                                      (1982a)
m[s.sub.4]       --       Male sterile
S[t.sub.2]       --       Fertile                   Hadley and Starnes
                                                      (1964)
s[t.sub.2]       --       Asynaptic sterile
S[t.sub.3]       --       Fertile                   Hadley and Starnes
                                                      (1964)
s[t.sub.3]       --       Asynaptic sterile
S[t.sub.4]       --       Fertile                   Hadley and Starnes
                                                      (1964)
s[t.sub.4]       --       Desynaptic sterile
[Y.sub.10]       --       Normal                    Probst (1950)
[Y.sub.10]       --       Greenish-yellow
                            seedlings
[Y.sub.18]       --       Normal                    Peterson and Weber
                                                      (1969)
[Y.sub.18]       --       Near lethal yellow        Sheridan and Palmer
                            seedlings                 (1975)

([dagger]) Loci not assigned to any linkage group based on linkage map
in Palmer and Shoemaker (1998).

Table 3. List of crosses used to generate segregating populations
of soybean.

Cross number  Genotypest ([dagger])

Clark T/T
1             [w.sub.1][w.sub.1] ad[h.sub.1] ad[h.sub.1] tt N/N x
                [W.sub.1][W.sub.1] Ad[h.sub.1] Ad[h.sub.1] TT T/T
2             [w.sub.1][w.sub.1] ad[h.sub.1]ad[h.sub.1]
                m[s.sub.1]m[s.sub.1] tt N/N x [w.sub.1][W.sub.1]
                Ad[h.sub.1]Ad[h.sub.1] M[s.sub.1]M[s.sub.1] TT T/T
3             [w.sub.1][w.sub.1] ad[h.sub.1]ad[h.sub.1]
                m[s.sub.6]m[s.sub.6] tt N/N x [W.sub.1][W.sub.1]
                Ad[h.sub.1]Ad[h.sub.1] M[s.sub.6]M[s.sub.6] TT T/T
4             [w.sub.1][w.sub.1] ad[h.sub.1]ad[h.sub.1]
                [y.sub.23][y.sub.23] tt N/N x [W.sub.1][W.sub.1]
                Ad[h.sub.1]Ad[h.sub.1] [Y.sub.23][Y.sub.23] TT T/T
5             [w.sub.1][w.sub.1] [y.sub.23][y.sub.23] tt
                S[t.sub.5]S[t.sub.5] T/T x [W.sub.1][W.sub.1]
                [Y.sub.23][Y.sub.23] TT S[t.sub.5]S[t.sub.5] N/N
6             [w.sub.1][w.sub.1] [Y.sub.11][Y.sub.11]
                m[s.sub.1]m[s.sub.1] tt N/N x [W.sub.1][W.sub.1]
                [Y.sub.11][Y.sub.11] M[s.sub.1]M[s.sub.1] TT T/T
7             [w.sub.1][w.sub.1] m[s.sub.1]m[s.sub.1] tt N/N x
                [W.sub.1][W.sub.1] M[s.sub.1]M[s.sub.1] TT T/T
8             [w.sub.1][w.sub.1] d[f.sub.2]d[f.sub.2]
                [Y.sub.11][y.sub.11] m[s.sub.1]m[s.sub.1] tt N/N x
                [W.sub.1][W.sub.1] D[f.sub.2]D[f.sub.2]
                [Y.sub.11][Y.sub.11] M[s.sub.1]M[s.sub.1] TT T/T
9             d[t.sub.1]d[t.sub.1] N/N x D[t.sub.1]D[t.sub.1] T/T
10            [w.sub.1][w.sub.1] epep M[s.sub.1]m[s.sub.1] tt N/N x
                [W.sub.1][W.sub.1] EpEp M[s.sub.1]M[s.sub.1] TT T/T
11            GG N/N x gg T/T
12            II N/N x ii T/T
13            [w.sub.1][w.sub.1] m[s.sub.1]m[s.sub.1] N/N x
                [W.sub.1][W.sub.1] M[s.sub.1]M[s.sub.1] T/T
14            [w.sub.1][w.sub.1] wmwm N/N x [W.sub.1][W.sub.1] WmWm T/T
15            [w.sub.1][w.sub.1] d[f.sub.2]d[f.sub.2]
                [Y.sub.11][y.sub.11] m[s.sub.1]m[s.sub.1] N/N x
                [W.sub.1][W.sub.1] D[f.sub.2]D[f.sub.2]
                [Y.sub.11][Y.sub.11] M[s.sub.1]M[s.sub.1] T/T
16            r[j.sub.1]r[j.sub.1] tt epep N/N x R[j.sub.1]R[j.sub.1]
                TT EpEp T/T
17            F[r.sub.1]f[r.sub.1] N/N x F[r.sub.1]F[r.sub.1] T/T
18            S[t.sub.2]s[t.sub.2] N/N x S[t.sub.2]S[t.sub.2] T/T
19            PbPb N/N x pbpb T/T
20            [K.sub.2][K.sub.2] N/N x [k.sub.2][k.sub.2] T/T
21            S[t.sub.3]s[t.sub.3] N/N x S[t.sub.3]S[t.sub.3] T/T
22            S[t.sub.4]s[t.sub.4] N/N x S[t.sub.4]S[t.sub.4] T/T
23            m[s.sub.3]m[s.sub.3] N/N x M[s.sub.3]M[s.sub.3] T/T
24            m[s.sub.4]m[s.sub.4] N/N x m[s.sub.1]m[s.sub.1] T/T
25            [y.sub.9][y.sub.9] N/N x [Y.sub.9][Y.sub.9] T/T
26            [y.sub.10][y.sub.10] N/N x [Y.sub.10][Y.sub.10] T/T
27            [y.sub.12][y.sub.12] N/N x [Y.sub.12][Y.sub.12] T/T
28            [y.sub.18][y.sub.18] N/N x [Y.sub.18][Y.sub.18] T/T
KS172-11-3
29            [w.sub.1][w.sub.1] ad[h.sub.1]ad[h.sub.1] TT N/N x
                [W.sub.1][W.sub.1] Ad[h.sub.1]Ad[h.sub.1] tt T/T
30            [w.sub.1][w.sub.1] ad[h.sub.1]ad[h.sub.1]
                m[s.sub.1]m[s.sub.1] TT N/N x [W.sub.1][W.sub.1]
                Ad[h.sub.1]Ad[h.sub.1] M[s.sub.1][M.sub.1] tt T/T
31            [w.sub.1][w.sub.1] ad[h.sub.1]ad[h.sub.1]
                [y.sub.23][y.sub.23 TT N/N x [W.sub.1][W.sub.1]
                Ad[h.sub.1]Ad[h.sub.1] [Y.sub.23][Y.sub.23] tt T/T
32            [w.sub.1][w.sub.1] [y.sub.23][y.sub.23]
                S[t.sub.5]S[t.sub.5] TT T/T x [W.sub.1][W.sub.1]
                [Y.sub.23][Y.sub.23] S[t.sub.5]s[t.sub.5] tt N/N
33            [w.sub.1][w.sub.1][y.sub.23][y.sub.23] TT N/N x
                [W.sub.1][W.sub.1] [Y.sub.23][Y.sub.23] tt T/T
34            [w.sub.1][w.sub.1] [Y.sub.11][y.sub.11]
                m[s.sub.1]m[s.sub.1] TT N/N x [W.sub.1][W.sub.1]
                [Y.sub.11][Y.sub.11]M[s.sub.1]M[s.sub.1] tt T/T
35            [w.sub.1][w.sub.1] d[f.sub.2]d[f.sub.2] TT N/N x
                [W.sub.1][W.sub.1]D[f.sub.2]D[f.sub.2] tt T/T
36            [w.sub.1][w.sub.1] m[s.sub.1]m[s.sub.1] TT N/N x
                [W.sub.1][W.sub.1] M[s.sub.1]M[s.sub.1] tt T/T
37            m[s.sub.1]m[s.sub.1] TT N/N x M[s.sub.1]M[s.sub.1] tt T/T
KS175-7-3
38            [w.sub.1][w.sub.1] TT N/N x [W.sub.1][W.sub.1] tt T/T
39            [w.sub.1][w.sub.1] m[s.sub.1]m[s.sub.1] TT N/N x
                [W.sub.1][W.sub.1] M[s.sub.1]M[s.sub.1] tt T/T
40            [w.sub.1][w.sub.1] [y.sub.23][y.sub.23] TT N/N x
                [W.sub.1][W.sub.1] [Y.sub.23][Y.sub.23] tt T/T
41            [w.sub.1][w.sub.1] [y.sub.23][y.sub.23]
                S[t.sub.5]S[t.sub.5] TT T/T x [W.sub.1][W.sub.1]
                [Y.sub.23][Y.sub.23] S[t.sub.5]s[t.sub.5] tt N/N
42            [w.sub.1][w.sub.1] [Y.sub.11][y.sub.11]
                m[s.sub.1]m[s.sub.1] TT N/N x [W.sub.1][W.sub.1]
                [Y.sub.11][Y.sub.11]M[s.sub.1]M[s.sub.1] tt T/T
43            [w.sub.1][w.sub.1] d[f.sub.2]d[f.sub.2] TT N/N x
                [W.sub.1][W.sub.1]D[f.sub.2]D[f.sub.2]
                [Y.sub.11][Y.sub.11] tt T/T
44            [w.sub.1][w.sub.1] m[s.sub.1]m[s.sub.1] TT N/N x
                [W.sub.1][W.sub.1] M[s.sub.1]M[s.sub.1] tt T/T
45            m[s.sub.1]m[s.sub.1] TT N/N x M[s.sub.1]M[s.sub.1] tt T/T
KS171-31-2
46            [W.sub.1][W.sub.1] D[f.sub.2]D[f.sub.2]
                [Y.sub.11][Y.sub.11] tt T/T x [w.sub.1][w.sub.1]
                d[f.sub.2]d[f.sub.2] [Y.sub.11][Y.sub.11] TT N/N
47            [w.sub.1][w.sub.1] [y.sub.23][y.sub.23] N/N x
                [W.sub.1][W.sub.1] [Y.sub.23][Y.sub.23] T/T
48            [w.sub.1][w.sub.1] m[s.sub.1]m[s.sub.1] TT N/N x
                [W.sub.1][W.sub.1] M[s.sub.1]M[s.sub.1] tt T/T
49            S[t.sub.5]S[t.sub.5] T/T x S[t.sub.5]s[t.sub.5] N/N
L75-0283-4
50            [w.sub.1][w.sub.1] d[f.sub.2]d[f.sub.2]
                [Y.sub.11][y.sub.11] m[s.sub.1]m[s.sub.1] N/N x
                [W.sub.1][W.sub.1] D[f.sub.2]D[f.sub.2]
                [Y.sub.11][Y.sub.11] M[s.sub.1]M[s.sub.1] T/T
51            EpEp TT N/N x epep tt T/T
52            [w.sub.1][w.sub.1] EpEp msymsy TT N/N x
                [W.sub.1][W.sub.1] epep m[s.sub.1]m[s.sub.1] tt T/T
53            [w.sub.1][w.sub.1] d[f.sub.2]d[f.sub.2]
                [Y.sub.11][y.sub.11] epep tt T/T x [W.sub.1][W.sub.1]
                D[f.sub.2]D[f.sub.2] [Y.sub.11][Y.sub.11] EpEp
                TT N/N
PI 189866
54            [Y.sub.11][y.sub.11] m[s.sub.1]m[s.sub.1] N/N x
                [Y.sub.11][Y.sub.11] M[s.sub.1]M[s.sub.1] T/T
55            CSCS tt N/N x CSCS TT T/T ([double dagger])
56            [Y.sub.11][Y.sub.11] N/N x [Y.sub.11][y.sub.11] T/T

([dagger]) N/N = homozygous for noninterchange (normal) chromosome; T/T
= homozygous for interchange chromosome.

([double dagger]) CS is a male-sterile, female-sterile mutant; not
tested for allelism with known male-sterile, female-sterile mutants.

Table 4. Percentage recombination values between loci and translocation
breakpoints calculated from [F.sub.2] progeny data in soybean.

                        Cross                   Z-SS
Tr-Locust ([dagger])    number             ([doubledagger])    Z-N

                            Clark T/T

Tr-Ad[h.sub.1] (#)      1-4                       374           325
Tr-D[f.sub.2]           8, 15                     625           319
Tr-D[t.sub.1]           9                         403           404
Tr-Ep                   10, 16                    271           258
Tr-G                    11                        277           284
Tr-I                    12                        107           106
Tr-M[s.sub.1]           6-8, 10, 13, 15          1876           112
Tr-M[s.sub.6]           3                         115           108
Tr-R[j.sub.1]           16                        503           484
Tr-T                    1-8, 10, 16              2509          2385
Tr-[W.sub.1]            1-8, 10, 13-15           3307           856
Tr-Wm                   14                        300           312
Tr-[Y.sub.11]
  ([double dagger])     6, 8, 15                  210           410
Tr-[Y.sub.23]           4, 5                      882           743
Tr-F[r.sub.1]           17                        276           247
Tr-Pb                   19                        554           502

                           KS172-11-3

Tr-Ad[h.sub.1]          29-31                     152            87
Tr-M[s.sub.1]           30, 34, 36, 37            137            75
Tr-T                    29-31                     407           340
Tr-[W.sub.1]            29-37                     545           278
Tr-[Y.sub.23]           31-33                     373           240

                           KS175-7-3

Tr-M[s.sub.1]           39, 42, 44, 45            257           191
Tr-S[t.sub.5]           41                         26            46
Tr-[W.sub.1]            38-44                    1007           860
Tr-[Y.sub.23]           40, 41                    720           561

                           KS171-31-2

Tr-[W.sub.1]            47                        184           138
Tr-[Y.sub.23]           47                        179           133

                                         No.        [chi square]
                                       [F.sub.2]    deviation
Tr-Locust ([dagger])    zzSS    zzN      pla nts    ([section])

                             Clark T/T

Tr-Ad[h.sub.1] (#)      121      96       916           0.3
Tr-D[f.sub.2]            93     102      1139          32.5
Tr-D[t.sub.1]           140     141      1088           0.0
Tr-Ep                    84      80       693           0.0
Tr-G                     87      84       732           0.1
Tr-I                     35      30       278           0.2
Tr-M[s.sub.1]           --      --       1988         188.5
Tr-M[s.sub.6]           --      --        233           0.2
Tr-R[j.sub.1]           171     150      1308           0.5
Tr-T                    832     771      6497           0.0
Tr-[W.sub.1]            867     702      5732         222.5
Tr-Wm                   105     123       840           0.7
Tr-[Y.sub.11]
  ([double dagger])     825     336      1718         447.4
Tr-[Y.sub.23]           314     274      2213           0.1
Tr-F[r.sub.1]            93      92       708           0.3
Tr-Pb                   185     176      1417           0.0

                            KS172-11-3

Tr-Ad[h.sub.1]           45      48       332           5.5
Tr-M[s.sub.1]           --      --        212          18.1
Tr-T                    120     103       970           0.1
Tr-[W.sub.1]             93     237      1153         141.3423
Tr-[Y.sub.23]            67     162       842          69.2

                            KS175-7-3

Tr-M[s.sub.1]           --      --        448           9.7
Tr-S[t.sub.5]           --      --         72           0.0
Tr-[W.sub.1]            160     189      2216           8.2
Tr-[Y.sub.23]           167      70      1518           3.8

                            KS171- 31-2

Tr-[W.sub.1]             57      32       411           0.7
Tr-[Y.sub.23]            62      37       411           0.7

                        % R [+ or -] SE
Tr-Locust ([dagger])    ([paragraph])

                            Clark T/T

Tr-Ad[h.sub.1] (#)              I
Tr-D[f.sub.2]           20.6 [+ or -] 2.1
Tr-D[t.sub.1]                   I
Tr-Ep                           I
Tr-G                            I
Tr-I                            I
Tr-M[s.sub.1]            7.0 [+ or -] 1.1
Tr-M[s.sub.6]                   I
Tr-R[j.sub.1]                   I
Tr-T                            I
Tr-[W.sub.1]            22.9 [+ or -] 1.0
Tr-Wm                   39.1 [+ or -] 6.9
Tr-[Y.sub.11]
  ([double dagger])     14.4 [+ or -] 0.0
Tr-[Y.sub.23]                   I
Tr-F[r.sub.1]                   I
Tr-Pb                           I

                           KS172-11-3

Tr-Ad[h.sub.1]          27.6 [+ or -] 5.2
Tr-M[s.sub.1]            3.2 [+ or -] 0.1
Tr-T                            I
Tr-[W.sub.1]            13.7 [+ or -] 1.5
Tr-[Y.sub.23]           16.1 [+ or -] 2.0

                            KS175-7-3

Tr-M[s.sub.1]           17.0 [+ or -] 3.7
Tr-S[t.sub.5]                   I
Tr-[W.sub.1]            31.0 [+ or -] 2.4
Tr-[Y.sub.23]                   I

                        KS171-31-2

Tr-[W.sub.1]                    I
Tr-[Y.sub.23]                   I

([dagger]) Tr represents translocation breakpoint.

([double dagger]) Z, z designate dominant and recessive alleles of each
gene, and N, SS designate normal and semisterile plants, respectively.

([section]) [chi square]'s are calculated to test deviations from 50%
recombination based on 1 df as presented by Hanson and Kramer (1950).

([paragraph]) R [+ or -] SE = recombination value [+ or -] standard
error, I = independent.

(#) Expected ratios for all loci are 3 Z-SS: 3 Z-N: 1 zzSS: 1 zzN.

([double dagger]) Segregation ratios for [y.sub.11] are 1[Y.sub.11]
[Y.sub.11]SS: 1[Y.sub.11][Y.sub.11]N: 2 [Y.sub.11][y.sub.11]SS:
2[Y.sub.11][y.sub.11]N.

Table 5. Percentage recombination values between loci and translocation
breakpoints calculated from [F.sub.23] progeny data in soybean.

                        Cross                      Z-SS
Tr-Locust ([dagger])    number                ([doubledagger])    ZzSS

                             Clark T/T

Tr-D[f.sub.2] (#)       8, 15                         97          214
Tr-I                    12                            27           85
Tr-M[s.sub.1]           2, 6-8, 10, 13, 15           151          731
Tr-T                    8, 10, 16                     50          138
Tr-[W.sub.1]            1-8, 10, 13-15               200          363
Tr-S[t.sub.5]           5                             67           37
Tr-[Y.sub.11]           6, 8, 15                     118          650
Tr-S[t.sub.2]           18                            41           79
Tr-[K.sub.2]            20                            42           82
Tr-S[t.sub.3]           21                            50          106
Tr-S[t.sub.4]           22                            55          118
Tr-M[s.sub.3]           23                           153          326
Tr-M[s.sub.4]           24                            66          128
Tr-[Y.sub.9]            25                            47          100
Tr-[Y.sub.10]           26                            46          117
Tr-[Y.sub.11]           27                            60          124
Tr-[Y.sub.18]           28                            86          176

                             KS172-11-3

Tr-D[f.sub.2]           35                            63          102
Tr-M[s.sub.1]           30, 34, 36, 37                65          334
Tr-S[t.sub.5]           32                            42          104
Tr-T                    30, 32, 34-37                225          418
Tr-[W.sub.1]            29-36                         90          300
Tr-[Y.sub.11]           34                           148          165
Tr-[Y.sub.23]           32                            27          119

                             KS175-7-3

Tr-D[f.sub.2]           43                            80           33
Tr-M[s.sub.1]           39, 42, 44, 45                64          201
Tr-S[t.sub.5]           41                            39           34
Tr-T                    43-45                         76          126
Tr-[W.sub.1]            43, 44                        96           80
Tr-[Y.sub.11]           42, 43                        69          222
Tr-[Y.sub.23]           41                            28           45

                             KS171-31-2

Tr-D[f.sub.2]           46                            99          114
Tr-M[s.sub.1]           48                            51          108
Tr-S[t.sub.5]           49                            42          104
Tr-T                    46, 48                        97          215
Tr-[W.sub.1]            46-48                        117          195
Tr-[Y.sub.11]           46                            32           28
Tr-[Y.sub.23]           47                            27          119

                             L75-0283-4

Tr-D[f.sub.2]           50, 53                       113          120
Tr-Ep                   51, 52, 53                   124           47
Tr-M[s.sub.1]           50, 52                        85          181
Tr-T                    51, 53                        65          144
Tr-[W.sub.1]            50, 52, 53                   116          105
Tr-[Y.sub.11]           50                            66          103

                             PI 189866

Tr-M[s.sub.1]           54                           132          299
Tr-CS ([double dagger]) 55                            10           36
Tr-T                    55                            39            7
Tr-[Y.sub.11]           54, 56                        34           67

                                     No.        [chi square]
                                   [F.sub.2]    deviation
Tr-Locust ([dagger])        ZzN      pla nts    ([section])

                         Clark T/T

Tr-D[f.sub.2] (#)            86       592          90.6
Tr-I                         77       214           0.0
Tr-M[s.sub.1]               160      1720         767.6
Tr-T                         82       304          11.9
Tr-[W.sub.1]                288      1230          76.2
Tr-S[t.sub.5]                22       170           2.8
Tr-[Y.sub.11]                92      1477         819.6
Tr-S[t.sub.2]                84       244           0.1
Tr-[K.sub.2]                 78       243           0.1
Tr-S[t.sub.3]                99       302           0.0
Tr-S[t.sub.4]               123       354           0.5
Tr-M[s.sub.3]               283       896           0.0
Tr-M[s.sub.4]               118       369           0.1
Tr-[Y.sub.9]                 93       288           0.1
Tr-[Y.sub.10]                96       308           1.2
Tr-[Y.sub.11]               120       359           0.1
Tr-[Y.sub.18]               150       489           0.1

                        KS172-11-3

Tr-D[f.sub.2]                60       301           7.9
Tr-M[s.sub.1]               151       639          41.8
Tr-S[t.sub.5]                55      2211           0.0
Tr-T                        225       998           0.0
Tr-[W.sub.1]                109       579          25.3
Tr-[Y.sub.11]               195       782          31.5
Tr-[Y.sub.23]                60       220          11.9

                        KS175-7-3

Tr-D[f.sub.2]                35       262           8.3
Tr-M[s.sub.1]               151       540          26.8
Tr-S[t.sub.5]                 4       100           0.0
Tr-T                        164       464           1.0
Tr-[W.sub.1]                 55       347           6.1
Tr-[Y.sub.11]                80       560         130.3
Tr-[Y.sub.23]                12       100           0.2

                        KS171-31-2

Tr-D[f.sub.2]               106       412           0.0
Tr-M[s.sub.1]                90       296           0.2
Tr-S[t.sub.5]                55       220           0.0
Tr-T                        182       596           1.6
Tr-[W.sub.1]                168       596           0.5
Tr-[Y.sub.11]                32       112           3.5
Tr-[Y.sub.23]                50       220           2.5

                        L75-0283-4

Tr-D[f.sub.2]               144       487           0.9
Tr-Ep                        50       348           0.0
Tr-M[s.sub.1]               168       516           0.0
Tr-T                        126       393           0.0
Tr-[W.sub.1]                124       453           1.4
Tr-[Y.sub.11]                85       317           0.2

                        PI 189866

Tr-M[s.sub.1]               267       827           0.4
Tr-CS ([double dagger])      25        91           5.3
Tr-T                         16        91           0.0
Tr-[Y.sub.11]                72       207           0.1

                            % R [+ or -] SE
Tr-Locust ([dagger])          ([paragraph])

                                Clark T/T

Tr-D[f.sub.2] (#)           18.0 [+ or -] 1.5
Tr-I                                I
Tr-M[s.sub.1]               8.9 [+ or -] 0.5
Tr-T                                I
Tr-[W.sub.1]                24.5 [+ or -] 1.4
Tr-S[t.sub.5]                       I
Tr-[Y.sub.11]               6.7 [+ or -] 0.4
Tr-S[t.sub.2]                       I
Tr-[K.sub.2]                        I
Tr-S[t.sub.3]                       I
Tr-S[t.sub.4]                       I
Tr-M[s.sub.3]                       I
Tr-M[s.sub.4]                       I
Tr-[Y.sub.9]                        I
Tr-[Y.sub.10]                       I
Tr-[Y.sub.11]                       I
Tr-[Y.sub.18]                       I

                               KS172-11-3

Tr-D[f.sub.2]               29.4 [+ or -] 3.7
Tr-M[s.sub.1]               23.8 [+ or -] 1.9
Tr-S[t.sub.5]                       I
Tr-T                                I
Tr-[W.sub.1]                26.2 [+ or -] 2.3
Tr-[Y.sub.11]               26.9 [+ or -] 1.9
Tr-[Y.sub.23]               33.1 [+ or -] 5.3

                                KS175-7-3

Tr-D[f.sub.2]               28.4 [+ or -] 3.7
Tr-M[s.sub.1]               25.7 [+ or -] 2.3
Tr-S[t.sub.5]                       I
Tr-T                                I
Tr-[W.sub.1]                31.3 [+ or -] 3.8
Tr-[Y.sub.11]               14.8 [+ or -] 1.8
Tr-[Y.sub.23]                       I

                               KS171-31-2

Tr-D[f.sub.2]                       I
Tr-M[s.sub.1]                       I
Tr-S[t.sub.5]                       I
Tr-T                                I
Tr-[W.sub.1]                        I
Tr-[Y.sub.11]                       I
Tr-[Y.sub.23]                       I

                               L75-0283-4

Tr-D[f.sub.2]                       I
Tr-Ep                               I
Tr-M[s.sub.1]                       I
Tr-T                                I
Tr-[W.sub.1]                        I
Tr-[Y.sub.11]                       I

                                PI 189866

Tr-M[s.sub.1]                       I
Tr-CS ([double dagger])     24.7 [+ or -] 5.3
Tr-T                                I
Tr-[Y.sub.11]                       I

([dagger]) Tr represents translocation breakpoint.

([double dagger]) Z, z designate dominant and recessive alleles of each
gene, and N, SS designate normal and semisterile plants, respectively.

([section]) [chi square]'s are calculated to test deviations from 50%
recombination based on 1 df as presented by Hanson and Kramer (1950).

([paragraph]) R [+ or -] SE = recombination value [+ or -] standard
error, I = independent.

(#) Expected ratios are 3 ZZSS: 2 ZzSS: 1 ZZN: 2 ZzN.

([double dagger]) CS is a male-sterile, female-sterile mutant,
not tested for allelism with known male-sterile, female-sterile
mutants.


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Brim, C.A., and M.F. Young. 1971. Inheritance of a male-sterile character in soybeans. Crop Sci. 11:564-566.

Buzzell, R.I., and B.R. Buttery. 1969. Inheritance of peroxidase activity in soybean seed coats. Crop Sci. 9:387-388.

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Byth, D.E., and C.R. Weber. 1969. Two mutant genes causing dwarfness in soybeans. J. Hered. 60:278-280.

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A. Assbi Mahama and Reid G. Palmer *

A.A. Mahama, Dep. of Plant Pathology, 351 Bessey Hall, and R.G. Palmer, USDA-ARS-CICGR and Dep. of Agronomy and Zoology/ Genetics, Iowa State Univ., Ames, IA 50011. Joint contribution J#19834 of the Iowa Agric. and Home Econ. Exp. Stn., Ames, IA, Project No. 3769 and the USDA-ARS, Corn Insects and Crop Genetics Research Unit, and supported by Hatch Act and State of Iowa. Received 29 July 2002. * Corresponding author (rpalmer@iastate.edu).
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Title Annotation:Crop Breeding, Genetics & Cytology
Author:Assibi Mahama, A.; Palmer, Reid G.
Publication:Crop Science
Date:Sep 1, 2003
Words:8058
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