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a USDA-ARS, Soybean and Alfalfa Research Lab., Beltsville, MD 20705 USA
b Dep. of Biology, Univ. of Utah, Salt Lake City, UT 84112 USA
c USDA-ARS-CICG, Dep. of Agronomy, Iowa State Univ., Ames, IA 50011 USA
d Biogenetic Services Inc., 801 32 Ave., Brookings, SD 57006 USA
e Yuzuncu Yil Univ., College of Agriculture, 65080-Van, Turkey
f USDA-ARS, Soil Drainage Unit, 590 Woody Hayes Drive, Columbus, OH 43210 USA
g Dep. of Horticulture and Crop Science, The Ohio State Univ. and the Ohio Agric. Res. and Dev. Center, Wooster, OH 44691 USA
h Dep. of Agronomy, Gyeong Sang National Univ., Chinju, South Korea 660-701
i Dep. of Agronomy, Univ. Nebraska, Lincoln, NE 68583-0915 USA
pcregan{at}nal.usda.gov
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ABSTRACT |
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 TOPNOTES ABSTRACT INTRODUCTIONMaterials and methodsResults and discussionREFERENCES |
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Abbreviations: AFLP, amplified fragment length polymorphism • BAC, bacterial artificial chromosome • CLG, classical linkage group • cM, centimorgan • MLG, molecular linkage group • NIL, near isogenic line • PCR, polymerase chain reaction • QTL, quantitative trait loci • RAPD, random amplified polymorphic DNA • RIL, recombinant inbred line • RFLP, restriction fragment length polymorphism • SSR, simple sequence repeat
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INTRODUCTION |
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TOPNOTESABSTRACTINTRODUCTIONMaterials and methodsResults and discussionREFERENCES |
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Using a mapping population derived from an interspecific G. max x G. soja cross, Shoemaker and Olson (1993) developed a molecular genetic linkage map that consisted of 25 linkage groups with about 365 RFLP, 11 RAPD (random amplified polymorphic DNA), three classical markers, and four isozyme loci. The current soybean classical marker map consists of 68 loci dispersed among 20 small linkage groups with a few loci each (Palmer and Shoemaker, 1998). A partial integration of the various marker types into a common linkage map was recently achieved by Shoemaker and Specht (1995). These authors used a soybean mapping population derived from a mating of near-isogenic lines of the cultivars Clark and Harosoy to create a linkage map that included 13 classical and 7 isozyme loci along with 110 RFLP and 8 RAPD loci. A set of anchoring RFLP loci that segregated in both the Clark x Harosoy and the G. max x G. soja mapping populations was used to identify linkage group homologies between the molecular and classical marker maps.
Two characteristics of RFLP markers in soybean tend to complicate the task of consolidating linkage maps from different mapping populations. First, only rarely have more than two alleles been identified at RFLP loci in soybean. Because these two alleles generally have asymmetric frequencies, e.g., p > 0.9, q < 0.1 (Keim et al., 1989; Keim et al., 1992), the likelihood that any two genotypes will be polymorphic at a particular RFLP locus is relatively low. This is particularly true when both parents of the mapping population come from adapted soybean germplasm pools (Apuya et al., 1988; Lark et al., 1993). For example, Muehlbauer et al. (1991) also observed that only one-third of the available RFLP probes tested on donor parent, near-isogenic line, or recurrent parent triplets were actually polymorphic between the two parents. Similarly, Shoemaker and Specht (1995) reported that only 118 of 365 RFLP markers polymorphic in the G. max x G. soja population were segregating in the Clark x Harosoy mapping population. Thus, a polymorphic fragment mapped in one population may not be segregating in another. A second factor that complicates the use of RFLP markers in soybean is the detection of multiple DNA fragments (i.e., multiple loci) with most probes. This may be the result of the tetraploid origin of soybean (Hymowitz and Singh, 1987). Because one fragment in a multiple banding pattern may segregate in one population and a different or an additional fragment in another, one must define a RFLP locus not only by the probe and restriction enzyme being used, but also by the molecular weight of the segregating fragments(s). Up to 19 independent loci have been mapped by specific RFLP probes (Mansur et al., 1996). The multiplicity of RFLP loci can make RFLP linkage maps ambiguous with respect to RFLP locus identity, and often precludes the use of such loci for the evaluation of linkage group homology among different maps.
One possible solution to this complexity is the use of single locus DNA markers with multiple alleles. In soybean, the highly polymorphic nature (i.e., multi-allelism) of simple sequence repeat (SSR) or microsatellite DNA markers is quite clear as shown by initial work of Akkaya et al. (1992) and Morgante and Olivieri (1993). Subsequent reports (Rongwen et al., 1995; Maughan et al., 1995; Powell et al., 1996; Diwan and Cregan, 1997) have described highly polymorphic microsatellite loci with as many as 26 alleles. A high level of allelic diversity makes it likely that a particular SSR locus will be polymorphic in many of the two-parent populations derived from the hybridization of adapted soybean genotypes. Multiple allele molecular markers are much more useful than dimorphic markers when tracking the fate of genomic segments in multi-parent pedigrees and in multi-parent mated populations. Another virtue of SSR markers is their simplicity. In the development of these markers, care is taken to select polymerase chain reaction (PCR) primers that produce one amplification product in an inbred soybean genotype (Cregan et al., 1994). Primer sets producing more than one product are discarded. Thus, the difficulty of the genetic interpretation of multiple banding patterns is eliminated. In addition, as reported in humans and other mammalian species, microsatellite loci in soybean seem to distribute fairly randomly throughout the genome, with minimal evidence of clustering (Akkaya et al., 1995).
While extensive SSR or microsatellite DNA based maps are now available and used by human and other mammalian geneticists (Hudson et al., 1995; Dietrich et al., 1994; Archibald et al., 1995), relatively smaller numbers of SSR markers have been developed and integrated into existing plant linkage maps. Bell and Ecker (1994) reported the assignment of 30 microsatellite loci to the linkage map of Arabidopsis. Akkaya et al. (1995) integrated 40 SSR markers into a soybean linkage map and Mansur et al. (1996) added another 22 loci shortly thereafter. Senior et al. (1996) mapped 42 distinct GenBank-derived SSR loci in maize (Zea mays L.). More recently, Sharon et al. (1997) mapped 25 SSR loci on a genetic linkage map of avocado (persea americana Mill.). Development and mapping of wheat (Triticum aestivum L.) microsatellite loci on a similar scale has been reported (Röder et al., 1995; and Bryan et al., 1997).
The availability of a molecular genetic map saturated with highly informative, PCR-based, single-locus, multiple-allele molecular markers would be of great utility to a wide range of soybean researchers. Therefore, the first objective of the work reported here was to develop a set of SSR markers that could be used under a standard set of amplification conditions. The second objective was to map those SSR loci in three existing mapping populations to attempt the alignment of homologous linkage groups and the identification of 20 consensus linkage groups corresponding to the 20 pairs of soybean chromosomes.
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Materials and methods |
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TOPNOTESABSTRACTINTRODUCTIONMaterials and methodsResults and discussionREFERENCES |
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Each selected primer pair was initially tested in two different PCR amplification reactions. The first used as template the plasmid containing the Williams soybean genomic insert from which the sequence data for the selection of the primer set was selected. The second reaction used genomic DNA of Williams soybean as template. When both amplification reactions yielded a single product of predicted size, the primer set was further tested on a set of 10 soybean genotypes in order to obtain an estimate of the level of SSR length polymorphism associated with each locus. The genotypes included the cultivars Clark (Maturity Group [MG] IV), Harosoy (MG II), Jackson (MG VII), Williams (MG III), Amsoy (MG II), Archer (MG I), Fiskeby V (MG 000), Minsoy (MG 0), Noir 1 (MG 0), and Tokyo (MG VII). Primer sets that produced multiple products in any of the 10 genotypes were discarded. The size of the alleles (i.e., base pair number in the PCR products) produced by each genotype was determined with DNA sequencing gels with a modified sequencing gel formulation (6% [w/v] acrylamide:bis-acrylamide [19:1], 5.6 M ultra-pure urea, and 30% [v/v] formamide in TBE buffer) as described by Cregan and Quigley (1997). These allele size estimates were used to calculate the gene diversity or informativeness of each SSR locus. This calculation was described by Anderson et al. (1993).
Soybean Mapping Populations and Genetic Marker Data
USDA/Iowa State University Population.
This is an F2-derived mapping population from the interspecific cross of the G. max Breeding Line A81-356022 and G. soja (wild soybean ) PI 468.916. This population currently consists of 59 F2 plant derivatives and has been described in detail (Shoemaker and Specht, 1995; Shoemaker and Olson, 1993). The extraction of DNA and the mapping of RFLP loci in this population was previously described by Keim et al. (1988). DNA isolation from the G. max x G. soja population and RFLP probe hybridization procedures were the same as previously described (Diers et al., 1992a, c; Keim et al., 1990).
University of Utah Recombinant Inbred Line (RIL) Population.
Originating from a cross of Minsoy x Noir I, this population consists of 240 F7-derived RIL and has been described previously (Mansur et al., 1996). The DNA isolation and RFLP analysis were described by Lark et al. (1993) and Mansur et al. (1996).
University of Nebraska Population.
This is an F2 -derived population from the cross of near isogenic lines (NILs) of the important cultivars Clark and Harosoy. Each of the Clark and Harosoy NILs used as the parents carries a number of pigmentation and/or morphological mutants thereby allowing the mapping of these classical genetic loci along with molecular loci. The population consists of derivatives of 57 F2 plants and previously was described by Shoemaker and Specht (1995). The isolation of DNA and the RFLP analysis was described earlier (Shoemaker and Specht, 1995).
The segregation of alleles at each SSR locus in each of the above populations was determined by amplifying template DNA from each RIL or F2 derivative followed by electrophoretic separation of the resulting products on DNA sequencing gels. These procedures are described in Cregan and Quigley (1997).
Genetic Mapping
MAPMAKER 3.0b (Lander et al., 1987; Lincoln and Lander, 1993) was used to group and order genetic loci within each of the three mapping populations. Marker loci were first grouped at LOD 5.0 and then ordered by repetitive use of the Ripple command of MAPMAKER with a window size of 6. The Kosambi centimorgan function was used with error detection on. The error detection probability level was set at 5% in the case of the USDA/Iowa State G. max x G. soja and University of Nebraska Clark x Harosoy populations and at 1% in the analysis of the Univ. of Utah Minsoy x Noir 1 population. The absence of heterozygous genotypes in the latter population made genotypic classification less prone to error.
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Results and discussion |
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TOPNOTESABSTRACTINTRODUCTIONMaterials and methodsResults and discussionREFERENCES |
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The total numbers of markers (SSR, RFLP, RAPD, AFLP, isozyme, and classical) mapped in any one of the three populations ranged from 523 to 1004 (Table 1) . The total number of unique loci in the three maps combined totaled 1423, which included 606 SSR, 689 RFLP, 79 RAPD, 11 AFLP, 10 isozyme, and 26 classical loci. Before the inclusion of the SSRs in the G. max x G. soja map, the last reported MAPMAKER analysis indicated a total of 25 linkage groups (Shoemaker and Specht, 1995). This number was reduced to 23 by the addition of the SSR loci. The addition of well over 300 new SSR loci more than doubled the number of markers on the Univ. of Utah map and reduced the number of linkage groups from 36 to 22. By aligning the linkage groups in these two maps with the linkage groups of the Clark x Harosoy population, on the basis of the presence of common SSR loci, a total of 20 linkage groups are now readily discernable (Fig. 1) . For example, the two separate G. max x G. soja linkage groups D1a and Q are very likely one linkage group given SSR loci they have in common with D1a+Q-U08 of the Minsoy x Noir 1 map (Fig. 1, Panel MLG D1a+Q). Indeed if the LOD threshold is reduced to 2.7, MAPMAKER joins D1a and Q. On the basis of a similar rationale, G. max x G. soja groups W and D1b (Fig. 1, Panel MLG D1b+W) were combined on the basis of alignment with the homologous Clark x Harosoy linkage groups. Similarly, Utah linkage groups U18 and U02 were joined (Fig. 1, Panel MLG E), as were U13a and U13b (Fig. 1, Panel MLG F) on the basis of alignments with linkage groups E-ISU and F-ISU, respectively, of the G. max x G. soja map. The alignment of linkage groups based upon the presence of common SSR loci across the three maps resulted in the establishment of 20 consensus linkage groups (Fig. 1). In the case of both the G. max x G. soja and the Clark x Harosoy maps, only one (Y-ISU) and two (CH54 and CH24) small linkage groups, respectively, could not be aligned with any of the 20 consensus groups. The fact that these remaining small groups contain no SSR loci (Fig. 1, Panel MLG Y + Unlinked) made their alignment particularly difficult.
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Positioning of SSR Markers within Linkage Groups
In previous work, Akkaya et al. (1995) indicated that the first 40 SSR loci mapped in soybean appeared to distribute randomly throughout the genome. However, these authors also observed one cluster of five SSR loci that would have a low probability of occurring by random chance if SSR loci were truly distributed randomly in the soybean genome. With the mapping of as many as 500 SSR loci in a single mapping population, it is still not clear if there is substantial clustering of SSRs. Portions of many linkage groups contain groups of SSR loci and similar groups of RFLP loci. Linkage group D1a (Fig. 1, Panel MLG D1a+Q) demonstrates this clustering especially in the case of the G. max x G. soja map. At the top of the linkage group is a group of 16 markers, 13 of which are RFLPs. Adjacent to these are groups of uninterrupted stretches of 11 and 18 SSR markers separated by a group of eight loci, seven of which are RFLPs. Linkage group E (Fig. 1, Panel MLG E) in the G. max x G. soja map has a very long stretch of mostly RFLP loci in which there is only one SSR (Sat_124). Flanking this region are two clusters of markers that are mostly SSRs. A similar cluster of RFLP loci occurs in linkage group G (Fig. 1, Panel MLG G) of the G. max x G. soja map between SSR loci Satt501 and Satt505. While this type of clustering does not occur in every linkage group, it is a fairly frequent occurrence.
As indicated above, it is not clear from observing the maps presented in Fig. 1 that clustering of markers is the result of the clustering of RFLP loci or of SSR loci or both. In the development of libraries from which SSR-containing genomic clones were selected, many different restriction enzymes and combinations of restriction enzymes were used to create genomic fragments in the 500 to 700 bp range (Cregan et al., 1994; Cregan et al., 1999). This was done to avoid duplicate clones and to sample different portions of the soybean genome. In contrast, one of the techniques in the development of the RFLP probes used here was the development and isolation of probes from PstI libraries (Keim and Shoemaker, 1988). The rationale for this approach is that methylation-sensitive enzymes such as PstI preferentially cut non-methylated regions which are presumed to contain less repetitive DNA. However, this approach may have produced sets of restriction fragments that were not randomly distributed throughout the genome. Thus, it is possible that clustering of markers we have noted here is more closely associated with RFLP than with SSR loci.
Regardless of the reason for the possible clustering of loci, the application of the set of SSR loci described here will be detrimentally affected by large intervals or gaps in which no markers are present. For example, it is obvious that quantitative trait loci (QTL) in genomic regions lacking markers will be undetectable. A total of 36 intervals of greater than 20 cM that do not contain an SSR locus are present in the Minsoy x Noir 1 map (Table 3) . Each of the 20 consensus linkage groups contains at least one such gap suggesting that such intervals are not confined to a specific subset of chromosomes. In many instances such as those in linkage group E-U18+02 and G-U05, clusters of RFLP loci are present in the intervals lacking SSRs. In an effort to place SSR loci in regions with only RFLP markers, we are attempting to use bacterial artificial chromosome (BAC) clones for targeted SSR marker development as described by Cregan et al. (1999). These BAC clones are being selected via hybridization with the RFLP probes that map to regions devoid of SSR loci. However, in 10 of the 36 intervals of greater than 20 cM listed in Table 3, no RFLP loci are present in the gaps flanked by the SSR loci. It may be difficult to develop markers that map to these intervals.
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The Integration of Classical Genetic Loci into the Soybean Map
Palmer and Shoemaker (1998) defined 20 classical linkage groups (CLG) that contain a total of 68 of the more than 250 classical pigmentation, morphological, isozyme, disease resistance, etc. genes that have been named. The inclusion of the Univ. of Nebraska population derived from the cross of NILs of Clark x Harosoy in this study is important because it segregated for 14 classical loci (Shoemaker and Specht, 1995). These classical loci and other classical loci linked to them are now integrated with molecular markers to form a comprehensive genetic map. Some of the linkage data were reported earlier (Shoemaker and Specht, 1995), but it is reproduced here for completeness. As shown in Fig. 1 and listed in Table 2, all but one CLG (CLG06, Fig. 1, Panel MLG Y+Unlinked) can now be associated with a molecular linkage group.
The integration of the classical, RFLP, and SSR markers into one comprehensive linkage map is a powerful tool for the advancement of soybean genetics. For those studies that have been based upon RFLP marker technology it provides an opportunity to convert to a PCR-based marker system. Because SSR markers are so commonly used by human geneticists, technologies for their use are readily available. For example, automated allele sizing is being applied to plant genetic studies (Kresovich et al., 1995; Diwan and Cregan, 1997). In the future, high throughput systems will provide the opportunity for genetic analysis that is appropriate to the needs of large scale plant improvement programs that routinely analyze thousands or hundreds of thousands of segregating progeny.
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ACKNOWLEDGMENTS |
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NOTES |
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TOPNOTESABSTRACTINTRODUCTIONMaterials and methodsResults and discussionREFERENCES |
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Received for publication January 12, 1998.
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REFERENCES |
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TOPNOTESABSTRACTINTRODUCTIONMaterials and methodsResults and discussionREFERENCES |
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D. M. Nichols, K. D. Glover, S. R. Carlson, J. E. Specht, and B. W. Diers Fine Mapping of a Seed Protein QTL on Soybean Linkage Group I and Its Correlated Effects on Agronomic Traits Crop Sci., February 24, 2006; 46(2): 834 - 839. [Abstract] [Full Text] [PDF]
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E. A. Kabelka, S. R. Carlson, and B. W. Diers Glycine soja PI 468916 SCN Resistance Loci's Associated Effects on Soybean Seed Yield and Other Agronomic Traits Crop Sci., February 1, 2006; 46(2): 622 - 629. [Abstract] [Full Text] [PDF]
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D. R. Walker, A. M. Scaboo, V. R. Pantalone, J. R. Wilcox, and H. R. Boerma Genetic Mapping of Loci Associated with Seed Phytic Acid Content in CX1834-1-2 Soybean Crop Sci., January 24, 2006; 46(1): 390 - 397. [Abstract] [Full Text] [PDF]
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C. D. Messina, J. W. Jones, K. J. Boote, and C. E. Vallejos A Gene-Based Model to Simulate Soybean Development and Yield Responses to Environment Crop Sci., January 24, 2006; 46(1): 456 - 466. [Abstract] [Full Text] [PDF]
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T. R. Stefaniak, D. L. Hyten, V. R. Pantalone, A. Klarer, and T. W. Pfeiffer Soybean Cultivars Resulted from More Recombination Events Than Unselected Lines in the Same Population Crop Sci., December 2, 2005; 46(1): 43 - 51. [Abstract] [Full Text] [PDF]
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S. G. Gordon, S. K. St. Martin, and A. E. Dorrance Rps8 Maps to a Resistance Gene Rich Region on Soybean Molecular Linkage Group F Crop Sci., December 2, 2005; 46(1): 168 - 173. [Abstract] [Full Text] [PDF]
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D. V. Charlson, T. B. Bailey, S. R. Cianzio, and R. C. Shoemaker Molecular Marker Satt481 is Associated with Iron-Deficiency Chlorosis Resistance in a Soybean Breeding Population Crop Sci., October 27, 2005; 45(6): 2394 - 2399. [Abstract] [Full Text] [PDF]
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V. S. Primomo, V. Poysa, G. R. Ablett, C.-J. Jackson, M. Gijzen, and I. Rajcan Mapping QTL for Individual and Total Isoflavone Content in Soybean Seeds Crop Sci., October 27, 2005; 45(6): 2454 - 2464. [Abstract] [Full Text] [PDF]
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E. A. Kabelka, S. R. Carlson, and B. W. Diers Localization of Two Loci that Confer Resistance to Soybean Cyst Nematode from Glycine soja PI 468916 Crop Sci., October 27, 2005; 45(6): 2473 - 2481. [Abstract] [Full Text] [PDF]
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 G. Zabala and L. O. Vodkin The wp Mutation of Glycine max Carries a Gene-Fragment-Rich Transposon of the CACTA Superfamily PLANT CELL, October 1, 2005; 17(10): 2619 - 2632. [Abstract] [Full Text] [PDF]
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D. Sandhu, K. G. Schallock, N. Rivera-Velez, P. Lundeen, S. Cianzio, and M. K. Bhattacharyya Soybean Phytophthora Resistance Gene Rps8 Maps Closely to the Rps3 Region J. Hered., September 1, 2005; 96(5): 536 - 541. [Abstract] [Full Text] [PDF]
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K. Van, E.-Y. Hwang, M. Y. Kim, H. J. Park, S.-H. Lee, and P. B. Cregan Discovery of SNPs in Soybean Genotypes Frequently Used as the Parents of Mapping Populations in the United States and Korea J. Hered., September 1, 2005; 96(5): 529 - 535. [Abstract] [Full Text] [PDF]
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J. L. Alt, W. R. Fehr, G. A. Welke, and D. Sandhu Phenotypic and Molecular Analysis of Oleate Content in the Mutant Soybean Line M23 Crop Sci., August 26, 2005; 45(5): 1997 - 2000. [Abstract] [Full Text] [PDF]
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D. R. Panthee, V. R. Pantalone, D. R. West, A. M. Saxton, and C. E. Sams Quantitative Trait Loci for Seed Protein and Oil Concentration, and Seed Size in Soybean Crop Sci., August 26, 2005; 45(5): 2015 - 2022. [Abstract] [Full Text] [PDF]
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K. Komatsu, S. Okuda, M. Takahashi, R. Matsunaga, and Y. Nakazawa QTL Mapping of Antibiosis Resistance to Common Cutworm (Spodoptera litura Fabricius) in Soybean Crop Sci., August 26, 2005; 45(5): 2044 - 2048. [Abstract] [Full Text] [PDF]
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A. H. A. Assar, R. Uptmoor, A. A. Abdelmula, M. Salih, F. Ordon, and W. Friedt Genetic Variation in Sorghum Germplasm from Sudan, ICRISAT, and USA Assessed by Simple Sequence Repeats (SSRs) Crop Sci., June 24, 2005; 45(4): 1636 - 1644. [Abstract] [Full Text] [PDF]
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M. E. Patzoldt, C. R. Grau, P. A. Stephens, N. C. Kurtzweil, S. R. Carlson, and B. W. Diers Localization of a Quantitative Trait Locus Providing Brown Stem Rot Resistance in the Soybean Cultivar Bell Crop Sci., May 27, 2005; 45(4): 1241 - 1248. [Abstract] [Full Text] [PDF]
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R. Takahashi, E. R. Benitez, H. Funatsuki, and S. Ohnishi Soybean Maturity and Pubescence Color Genes Improve Chilling Tolerance Crop Sci., May 27, 2005; 45(4): 1387 - 1393. [Abstract] [Full Text] [PDF]
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G. Zhao, G. R. Ablett, T. R. Anderson, I. Rajcan, and A. W. Schaafsma Inheritance and Genetic Mapping of Resistance to Rhizoctonia Root and Hypocotyl Rot in Soybean Crop Sci., May 27, 2005; 45(4): 1441 - 1447. [Abstract] [Full Text] [PDF]
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M. E. Patzoldt, S. R. Carlson, and B. W. Diers Characterization of Resistance to Brown Stem Rot of Soybean in Five Accessions from Central China Crop Sci., May 6, 2005; 45(3): 1092 - 1095. [Abstract] [Full Text] [PDF]
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M. Ulloa, S. Saha, J. N. Jenkins, W. R. Meredith Jr., J. C. McCarty Jr., and D. M. Stelly Chromosomal Assignment of RFLP Linkage Groups Harboring Important QTLs on an Intraspecific Cotton (Gossypium hirsutum L.) Joinmap J. Hered., March 1, 2005; 96(2): 132 - 144. [Abstract] [Full Text] [PDF]
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E. R. Benitez, H. Funatsuki, Y. Kaneko, Y. Matsuzawa, S. W. Bang, and R. Takahashi Soybean Maturity Gene Effects on Seed Coat Pigmentation and Cracking in Response to Low Temperatures Crop Sci., November 1, 2004; 44(6): 2038 - 2042. [Abstract] [Full Text] [PDF]
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S. K. Stombaugh, J. H. Orf, H. G. Jung, K. Chase, K. G. Lark, and D. A. Somers Quantitative Trait Loci Associated with Cell Wall Polysaccharides in Soybean Seed Crop Sci., November 1, 2004; 44(6): 2101 - 2106. [Abstract] [Full Text] [PDF]
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V. C. Concibido, B. W. Diers, and P. R. Arelli A Decade of QTL Mapping for Cyst Nematode Resistance in Soybean Crop Sci., July 1, 2004; 44(4): 1121 - 1131. [Abstract] [Full Text] [PDF]
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V. A. Fasoula, D. K. Harris, and H. R. Boerma Validation and Designation of Quantitative Trait Loci for Seed Protein, Seed Oil, and Seed Weight from Two Soybean Populations Crop Sci., July 1, 2004; 44(4): 1218 - 1225. [Abstract] [Full Text] [PDF]
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 Z. W. Luo, R. M. Zhang, and M. J. Kearsey Theoretical basis for genetic linkage analysis in autotetraploid species PNAS, May 4, 2004; 101(18): 7040 - 7045. [Abstract] [Full Text] [PDF]
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B.-K. Ha, J. B. Bennett, R. S. Hussey, S. L. Finnerty, and H. R. Boerma Pedigree Analysis of a Major QTL Conditioning Soybean Resistance to Southern Root-Knot Nematode Crop Sci., May 1, 2004; 44(3): 758 - 763. [Abstract] [Full Text] [PDF]
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E. A. Kabelka, B. W. Diers, W. R. Fehr, A. R. LeRoy, I. C. Baianu, T. You, D. J. Neece, and R. L. Nelson Putative Alleles for Increased Yield from Soybean Plant Introductions Crop Sci., May 1, 2004; 44(3): 784 - 791. [Abstract] [Full Text] [PDF]
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K. D. Glover, D. Wang, P. R. Arelli, S. R. Carlson, S. R. Cianzio, and B. W. Diers Near Isogenic Lines Confirm a Soybean Cyst Nematode Resistance Gene from PI 88788 on Linkage Group J Crop Sci., May 1, 2004; 44(3): 936 - 941. [Abstract] [Full Text] [PDF]
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M. D. Smalley, W. R. Fehr, S. R. Cianzio, F. Han, S. A. Sebastian, and L. G. Streit Quantitative Trait Loci for Soybean Seed Yield in Elite and Plant Introduction Germplasm Crop Sci., March 1, 2004; 44(2): 436 - 442. [Abstract] [Full Text] [PDF]
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J.M. Narvel, T.E. Carter Jr., L.R. Jakkula, J. Alvernaz, M.A. Bailey, M.A.R. Mian, S.H. Lee, G.J. Lee, and H.R. Boerma Registration of NC113 Soybean Mapping Population Crop Sci., March 1, 2004; 44(2): 704 - 706. [Full Text]
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 C. Wu, S. Sun, P. Nimmakayala, F. A. Santos, K. Meksem, R. Springman, K. Ding, D. A. Lightfoot, and H.-B. Zhang A BAC- and BIBAC-Based Physical Map of the Soybean Genome Genome Res., February 1, 2004; 14(2): 319 - 326. [Abstract] [Full Text] [PDF]
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T. E. Devine The Pd2 and Lf2 Loci Define Soybean Linkage Group 16 Crop Sci., November 1, 2003; 43(6): 2028 - 2030. [Abstract] [Full Text] [PDF]
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N. Reyna, B. Cornelious, J. G. Shannon, and C. H. Sneller Evaluation of a QTL for Waterlogging Tolerance in Southern Soybean Germplasm Crop Sci., November 1, 2003; 43(6): 2077 - 2082. [Abstract] [Full Text] [PDF]
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K. K. Kato and R. G. Palmer Molecular Mapping of the Male-Sterile, Female-Sterile Mutant Gene (st8) in Soybean J. Hered., September 1, 2003; 94(5): 425 - 428. [Abstract] [Full Text] [PDF]
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A. A. Mahama and R. G. Palmer Translocation Breakpoints in Soybean Classical Genetic Linkage Groups 6 and 8 Crop Sci., September 1, 2003; 43(5): 1602 - 1609. [Abstract] [Full Text] [PDF]
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K. D. Burnham, A. E. Dorrance, T. T. VanToai, and S. K. St. Martin Quantitative Trait Loci for Partial Resistance to Phytophthora sojae in Soybean Crop Sci., September 1, 2003; 43(5): 1610 - 1617. [Abstract] [Full Text] [PDF]
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V. A. Fasoula, D. K. Harris, M. A. Bailey, D. V. Phillips, and H. R. Boerma Identification, Mapping, and Confirmation of a Soybean Gene for Bud Blight Resistance Crop Sci., September 1, 2003; 43(5): 1754 - 1759. [Abstract] [Full Text] [PDF]
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J. J. Zou, R. J. Singh, and T. Hymowitz Association of the Yellow Leaf (y10) Mutant to Soybean Chromosome 3 J. Hered., July 1, 2003; 94(4): 352 - 355. [Abstract] [Full Text] [PDF]
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J. Abe, D. Xu, A. Miyano, K. Komatsu, A. Kanazawa, and Y. Shimamoto Photoperiod-Insensitive Japanese Soybean Landraces Differ at Two Maturity Loci Crop Sci., July 1, 2003; 43(4): 1300 - 1304. [Abstract] [Full Text] [PDF]
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J. Chung, H. L. Babka, G. L. Graef, P. E. Staswick, D. J. Lee, P. B. Cregan, R. C. Shoemaker, and J. E. Specht The Seed Protein, Oil, and Yield QTL on Soybean Linkage Group I Crop Sci., May 1, 2003; 43(3): 1053 - 1067. [Abstract] [Full Text] [PDF]
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T. Tischner, L. Allphin, K. Chase, J. H. Orf, and K. G. Lark Genetics of Seed Abortion and Reproductive Traits in Soybean Crop Sci., March 1, 2003; 43(2): 464 - 473. [Abstract] [Full Text] [PDF]
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Y. L. Zhu, Q. J. Song, D. L. Hyten, C. P. Van Tassell, L. K. Matukumalli, D. R. Grimm, S. M. Hyatt, E. W. Fickus, N. D. Young, and P. B. Cregan Single-Nucleotide Polymorphisms in Soybean Genetics, March 1, 2003; 163(3): 1123 - 1134. [Abstract] [Full Text] [PDF]
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J. A. Hoeck, W. R. Fehr, R. C. Shoemaker, G. A. Welke, S. L. Johnson, and S. R. Cianzio Molecular Marker Analysis of Seed Size in Soybean Crop Sci., January 1, 2003; 43(1): 68 - 74. [Abstract] [Full Text] [PDF]
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K. D. Burnham, A. E. Dorrance, D. M. Francis, R. J. Fioritto, and S. K. St. Martin Rps8, A New Locus in Soybean for Resistance to Phytophthora sojae Crop Sci., January 1, 2003; 43(1): 101 - 105. [Abstract] [Full Text] [PDF]
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I. M. Tasma and R. C. Shoemaker Mapping Flowering Time Gene Homologs in Soybean and Their Association with Maturity (E) Loci Crop Sci., January 1, 2003; 43(1): 319 - 328. [Abstract] [Full Text] [PDF]
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A. A. Mahama, K. S. Lewers, and R. G. Palmer Genetic Linkage in Soybean: Classical Genetic Linkage Groups 6 and 8 Crop Sci., September 1, 2002; 42(5): 1459 - 1464. [Abstract] [Full Text] [PDF]
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 H. C. Karakaya, Y. Tang, P. B. Cregan, and H. T. Knap Molecular mapping of the fasciation mutation in soybean, Glycine max (Leguminosae) Am. J. Botany, April 1, 2002; 89(4): 559 - 565. [Abstract] [Full Text] [PDF]
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K. D. Burnham, D. M. Francis, A. E. Dorrance, R. J. Fioritto, and S. K. St. Martin Genetic Diversity Patterns among Phytophthora Resistant Soybean Plant Introductions Based on SSR Markers Crop Sci., March 1, 2002; 42(2): 338 - 343. [Abstract] [Full Text] [PDF]
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Z. Li, R. F. Wilson, W. E. Rayford, and H. R. Boerma Molecular Mapping Genes Conditioning Reduced Palmitic Acid Content in N87-2122-4 Soybean Crop Sci., March 1, 2002; 42(2): 373 - 378. [Abstract] [Full Text] [PDF]
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B. K. Cornelious and C. H. Sneller Yield and Molecular Diversity of Soybean Lines Derived from Crosses of Northern and Southern Elite Parents Crop Sci., March 1, 2002; 42(2): 642 - 647. [Abstract] [Full Text] [PDF]
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T. MacGregor, M. Bhattacharyya, B. Tyler, R. Bhat, A. F. Schmitthenner, and M. Gijzen Genetic and Physical Mapping of Avr1a in Phytophthora sojae Genetics, March 1, 2002; 160(3): 949 - 959. [Abstract] [Full Text] [PDF]
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S. C. Jeong, S. Kristipati, A. J. Hayes, P. J. Maughan, S. L. Noffsinger, I. Gunduz, G. R. Buss, and M. A. S. Maroof Genetic and Sequence Analysis of Markers Tightly Linked to the Soybean mosaic virus Resistance Gene, Rsv3 Crop Sci., January 1, 2002; 42(1): 265 - 270. [Abstract] [Full Text] [PDF]
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J. Yuan, V. N. Njiti, K. Meksem, M. J. Iqbal, K. Triwitayakorn, My. A. Kassem, G. T. Davis, M. E. Schmidt, and D. A. Lightfoot Quantitative trait loci in Two Soybean Recombinant Inbred Line Populations Segregating for Yield and Disease Resistance Crop Sci., January 1, 2002; 42(1): 271 - 277. [Abstract] [Full Text] [PDF]
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J. M. Narvel, D. R. Walker, B. G. Rector, J. N. All, W. A. Parrott, and H. R. Boerma A Retrospective DNA Marker Assessment of the Development of Insect Resistant Soybean Crop Sci., November 1, 2001; 41(6): 1931 - 1939. [Abstract] [Full Text] [PDF]
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C. Weng, K. Yu, T. R. Anderson, and V. Poysa Mapping Genes Conferring Resistance to Phytophthora Root Rot of Soybean, Rps1a and Rps7 J. Hered., September 1, 2001; 92(5): 442 - 446. [Abstract] [Full Text] [PDF]
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M. E. Gardner, T. Hymowitz, S. J. Xu, and G. L. Hartman Physical Map Location of the Rps1-k Allele in Soybean Crop Sci., September 1, 2001; 41(5): 1435 - 1438. [Abstract] [Full Text] [PDF]
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P. Yue, D. A. Sleper, and P. R. Arelli Mapping Resistance to Multiple Races of Heterodera glycines in Soybean PI 89772 Crop Sci., September 1, 2001; 41(5): 1589 - 1595. [Abstract] [Full Text] [PDF]
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A. Demirbas, B. G. Rector, D. G. Lohnes, R. J. Fioritto, G. L. Graef, P. B. Cregan, R. C. Shoemaker, and J. E. Specht Simple Sequence Repeat Markers Linked to the Soybean Rps Genes for Phytophthora Resistance Crop Sci., July 1, 2001; 41(4): 1220 - 1227. [Abstract] [Full Text] [PDF]
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T. T. VanToai, S. K. St. Martin, K. Chase, G. Boru, V. Schnipke, A. F. Schmitthenner, and K. G. Lark Identification of a QTL Associated with Tolerance of Soybean to Soil Waterlogging Crop Sci., July 1, 2001; 41(4): 1247 - 1252. [Abstract] [Full Text] [PDF]
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P. B. Cregan, K. P. Kollipara, S. J. Xu, R. J. Singh, S. E. Fogarty, and T. Hymowitz Primary Trisomics and SSR Markers as Tools to Associate Chromosomes with Linkage Groups in Soybean Crop Sci., July 1, 2001; 41(4): 1262 - 1267. [Abstract] [Full Text] [PDF]
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