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a Center for Excellence in Soybean Research, Teaching and Outreach, Department of Plant, Soil, and Agricultural Systems, Southern Illinois University at Carbondale, Carbondale, IL 62901-4415
b Department of Biological Sciences, Kean University, Union, NJ 07083
* Corresponding author (ga4082{at}siu.edu)
A genetic map of soybean [Glycine max (L.) Merr.] constructed with a recombinant inbred line (RIL) mapping population (Reg. no. MP-2, NSL 431663 MAP) from the cross Essex MAP (PI 636326 MAP) by Forrest MAP (PI 636325 MAP) has been used extensively worldwide. Essex was registered by Smith and Camper (1973) and Forest by Hartwig and Epps (1973). Since most morphological traits do not vary greatly, the RIL population has been used extensively worldwide to map genes underlying biochemical and physiological traits (Table 1). The genetic marker data encompass thousands of polymorphic markers and tens of thousands of sequence-tagged site (STS) that were collected at SIUC by Dr. Lightfoot's group (Table 2). Several genetic maps of ExF94 have been constructed (Chang et al., 1997; Iqbal et al., 2001; Kassem et al., 2004b) and will continue to be developed.
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Parents
Forrest is an F5derived line from the cross Dyer x Bragg and was developed by USDA-ARS and Tennessee Agricultural Experiment Stations (Hartwig and Epps, 1973). It is late group V in maturity and is characterized by determinate growth habit, white flowers, tawny pubescent, tan pods, yellow seed coats, and black hila. It was released for resistance to soybean cyst nematode (HG type 0; races 3, Heterodera glycines Ichinohe), root-knot nematode [Meloidogyne incognita (Kofoid & White) Chitwood], bacterial pustule [caused by Xanthomonas axonopodis pv. glycines (Nakano 1919) Vauterin, Hoste, Kersters & Swings 1995], wildfire [caused by Pseudomonas syringae pv. tabaci (Wolf & Foster 1917) Young, Dye & Wilkie 1978], target spot [caused by Corynespora cassiicola (Berk. & M.A. Curtis) C.T. Wei], and moderate resistance to Phytophthora root rot (caused by Phytophthora sojae Kaufmann & Gerdemann). Forrest was subsequently found to be resistant (disease index, DX < 1%) to foliar SDS (Hnetkovsky et al., 1996), root infection (infection severity, IS > 20%) by F. solani f. sp. glycines (Njiti et al., 1997a), and F. virguliforme and F. tucumaniae (Aoki et al., 2003). Forrest has excellent pod-shatter resistance (Hartwig and Epps, 1973).
Essex is an F7derived line from the cross Lee x S55-7075 and released by the Virginia Experiment Station (Smith and Camper, 1973). It was initially released for high seed yield and seed quality. It is maturity group V and is characterized by a determinate growth habit, purple flower, gray pubescent, yellow seed coat, and buff hila. It is resistant to bacterial pustule, several races of downy mildew [caused by Peronospora manshurica (Naumov) Syd. In Gaum.], frogeye leaf spot (caused by Cercospora sojina K. Hara), and purple seed strain disease [caused by Cercospora kikuchii (Mastsumoto & Tomoyasu) M.W. Gardner]. It is moderately resistant to Phytophthora root rot, as is Forrest. It has been found to be susceptible (DX > 10%) to SDS (Hnetkovsky et al., 1996), root infection (IS > 40%) by F. solani f. sp. glycines (Njiti et al., 1997a), and F. tucumanes (unpublished).
Development of the Population
The cross was made in 1983 at Southern Illinois University at Carbondale using seed obtained from the originating programs. About 4500 F2 plants were inbred to the F5 generation by modified single seed descent (Brim, 1966). In 1988, 500 F5 plants were harvested of which 150 were randomly selected, intentionally excluding a few undesirable extremes. Seeds of each plant were planted in a progeny row and in 1989, 100 recombinant inbred lines were retained for evaluation of morphological and agronomic traits, stress resistance, and 94 were used for construction of genetic map and QTL discovery. The population is now at the F5:15 generation. The RILs are identified as ExF1 to ExF100. ExF78 has been released as disease resistant germplasm under the name LS-G96 (Schmidt et al., 1999). Ninety-four of these 100 lines have been officially released by Southern Illinois University at Carbondale and all individuals interested in studying these lines may request seeds from Dr. Lightfoot at the above institution for small samples of seeds from the above institution for the next 5 yr. Large samples are available each biennium as the seed source is refreshed. For security lines are stored at National Center for Genetic Resources Preservation, 1111 S. Mason St. Fort Collins, CO 80521-4500.
Description of the Population
The population has a 7-d spread in maturity (105112 d after planting). Forty-six RILs produce white flowers, 52 produce purple flowers, and two are heterogeneous for flower color. Fifty-three RILs produce gray pubescence, 44 produce tawny pubescence, and three are heterogeneous for pubescence color. All RILs have a determinate growth habit. Plant height ranges from 71 to 106 cm for full season planting. All RILs produce seeds with yellow seed coat, 46 have black seed hila, 52 have buff seed hila, and two are heterogeneic for seed hila color. Seed yield ranges from 2.9 to 3.9 Mg ha1. The RILs show differential response to the SCN Hgtype 0 (race 3) isolate AP3, with 22 resistant [index of parasitism (IP) 10], 65 susceptible (IP > 10), and 13 heterogeneic (based on individual plant IP). The RILs show differential response to SDS, with disease incidence ranging from 5 to 100% and disease index (DX) ranging from 1 to 24%. The population segregates for resistance to frogeye leaf spot, Phytophthora root rot, Mn toxicity, salt tolerance, water deficit tolerance, seed isoflavone (daidzein, genistein, and glycitein) content, and trigonelline content. The population may have the potential to segregate for protein, oil, and carbohydrate content and quality. The population may segregate for phytate and other dietary micronutrients and anti-nutritional factors.
Genetic and Physical Maps
The ExF RIL genetic map currently contains 368 linked markers, but 231 are high quality microsatellite markers that have been scored repeatedly (Kassem et al., 2004b). The average distance between markers is 17 cM, and there are about 23 markers per linkage group and 79 markers unlinked. The polymorphic markers include 90 RAPD markers, 27 RFLP markers, 20 AFLP markers, and 231 microsatellite markers. All are publicly available on the web at Soybase http://soybase.agron.iastate.edu/ (verified 7 March 2005), on the physical map at http://bioinformatics.siu.edu (verified 7 March 2005) and their sequences are deposited in Genbank at http://www.ncbi.nlm.nih.gov/ (verified 7 March 2005).
Two soybean large-insert libraries have been constructed from Forrest in the pCLD04541 (V41) binary vector after partial digestion of genomic high-molecular-weight DNA with BamHI or HindIII (Meksem et al., 2000). The libraries contain 76800 clones with an average insert size of 125 kbp, and therefore represent 9.5-fold haploid genome equivalents. Another soybean BAC library was constructed from Forrest in pBELOBACII vector after digestion with EcoRI. The library contains 38400 clones with inserts of 153 kbp representing 5.8 haploid genome equivalents (Wu et al., 2004a, 2004b). These BAC libraries were used to generate a physical map containing anchored microsatellite markers. Updates to the physical map, the BAC clones, gene fingerprints, and the libraries are publicly available through request at http://bioinformatics.siu.edu; verified 25 March 2005.
QTL Mapped in the ExF RIL Population
Twenty-five published QTL among six traits identified in this population include six QTL for resistance to SDS (Hnetkovsky et al., 1996; Chang et al., 1996, 1997; Iqbal et al., 2001), three QTL for high yield in Southern Illinois (Yuan et al., 2002), eight QTL for seed isoflavone content (Kassem et al., 2004a), two QTL for trigonelline content and water deficit tolerance (Cho et al., 2002), four QTL for resistance to manganese toxicity (Kassem et al., 2004b), and two for bigenic resistance to SCN (Meksem et al., 2001c). Among these QTL, we have verified in a second population the positions of four of six SDS resistance QTL (Njiti et al., 1998), one of two yield QTL (Yuan et al., 2002), and both SCN resistance genes (Webb et al., 1995; Meksem et al., 1999). The remaining QTL have not been tested to verify to date.
ACKNOWLEDGMENTS
The authors acknowledge diverse contributions toward the researches reported in this registration notice: For funding from 19912004 the ISPOB, USB, IMBA, NSF, and NCSRB. For assistance Dr. O. Myers Jr., Dr. M.E. Schmidt and all members of the SIUC field teams from 1983 to 2003. This material is based on work supported by the National Science Foundation under Grant No. 9872635. Any opinions, findings, and conclusions or recommendations expressed in this material are those of the author(s) and do not necessarily reflect the views of the National Science Foundation.
NOTES
Accepted for publication December 31, 2004.
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