Capsicum rootstock

Abstract

The present invention relates to a plant from the Capsicum family suitable for use as rootstock, capable of being obtained from the initial crossing of a Capsicum plant, preferably a C. annuum plant, with a Capsicum baccatum plant used as female parent, optionally followed by one or more additional crossing(s) with a Capsicum baccatum plant. Another subject of the present invention is a grafted plant constituted of a rootstock according to the invention and a graft. A further subject of the present invention is a seed, a cell or a part of a rootstock plant according to the invention. A final subject of the present invention is the use of a plant according to the invention as rootstock and a process for obtaining a grafted plant.

Claims

1. A Capsicum baccatum plant suitable for use as a rootstock, obtained from the initial crossing of a Capsicum annuum plant, used as the male parent, with a Capsicum baccatum plant used as the female parent, wherein the initial crossing is followed by at least one additional crossing with a Capsicum baccatum plant, and wherein said Capsicum baccatum plant suitable for use as a rootstock has at a given crop stage, a root volume, a root mass or a hypocotyl height that is 10% greater than those of the TECNICO variety at the same stage, cultivated under identical conditions.

2. A Capsicum baccatum plant suitable for use as a rootstock, obtained from the initial crossing of a Capsicum annuum plant, with a first Capsicum baccatum plant, wherein the initial crossing is followed by at least one additional crossing with a Capsicum baccatum plant, and wherein said Capsicum baccatum plant suitable for use as a rootstock has at a given crop stage, a root volume, a root mass or a hypocotyl height that is 10% greater than those of the TECNICO variety at the same stage, cultivated under identical conditions.

3. A plant according to claim 1, wherein the Capsicum baccatum plant used in said at least one additional crossing is identical to the Capsicum baccatum plant used in the initial crossing.

4. A plant according to claim 1, wherein the Capsicum baccatum plant used in said at least one additional crossing is different from the Capsicum baccatum plant used in the initial crossing.

5. A plant according to claim 1, wherein said Capsicum annuum plant comprises at least one trait of agronomic interest, and wherein said at least one crossing is followed by a step of selecting plants comprising said trait of agronomic interest.

6. A plant according to claim 5, wherein said trait of agronomic interest is resistance to a pathogen.

7. A plant according to claim 6, wherein said pathogen is Ralstonia solanacearum, Rhizoctonia solani, Pythium spp., Fusarium oxysporum, Phytophthora capsici, Sclerotium rolfsii, Verticillium albo-atrum, Verticillium dahliae, Meloidogyne incognita, Meloidogyne arenaria, Meloidogyne javanica, Meloidogyne hapla or Meloidogyne enterolobii; or a PMMV, TMV, TSWV or PVY virus.

8. A plant according to claim 1, wherein said Capsicum baccatum plant used in said crossing(s) comprises at least one trait of agronomic interest, and wherein said at least one crossing is followed by a step of selecting plants comprising said trait of agronomic interest.

9. A plant according to claim 5, wherein the selection of plants comprising said trait of agronomic interest is performed by means of a phenotypic selection or by the use of molecular markers.

10. A Capsicum baccatum plant according to claim 1, further comprising intermediate resistance to Phytophthora capsici and intermediate resistance to Meloidogynes wherein said resistances are traits derived from Capsicum annuum.

11. A seed of a plant according to claim 1.

12. A cell of a plant according to claim 1.

13. A part of a plant according to claim 1.

14. A method of obtaining a grafted plant comprising using a Capsicum baccatum plant according to claim 1 as a rootstock and obtaining the grafted plant from said Capsicum plant.

15. The method of claim 14, further comprising grafting a graft on to the Capsicum baccatum plant according to claim 1.

16. The method of claim 15, wherein said graft is a plant of the Capsicum genus.

17. A method for obtaining a Capsicum baccatum plant suitable for use as a rootstock according to claim 1, or a part, cell or seed thereof, said method comprising the initial crossing of a Capsicum annuum plant used as the male parent, with a Capsicum baccatum plant used as the female parent, wherein the initial crossing is followed by at least one additional crossing with a Capsicum baccatum plant, and recovering said plant or part, cell or seed thereof, wherein said Capsicum baccatum plant suitable for use as a rootstock has at a given crop stage, a root volume, a root mass or a hypocotyl height that is 10% greater than those of the TECNICO variety at the same stage, cultivated under identical conditions.

Description

FIGURES

(1) FIG. 1Capsicum flowers

(2) ACapsicum annuum flower (origin: Central America, root weakness, resistance to Phytophthora and nematodes)

(3) BCapsicum baccatum flower (origin: Amazon Basin, root strength (R asphyxiation), possible resistance to Verticillium and Ralstonia)

(4) CResult of hybridization between C. annuum and C. baccatum. Crossings between C. annuum and about one hundred accessions of C. baccatum were produced, twenty hybrids were harvested and seven hybrids were verified (including by marking). On the left: flower of the C. annuum male parent (hairy plant and white flower); on the right: flower of the C. baccatum female parent (glabrous plant and flower with spot); in the centre, flower of the F1 hybrid obtained by crossing two parents (slightly hairy plant and flower with light spots).

(5) FIG. 2Root volume and architecture of non-grafted plants

(6) AC. baccatum*PBC1405 hybrid: root volume=5

(7) BC. annuum TECNICO control: root volume=2

(8) CC. annuum SNOOKER control: root volume=2

(9) DC. annuum TRESOR control: root volume=1

(10) FIG. 3A-3ERoot volume and architecture of grafted plants (trial 2)

(11) 3ASNOOKER rootstock: root volume=2; root architecture=2

(12) 3BTRESOR rootstock: root volume=2; root architecture=1

(13) 3CTECNICO rootstock: root volume=3; root architecture=2

(14) 3DROBUSTO rootstock: root volume=4; root architecture=2

(15) 3ERootstock obtained from crossing PI 413669 01 SD*PBC1405: root volume=5; root architecture=5.

EXPERIMENTAL PART

1. Selection and Use of C. baccatum Plants as Rootstocks

(16) The inventors performed three sequential steps.

(17) First, they identified, outside of the C. annuum species, within the C. baccatum species, genetic resources making it possible to improve the productivity of the current rootstocks, in particular according to their significant root vigour.

(18) They then confirmed that these resources had valuable behaviours with regard to grafting sites (good germination and sufficient hypocotyl length).

(19) The inventors finally formed a grafted plant using a C. baccatum rootstock, in particular hybrid rootstocks utilizing the heterosis between different C. baccatum accessions and having valuable characteristics in the nursery (good germination and hypocotyl length) and in cultivation (gain in vigour and productivity).

(20) It should be noted that the improvement in the C. baccatum hybrid plant according to the third step may be optional, according to characteristics already present in the C. baccatum plant chosen as the rootstock. Indeed, while, in general, one may expect that the rootstock is preferably a hybrid plant obtained by crossing two C. baccatum lines utilizing heterosis, it cannot be denied that a baccatum plant line has enough traits of interest to, alone, be an acceptable rootstock.

(21) First, the inventors identified, from a representative collection of C. baccatum plants, the best accessions in inherent value and in hybrid form on the basis of their behaviour in the nursery and in the field, with or without grafting according to the following: root development, vigour of the grafted plant, germination, vigour and height of the plant in the nursery.

(22) The results presented below relate to four grafting trials performed over two consecutive years and in two different locations.

(23) 1.1 Comparison of the Root Volume of C. annuum and C. baccatum Plants

(24) The inventors compared the root volume of six different C. annuum control plants with that of 90 different C. baccatum plants, under identical culture conditions, using a scale ranging from 1 (very weak) to 5 (highly developed).

(25) The planting was performed in mid-May and the evaluation of the root volume was performed at the beginning of November.

(26) The results are presented in table 1 and show that the average root volume of the C. baccatum plants is 4.2, i.e. on average two times greater than that of the C. annuum control plants (2.1). The root volume of an HF-1 hybrid obtained by crossing accessions USDA PI 413669 01 and PBC1405 was also determined to be equal to 4.

(27) The roots of some of the plants obtained are presented in FIG. 2.

(28) TABLE-US-00001 TABLE 1 Root volume Root Plants (not grafted) Number volume C. annuum (average) 6 2.1 C. baccatum (average) 90 4.2 PG-TRESOR C. annuum 1.0 PG-SNOOKER C. annuum 2.0 TECNICO C. annuum 2.0 USDA PI 413669 01 * PBC1405 (HF1) 4.0 C. baccatum Root volume: 1 = very weak; 2 = weak; 3 = average; 4 = developed; 5 = highly developed
1.2 Behaviour in the Nursery

(29) Prior to the grafting, the inventors compared the germination percentage in the nursery and the hypocotyl length just before grafting different C. annuum plants, with that of plants obtained from crossing different C. baccatum accessions with the C. baccatum accession PBC1405.

(30) The results are presented in table 2 and show that the C. baccatum seeds have satisfactory germinations, their average germination percentage even being greater than that of the C. annuum plants. This result is unexpected insofar as, with relatively undomesticated plants, the risk of poor germination is considered to be high.

(31) The hypocotyl length of the C. baccatum plants is greater than that of the C. annuum plants. This characteristic is particularly advantageous in the context of use as a rootstock.

(32) TABLE-US-00002 TABLE 2 Germination percentage and hypocotyl length Hypocotyl % length Trial Plant Number Germination (in mm) Trial C. annuum (average) 30 30.4 1 C. baccatum (average) 18 37.3 PG-TRESOR C. annuum 29 PG-SNOOKER C. annuum 37 TECNICO C. annuum 34 USDA PI 413669 01 37 SD*PBC1405 C. baccatum Trial C. annuum (average) 28 78% 21.3 2 C. baccatum (average) 15 80% 29.9 PG-TRESOR C. annuum 80% 23.3 PG-SNOOKER C. annuum 90% 28 ROBUSTO C. annuum 72% 17.7 TECNICO C. annuum 60% 26.5 USDA PI 413669 01 100% 33.9 SD*PBC1405 C. baccatum Trial C. annuum (average) 56 77% 29.2 3 C. baccatum (average) 35 75% 31.5 PG-TRESOR C. annuum 100% 27 PG-SNOOKER C. annuum 80% 31 TECNICO C. annuum 100% 27 USDA PI 413669 01 100% 35.4 SD*PBC1405 C. baccatum Trial C. annuum (average) 28 73% 26.3 4 C. baccatum (average) 19 82% 34.1 PG-TRESOR C. annuum 96% 30 PG-SNOOKER C. annuum 72% 29.5 ROBUSTO C. annuum 68% 19.8 TECNICO C. annuum 94% 26.4 USDA PI 413669 01 96% 35.5 SD*PBC1405 C. baccatum
1.3 Obtaining Grafted Plants

(33) The inventors grafted a C. annuum plant on different C. annuum rootstock plants and different C. baccatum rootstock plants obtained from crossings. The same C. annuum graft, Eppo, was used for all of the grafted plants. The graft technique used is the Japanese graft.

(34) The characteristics of the grafted plants were then determined. The results are presented in table 3. The roots of the plants of trial 2 are presented in FIGS. 3A-3E.

(35) The grades for vigour and early fruiting make it possible to determine whether the growth of the plant is balanced, i.e. if the balance between vegetative and generative development is respected. A low earliness grade indicates excessive vegetative development, while a low vigour grade indicates excessive generative development. The ideal is to have high earliness (good fruiting) with high vigour. The results obtained show that there was no imbalance after grafting, even though this is often the case during grafting with a C. annuum rootstock (excessively generative) and there is a risk of obtaining an excessively vegetative grafted plant due to the use of a highly vigorous rootstock plant.

(36) The affinity grade characterizes the quality of the junction (connection of vessels, etc.) between the stem of the graft and that of the rootstock. A high grade corresponds to a good alignment and an absence of enlargement at the graft point. The results obtained show that the compatibility between the C. annuum graft and the C. baccatum rootstocks is approximately the same as that of the graft with the C. annuum rootstocks.

(37) The grades for root volume and extent of root hair make it possible to evaluate the quality of the root system. The results obtained show that the root system of the grafted plants for which the rootstocks are C. baccatum plants is significantly superior to that of the plants grafted using C. annuum rootstocks. Consequently, the use of C. baccatum rootstocks makes it possible to obtain grafted plants of which the root system is more developed and denser, thereby allowing for improved productivity.

(38) TABLE-US-00003 TABLE 3 Characteristics of grafted plants Root Root Trial Plant Number Earlines Vigour Affinity Vol. cap. Trial 1 C. annuum (average) 30 3.1 3.2 C. baccatum (average) 18 4.0 3.0 PG-SNOOKER C. annuum 3.0 3.8 PG-TRESOR C. annuum 3.0 3.2 TECNICO C. annuum 2.5 2.8 USDA PI 413669 01 4.0 3.8 SD*PBC1405 C. baccatum Trial 2 C. annuum (average) 28 3.1 3.4 2.3 3.0 2.5 C. baccatum (average) 14 3.5 3.6 3.1 3.7 4.1 PG-SNOOKER C. annuum 4.0 3.8 2.8 2.0 2.0 PG-TRESOR C. annuum 3.3 3.5 3.0 2.0 1.0 ROBUSTO C. annuum 3.0 3.3 2.0 4.0 2.0 TECNICO C. annuum 3.5 3.3 3.5 3.0 2.0 USDA PI 413669 01 3.3 4.3 4.0 5.0 5.0 SD*PBC1405 C. baccatum Trial 4 C. annuum (average) 28 2.7 3.4 3.5 3.7 3.6 C. baccatum (average) 19 2.5 3.2 2.5 4.0 3.8 PG-SNOOKER C. annuum 3.0 2.8 3.5 3.0 2.5 PG-TRESOR C. annuum 1.5 3.7 4.0 3.5 3.5 ROBUSTO C. annuum 3.5 2.8 3.5 2.0 3.0 TECNICO C. annuum 2.2 3.2 4.2 4.2 4.0 USDA PI 413669 01 3.0 2.7 3.0 4.0 5.0 SD*PBC1405 C. baccatum Earliness: early fruiting or setting; vigour: aerial development of the plant (foliar mass); affinity: quality of the junction between the graft and the rootstock; root vol.: root volume; root cap.: extent of root hair (root mass); 1 = very low; 2 = low; 3 = medium; 4 = high; 5 = very high

2. Transfer of Resistance from C. annuum to C. baccatum

(39) Two crossings were performed between the accessions of C. annuum CM334 comprising genes with P5 resistance (partial resistance to Phytophthora) and Mel (partial resistance to nematodes), and C. baccatum PBC1405. For the first crossing, the female parent was the accession PBC1405 and for the second crossing, the female parent was the accession CM334. The first crossing resulted in the presence of fruit, but the second did not.

(40) The inventors chose to use the C. annuum accession CM334 as the male parent and crossed it in a conventional manner with 95 different C. baccatum accessions used as the female parents.

(41) The presence of fruit after hybridization, the presence of seeds in the hybridized fruit and the number of seeds harvested by crossing were then determined. A maximum of 10 seeds were then sown for each crossing having enabled the seed harvest. The capacity of sown F1 (HF1) hybrid seeds leading to germination and the number of F1 hybrid plants obtained were also determined. Of the 95 crossings performed, 46 resulted in fruit, 24 enabled hybrid seeds to be obtained, and the seeds from 10 of these crossings led to germination. These results confirm that it is possible, without using an in vitro technique, to obtain F1 hybrid seeds using C. annuum as the male plant and C. baccatum as the female parent.

(42) The inventors used two molecular markers and morphological observations to confirm the interspecific nature of the HF1 hybrids obtained. In particular, the scar marker CD (Djian-Capolalino et al., 2007 (10)) was used for resistance to nematodes and an internal marker was used for resistance to Phytophthora.

(43) The Capsicum annuum plants used in these trials have a hairy stem and white flowers, while the Capsicum baccatum plants used have a glabrous stem and white flowers with yellow spots. The plants obtained after these crossings have a slightly hairy stem and flowers with light yellow spots (see FIG. 1), which confirms their hybrid character.

(44) The inventors also determined that seven of the plants obtained from the crossings performed made it possible to obtain HF1 hybrids that combine the markers of resistance to nematodes with those to Phytophthora obtained from the accession CM334.

(45) The results relating to three of the hybrids obtained are presented in table 4.

(46) TABLE-US-00004 TABLE 4 HF1 hybrids obtained by crossing different C. baccatum plants (female parent) with C. annuum CM334 (male parent) and marking to determine their trait of resistance or susceptibility to nematodes and Phytophthora Presence of Presence of Average number Germintation Number of plants fruit after seeds in the of HF1 seeds of the HF1 obtained from R nematodes R Crossing hybridization hybrid fruit harvested per plant seeds HF1 seeds (scar CD) Phytophthora Interpretation USDA PI X X 3.00 X 2.00 R heterozygote hybrid 413669 01 SD*CM334 USDA PI X X 30.00 X 4.00 R heterozygote hybrid 159252 01 SD*CM334 PEN79*CM334 X X 80.00 X 2.00 R heterozygote hybrid

(47) The inventors then crossed each of the seven hybrids obtained with either the recurrent C. baccatum parent (backcrossing) or the C. baccatum accession PBC1405 also identified as being valuable in particular in that it has excellent root vigour (level 5).

(48) The seeds obtained from each crossing were harvested and BC1 embryos were placed in culture. The embryo culture was performed according to the protocols of Dumas de Vaulx, 1992 (Dumas Culture d'embryons et d'ovaires fconds. In: Jahier J. (ed) Techniques de cytogntique vgtale, INRA editions, page 141).

(49) The number of seeds harvested by crossing and the number of plantlets obtained from these seeds were then determined. The inventors then produced a molecular marker screen for certain of the plants obtained for resistance to Phytophthora and nematodes. Four BC1 plants having a double resistance to Phytophthora and nematodes were obtained.

(50) The results relating to three of the hybrids are presented in tables 5 and 6.

(51) TABLE-US-00005 TABLE 5 Characteristics of the Plants (BC1) obtained after crossing of the F1 Hybrids with either the C. baccatum plant used as the parent in the initial crossing or C. baccatum PBC1405 Number of Number of Number of Number of R Number of R Number of Crossing seeds plantlets obtained plantlets tested nematodes Phytophthora double R (USDA PI 413669 01 12 1 3 2 3 2 SD*CM334)*PI 413669 (USDA PI 159252 01 4 2 SD*CM334)*PI 159252 (USDA PI 159252 01 29 18 11 7 3 2 SD*CM334)*PBC 1405 (PEN79*CM334)*PEN79 15 8 4 3 0 0 (R = resistance)

(52) TABLE-US-00006 TABLE 6 Marking of plants resulting from the first backcrossing (BC1) R. nematode R. (Me7) phytophthora No. Variety marking marking 1 (PEN79*CM334)*PEN79 R susceptible 2 (PEN79*CM334)*PEN79 R susceptible 3 (PEN79*CM334)*PEN79 R susceptible 14 (PEN79*CM334)*PEN79 S susceptible 4 (USDA PI 413669 01 S Hetero. R SD*CM334)*USDA PI 413669 01 SD 5 (USDA PI 413669 01 R Hetero. R SD*CM334)*USDA PI 413669 01 SD 16 (USDA PI 413669 01 R Hetero. R SD*CM334)*USDA PI 413669 01 SD 7 (USDA PI 159252 01 S susceptible SD*CM334)*PBC1405 8 (USDA PI 159252 01 S susceptible SD*CM334)*PBC1405 9 (USDA PI 159252 01 R susceptible SD*CM334)*PBC1405 10 (USDA PI 159252 01 R susceptible SD*CM334)*PBC1405 11 (USDA PI 159252 01 ? Hetero. R SD*CM334)*PBC1405 12 (USDA PI 159252 01 R Hetero. R SD*CM334)*PBC1405 17 (USDA PI 159252 01 R susceptible SD*CM334)*PBC1405 18 (USDA PI 159252 01 R Hetero. R SD*CM334)*PBC1405 19 (USDA PI 159252 01 R susceptible SD*CM334)*PBC1405 20 (USDA PI 159252 01 S susceptible SD*CM334)*PBC1405 21 (USDA PI 159252 01 R susceptible SD*CM334)*PBC1405

(53) The inventors then performed a step of an additional crossing between BC1 plants identified as being sensitive or resistant to Phytophthora and nematodes with C. baccatum (BC2) plants followed by self-pollinations (BC1S1). The inventors produced a molecular marker screen for certain of the plants obtained for resistance to Phytophthora and nematodes. In addition to the marker already mentioned above, a second co-dominant scar marker, adapted to the marker described in Wang et al., 2009 (11) was also used for nematode resistance. The results obtained show that the two markers are related (3% of recombinations).

(54) The segregations obtained are compatible with those expected on the basis of the level of resistance of the BC1 plants of origin and the type of progeny (BC2 or BC1S1). The results are shown in tables 7, 8 and 9.

(55) TABLE-US-00007 TABLE 7 BC2 and BC1S1 data for the Phytophthora marking BC1 prior number of plants % of plants Code State Crossing origin results R_R R_S S_S % R:R % R:S % S:S 10A6001- BC2 (PEN79*CM334)*PEN79*PEN79 1-D susceptible 1 0% 0% 100% 1DP1*10A5439 10A6002- BC2 (PEN79*CM334)*PEN79*PEN79 2-D susceptible 1 0% 0% 100% 2DP1*10A5439 10A6003- BC2 (PEN79*CM334)*PEN79*PBC1405 3-D susceptible 2 0% 0% 100% 3DP1*10A5421 10A6006- BC2 (USDA PI 159252 01 9-L susceptible 42 0% 0% 100% 9LP1*10A5421 SD*CM334)*PBC1405*PBC1405 10A6006- BC2 (USDA PI 159252 01 9-L susceptible 52 0% 0% 100% 9LP1*10A5536 SD*CM334)*PBC1405*USDA PI 159252 01 SD 10A6007- BC2 (USDA PI 159252 01 10-M susceptible 38 0% 0% 100% 10MP1*10A5421 SD*CM334)*PBC1405*PBC1405 10A6007- BC2 (USDA PI 159252 01 10-M susceptible 52 0% 0% 100% 10MP1*10A5536 SD*CM334)*PBC1405*USDA PI 159252 01 SD 10A6012- BC2 (USDA PI 159252 01 19-V susceptible 10 0% 0% 100% 19YP1*10A5421 SD*CM334)*PBC1405*PBC1405 10A6013- BC2 (USDA PI 159252 01 21-A susceptible 50 0% 0% 100% 21ABP1*10A5421 SD*CM334)*PBC1405*PBC1405 Total 0 0 248 0% 0% 100% 10A6004- BC2 (USDA PI 413669 01 4-H hetero 11 0% 0% 100% 4HP1*10A5421 SD*CM334)*USDA PI 413669 01 SD*PBC1405 10A6004- BC2 (USDA PI 413669 01 4-H hetero 1 2 0% 33% 67% 4HP1*10A5440 SD*CM334)*USDA PI 413669 01 SD*USDA PI 413669 01 SD 10A6009- BC2 (USDA PI 413669 01 16-V hetero 6 9 0% 40% 60% 16VP1*10A5421 SD*CM334)*USDA PI 413669 01 SD*PBC1405 10A6009- BC2 (USDA PI 413669 01 16-V hetero 2 3 0% 40% 60% 16VP1*10A5440 SD*CM334)*USDA PI 413669 01 SD*USDA PI 413669 01 SD Total 0 9 25 0% 26% 74% 10A6009-16VP1 BC1S1 F2[(USDA PI 413669 01 16-V hetero 2 6 25% 75% 0% SD*CM334)*USDA PI 413669 01 SD] 10A6011-18YP1 BC1S1 F2[(USDA PI 159252 01 18-V hetero 17 22 13 33% 42% 25% SD*CM334)*PBC1405] Total 19 28 13 32% 47% 22%

(56) TABLE-US-00008 TABLE 8 BC2 and BC1S1 data for the nematode marking (marker 1) BC1 prior number of plants % of plants Code State Crossing origin results R S % R % S 10A6004- BC2 (USDA PI 413669 01 4-H susceptible 14 0% 100% 4HP1*10A5421 SD*CM334)*USDA PI 413669 01 SD*PBC1405 10A6004- BC2 (USDA PI 413669 01 4-H susceptible 3 0% 100% 4HP1*10A5440 SD*CM334)*USDA PI 413669 01 SD*USDA PI 413669 01 SD Total 0 17 0% 100% 10A6001- BC2 (PEN79*CM334)*PEN79*PEN79 1-D resistant 1 0% 100% 1DP1*10A5439 10A6002- BC2 (PEN79*CM334)*PEN79*PEN79 2-D resistant 1 0% 100% 2DP1*10A5439 10A6003- BC2 (PEN79*CM334)*PEN79*PBC1405 3-D resistant 1 1 50% 50% 3DP1*10A5421 10A6006- BC2 (DSDA PI 159252 01 9-L resistant 26 19 58% 42% 9LP1*10A5421 SD*CM334)*PBC1405*PBC1405 10A6006- BC2 (USDA PI 159252 01 9-L resistant 20 32 38% 62% 9LP1*10A5536 SD*CM334)*PBC1405*USDA PI 159252 01 SD 10A6007- BC2 (USDA PI 159252 01 10-M resistant 15 25 38% 63% 10MP1*10A5421 SD*CM334)*PBC1405*PBC1405 10A6007- BC2 (USDA PI 159252 01 10-M resistant 18 34 35% 65% 10MP1*10A5536 SD*CM334)*PBC1405*USDA PI 159252 01 SD 10A6009- BC2 (USDA PI 413669 01 16-V resistant 11 4 73% 27% 16VP1*10A5421 SD*CM334)*USDA PI 413669 01 SD*PBC1405 10A6009- BC2 (USDA PI 413669 01 16-V resistant 2 3 40% 60% 16VP1*10A5440 SD*CM334)*DSDA PI 413669 01 SD*USDA PI 413669 01 SD 10A6012- BC2 (USDA PI 159252 01 19-V resistant 7 5 58% 42% 19YP1*10A5421 SD*CM334)*PBC1405*PBC1405 10A6013- BC2 (USDA PI 159252 01 21-A resistant 25 26 49% 51% 21ABP1*10A5421 SD*CM334)*PBC1405*PBC1405 Total 125 151 45% 55% 10A6009-16VP1 BC1S1 F2[(USDA PI 413669 01 16-V resistant 4 4 50% 50% SD*CM334)*USDA PI 413669 01 SD] 10A6011-18YP1 BC1S1 F2[(USDA PI 159252 01 18-V resistant 35 15 70% 30% SD*CM334)*PBC1405] Total 39 19 67% 33%

(57) TABLE-US-00009 TABLE 9 BC2 and BC1S1 data for the nematode marking (marker 2) BC1 prior number of plants % of plants Code State Crossing origin results R_R R_S S_S % R:R % R:S % S:S 10A6004- BC2 (USDA PI 413669 01 4-H susceptible 13 0% 0% 100% 4HP1*10A5421 SD*CM334)*USDA PI 413669 01 SD*PBC1405 10A6004- BC2 (USDA PI 413669 01 4-H susceptible 3 0% 0% 100% 4HP1*10A5440 SD*CM334)*USDA PI 413669 01 SD*USDA PI 413669 01 SD Total 0 0 16 0% 0% 100% 10A6001- BC2 (PEN79*CM334)*PEN79*PEN79 1-D resistant 1DP1*10A5439 10A6002- BC2 (PEN79*CM334)*PEN79*PEN79 2-D resistant 2DP1*10A5439 10A6003- BC2 (PEN79*CM334)*PEN79*PBC1405 3-D resistant 1 1 0% 50% 50% 3DP1*10A5421 10A6006- BC2 (USDA PI 159252 01 9-L resistant 26 18 0% 59% 41% 9LP1*10A5421 SD*CM334)*PBC1405*PBC1405 10A6006- BC2 (USDA PI 159252 01 9-L resistant 20 31 0% 39% 61% 9LP1*10A5536 SD*CM334)*PBC1405*USDA PI 159252 01 SD 10A6007- BC2 (USDA PI 159252 01 10-M resistant 14 22 0% 39% 61% 10MP1*10A5421 SD*CM334)*PBC1405*PBC1405 10A6007- BC2 (USDA PI 159252 01 10-M resistant 19 27 0% 41% 59% 10MP1*10A5536 SD*CM334)*PBC1405*USDA PI 159252 01 SD 10A6009- BC2 (USDA PI 413669 01 16-V resistant 9 4 0% 69% 31% 16VP1*10A5421 SD*CM334)*USDA PI 413669 01 SD*PBC1405 10A6009- BC2 (USDA PI 413669 01 16-V resistant 2 1 0% 67% 33% 16VP1*10A5440 SD*CM334)*USDA PI 413669 01 SD*USDA PI 413669 01 SD 10A6012- BC2 (USDA PI 159252 01 19-V resistant 4 4 0% 50% 50% 19YP1*10A5421 SD*CM334)*PBC1405*PBC1405 10A6013- BC2 (USDA PI 159252 01 21-A resistant 4 15 13 13% 47% 41% 21ABP1*10A5421 SD*CM334)*PBC1405*PBC1405 Total 4 110 121 2% 47% 51% 10A6009-16VP1 BC1S1 F2[(USDA PI 413669 01 16-V resistant 1 3 4 13% 38% 50% SD*CM334)*USDA PI 413669 01 SD] 10A6011-18YP1 BC1S1 F2[(USDA PI 159252 01 18-V resistant 12 22 12 26% 48% 26% SD*CM334)*PBC1405] Total 13 25 16 24% 46% 30%

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