Identification of stem soft rot (Erwinia carotovora subspeciesatroseptica) resistance in potato

Development of potato cultivars resistant to Erwinia species (Erwinia carotovora subspeciesatroseptica, E. carotovora subspeciescarotovora, andErwinia chrysanthemi) causing soft rot of stems and tubers involves identification and transfer of the resistance to breeding lines possessing desirable agronomic traits. In total, 363 accessions of six wildSolanum species, three somatic hybrids and nine sexual progeny of somatic hybrids were screened for stem soft rot resistance. Thirty-one, 50, and 67 percent of accessions ofSolanum boliviense, Solanum chacoense, and Solanumsanctarosae, respectively, were resistant or highly resistant, whereas, 100, 97, and 95 percent of accessions ofSolanum canasense, Solanum tarijense, andSolanum spegazzinii were susceptible or highly susceptible, respectively. Two tuber soft-rot resistant somatic hybrids, A937 and T355-11 produced earlier by the fusion ofSolanum brevidens (PI 218228) andSolanum tuberosum (PI203900) orS. tuberosum cv. Russet Burbank, respectively, and the sexual progeny of A937 andS. tuberosum cv. Katahdin were also highly resistant. Transfer of resistance fromS. brevidens to somatic hybrid (A937) and subsequently to the progeny suggests that the resistance is simply inherited. In addition to the identification of 65 resistant or highly resistant clones of wildSolanum species, this study has identified three tuber soft rot-resistant clones that are also resistant to stem soft rot. Potato cultivars showed a range of responses to the pathogen.     

Erwinia resistance in backcross progenies ofSolanum tuberosum x S. tarijense andS. tuberosum (+) S. commersonii hybrids

Summary The resistance to tuber soft rot caused byErwinia carotovora ofSolanum tuberosum x S. tarijense andS. tuberosum (+) S. commersonii hybrids and their backcrosses is reported. A number of resistant diploid sexual hybrids and tetraploid/hexaploid somatic hybrids were selected. Backcross progenies were obtained through 2x×4x crosses involving a resistant diploid hybrid and tetraploidS. tuberosum, and through 4x×4x crosses between a resistant somatic hybrid andS. tuberosum. The hybrids showed high variability interms of resistance to tuber soft rot. The resistance of progeny from 2x×4x backcrosses was similar to that of the parental sexual hybrid. By contrast, the resistance of genotypes deriving from 4x×4x backcrosses was reduced compared with the resistant somatic hybrid. In general, tuber characteristics of the backcross hybrids improved considerably as compared with their parents, and tuber yield per plant was good.     

Sexual progeny of somatic hybrids between potato andSolanum brevidens: Potential for use in breeding programs

Hexaploid somatic hybrids have been obtained by fusion of protoplasts fromSolanum brevidens (PI 218228, 2x = 2n = 24) andS. tuberosum (PI 203900 or cv. Russet Burbank; 2x = 4n = 48). In the work reported here, pentaploid progeny derived from sexual crosses between the somatic hybrids and the potato cultivar, Katahdin were assessed for transfer of disease resistances and improvement of agronomic traits. Segregation was noted in disease resistances to race 0 ofPhytophthora infestans and to tuber soft rot caused byErwinia spp., indicating that sexual transfer of traits captured by protoplast fusion had occurred. The pentaploid progeny showed substantial variation in and often improvement of some agronomic traits as compared to the somatic hybrids. The results indicate that interspecific somatic hybridization can produce material with potential for use in a potato breeding program     

Resistance to blackleg and tuber soft rot in sexual and somatic interspecific hybrids with different genetic background

Diploid interspecific hybrids from crosses betweenSolanum tuberosum haploids andS. canasense, S. multidissectum, andS. tarijense, along with tetraploid and hexaploid somatic hybrids S.tuberosum (+)S. commersonii, were screened for resistance to blackleg and tuber soft rot byErwinia carotovora subsp.atroseptica andE. carotovora subsp.carotovora. Among the diploid hybrids, those involving clone mlt la of S.multidissectum and tar 2b ofS. tarijense yielded the highest number of genotypes resistant to both blackleg and tuber soft rot. By contrast, all the hybrids involving clone tar 1lb ofS. tarijense were susceptible to both bacterial diseases. As far as resistance of somatic hybrids is concerned, the most interesting genotype was the tetraploid one which showed resistance to both blackleg and tuber soft rot Significant correlations were found betweenE. carotovora subsp.atroseptica andE. carotovora subsp.carotovora both in resistance to blackleg and in resistance to tuber soft rot, suggesting that resistance to the two subspecies is controlled by the same gene/s or that the relevant genes are linked. On the other hand, correlations between blackleg and tuber soft rot were never significant, indicating that different mechanisms may control resistance in tuber and stems. Alternatively, the resistance could be controlled by the same gene/s under a different spatial/temporal expression pattern. Twelve diploid sexual hybrids and one tetraploid somatic hybrid were selected for resistance to blackleg and/or tuber soft rot and for tuber characteristics.     

Resistance to soft rot (Erwinia carotovora subsp.atroseptica) in tetraploid potato families obtained from 4x-2x crosses

Diploid potato clones, interspecific hybrids ofSolanum species, having in their originS. tuberosum,S. chacoense, S. yungasense, S. phureja, S. gourlayi, andS. demissum, with resistance to soft rot, were crossed to tetraploid potato clones in 4x-2x crosses. The 24 tetraploid families obtained in a North Carolina II design were examined for tuber resistance to soft rot in a laboratory test and for basic agronomic traits in field trials conducted for two consecutive years. In addition, one family originating from a 4x-2x cross of two susceptible parents was tested. Statistical analysis of the data revealed significant general (GCA) and specific combining ability (SCA) effects, year, GCA (female) × year, GCA (male) × year, and SCA × year upon the inheritance of resistance to soft rot. About 35% of the progeny was selected as resistant to tuber soft rot, and of these 11% showed high resistance combined with good tuber yield, tuber weight, and tuber appearance. The relationships between resistance to soft rot and chosen agronomic traits were not noted or were weakly significant and sporadic. The resistance to tuber soft rot found in diploid potato hybrids can be transferred to the cultivated tetraploid pool through 4x-2x crosses, and a high frequency of offspring posses resistance.     

The Effect of Long Term Storage on Bacterial Soft Rot Resistance in Potato

Bacterial soft rot is a serious disease in potato (Solanum tuberosum L.), causing rapid tuber tissue maceration and, consequently, marketable yield loss. Soft rot bacteria, including Pectobacterium carotovorum subsp. carotovorum (Pbc), are favored by moist conditions, which are prevalent in large potato storage facilities. However, although most potatoes in North America are stored before use, there are no published surveys of soft rot resistance in cultivars exposed to long-term storage conditions. Thus, we tested 65 cultivars and 13 breeding lines for soft rot resistance after 6 months of storage. There was a significant effect of cultivar and production environment on soft rot resistance score. During 6 months of storage, tuber soft rot resistance in resistant clones did not change, while it changed in susceptible clones. The three most resistant cultivars to soft rot were Freedom Russet, Anett, and Alaska Red Eye.     

Production and characterization of “second generation” somatic hybrids derived from protoplast fusion between interspecific somatohaploid and dihaploidSolanum tuberosum L.

Protoplasts were fused to produce somatic hybrids between a triploid (2n=3x=32-34) interspecific somatohaploid betweenSolanum brevidens Phil. andS. tuberosum L., and a dihaploid (2n=2x=24) anther-derived line ofS. tuberosum cv. Van Gogh. A total of 265 plants were regenerated from protoplast fusion derived calli and their hybridity was verified using fusion partner specific RAPD markers. These “second generation” somatic hybrids were aneuploid pentaploids (2n=5x=51-65) with a 2C DNA content ranging from 3.36 to 4.43 pg, which corresponded to the sum of the 2C values of each of the fusion partners (somatohaploid: 2.22 pg; and the dihaploid line of cv. Van Gogh: 1.87 pg). Most of the “second generation” somatic hybrids were vigorous, but variable in morphology. They were extremely resistant to PLRV and they had tolerance to PVY infection derived from the somatohaploid fusion partner. Even though most of the “second generation” hybrids tuberized, the tuber morphology was variable and most were poorly shaped. InErwinia soft rot resistance tests, the tubers showed higher level of resistance than the tetraploidS. tuberosum cultivars, the dihaploidS. tuberosum fusion partners and the hexaploid somatic hybrids betweenS. brevidens andS. tuberosum. The “second generation” somatic hybrids were all male sterile and failed to produce berries or seeds.     

The inheritance of resistance to soft rot (Erwinia carotovora subsp.atroseptica) in diploid potato families

Soft rot of potato tubers, caused byErwinia (Pectobacterium), is a serious disease affecting potato crops during storage. Studies on the inheritance of resistance to tuber soft rot were undertaken on diploid potato hybrids. A total of 480 clones, derived from 12 families, were examined for resistance to soft rot in laboratory tests over 3 years and for basic agronomic traits in field experiments over 2 years. General (GCA) and specific (SCA) combining abilities were significant in the inheritance of resistance to soft rot; however, GCA for female parents were significant in 1 or 2 years during 3 years of evaluation and SCA was significant for eight, two, and six families out of 10 tested in 1994, 1995, and 1996, respectively. There were also significant differences between years. Moreover, marked interactions for GCA_females × year and SCA × year in the variation of soft rot resistance were found. Broad-sense and narrowsense heritability of resistance, measured as diameter of rotten tissue, was estimated as 0.92 and 0.89, respectively. The maternal effect, evaluated in two sets of reciprocal crosses, was not significant for the genetic determination of resistance to soft rot. No significant relationships were found between resistance to soft rot and the main agronomic traits in the tested diploid families. These results suggest that diploid potato clones with resistance to soft rot can be selected after being evaluated over a few seasons. It is also possible to select clones combining good resistance to soft rot with high yield, superior tuber characteristics, and acceptable starch content. These resistant diploids can be used as male parents in 4x-2x crosses in breeding tetraploid potato resistant to soft rot.     

Breeding Potential of Potato Somatic Hybrids: Evaluations for Adaptability,Tuber Traits,Late Blight Resistance,Keeping Quality and Backcross (BC<Subscript>1</Subscript>) Progenies

Ten tetraploid interspecific potato somatic hybrids developed earlier through protoplast fusion between the dihaploid Solanum tuberosum L. ‘C-13’ and the diploid wild species Solanum pinnatisectum Dun. were used in this study. Somatic hybrids and standard control cultivars were evaluated for adaptability, tuber traits, late blight resistance and keeping quality attributes (dormancy, sprouting, weight loss and appearance after 75 days of storage) during two successive winter crop seasons (short-days) in the sub-tropical plains of India, where nearly 90% of India’s potatoes are grown. Somatic hybrids showed medium to good plant vigour and late to very late plant maturity. Though none of the somatic hybrids produced equal or higher tuber yield than the control cultivars, except clone P7, all other somatic hybrids produced significantly higher marketable and total tuber yield after a 90-day growth than the parent C-13. Most of the somatic hybrids possessed higher tuber dry matter concentration, resistance to late blight and better keeping quality attributes than the control cultivars. Notably, the tubers of somatic hybrids showed a tendency for colour change from white to purple on exposure to sunlight. Breeding potential of somatic hybrids was further assessed by crossing them with commonly grown local potato cultivars for the development of backcross (BC₁) progenies. In the hybridization programme, berries were formed while somatic hybrids were used either as male or female parents with common cultivars. Our results revealed that the use of bulk pollen of somatic hybrids not only resulted in higher pollination success but also helped to achieve higher numbers of true potato seeds (TPS) per berry. Despite the formation of berries, TPS was absent in 22 crosses using somatic hybrids as a female or male parent which may be due to ovule or embryo abortions. Segregating BC₁ generations were evaluated under the same sub-tropical conditions for further exploitation in potato breeding. Based on our study, the somatic hybrids P4, P8 and P10 can be utilized for the introgression of important characters such as high tuber dry matter concontration, resistance to late blight and excellent keeping quality attributes into the cultivated potato via conventional breeding methods for cultivar development in the sub-tropical plains of India.     

Evaluation for chipping and tuber soft rot (Erwinia carotovora) resistance in potato clones from unilateral sexual polyploidization (2x × 4x)

The aim of this work is the evaluation of tetraploid potato clones with germplasm fromS. chacoense andS. phureja for chipping and resistance to tuber soft rot (Erwinia carotovora subsp.carotovora). Tetraploid potato clones were obtained via 2n eggs through 2x–4x crosses. Cytological mechanisms were investigated in the diploid hybrid 2n-egg producer used as parent in the unilateral sexual polyploidization scheme; the omission of the second meiotic division and cytokinesis failure at the end of meiosis were identified with a genetic consequence equivalent to second division restitution (SDR). Tuber marketable yield, specific gravity and chip-processing ability evaluated at harvest and after cold storage with and without reconditioning were higher in some clones as compared to theS. tuberosum tetraploid parent. Tuber soft rot resistance, contributed byS. chacoense, was successfully transferred to the4x level. After inoculation withErwinia carotovora subsp.carotovora and estimation of the rotted area, one tetraploid clone proved to be resistant and three partially resistant. The genetic value of exotic germplasm and sexual polyploidization strategy are discussed in relation to potato improvement.     

Tuber quality and soft rot resistance of hybrids betweenSolarium tuberosum and the incongruent wild relativeS. commersonii

Tetraploid BC₃ hybrids between sexually incompatible speciesSolanum commersonii andS. tuberosum were characterized for tuber quality (stolon length, tuber eye depth, chipping color from 3-month storage at 5 C, and specific gravity) and soft rot resistance traits. For all analysed morphological traits, except stolon length, genotypes resembled theS. tuberosum phenotype. On average, the specific gravity of selected genotypes was quite high (1.087), and 12 of them had a value >1.090. As for chipprocessing ability, 4.8% of genotypes chipped well directly from cold storage, and 14.3% after reconditioning. All but one of these genotypes also possessed a high specific gravity of tubers. Variability was found for resistance toErwinia carotovora subsp.carotovora. In particular, 24% of hybrids had a rotted area diameter ≤ 4 mm and were classified as resistant. The average percentage ofcommersonii-specific markers still present in the BC₃ hybrids was 22%, ranging from 9% to 37%. Genotypes combining good qualitative tuber traits with a wild genome content lower than the average value of the population were selected for further breeding efforts. The importance of molecular-assisted selection is discussed.     

Comparison of the effectiveness of different methods of screening for bacterial soft rot resistance of potato tubers

Tubers from eight potato cultivars (cvs) grown at two different locations in Wisconsin were tested for bacterial soft rot resistance using different inoculation techniques. The procedures included 1) point inoculations of tubers with different inoculum levels followed by incubation in ambient or low oxygen condtions, 2) inoculation of mechanically bruised tubers followed by incubation in a mist chamber, and 3) a standard slice inoculation method. The point titration test followed by incubation in dew chamber and the mist chamber-bruise test showed similar patterns of resistance for cultivars that were used in these experiments. These two methods are considered to be effective for screening potato tubers for bacterial soft rot resistance. Point titration methods are very useful if only limited numbers of tubers are available. The mist chamber-bruise test is simpler than the other procedures; however, to obtain reproducible results large numbers of tubers are required. Because of the great variability of the results obtained in inoculation of slices, the reliability of this approach can be questioned as a standardized method for evaluation of resistance. Tubers of somatic hybrids ofS. brevidens andS. tuberosum and their sexual progeny were significantly more resistant to bacterial soft rot than tubers of moderately resistant cultivars when evaluated by each of the assay procedures.     

Resistance toErwinia spp. in diploid potato with a high starch content

Summary Tubers of diploid interspecific hybrids, hexaploid somatic hybrids ofSolanum tuberosum andSolanum brevidens, a fusion parental line (S. tuberosum) and several Polish and American commercial cultivars were screened for resistance toErwinia carotovora subsp.atroseptica andErwinia chrysanthemi, in laboratory tests. Both groups of hybrids showed a high level of resistance to soft rot and gave a similar type of resistant reaction, which distinguished them from the cultivars and fusion parent. Tubers of the diploid interspecific hybrids had a high starch content. Our results indicate the possibility of obtaining breeding material with a high starch content and resistance to bacterial soft rot.     

利用体细胞杂交获取马铃薯软腐病的抗性

对马铃薯四倍体栽培种和二倍体野生种Solanumbrevidens的体细胞杂种通过叶片离体培养获得的四倍体植株,以及用四倍体栽培种进行回交获得的五倍体植株的块茎对软腐病的抗性进行了测定。结果表明,由体细胞杂种通过叶片组织离体培养再生植株中,有一个株系SC107对软腐病菌Erwiniacarotovora具有较强的抗性。在用不抗软腐病的马铃薯栽培种对体细胞杂种进行回交获得的杂种后代中,大部分株系对软腐病菌具有高水平的抗性,从而说明Solanumbrevidens对软腐病的抗性基因已转移到马铃薯栽培种。     

Soft rot and blackleg reactions in diploid potato hybrids inoculated withErwinia spp.

From 1993 to 1996 three groups of potato genotypes were evaluated for resistance toErwinia spp.: (1) 31 interspecific diploid hybrids (28 resistant and three susceptible), (2) five hexaploid or pentaploid somatic hybrids ofSolanum tuberosum (tbr) ×S. brevidens (brd), and (3) eight cultivars. Two evaluation methods were applied: tuber point inoculation with eitherErwinia carotovora ssp.atroseptica (Eca) orE. chrysanthemi (Ech) to test tuber soft rot resistance and stem base inoculation with Eca to test blackleg resistance. Some resistant diploid hybrids and somatic hybrids oftbr × brd were superior to cultivars for both tuber and stem resistance. Tuber and stem resistance toErwinia spp. in the most resistant diploid hybrids were equal to the highly resistant somatic hybrids oftbr xbrd. Tuber resistance to Eca was highly correlated to tuber resistance to Ech (r=0.815***). In two years of evaluation for stem resistance, three diploid hybrids and a derivative of one of the somatic hybrids (USA M 264) failed to develop symptoms of blackleg following inoculation with Eca, Analyses of variance for tuber and stem resistance indicated significant effects of genotype, year and genotype × year interaction. A positive relationship between tuber and stem resistance toErwinia spp. has been found, however the genetic control of resistance in tuber and stem is partially independent. In the case of Eca the correlation coefficient was r = 0.725***. Therefore it should be possible to obtain resistant genotypes to both blackleg and tuber soft rot. Several resistant diploid hybrids were selected from among those tested, which also have several other characters desirable for potato breeders.     

Germplasm transfer from the wild tetraploid speciesSolanum acaule bitt. to the cultivated potato,S. tuberosum L. using 2N eggs

Solanum acaule Bitt. (acl ¹, 2n=4x=48 and 2n=6x=72) is a wild potato species in which resistance to adverse biotic and abiotic agents has been found. The tetraploid subspeciesacaule andaemulans cannot be easily crossed with the common potato,S. tuberosum L. Grouptuberosum (tbr, 2n=4x=48) because the development of the endosperm in the hybrid seed is abnormal. Since 4xacl behaves as a diploid in intra and interploidy crosses, it was postulated that the hybridization barrier withtbr could be overcome if the wild species produced 2n gametes. One hundred and ninety-seven plants of 4xacl (22 introductions) were screened for pollen of heterogeneous size; only two plants produced this type of pollen, with 3% and 5% of pollen in the large class, respectively. On the other hand, 427 plants (47 introductions) were screened for 2n eggs through controlled crosses withtbr. Forty-one plants (13 introductions) produced 219 berries of which 46 contained at least one plump seed. Seed from berries with low seed set gave rise to hexaploid hybrids possibly through the functioning of 2n eggs from 4xacl. A sample of these hybrids was successfully backcrossed as female parents totbr, giving rise to 5x BC₁ progenies. These BC₁ progenies could be easily back-crossed totbr, yielding tetraploid or near tetraploid BC₂ plants.     

Inheritance of a pectate lyase mediated soft rot resistance in crosses between transgenic potato lines of cv. désirée andS. tuberosum cultivars

Summary Expression of the pectate lyase (PL) isoenzyme 3 in transgenic potato lines of cv. Désirée mediates an enhanced resistance of tuber tissue toErwinia carotovora (Ec) soft rot due to a pre-activation of plant defence mechanisms. Therefore, theSolanum tuberosum cvs Agave and Adretta with a moderate level of soft rot resistance were crossed with such PL-expressing potato lines. The resulting progenies were assessed with respect to plant/tuber characteristics over a period of four years and then tested for PL3-expression as well as for soft rot resistance. 71% of the selected progeny lines exhibited a stable production of the PL3 enzyme. Statistical analysis revealed differences between the transgenic and the non-transgenic progeny concerning the soft rot resistance of tuber tissue. Compared with the PL-inactive progeny, extension of Ec-rotting on the wound surface of PL-transgenic potatoes was diminished on average by 51.8%. Similarly, the degree of cell lysis caused by bacterial maceration was significantly (P<0.05) reduced in tuber tissue of PL-expressing progeny lines. The latter also revealed an enhanced PPO and PAL activity in their tuber tissue indicating an active plant defence. It is concluded therefore that the PL-mediated soft rot resistance introduced into potatoes by means of molecular techniques is heritable.     

Evaluation of the reliability of determining soft rot resistance in potatoes by the tuber slice method

Soft rot resistance in potato (Solanum tuberosum L.) tubers can be determined by inoculating tuber slices with soft rotErwinia species. Questions have been raised in the literature concerning the reliability of the tuber slice method. The objectives of this study were to 1) examine the statistical assumptions underlying the analysis of variance for different response variables as measures of soft rot resistance using the tuber slice method of evaluation; 2) estimate the sample sizes necessary to detect specified differences with power 0.83; and, 3) choose the “best” variable for measuring resistance to soft rot based on valid statistical analysis and minimal sample size. Slices from fifteen tubers from each of three cultivars (Atlantic, Norchip, Superior) were inoculated withErwinia carotovora subsp.atroseptica, E. carotovora subsp.carotovora orE. chrysanthemi and incubated at two different temperatures (20 C and 30 C) for 48 hrs. The test was conducted on two dates. Tuber samples were sliced and weighed prior to inoculation and after the macerated tissue was removed following a 48 hr incubation period. The maximum diameter of macerated tissue, actual weight loss, four measures of proportional weight loss, and various transformations of these variables were analyzed. The “best” response variables for measuring resistance to soft rot by the tuber slice method were the diameter of the macerated tissue and the square root transformation of one of the measures of proportional weight loss. No differences were found among the cultivars nor theErwinia subsp. for either of these response variables, and the cultivar ×Erwinia subsp., cultivar × temperature and cultivar ×Erwinia subsp. × temperature interactions were not significant. However, there were significant differences between the incubation temperatures, and theErwinia subsp. × temperature interaction was significant for both response variables. Estimates of sample sizes necessary to find a 20% difference in main effect and interaction effect means were calculated.     

Resistance toErwinia carotovora subsp.atroseptica in second backcross populations of somatic hybrids betweenSolanum tuberosum andS. brevidens

Summary Eleven clones obtained from a cross between cv. Katahdin and fusion products betweenSolanum tuberosum and the non tuber-bearing speciesS. brevidens, were backcrossed again withS. tuberosum (clone AR80-127-5). Small tubers harvested from 583 seedlings of these second backcross (BC2) populations were screened for tuber tissue resistance toErwinia carotovora subsp.atroseptica under aerobic conditions. After multiplication in the field. BC2 clones again were screened for soft rot resistance under aerobic as well as anaerobic conditions. In general, the resistance of the BC2 populations was reduced in comparison with the BC1. Variation for resistance was found within the BC2 populations, but there was no correlation between the results of the three tests performed. Four BC2 populations were also screenned for resistance to blackleg in the field. Significant differences were found between populations for mean percentage of diseased plants, but these differences could not be explained by the resistance of the parental clones.     

Leptines and other glycoalkaloids in tetraploidSolanum tuberosum xSolanum chacoense F2 Hybrid and Backcross Families

The leptine glycoalkaloids fromSolanum chacoense Bitter are potential resistance factors to Colorado potato beetle,Leptinotarsa decemlineata Say. A selected sample of 17 F₁ hybrids (4x) from crosses of a 4xS. chacoense clone that synthesizes leptines (8380-1) with 4xS. tuberosum L. lines were selfed to generate F₂ hybrids (4x) and were also backcrossed toS. tuberosum. Glycoalkaloid (GA) content in foliage and tubers was measured for 786 sibling genotypes from the two generations in the field at Beltsville, Maryland. Leptines were found in the foliage of all F₂ hybrids and in 98% of the backcross genotypes. Leptines were not detected in tubers from either generation. Foliage concentrations of leptines for the F₂ hybrids averaged 156 mg/100g fresh weight (FW). The proportion of the total GA content consisting of leptines averaged 42%. By backcrossing the F₁ hybrids toS. tuberosum the average concentration of leptines was reduced to 44 mg/100g FW. The proportion of leptines averaged 16%. The tuber contents of solanine plus chaconine glycoalkaloids averaged 52 mg/100g FW for the F₂ hybrids and 27 mg/100g for the backcross genotypes.