Recurrent selection for potato leafhopper resistance in potato,Solanum tuberosum L. Gp. Tuberosum

ASolanum tuberosum L. Gp. Tuberosum population was recurrently selected for resistance to potato leafhopper (PLH) (Empoasca fabae, Harris). Primary selection was for low levels of nymphal infestation under natural infestation in field tests. The selection procedures allowed completion of a selection generation per year. The original population and seven selection-generation populations were tested for PLH resistance for three years at two locations. Averaged over six tests, seven generations of selection reduced the level of nymphal infestation by 71% and severity of plant damage (hopperburn) by 45%. The rate of reduction slowed in later generations.     

Glycoalkaloid contents in tubers fromSolanum tuberosum populations selected for potato leafhopper resistance

Solanum tuberosum L. Gp. Tuberosum populations that had been improved for resistance to potato leafhopper,Empoasca fabae (Harris), by recurrent selection, were analyzed for tuber contents of the glycoalkaloids α-solanine and α-chaconine. The mean content of both glycoalkaloids was significantly higher in the more resistant populations compared with the unselected population. In a population generated from seven cycles of selection for resistance, the mean content of solanine + chaconine was 9.5 mg/100 g fresh weight compared with 4.0 mg/100 g in the original, unselected population; a 137% increase. Kennebec, in the same experiment, averaged 10.9 mg/100 g; Katahdin averaged 4.6 mg/100 g.     

Total foliar glycoalkaloids and resistance of wild potato species toEmpoasca fabae (Harris)

Levels of total glycoalkaloids (TGA) in foliage of 10 wild, tuberbearingSolanum (Tourn.) L. species differentially resistant to infestation by the potato leafhopper,Empoasca fabae (Harris), were determined. Levels of TGA ranged from a high of 688 mg/100 g fresh wt. in a resistant species,S. polyadenium Greenm. to a low of 13 mg/100 g fresh wt. in a susceptible species,S. bulbocastanum Dun. Foliar concentration of TGA and nymphal infestation by the potato leafhopper were highly correlated (r = ?0.75, p = 0.01). The significant correlation of TGA levels and potato leafhopper resistance suggests that foliar TGA may be a significant factor in the defense of wild potato species against this pest.     

Selection for resistance to potato leafhopper in potatoes. II. Progress after two cycles of phenotypic recurrent selection

ASolatium tuberosum L. gp.tuberosum population was subjected to 2 cycles of phenotypic recurrent selection for resistance to the potato leafhopper,Empoasca fabae (Harris), as expressed by low levels of nymph infestation. Samples of lines from the original population and from the cycle 2 population were tested for infestation levels at 2 locations for 2 years. Infestation in cycle 2 lines was 54% lower than that in lines from the original, unselected population.     

Selection for resistance to potato leafhopper in potatoes: III. Comparison of two selection procedures

A comparison was made of two breeding procedures for recurrently selecting potato,Solanum tuberosum L., for resistance to infestation by potato leafhopper,Empoasca fabae (Harris). On the basis of selection advance per year, the more efficient procedure involved randomly intermating a group of clones, planting seedling tubers in a field test to measure resistance to infestation, and then bulking the seed from the most resistant clones to complete the selection cycle in one year. A population developed by five cycles of selection (five years) for resistance to infestation using this procedure resulted in a decrease of 57% in the level of infestation from the original population.     

Tuber yields and specific gravities inSolanum tuberosum populations recurrently selected for resistance to potato leafhopper

Solanum tuberosum genotypes from the 3rd, 5th, and 7th generations of recurrent selection for resistance to potato leafhopper were compared with genotypes from the base, unselected population for tuber yield and specific gravity over a five year period. Mean tuber yields of the genotypes in the 3rd and 5th generations were equal to the yield of those in the base population. The mean tuber yield in the 7th generation was significantly lower than the mean in the base population and the means in the other selection generations. However, the mean tuber specific gravities of the 5th and 7th generation genotypes were slightly higher than the mean in the base population. The variability of tuber yield among genotypes in the 7th selection generation was lower than the variability in the base population and in the 3rd and 5th selection generations. It was concluded that there was moderate genetic potential for selecting genotypes with resistance to potato leafhopper that had acceptable tuber yields in the 3rd and 5th selection generations, but by the 7th generation the potential was considerably lower. The genetic potential for selecting resistant genotypes with acceptable tuber specific gravities remained about the same in each of the selection generations.     

Genetic variation in a population of tetraploid potatoes: Response to the potato leafhopper and the potato flea beetle

Progenies from a group of tetraploid parental clones from the USDA potato breeding program were used to investigate variation in resistance to the potato leafhopper,Empoasca fabae (Harris), and the potato flea beetle,Epitrix cucumeris (Harris). The study utilized two mating designs: i) selfing and testing both the parents and the S₁ progeny; and ii) nine clones, used as males, were each crossed to three different clones, and progenies from the resulting families were tested. Statistically significant differences between families were measured in each test for both insects. Non-additive genetic variance was larger than additive genetic variance in progeny reaction to leafhopper infestation and hopperburin, but was smaller in progeny reaction to flea beetles. Environmental variation contributed heavily to the total variation of plant reaction to both insect species. Selecting individuals was indicated to be slightly more effective than selecting males on half-sib progeny performance but not as effective as selecting clones on S₁ progeny performance. Because of the large environmental variance and small additive variance for both the leafhopper and flea beetle, slow progress in increasing the level of resistance to these two species in this sample population was predicted. Resistance to leafhopper infestation was genetically quite highly correlated (positive) with resistance to hopperburn, but phenotypically the correlation was considerably smaller. Negative genotypic, phenotypic, and environmental correlations between leafhopper infestation and flea beetle infestation suggest that selecting for resistance to one of these species, in the population sampled for these tests, would tend to increase susceptibility to the other.     

Segregation of leptines and other glycoalkaloids inSolanum tuberosum (4x) ×S. chacoense (4x) crosses

A clone, 8380-1, selected fromSolanum chacoense (PI 458310) for its high foliage content of the leptine glycoalkaloids, a factor in resistance to Colorado potato beetle, was doubled in chromosome number from 2n=2x=24 to 2n=4x=48. Three 4x clones were crossed with sixS. tuberosum (4x) clones. Foliage glycoalkaloid contents were measured for 452 F1 hybrids from 15 crosses. The 4xchacoense parental clones were not different in respect to glycoalkaloid contents and were similar to the original 2x clone. All F1 hybrids synthesized foliage leptines ranging from 9 to 369 mg/100 g fresh weight (fw) with a mean content of 113 mg/100 g fw. The proportion of leptines in the total glycoalkaloid content ranged from 1% to 62% with a mean of 25%. The 4xchacoense parent mean leptine content was 1482 mg/ 100 g fw which was 90% of the total glycoalkaloid content. Tubers from 136 hybrids and the threechacoense parental clones were tested for glycoalkaloid contents. The tuber solanine + chaconine contents of the 136 hybrids ranged from 30 to 180 mg/100 g fw with a mean of 79. The mean tuber content of the threechacoense parental clones was 157 mg/100 g fw. Leptines were not found in any of the tubers.     

Glycoalkaloid content in tubers of hybrid and backcross populations from aSolanum tuberosumx s. chacoense cross

The glycoalkaloid contents of tubers were measured in hybrid and back-cross populations derived from a cross ofSolarium chacoense Bitter (PI 472810, 2n=2x=24) withSolanum tuberosum L. (2n=4x=48). In tubers sampled from near the median size in each population and parent, the mean total glycoalkaloid (TGA) contents of theS. chacoense andS. tuberosum parent genotypes, and the F₂ and F₄ hybrid genotypes were 245, 4, 39, and 35 mg/ 100 g fresh weight (fw), respectively. Backcrossing the F₂ genotypes toS. tuberosum reduced the mean TGA content to 15 mg/100 g fw. A second backcross further reduced the content to 9 mg/100 g fw. However, some genotypes in both backcross populations (25% and 10%) had TGA contents above the generally accepted upper limit of 20 mg/100 g fw. Tuber size was significantly smaller in theS. chacoense parent and the hybrid populations compared with the size in theS. tuberosum parent and backcross populations. In a second sampling of tubers from each population, intentionally selected to equalize tuber size among populations, the mean tuber size for populations was reduced by 79%. The mean population TGA content increased by 44%. However, the differences and rankings among parent, hybrid, and backcross populations were the same for both tuber samplings.     

Glycoalkaloids as pest resistance factors

Potato glycoalkaloids, a class of steroid glycosides, possess antimicrobial and pesticidal properties. These phytochemicals have been implicated as natural resistance factors inSolanum species against the Colorado potato beetle,Leptinotarsa decemlineata Say, and the potato leafhopper,Empoasca fabae (Harris). Adverse effects of these secondary metabolites on insect behavioral and developmental biology are described, as well as considerations in manipulation of foliar glycoalkaloids in breeding for varietal resistance to insects.     

Performance of populations derived by selecting for resistance to potato leafhopper in A 4XSolanum Tuberosum X 2XSolanum Chacoense cross

Intercross and backcross populations derived from aSolanum tuberosum L. (2n = 2x = 48) ×Solanum chacoense Bitt. (2n = 2x = 48) cross by selecting for resistance to potato leafhopper (PLH) (Empoasca fabae Harris), were tested in the field. In F2 and F4 populations, the levels of nymphal infestation and leaf necrosis from PLH feeding were considerably lower and tuber yields were much lower compared with those measured on the susceptible cultivar Kennebec. No differences were found between the two populations. Backcrosses of the F2 and F4 populations toS. tuberosum decreased resistance to PLH. Resistance to infestation was still greater than that of Kennebec; severity of leaf necrosis equaled that of Kennebec. Tuber yields increased, but were less than Kennebec’s yield. Foliage types shifted closer to those ofS. tuberosum. Three generations of selection and intercrossing within the backcross populations did not change the level of resistance to PLH but did tend to lower tuber yields. Foliage types remained primarilyS. tuberosum in appearance although distinctive types segregated with increased frequency. A second backcross toS. tuberosum did not change the level of resistance to PLH or the level of tuber yield relative to those of the first backcross or Kennebec.     

Selection for resistance to potato leafhopper in potatoes I.selection methods

A randomly mated potato (Solarium tuberosum L.) population (3 generations, without selection) derived from clones from several breeding programs, displayed considerable resistance in the field to potato leafhopper,Empoasca fabae (Harris). Leafhopper population levels on 161 clones ranged from 2 to 28 nymphs per 45-sec count (mean = 12), whereas a susceptible check averaged 20 nymphs per count. Clones selected for low leafhopper counts in 1971 were compared with the population in 1972 to evaluate the effectiveness of selecting from small plots (three-hill) with minimum replication. The selected clones averaged 30% fewer leafhoppers than the population average in 1971 and 10% fewer in 1972. The average heritability (broad sense) for one plot per clone per location, two locations, and two 45-sec nymph counts per plot equaled 36%.     

Glandular pubescence,glycoalkaloid composition,and resistance to the green peach aphid,potato leafhopper,and potato fleabeetle inSolanum berthaultii

Fifteen accessions ofSolarium berthaultii Hawkes andS. berthaultii xS. tarijense Hawkes were assessed for resistance to field infestations of the green peach aphid,Myzus persicae (Sulzer), potato leafhopper,Empoasca fabae (Harris), and potato fleabeetle,Epitrix cucumeris (Harris). Accessions bearing both Type A and Type B glandular trichomes were much more resistant to the green peach aphid and potato leafhopper than accessions bearing Type A hairs alone. All accessions had significantly smaller populations of these 3 pests than S.tuberosum cultivars. Total glycoalkaloid (TGA) content of foliage and tubers was not correlated with insect populations. Foliar TGA levels of field-grown plants varied among accessions, ranging from < 2–240 mg/100 g fresh wt. Solasonine and solamargine were the major foliar glycoalkaloids while solamarines predominated in tubers.     

Glycoalkaloid content inSolanum species and hybrids from a breeding program for resistance to late blight (Phytophthora infestans)

As part of an effort to study the relationship between the “glycoalkaloid trait” and genetic resistance to late blight (Phytophthora infestans), glycoalkaloid content in tuber and foliar tissues from a wide genetic background withinSolanum, includingS. demissum, S. iopetalum and 15 hybrids selected from a backcrossing breeding scheme was determined. Analysis of variance indicated significant genotypic effect on total glycoalkaloid, solanidine, α-solanine, and α-chaconine content in both tubers and leaves. Tubers from wild potato species commonly contain glycoalkaloids in concentrations that exceed international health regulations for human consumption (20 mg/100 g fresh weight). In this study,S. demissum andS. iopetalum were highest in total tuber glycoalkaloids among all materials tested, with 70.4 and 76.2 mg/100 g fresh weight, respectively. In contrast, both commercial cultivars had the lowest concentration, both below the safety limit. Solanine was more abundant than chaconine in all but one genotype. All hybrids were intermediate to low in total glycoalkaloids. Except for the two wild species, glycoalkaloid concentration in leaves of all genotypes studied was at least twice that in tubers, with glycosylated forms accounting for more than 80% total glycoalkaloid content. Correlation between tuber and foliage alkaloids was poor. In view of the observed field resistance to late blight, it was concluded that tuber glycoalkaloid content may not be responsible for such resistance.     

High level of metaflumizone resistance and multiple insecticide resistance in field populations of Spodoptera exigua (Lepidoptera: Noctuidae) in Guangdong Province,China

Spodoptera exigua is a phytophagous pest that causes critical economic losses in vegetable crops, and insecticides are commonly used against it in vegetable growing areas. However, excessive and frequent applications of insecticides cause resistance in S. exigua. The current resistance in field populations of S. exigua collected from Huizhou, Guangdong Province, China to 12 insecticides was investigated. S. exiguahad developed very high resistance to lambda cyhalothrin (2925- to 3449-fold), chlorpyrifos (>1786-fold), emamectin benzoate (174- to 867-fold), and metaflumizone (60.3- to 942-fold). High resistance to tebufenozide (51.5- to 75.4-fold) and chlorfluazuron (60.4- to 63.0-fold) was also found. Synergism assays revealed that the resistance to metaflumizone and lambda cyhalothrin was associated with esterase and microsomal oxidases, respectively. The resistance to emamectin benzoate was not affected by detoxification enzymes inhibitors and might be conferred by other mechanisms. The selection of the field population by metaflumizone for 10generations in the laboratory resulted in a 6.1-fold increase in metaflumizone resistance but did not lead to increases in resistance to other insecticides. After metaflumizone selection, susceptibilities to spinosad and endosulfan did not change, and the susceptibilities to indoxacarb, methomyl, pyridalyl, tebufenozide, chlorfluazuron, emamectin benzoate and lambda cyhalothrin decreased slightly. However, no statistically significant differences in the resistance levels were observed among the selected population, its starting strain and the unselected strain. The resistance to chlorantraniliprole noticeably decreased in unselected strain and the strain subjected to selection for 10 generations compared with their starting strains. Lack of cross-resistance to tested insecticides suggested the involvement of multiple mechanisms of resistance and the need for wise application of these insecticides for the management of S. exigua.     

Tolerance to potato leafhopper inSolanum tuberosum L. gp. Tuberosum clones

We identified clones inSolanum tuberosum L. gp. Tuberosum breeding populations that, although susceptible to infestation by potato leafhopper,Empoasca fabae (Harris), exhibited a tolerance which was manifested by the ability to produce higher than expected tuber yields when heavily infested with leafhoppers. Such tolerance was noted in a number of clones in varying degrees, but one, B6930-1, was exceptional. It sustained only a 19% loss in tuber yield compared to ‘Katahdin,’ a susceptible cultivar, that averaged 33% lower tuber yields under similar infestation levels. We studied the seasonal growth patterns of this clone and compared them to those of Katahdin, and B6705-10, a breeding clone with known resistance to leafhopper infestation. We found that much of B6930-l’s tolerance results from its early, vigorous vine growth and its early and rapid rate of tuber bulking, enabling this clone to escape some of the effects of leafhopper feeding, whereas Katahdin was usually just beginning to tuberize at the height of leafhopper infestation.     

Effect of plant age on potato leafhopper infestation of resistant and susceptible potato clones

The effect of age of resistant potato clones,Solarium tuberosum gp.tuberosum, on infestation by potato leafhopper,Empoasca fabae (Harris), was investigated by varying the date of planting in a naturally infested fieldtest Significant differences (P < .05) in infestation levels were found among clones and among dates of planting. The responses of resistant and susceptible clones to infestation differed with date of planting. But lower levels of infestation occurred on all clones in the first 2–3 weeks of leafhopper migration on the early plantings, indicating that plant age in potatoes is a factor in host preference of potato leafhopper     

Resistance of Spodoptera litura (Lepidoptera: Noctuidae) to profenofos: Relative fitness and cross resistance

Profenofos, an acetylcholine esterase inhibitor insecticide, has been used for the management of various lepidopteron pests of many crops in Pakistan. In the present study, we investigated the impact of insecticide resistance on fitness cost in Spodoptera litura, and evaluated cross resistance to other insecticides. The effect of profenofos on different life history parameters including survival rate, female ratio, fecundity and hatchability, intrinsic rate of population increase and biotic potential was determined. Significant differences associated with fitness costs were revealed. A field collected S. litura strain was selected by exposure to profenofos in the laboratory (Profen-SEL) and after 14 generations of selection it developed a 52-fold resistance to this insecticide. The Profen-SEL strain showed high cross-resistance to chlorpyrifos (62-fold), but very low to no cross resistance to lambda-cyhalothrin (2.34-fold) or methomyl (0.80-fold), respectively. The resistant strain had a relative fitness of 0.38 with a low larval survival rate, longer larval duration, longer male pupal duration, longer development time, low emergence rate of healthy adults, fecundity and hatchability compared with an unselected strain. The intrinsic rate of natural population increase, mean relative growth rate and biotic potential were lower for the selected strain compared with an unselected strain. Development of resistance may cause fitness costs for the resistant strain. This study provided important information for understanding profenofos resistance and facilitating a better strategy for the management of resistance.     

Glycoalkaloid content in foliage of hybrid and backcross populations from aSolanum tuberosum X S. chacoense cross

The glycoalkaloid contents of foliage were measured in populations derived from a cross ofSolanum chacoense Bitter (PI 472810) with a composite sample of genotypes from a randomly matedSolanum tuberosum L. population. The mean total glycoalkaloid contents of the parent PI 472810 andtuberosum populations, and the F₂ and F₄ populations were 856, 121, 286, and 279 mg/100g fresh weight, respectively. The total glycoalkaloid content of the first backcross population, F₂ xtuberosum, was 290 mg/100 g, close to the content of the F₂. In a second backcross totuberosum, the mean total content was 148 mg/100 g, close to the content of the parenttuberosum population. Only the glycoalkaloids α-solanine and α-chaconine were found in PI 472810, whereas in the parenttuberosum, hybrid, and backcross populations, the glycoalkaloids α & β-solamarine also were found in a small number of genotypes. The ratios of chaconine to solanine contents were significantly (P≤.05) different in the parent PI 472810 andtuberosum populations, 2.55 and 2.12, respectively. The ratios in the F₂ and F₄ populations were similar to that of PI 472810, whereas the ratios in the backcross populations were closer to that of thetuberosum parent. The levels of foliage glycoalkaloids in the hybrid and backcross populations paralleled the levels of potato leafhopper,Empoasca fabae Harris, resistance measured in a previous study.     

Evaluation of wild tuber-bearingSolanum accessions for foliar glycoalkaloid level and composition

Six hundred forty-five accessions of 70Solanum species and six natural hybrids were screened for foliar glycoalkaloids. Total foliar glycoalkaloid (TFGA) level and glycoalkaloid composition of one or more accessions of each species were determined by TLC and GLC analyses. Of the more than 12 different glycoalkaloids found in the wild species in this survey, solanine and chaconine account for more than 60% of the TFGA found in the foliage of these plants. There was wide variation in the total glycoalkaloid levels among the species.S. neocardenasii had the highest average TFGA value of the 70 species, 222 mg/100 g fresh wt (222 mg %) and an accession ofS. chacoense had the highest TFGA value recorded in the survey, 486 mg %. Of the 70 species, only 11 had average TFGA levels of more than 100 mg %. However, one or more accessions in 27 of the species synthesized more than 100 mg %. Forty-five species synthesized an average level of less than 50 mg %. The level usually found in the foliage of commercial potato cultivars is about 50 mg %. Therefore, most of theSolanum species in the potato germplasm collection can be considered low glycoalkaloid species.