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.     

Foliar solanidine glycoside levels inSolanum tuberosum populations selected for potato leafhopper resistance

Solanum tuberosum L. Gp. Tuberosum populations that had been recurrently selected for resistance to potato leafhopper,Empoascafabae (Harris), were tested for levels of foliar solanidine glycosides. Leafhopper infestation level was 57% lower than for the unselected population after seven generations of selection. The mean concentrations of foliar solanidine glycosides significantly (P<05) increased from 40 mg/100 g fresh weight (fw) in the unselected population to 65 mg/100 g fw in the population developed from seven generations of selection for leafhopper resistance. The linear relationship between the level of leafhopper infestation and the concentration of foliar solanidine glycosides among population means was significant (P<01) and negative (r=?0.85).     

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.     

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.     

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.     

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.     

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.     

Segregation of leptine glycoalkaloids and resistance to Colorado potato beetle (Leptinotarsa decemlineata (Say)) in F2Solanum tuberosum (4x) ×S. chacoense (4x) potato progenies

Solanum chacoense Bitter is resistant to the Colorado potato beetle (CPB),Leptinotarsa decemlineata (Say). Resistance has been associated with the presence of a rare class of glycoalkaloids, the leptines. In this study, seven tetraploid, F2S. tuberosum xS. chacoense families were evaluated for foliar production of leptines I and II, leptinines I and II, and α-solanine and α-chaconine; and screened for resistance to CPB in the laboratory and field. Resistance was correlated with the concentrations of glycoalkaloids on a family and an individual basis. Leptine concentrations ranged from undetectable to a high of 18.0 mg/g dry weight. All of the progeny produced solanine and chaconine. Family 9623 had the highest mean leptine concentration and the lowest mean leaf disk feeding and CPB defoliation levels. Family 9616 had the lowest mean glycoalkaloid concentration and ranked as one of the most susceptible families. Regression analyses of solanine + chaconine, leptine I and II, and leptinine I and II foliar concentrations versus leaf disk consumption and field defoliation revealed that only increased foliar levels of leptines resulted in decreased CPB feeding. The regression models for leptines versus leaf disk consumption and field defoliation were highly significant, accounting for 17% and 26% of the variation in consumption and defoliation, respectively. To the best of our knowledge, this is the first work reporting the impact of leptine and leptinine concentrations on CPB feeding in tetraploid,S. tuberosum xS. chacoense potato hybrids. Results are discussed within the context of breeding for resistance to CPB.     

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.     

Determination of Glycoalkaloids in Potatoes and Potato Products by Microwave Assisted Extraction

Potato glycoalkaloids can reach levels that are harmful to human health. A rapid and reliable microwave assisted extraction method for quantitative analysis of α-solanine and α-chaconine content in raw potato and potato based products is presented. A chemical microwave was used to determine optimal temperature and pressure conditions for the extraction of α-solanine and α-chaconine from Idaho grown tubers and six commercially available mashed potato products. Recovery efficiency of glycoalkaloids was 37% greater by microwave assisted extraction (19.92 mg/kg glycoalkaloid) as compared to conventional solid/liquid methods (12.51 mg/kg glycoalkaloid). Optimal extraction of glycoalkaloids from potato samples dissolved in methanol was achieved using a microwave reactor set to 90 °C for ten minutes. The interior of Idaho grown tubers was determined to contain lower levels of glycoalkaloids (19.92 mg/kg dry weight; 6.5 ± 1.78 mg α-solanine and 13.40 ± 1.65 mg α-chaconine), as compared to commercial potato products (33.86–81.59 mg/kg).     

Varietal and environmental effect on the glycoalkaloid content of potato (Solanum tuberosum L.)

The tubers of 14 potato varieties were analysed for glycoalkaloids. The level of glycoalkaloids in tubers of all the varieties analysed are within the safe limits for human consumption. The peels of tuber contained about 60–70% of the total glycoalkaloids present in the whole tuber. The levels of glycoalkaloids in leaves and tubers were correlated (r=0.865). There was a significant increase in the content of glycoalkaloids in peels of tubers exposed to sunlight. Glycoalkaloid contents increased at the rate of 1.9mg/100g fresh weight per day in peels of Kufri Jyoti tubers exposed to diffused sunlight.     

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.     

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%.     

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.     

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.     

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.     

Stability,cross-resistance and fitness costs of resistance to emamectin benzoate in a re-selected field population of the beet armyworm,Spodoptera exigua (Lepidoptera: Noctuidae)

Spodoptera exigua (Hübner) has a worldwide distribution with a high capacity for damaging a wide range of food, forage and fiber crops. It has been reported extensively from all over the world that populations of this pest species have developed field resistance against many insecticides. The objectives of this study were to determine whether an emamectin benzoate resistant field population of S. exigua re-selected with emamectin benzoate in the laboratory (Ema-SEL) showed cross-resistance to other insecticides, whether resistance was stable under laboratory conditions, and whether there were fitness costs associated with emamectin benzoate resistance. Bioassays at G₁ for the field population, gave resistance ratios (RRs) of 220, 149 and 38-fold for emamectin benzoate, spinosad and lufenuron, respectively, compared with a susceptible laboratory population (Lab-PK). Resistance ratios were increased by 526-fold and 6-fold compared with Lab-PK and the unselected field population (Ema-UNSEL, G₆), respectively after selection with emamectin benzoate (Ema-SEL) for five generations (G₆). Selection with emamectin benzoate had no apparent effect on susceptibility of Ema-SEL to spinosad and lufenuron, instead toxicity to the latter insecticides reduced, suggesting there was no cross-resistance between these compounds. Analysis of various life history traits suggested that the Ema-SEL population had a lower overall fitness (0.38) compared with the Lab-PK (1.0). Lack of cross-resistance and the apparent instability of resistance to emamectin benzoate suggest that spinosad and lufenuron are suitable alternatives for use with emamectin benzoate in resistance management. In addition, the high relative fitness costs observed suggests that emamectin benzoate-resistant insects are at a considerable disadvantage to susceptible populations in the absence of selection pressure although this remains to be tested under field conditions.     

Glycoalkaloids in potato tubers grown under controlled environments

Tuber content of α-solanine, α-chaconine, and total glycoalkaloids (TGA) was determined for the potato cultivars, Norland, Russet Burbank, and Denali grown under different environmental conditions in growth chambers. The lowest TGA concentrations (0.30 to 0.35 mg g¹ dry tissue) were found in the cv. Norland with 400 μnol m^-2 s^-1 photosynthetic photon flux (PPF), 12 h day length, 16 C temperature, and 350 μmol mol^-1 carbon dioxide. The ratio of α-chaconine to α-solanine was close to 60:40 under all growing conditions, except that it was 50:50 under the low temperature of 12 C. Cultivars responded similarly to environmental conditions although TGA was about 20% greater in cv. Russet Burbank and about 30% greater in Denali compared to Norland. The largest changes in TGA occurred with changes in temperature. In comparison to 16 C, TGA were 40% greater at 12 C, 80% greater at 20 C, and 125% greater at 24 C (0.70 mgg ^-1 dry weight). The TGA concentratation increased from 10 to 25% with an increase in light from 400 to 800 μmol m^-2 s^-1 PPF for all three cultivars. TGA increased 20% with extension of the day length from 12 to 24 hr and also increased 20% when carbon dioxide was increased from 350 to 1000 umol mol^-1. TGA concentrations were not influenced by changes in relative humidity from 50 to 80%. TGA concentrations decreased only slightly in harvests made from 9 to 21 weeks after planting. Variations in TGA among the different growing conditions and cultivars were below 20 mg/100 g fresh weight (= 1.0 mg g^-1 dry weight) recognized as the upper concentration for food safety. However the results suggest that TGA should be considered when potatoes are grown at temperatures above 20 C.     

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.     

Cross-resistance, inheritance and stability of resistance to acetamiprid in cotton whitefly, Bemisia tabaci Genn (Hemiptera: Aleyrodidae)

The cotton whitefly Bemisia tabaci, (Genn.) is an important pest of field crops, vegetables and ornamentals worldwide. Neonicotinoids are considered an important group of insecticides being used against B. tabaci for several years. B. tabaci has developed resistance to some of the compounds of the group. This study was designed to investigate if the selection of B. tabaci with acetamiprid would give a broad-spectrum of cross-resistance and to genetically classify the resistance. At G₁ a low level of resistance to acetamiprid, imidacloprid, thiamethoxam, thiacloprid and nitenpyram was observed with resistance ratios of 3-fold, 8-, 9-, 6- and 5-fold, respectively, compared with a laboratory susceptible population. After selection for eight generations with acetamiprid, resistance to acetamiprid increased to 118-fold compared with the laboratory susceptible population. Selection also increased resistance to imidacloprid, thiamethoxam, thiacloprid, nitenpyram, endosulfan and bifenthrin but no change in susceptibility to fipronil was observed. Furthermore resistance in a field population was stable in the absence of acetamiprid selection pressure. Genetic crosses between resistant and susceptible populations indicated autosomal and incompletely recessive resistance. Further genetic analysis suggested that resistance could be controlled by a single factor. The high level of cross-resistance and stability of incomplete resistance in the field population is of some concern. However, lack of cross-resistance between acetamiprid and fipronil or unstable resistance in the resistant population could provide options to use alternative products which could reduce acetamiprid selection pressure.