Determination of the glycoalkaloids α-solanine and α-chaconine levels in 18 varieties of potato (Solarium tuberosum L.) grown in Oman

Summary The glycoalkaloid content in 18 potato varieties grown in Oman was measured by HPLC using peel and flesh tissues. The total glycoalkaloid content of the tubers was found to be variety dependent and varied greatly between the 18 varieties. The total glycoalkaloid in the tubers of seven varieties was within the safe limit of ≤200 mg kg^{− 1} FW recommended for food safety, whereas the remaining 11 varieties exceeded this upper limit. The principal glycoalkaloids α-chaconine and σ-solanine were present in higher concentration in the peel than in the flesh of all varieties with the former being the principal alkaloid in the peel. The flesh of all of the 18 potato varieties contained total glycoalkaloid levels lower than the upper safety limit. The leaves of the 18 potato varieties were also analyzed for glycoalkaloids. The levels of glycoalkaloids in expanding leaves were higher than that of expanded leaves with the exception of six varieties. The principal glycoalkaloid in expanding and expanded leaves was α-chaconine. No correlation was observed between the glycoalkaloids levels of in the leaves and 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.     

Production of phytoalexins, glycoalkaloids and phenolics in leaves and tubers of potato cultivars with different degrees of field resistance after infection withPhytophthora infestans

Summary The kinetics of accumulation of phytoalexins, glycoalkaloids and phenolics was studied in two potato cultivars differing in their degrees of field resistance when infected withPhytophthora infestans. Tuber slices and leaves of cvs Pampeana INTA (high degree of field resistance, free of R genes) and Bintje (susceptible) were infected with race C (complex race 1, 3, 5, 7, 11) ofPhytophthora infestans. Phytoalexins and phenolics accumulated in tuber and leaf tissues which had been inoculated. The levels of these compounds in the susceptible cv. Bintje were relatively low and similar to those found before inoculation. Leaves of cv. Pampeana INTA had a very high glycoalkaloid content, suggesting that glycoalkaloids may play a role in protection of leaves against the fungus. However, we could find no correlation between resistance and glycoalkaloid content of tubers. Our results suggest a major role of phytoalexins, phenolics and glycoalkaloids in the complex mechanisms of field resistance.     

Glycoalkaloids and resistance to the Colorado potato beetle inSolanum chacoense Bitter

Resistance of 20 F₂ Solarium chacoense Bitter clones to the Colorado potato beetle,Leptinotarsa decemlineata (Say), was measured in a field test. Levels of total glycoalkaloids (TGA) and composición of the glycoalkaloid mixtures in foliage of the clones were also determined. Clones with either commersonine or dehydrocommersonine as the major foliar glycoalkaloid were significantly more resistant (lower damage ratings, fewer larvae and adult insects) than clones with solanine and chaconine. Damage ratings were negatively correlated (r = -0.67, p = 0.01) with foliar TGA levels. The results indicate that the types of glycoalkaloids present in the foliage ofS. chacoense may be as important as the level of TGA in limiting damage and numbers of insects.     

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.     

Resistance to late blight and soft rot in six potato progenies and glycoalkaloid contents in the tubers

Glycoalkaloids are anti-nutritional compounds commonly found in wildSolanum species used as resistance sources to major potato pathogens. It is therefore important for breeding purposes to know whether selecting for resistance using such species necessarily selects also for high glycoalkaloid contents in the tubers. To test this hypothesis, we used six partial progenies from crosses betweenSolanum tuberosum and accessions ofS. andigena, S. berthaultii, S. phureja, andS. vernei to investigate the possible correlation between resistance toPhytophthora infestans and/or toErwinia carotovora subsp.atroseptica and the concentration of glycoalkaloids in tubers. Concentrations of α-solanine and α-chaconine in the tubers segregated in each progeny, as did resistance to each pathogen. Some, but not all, clones from each progeny showed hypersensitive reactions to the isolate ofP. infestans used. Furthermore, clones within each progeny also differed for components of partial resistance toP. infestans, suggesting that all four wild species could be used as sources of both race-specific and partial resistance to late blight. With the exception of low, but statistically significant, correlations between concentration of α-solanine and two late blight resistance components (incubation period and spore production per unit lesion area) in progenies derived fromS. vernei, and despite a trend towards higher glycoalkaloid concentrations in the tubers of the clones most resistant to soft rot within progenies derived fromS. berthaultii andS. vernei, no consistent relationship between resistance to either disease and concentrations of α-solanine and/or α-chaconine was observed. These results indicate that neither race-specific nor partial resistance to late blight and soft rot in the accessions used as progenitors of resistance depend on high solanine or chaconine concentrations. These resistance sources could thus prove useful in breeding programs for improved behaviour againstP. infestans and/orE. carotovora.     

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).     

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.     

Change in Some Glycoalkaloids of Potato under Different Storage Regimes

The potato (Solanum tuberosum L.) is an important food crop worldwide. Potato tubers can be stored to provide continuity of supply between production seasons, but it is important that they be stored under appropriate conditions as incorrect storage will result in deterioration in end user quality and may increase glycoalkaloid levels. We have investigated the effects of types of household storage on potato glycoalkaloid content (total glycoalkaloids [TGA]; α-solanine; α-chaconine) in Turkey. Tubers of potato cultivars (cvs.) Agria and Bettina were stored under four types of storage conditions (indirect sunlight for 10 h per day and dark storage for the remaining 14 h per day; storage under continuous fluorescent light; storage in constant darkness; storage in the dark in a refrigerator) for 56 days. Samples of tubers were taken at the beginning of the storage period and after 14, 28 and 56 days of storage for tuber glycoalkaloid measurement. Tubers stored in the three light environments showed an increase in glycoalkaloid levels; however, none of the cv. Agria tubers reached the critical level of 200 mg/kg tuber. On day 56 the TGA level of cv. Bettina tubers stored under fluorescent light reached 234.31 mg/kg.     

Glycoalkaloid composition of potatoes: Diversity and biological implications

Glycoalkaloids are important potato quality factors. Tuber-bearingSolanum species exhibit wide quantitative and qualitative diversity in glycoalkaloid contents. Some species, of interest to breeders, contain extremely high total glycoalkaloid (TGA) levels and/or unusual types of glycoalkaloids of ill-defined or unknown toxic and teratogenic properties. Characterisation of glycoalkaloid content is a dynamic and, in some cases, controversial subject. Problems associated with analytical methodology and, to a lesser extent, with environmental effects and intraspecific heterogeneity have probably been the major causes of disagreements in the literature. Recent methodological developments have facilitated more accurate and sensitive measurements of TGA levels and of individual glycoalkaloid compositions. Such measurements should be used to analyze parental material and selected hybrid progeny in breeding programs. This is particularly important when certain wildSolanum species are being exploited as undesirable levels and types of glycoalkaloid can be genetically transmitted from the wild species to hybrid progeny. Absence of glycoalkaloid assessment from a breeding program can result in wasted effort, time and money.     

The effect of light intensity on Colorado potato beetle resistance and foliar glycoalkaloid concentration of fourSolanum chacoense clones

Controlled environment experiments were used to determine the effect of light intensity on levels of glycoalkaloids and resistance to the Colorado potato beetle (CPB),Leptinotarsa decemlineata Say, inSolanum chacoense Bitter clones that synthesized acetylated glycoalkaloids, the leptines. Plants were grown under two light intensities, 50 or 250 ?mol m^-2S^-1, and bioassayed for CPB resistance by placing neonate larvae on excised foliage. Larval development stage and mortality were recorded after four days. The leptine-synthesizing clones responded to the increase in light intensity with a 2–4 fold increase in level of foliar leptines. Larval development rate on foliage from plants grown under high light intensity was reduced and larval mortality increased. Results showed a significant (p>0.05) effect of light on glycoalkaloid synthesis and CPB resistance.     

Photoinduction of glycoalkaloids in cured potatoes

Curing of Sebago potatoes for 10 days at 25°C prior to common storage at 5°C, reduced the responsiveness of tubers to photoinduced glycoalkaloid synthesis. Continuous illumination with 15 and 25-Watt incandescent light for 10 days increased glycoalkaloid content of peelings (12–14% of tuber weight) in uncured potatoes by a factor of 3.2 and 2.8, respectively, while the corresponding factor for cured tubers was only 1.8 for both lights. The peeled tuber portion (86–88% of tuber weight) had negligible amounts of glycoalkaloids, averaging about 1 mg per 100 g of fresh weight. The rise of glycoalkaloid levels in peels of uncured tubers was nearly linear to 164.7 mg/100g (15W light) with no indication of levelling off. In peels of cured tubers, the rise began only after the 4th day of light exposure with an apparent maximum and levelling off at 94.7 mg or approximately 43% lower than the final TGA levels in uncured tubers.     

A method for large scale laboratory inoculation of potato tubers with late blight fungus

As part of a project to investigate the possibility that teratogenic factors are formed in potato tubers infected with the late blight fungus,Phytophthora infestans, a method was devised which allowed quick, efficient inoculation of large numbers of potato tubers. Tubers were placed in specially constructed trays, wounded by puncturing the periderm with flower “pin frogs”, and sprayed with a zoospore suspensionof P. infestans prepared from cultures grown on rye agar at 20C (68F) for 10 days. Approximately 95% of Katahdin and 93% of Russet Sebago tubers inoculated by this method exhibited typical late blight lesions after two weeks incubation at 20C, 80–90% RH. Eight tons of potatoes were processed in this manner.     

Laboratory testing of potato tubers for multigenic resistance to late blight

Resistance in the tubers of potato clones with various levels of foliar multigenic resistance toPhytophthora infestans was measured to detect possible correlations between tuber and foliar resistance. A highly significant correlation was found using wound-healed tuber tissues. High levels of tuber resistance were detected in most foliar-resistant clones when inoculations with race 1,2,3,4 were made after 24–48 hours of wound periderm formation at 20 C. The resistance of wound-healed tissues increased as the time between wounding and inoculation increased. This increase in resistance was significantly greater for clones with foliar resistance than for clones with susceptible foliage.     

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.     

Potato glycoalkaloids: Increases and variations of ratios in aged slices over prolonged storage

Four commercial cultivars of potatoes were maintained under normal storage conditions at 44 F for 34 weeks. Except for a final 10 week interval tubers were withdrawn at 6 week intervals. After slicing, a portion of the slices was immediately analyzed for total glycoalkaloid content. The remaining slices were aged for four days in the dark at room temperature, then similarly analyzed. The total glycoalkaloid content of the aged slices increased dramatically on aging. This increase on aging of slices reached a maximum early in storage then decreased gradually over the storage period. In determining the individual glycoalkaloids, α-solanine and α-chaconine both increased in these slices, but the greatest increase was in the former. Appearing solely in the aged slices of the Kennebec variety, α-and β-solamarine appeared early in the storage period and gradually decreased over the storage period. Analyses of the unaged slices indicated that the glycoalkaloid content and composition of the potato tubers was little affected by storage. Aging of potato sprouts did not change their glycoalkaloid content.     

Der Einflu\ variierter Düngung, einer Pflanzenschutzmittel-Anwendung und der Lagerung auf den Glykoalkaloid-Gehalt in Kartoffelknollen zweier Sorten

Zusammenfassung Es wurden Feldversuche in drei Jahren mit zwei Speisekartoffelsorten, verschiedenen Mengen an Mineraldüngern und der Anwendungen eines Pflanzenschutzmittelns gegenPhytophthora infestans angelegt. Die Knollen wurden direkt nach der Ernte sowie nach einer sechsmonatigen Lagerung auf Glykoalkaloidgehalt untersucht. Zwischen den Sorten treten statistisch gesicherte Unterschiede im Gehalt an Glykoalkaloiden auf. Die Anwendung von Pflanzenschutzmitteln w?hrend der Vegetation erh?ht den Gesamtgehalt an Glykoalkaloiden (TGA) in den Kartoffelknollen kurz nach der Ernte. Eine variierte Mineraldüngung hat keinen statistisch signifikanten Einfluss auf den TGA-Gehalt in den Proben kurz nach der Ernte, N- und K-Düngung ver?ndern diesen jedoch im Laufe der Lagerung. Unabh?ngig von den übrigen Bedingungen erh?ht sich der TGA-Gehalt in diesem Versuch w?hrend der Lagerung im Mittel von 5,8 auf 7,7 mg/100g Frischsubstanz.

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Summary Field experiments were carried out for three years to study the effect of various magnesium, potassium and nitrogen fertilization rates on two medium-early edible potato cultivars. Additionally, one of the experimental areas was sprayed againstPhytophthora infestans to ascertain the effect of fungicides on the content of total glycoalkaloids (TGA) in potato tubers after harvest and after six months of storage. The applied fungicide treatments increased total glycoalkaloids (TGA) of potato tubers (Fig. 4). Fertilization with mineral nitrogen caused different reactions in both cultivars. There was a tendency for cv. Beryl to accumulate more glycoalkaloids with increasing nitrogen fertilization rates, while the opposite was observed for cv. Mila (Fig. 7). Fertilization with potassium and magnesium did not affect the results. On the basis of these results there was no conclusive evidence that the TGA content in the tubers was dependent on cultivar or on the weather during the growing season (Figs 1,2 and 3). The TGA content increased during storage, independently of previous treatments, (Figs 4–7).

     

Alaska Frostless,an inherently frost resistant potato variety

A potato clone with frost resistant foliage designated Alaska Frostless has been selected from the progeny of crosses involvingSolanum acaule and commercial varieties. Its chromosome number is (2N=5X=60). Nearly mature vines have withstood field frosting at?3 C for 2 hours and have recovered from several frosts of?2 C. Yields in cwt per acre of tubers 2–3.5 inch in diameter have equaled those of Alaska's standard varieties. Dry matter of 21.7% is equivalent to that of the Green Mountain variety. Tubers are flattened, round, smooth and white with a nonuniform faintly purple blush after exposure to light. Culinary quality, flavor, and texture are very good.     

Diploid and tetraploidSolanum chacoense genotypes that synthesize leptine glycoalkaloids and deter feeding by Colorado potato beetle

A selection (8380-1) fromSolatium chacoense Bitter (2n=2x=24) accession PI 458310 that synthesizes the leptine glycoalkaloids was compared in growth chambers with tetraploid (2n=4x=48) genotypes derived from tissue culture of 8380-1 leaf expiants for plant growth habit, leaf glycoalkaloid content, and effect on the development of Colorado potato beetle [Leptinotarsa decemlineata (Say)] larvae. The plants of the 4x regenerant genotypes were more vigorous with larger, more oval shaped leaflets than 8380-1 plants. The leaf concentrations of leptines and total glycoalkaloids were significantly lower (about 34%) in the 4x genotypes than in 8380-1. The proportion of leptines in the total glycoalkaloid content was nearly the same (about 80%) in both ploidy groups. In leaf-disk feeding tests, the development of Colorado potato beetle neonate larvae was not significantly different for the 2x and 4x genotypes. Both groups significantly slowed development compared with development on cv. Kennebec leaf disks. The 8380-1 selection and a group of 4x 8380-1 regenerant genotypes are maintained in the Vegetable Laboratory, Beltsville Agricultural Research Center, Beltsville, MD 20705 and are available for distribution.