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.     

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.     

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     

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).     

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.     

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.     

Loss of yield in potato selections infested with potato leafhoppers

The loss of yield resulting from infestation with potato leafhoppers,Empoasca fabae (Harris), was measured in a group of potato selections (Solatium tuberosum subsp.tuberosum). For 6 years in which 300 different clones were tested, the average loss of yield was 39%. The correlation over clones between level of infestation and percent hopperburn equaled 0.56; between level of infestation and loss of yield 0.36; and between percent hopperburn and loss of yield 0.41. Each coefficient is significant at P=0.01. Significant differences were found in levels of infestation and in percent hopperburn among the selections, but even those selections with the most resistant foliage had greatly reduced yields when infested. No selections were found that consistently produced a normal yield when infested with leafhoppers. The low level of infestation on the more resistant clones, however, may make chemical control at lower rates more effective.     

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.     

Potato (Solanum spp.) in the hot tropics. VI. Plant population effects on soil temperature,plant development and tuber yield

Nine experiments were run at three hot tropical sites (5–12°S, 180–800 m) within Peru to quantify the influence of plant population on soil temperature and growth and yield of the potato.Radiation interception was greatest at the highest plant populations and soil cooling was directly proportional to the amount of crop cover over the soil, but no appreciable effect on the timing of tuber initiation was apparent. More stems per unit land area leading to a higher leaf area index (lai) were primarily responsible for greater interception of radiation at the higher plant populations, although some compensation in stem number per unit land area and in lai at lower populations was evident later in the season.In general, tuber yield increased linearly with increases in planted population over the range studied (2.7–12.5 plants m⁻², and in one experiment to 31.7 plants m⁻²), and was proportional to increases in the amount of intercepted radiation. Tuber yields ranged from 8 to 60 t ha⁻¹ over sites and populations. Vigorous clones with Solanum tuberosum spp. andigena in their genetic background constituted the exceptions to this linear trend, and for these clones yields declined at the highest populations, particularly when the rectangularity of planting vastly deviated from square patterns. Tuber yield of Solanum tuberosum spp. tuberosum and Neotuberosum (S. tuberosum spp. andigena selected for tuberization under long-day conditions) clones did not respond to variations in rectangularity of planting and, probably due to their small stature and early maturity, did not demonstrate signs of intense between-plant competition for tuber yield as measured with the Kira competition density index. In contrast, for clones with Solanum tuberosum ssp. andigena in their genetic background, maximum tuber yield at populations greater than 5.5 plants m⁻² was dependent on the rectangularity of planting, and declined as the latter deviated from squareness.Since the proportion of marketable tubers was scarcely affected by the planting densities, plant population of S. tuberosum ssp. tuberosum clones planted in hot climates should be as close as possible without limiting the amount of soil available for hilling-up.     

Glandular trichomes: Enzymic browning assays for improved selection of resistance to the green peach aphid

Polyphenoloxidase (PPO) activity in the glandular trichomes of 32Solanum tuberosum ×S. berthaultii F₃ hybrid clones differentially resistant to the green peach aphid,Myzus persicae (Sulzer) was determined. Within this population of segregating clones, aphid infestation levels varied 8-fold, while a 5-fold difference in glandular trichome PPO activity was observed. PPO activity and aphid resistance at three sample dates were correlated as follows: r = 0.38 (P = 0.05), r = 0.31 (P = 0.10), and r = 0.55 (P = 0.01), respectively. The total browning potential test (TBP), a modification of the PPO assay, was developed for use in screening for aphid resistance. The TBP procedure is rapid, inexpensive, and highly effective in identifying aphid resistant clones.     

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.     

Pollen-pistil incompatibility in a diploid hybrid potato population with cultivated and wild germplasm

The basis of low seed set was investigated in a diploid hybrid population with germplasm from the cultivated speciesSolanum tuberosum spp.tuberosum andS. tuberosum ssp.phureja and the wild speciesS. chacoense. Controlled crosses were performed following an incomplete diallel mating design which included the hybrid population and the parental species. Pollen grain germination and pollen tube growth were observed with fluorescence microscopy in 174 intrapopulation and interspecific combinations of genotypes. Fifty percent of the combinations of genotypes within the hybrid population and 46% of those between this population and the parental species were incompatible; overall, 71% of the interspecific combinations of genotypes were compatible when plants of the hybrid population were used as females, vs. 5% when used as males. Although the site of reaction could vary in a given intrapopulation or interspecific cross with the combinations of genotypes, pollen tube inhibition occurred in the first third of the style in 42% of them. Since the gene pool of the hybrid population had been widened with germplasm from other geographic areas and the genotypes ofS. tuberosum ssp.tuberosum, S. chacoense andS. tuberosum ssp.phureja used in this study were not the ones involved in the original crosses, it is concluded that the S-locus is not controlling the incompatibility reaction but rather that a cross-incompatibility system, possibly governed by more than one locus, is acting.     

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%.     

The heritability of resistance to early blight in diploid potatoes (Solanum tuberosum,subsp.Phureja andStenotomum)

The heritability of resistance to early blight caused byAlternaria solani (Ell. & Martin) Son was estimated for field grown, diploid potatoes at Fletcher, North Carolina in 1981. Fourteen breeding lines, selected from a random mated hybrid population ofSolanum tuberosum, subsp.phureja andstenotomum, were used. Resistance was measured as the slope obtained by regressing the logit of the disease proportion Y, measured as % defoliation, on time. A wide range of resistance was found. Narrow-sense heritability, as determined by offspring-midparent regression, on an individual plant basis, was calculated to be 0.825.     

Susceptibility of potato varieties and advanced breeding lines (Solanum tuberosum L.) toPhytophthora infestans (Mont.) de Bary in greenhouse screenings

Late blight (Phytophthora infestons (Mont.) de Bary) has re-emerged as an important pathogen of the cultivated potato (Solanum tuberosum subsp.tubero-sum L.) in North America. The purpose of this study was to evaluate the relative susceptibility of potato germplasm in the greenhouse in order to initiate a breeding program for resistance to the US-8/A2 mating type which is the more aggressive and prevalent strain of late blight. Whole plants of 147 cultivars and breeding lines were evaluated. Percent plant area infection was visually assessed. Seven days after inoculation, infection ranged from 0 to 100% and the overall mean was greater than 50%. Two-thirds of the cultivars and breeding lines tested were very susceptible to the US-8 genotype. The highest resistance was identified in the somatic hybrids betweenS. tuberosum andS. bulbocastanum and their backcross derivatives. Pike and Snowden were less susceptible than the other North American cultivars. Zarevo was most resistant among the European cultivars. Seven of the advanced breeding lines were equivalent to Zarevo in infection levels. The host plant resistance identified among the material tested in this study can be used by breeding programs to develop improved cultivars with resistance to US-8 genotypes of late blight.     

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.     

Effect of random mating on yield and specific gravity in twoSolanum tuberosum subsp.tuberosum populations

A group of biparental crosses of breeding clones and varieties (Solanum tuberosum subsp.tuberosum) from the USDA potato breeding program, Beltsville, MD, was used to establish two randomly mated populations. Random mating without selection was continued for seven generations, then cycles 1, 3, 5, and 7 were tested for yield and specific gravity. There was a small but non-significant decrease in tuber weight per hill, and a small but non-significant increase in tuber number from cycle 1 to cycle 7. Mean tuber weight significantly (P < 0.01) decreased (12%) from cycle 1 to cycle 7. Specific gravity also significantly (P < 0.10) decreased over cycles. The variance of clones within cycles did not significantly change over cycles for any of the traits measured.     

Immunodetection of Polyphenol oxidase in glandular trichomes ofS. berthaultii,S. tuberosum and their hybrids

Type A glandular trichomes confer insect resistance to the wild potato,S. berthaultii Hawkes by oxidatively polymerizing their contents upon breakage, resulting in insect entrapment. Polyphenol oxidase (PPO) is responsible for the oxidative polymerization of Type A glandular trichome exudate.S. tuberosum L. trichomes possess low amounts of PPO and lack the ability to either polymerize their contents or entrap insects. Polyclonal antibodies raised against purifiedS. berthaultii glandular trichome PPO were used to develop an enzyme-linked immunosorbent assay (ELISA) to quantify PPO content ofS. berthaultii, S tuberosum, and hybrid type A glandular trichomes. The PPO ELISA and immunoblots were used to quantify PPO expression as a function of leaf development and genetic background. PPO accumulates to its maximal levels in trichomes prior to completion of leaflet expansion. In advanced breeding generations, trichome PPO is expressed at levels varying from that found inS. tuberosum to that found inS. berthaultii. By serving as an initial screen for trichome PPO, this ELISA may be useful to assist rapid introgression of this insect resistance trait into adaptedS. tuberosum backgrounds.     

Screening for gibberellin deficiency mutants inSolanum tuberosum ssp.andigena

A GA deficiency mutant was previously described in a single population ofSolanum tuberosum ssp.andigena (adg). A representative sample of 120 adg populations in the US Potato Genebank was screened for the dwarfing mutant. Up to 500 seedlings of each adg population were evaluated. Dwarfs were detected in 14 populations from a minimum of 0.2% to a maximum of 27.7%, indicating that the allele is not particularly rare or at risk of loss from the Genebank.     

Cytoplasmic substitution inSolanum. II. Tuber characteristics of reciprocal backcross progeny

Summary Recurrent backcrosses (BC) were used to introduce nuclear factors ofS. tuberosum L. ssp.tuberosum into the cytoplasm of Andean potatoesS. tuberosum ssp.andigena (Juz. and Buk.) Hawkes andS. phureja Juz. and Buk. Each generation BC was produced in reciprocal directions to test the effect of cytoplasmic substitution on yield. This report analyses tuber number, tuber weight, and tuber length of BC₂ and BC₃ substitution into ssp.andigena cytoplasm and BC₂ and BC₂ substitutions intoS. phureja cytoplasm. Direction of cross had no consistent significant effect on yield components of ssp.andigena substitution lines. InS. phureja reciprocal progenies direction of cross was responsible for significant differences of some reciprocal sets. When there were significant differences the higher yield usually occurred when ssp.tuberosum was the pistillate parent, but in some progenies when ssp.tuberosum was the pistillate parent the yield was significantly lower than in the reciprocals. It is suggested that differences between reciprocals resulted from chromosomally encoded gene action, resulting from maternal and/or paternal effects, rather than cytoplasmic factors. Authorized for publication as paper No. 7494 in the Journal Series of The Pennsylvania Agriculture Experiment Station.