Green peach aphid resistance by glandular trichomes inSolanum tuberosum xS. berthaultii hybrids

Three interspecific F₃ hybrid clones ofSolarium tuberosum L. x 5.berthaultii Hawkes bearing both Type A and Type B glandular trichomes were assessed for resistance to the green peach aphid,Myzus persicae (Sulzer). In glasshouse trials, aphid mortality and population development on hybrid clones were similar to those on the aphid-resistant parent,S. berthaultii. Trichome exudate of the hybrid, B227-63, was as effective in aphid immobilization as that ofS. berthaultii. Field populations of the green peach aphid on hybrids were intermediate between those onS. berthaultii and susceptibleS. tuberosum cultivars.     

Mechanisms of resistance to potato leafhopper,Empoasca fabae (Harris), in potato

Potato genotypes were evaluated for resistance to natural infestations of the potato leafhopper,Empoasca fabae (Harris), in the field. Characterization of Type A and B glandular trichomes, phenolic properties of Type A trichomes, glycoalkaloid content and composition,E. fabae adult and nymph infestation, leaf damage (hopperburn), yield, and yield reduction were determined on genotypes PI 473331 and PI 473334 (accessions ofSolatium berthaultii (Hawkes)), NY123 and Q174-2 (advanced hybrids), and Elba and Allegany (commercial cultivars). Regarding the Type A trichome, PI 473334 was most pubescent, followed by PI 473331, both had high levels of enzymatic browning; on PI 473334 no glycoalkaloids were detected, while on PI 473331 solamargine and solasonine were present. Those genotypes were the least infested by adults; moreover, nymphs were never found on their leaves. PI 473331 and Q174-2 possess Type B trichomes, the latter had levels of Type A trichome comparable to the first and to NY123, while displaying high levels of phenolic oxidation of Type A and very low levels of glycoalkaloids. Although Q174-2 had intermediate infestation ofE. fabae, it suffered a yield reduction comparable to the control cultivar Allegany. The density of Type A trichomes of NY123 was relatively low, even though the browning assay results were significantly elevated. Foliage of this genotype was found to contain the four types of glycoalkaloids studied at high levels.Empoasca fabae infestation on NY123 was intermediate, and it yielded well on infested plots, but suffered an intermediate yield reduction. Elba presented densities of Type A similar to NY123. It suffered the lowest yield reduction despite its high levels of leafhopper infestation and hopperburn. Its tolerance might be associated with its phenology, since it is a late-maturing cultivar. High scores of the modified enzymatic browning assay were correlated with high infestation and high leaf damage. Results indicated a significant correlation between leafhopper infestation and hopperburn.     

Evaluation of six insect pests for transmission of potato spindle tuber viroid

Six insect pests of potato, green peach aphid,Myzus persicae (Sulzer); potato leafhopper,Empoasca fabae (Harris); Colorado potato beetle,Leptinotarsa decemlineata (Say); tarnished plant bug,Lygus lineolaris (Palisot de Beauvois); red-legged grasshopper,Melanoplus femurrubrum (DeGeer); and southern army worm,Prodenia eridania (Cramer), were tested for their ability to transmit potato spindle tuber viroid (PSTV). PSTV infection was determined by a gel electrophoretic assay made 6 weeks after transfer of insects from infected to non-infected plants of the cultivar, ‘Katahdin.’ Plants grown from tubers of the original non-infected hosts were also assayed. PSTV was detected in only 2 of 183 test plants; these 2 plants had been infested with the tarnished plant bug. These results suggest that the insect species studied are relatively insignificant as vectors of PSTV.     

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.     

The effect of an alternate weed host,hairy nightshade,Solanum sarrachoides (Sendtner) on green peach aphid distribution and Potato leafroll virus incidence in potato fields of the Pacific Northwest

Hairy nightshade, Solanum sarrachoides (Sendtner) is an annual solanaceous weed prevalent in potato farmlands of the Pacific Northwest. S. sarrachoides is also a reservoir for Potato leafroll virus (PLRV) and its most important vector, the green peach aphid, Myzus persicae (Sulzer). Green peach aphids prefer S. sarrachoides than potato, Solanum tuberosum (L.), and produce more nymphs on S. sarrachoides than on potato. Increased green peach aphid preference for S. sarrachoides and greater fecundity on this plant could lead to overcrowding and movement toward neighboring potato plants. To test our hypothesis, field trials were conducted at Kimberly, Idaho during the summers of 2003 and 2004. A row of S. sarrachoides seedlings was planted in the middle of potato plots and naturally occurring green peach aphid populations were monitored weekly on potato plots with and without S. sarrachoides. More aphids were consistently found on S sarrachoides plants than on potato plants. More aphids were found on potato plants in plots with S. sarrachoides plants than without S. sarrachoides. Targeted insecticide applications on S. sarrachoides rows alone reduced the number of aphids on potato plants in those plots, suggesting that insecticide sprays prevented aphid movement from S. sarrachoides to potato. Testing of potato tuber sprouts 90 days after harvest by double antibody sandwich ELISA indicated a greater PLRV infection rate on plots with S. sarrachoides than plots without S. sarrachoides. Therefore, the presence of S. sarrachoides in potato plots will likely increase aphid populations and PLRV incidence on neighboring potato plants.     

Mechanical interaction between larvae of the Colorado potato beetle and glandular trichomes ofSolanum berthaultii Hawkes

The rate of accumulation of glandular trichome exúdate ofSolanum berthaultii on the tarsi of Colorado potato beetle (Leptinotarsa decemlineata) larvae depended on larval size (instar), with large larvae bearing the hardened, darkened exudate on more of their tarsi than did small larvae. In greenhouse studies, third instar larvae accumulated large amounts of exudate regardless of which of 2S. berthaultii accessions they were exposed to, but exudate accumulation by first instars was influenced significantly by plant accession. Tarsal accumulation of exudate was lower in field plots of bothS. berthaultii accessions, but tended to increase with time of exposure. The significant differences in exudate accumulation among larvae exposed to the 2 accessions in the greenhouse were not evident in the field. The density of type A glandular trichomes on the abaxial leaflet surface differed significantly between the 2 accessions in both greenhouse and field. Despite these effects of host and environment on the accumulation of readily observed amounts of trichome exudate on larval tarsi, no larvae were trapped by the adhesive exudate.     

Evolutionary pathway of T-type Chloroplast DNA in potato

Potato was domesticated in the Andes of South America. However, the presently worldwide-grown potato (Solanum tuberosum L. ssp.tuberosum, 2n=4x=48) has characteristic T-type chloroplast DNA that was introduced after late blight epidemics in the mid-19th century from the Chilean potato (2n=4x=48) grown in the southern coastal regions in Chile. Among many wild potato species, the same chloroplast DNA was found only in some populations of a diploid speciesS. tarijense Hawkes (2n=2x=24), which ranges from central Bolivia to northwest Argentina. To elucidate an evolutionary pathway of T-type chloroplast DNA fromS. tarijense to Chilean potato, 215 accessions ofS. stenotomum Juz. et Buk., considered to be the most primitive, diploid cultivated potato species, from which all the Andean cultivated species evolved, and 286 accessions of the most widely grown, Andean tetraploid cultivated speciesS. tuberosum L. ssp.andigena Hawkes (2n=4x=48) were examined in this study. No accession ofS. stenotomum had T-type chloroplast DNA, while nine accessions, mostly from northwest Argentina, ofS. tuberosum ssp.andigena had T-type chloroplast DNA. Therefore, I conclude that some populations ofS. tarijense having T-type chloroplast DNA were naturally crossed as female withS. tuberosum ssp.andigena from which the Chilean potato was selected.     

The germplasm release of NYL 235–4, a clone with resistance to the Colorado potato beetle

Clones selected from the wild diploid species,Solanum berthaultii, have been shown to possess valuable levels of resistance to the Colorado potato beetle (Leptinotarsa decemlineata), as well as to insects such as aphids, flea beetles, leafhoppers and the potato tuber moth. Resistance is associated with the presence of glandular trichomes on the foliage ofS. berthaultii (2). Six generations after producing a hybrid between the diploid species and a tetraploid variety, a clone with good beetle and leafhopper resistance, as well as reasonable agronomic characteristics has been produced.     

Populations of aphids on named varieties of Irish potatoes

Populations of the green peach aphid,Myzus persicae (Sulzer), and the potato aphid,Macrosiphum euphorbiae (Thomas), were observed on 27 named varieties of Irish potatoes in the Yakima Valley of Washington intermittently from 1947 through 1968. The Epicure, Great Scot, and B5141-6 varieties produced relatively fewer green peach aphids than other varieties tested; Bliss Triumph and Red Pontiac produced relatively fewer potato aphids. Except for Doebay Red, the red-skinned varieties, Bliss Triumph, Red McLure, Norland, and Earnly Rose, were extremely to moderately productive of the green peach aphid. No correlation was apparent between abundance of aphids and the specific gravity or yield of tubers.     

Insecticides for control of potato tuberworm and green peach aphid on potatoes in California

Six insecticides were evaluated in a replicated field experiment at Gonzales, California, for control of the potato tuberworm,Phthorimaea operculella (Zeller), and the green peach aphid,Myzus persicae (Sulzer). Azodrin^®, methomyl, Stauffer N 4543 (O-isobutyl ethylphosphonodithioate S-ester with N-(mercaptomethyl) phthalimide) and azinphosmethyl were applied as foliage sprays by ground sprayer at 2-week intervals. The systemic insecticides, aldicarb (Temik^®) and carbofuran (Furadan^®) formulated as 10% granules, were applied to the soil as side dress treatments on the same day that the spray treatments were begun. Aldicarb granules were also applied topically so that the granules fell on the young plants and on the soil surface. Azinphosmethyl, Stauffer N 4543, Azodrin^® and methomyl effectively controlled the tuberworm larvae in the foliage but effective control of tuberworms in the harvested tubers occurred only where azinphosmethyl, methomyl and Stauffer N 4543 were used. The granule applications of carbofuran and aldicarb were totally ineffective in controlling the potato tuberworm. Effective green peach aphid control was obtained with Azodrin^® spray and the two granular aldicarb treatments. The side dressed application of aldicarb appeared to provide better green peach aphid control than did the soil surface application.     

Genetic Resistances to Potato Leafroll Virus, Potato Virus Y, and Green Peach Aphid in Progeny ofSolanum etuberosum

Increasing prevalence of potato leafroll virus (PLRV) and potato virus Y (PVY) has been reported in seed and commercial potato production, resulting in the rejection of potatoes for certification and processing. Host plant resistance to PLRV and PVY and their primary vector, green peach aphid,Myzus persicae, could limit the spread of these viruses. Host plant resistance to PLRV, PVY, and green peach aphid has been identified in non-tuber-bearingSolanum etuberosum (PI 245939) and in its backcross 2 (BC₂) progeny. Resistance to green peach aphid involved a reduction in fecundity and adult aphid size. In addition, one BC₂ individual was identified as possessing a genetic factor that was detrimental to nymph survival. PVY resistance was identified in all five BC₂ progenies evaluated in a field screening under intense virus pressure. PLRV resistance was identified in two of the five BC₂ progeny. This resistance was stable in field and cage evaluations with large populations of viruliferous aphids. Based on the segregation of virus resistances in the BC₂ , PVY and PLRV resistances appear to result from the action of independent genetic mechanisms that reduce the levels of primary and secondary virus infection. Two BC₂ individuals, Etb 6-21-3 and Etb 6-21-5 were identified as having multiple resistances to PLRV, PVY, and green peach aphid derived fromS. etuberosum. This germplasm could prove useful to potato breeders in the development of virus-resistant cultivars.     

Early generation selection of insect resistance in potato

Methods used in breeding and selecting potatoes for resistance to Colorado potato beetle, green peach aphid, potato aphid, and potato leafhopper are reviewed. Sources of resistant germplasm, the nature of resistance, breeding behavior of resistance factors, and methods used to measure resistance are discussed. Specific problems encountered in breeding insect-resistant potatoes are discussed in relation to early-generation selection procedures.     

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.     

Effect of time of inoculation with potato leafroll virus on development of net necrosis and stem-end browning in potato tubers

The Green Mountain cultivar was used in field tests to determine the effects of inoculating potato plants at various times with the potato leafroll virus (PLRV) on development of internal necrosis of tuber tissue. Viruliferous apterae of the green peach aphid,Myzus persicae (Sulz.), were placed on each stem in all hills to be inoculated in each 3.0 m single-row plot. Planting and inoculation dates were varied in all field experiments and, in one, several vine-killing dates were also included. All harvested tubers were stored for approximately four months at 10°C to enhance development of internal necrosis prior to examination. Similar but smaller greenhouse studies involving both apterous and alate green peach aphids were also conducted using Green Mountain, Irish Cobbler, and Russet Burbank cultivars. All results showed that as inoculation was delayed relative to plant development, more net necrosis (NN) occurred. Conversely, when plants were inoculated early, stem-end browning (SEB) rather than NN predominated. A high percentage of naturally occurring SEB tubers (cv. Russet Burbank) were found by ELISA to contain PLRV. Plants produced by these tubers only rarely developed leafroll symptoms. These findings suggest a previously unsuspected causal relationship between SEB and PLRV. Implications of this apparent relationship on the epidemiology of potato leafroll in Maine are discussed.     

Fine screeningSolanum (potato) germplasm accessions for resistance to Colorado potato beetle

This study was conducted to investigate the utility of systematic screening at the genotype level withinSolanum accessions highly resistant to the Colorado potato beetle. Evaluations of clonally replicated genotypes showed that most accessions reported to be uniform when screened as populations contained small but significant variation among genotypes for resistance to oviposition, larvae, and defoliation, differences for numbers of larvae being most common. Adult counts and percentage defoliation were not as useful in evaluating among-genotype variability in beetle resistance. Genotypes ofS. pinnatisectum WRF 343 andS. tarijense PI 473227 were the most uniformly and highly resistant to Colorado potato beetle. Genotypes of S.berthaultii PI 473331,S. chacoense PI 473405, andS. tarijense PI 473336 were moderately to highly resistant, and genotypes ofS. bukasovii PI 473494 andS. canasense PI 230511 were uniformly susceptible to Colorado potato beetle. Nonparametric correlation analyses indicated that number of egg masses, small larvae, large larvae, and defoliation scores were positively correlated, negatively correlated, or not correlated, depending on the species. One generation of selection attempting to segregate resistance and susceptibility in nearly uniform and highly resistantS. pinnatisectum WRF 343 resulted in “divergent” populations that could not be distinguished from each other or the base population. Thus, through genotype (“fine”) screening and selection, we showed that some existing populations are virtually pure for extreme resistance. Use of individuals from such families would make screening breeding populations more efficient, and reduce the risk of losing resistance genes that have non-dominant effects. Fine screening, recurrent selection, and maintenance of such elite populations is recommended as an extension of population-based evaluation usually done by genebanks.     

Comparative analysis of aphid vector behavior in response to potato plants grown from field tubers, minitubers or plantlets

The greater sensitivity to aphid vectored diseases of potato plants originating from plantlets or minitubers compared to plants originating from field grown tubers could not be explained by changes in probing behavior, by increased aphid movement between plants or by greater levels of aphid colonization favorable to virus spread. The number of seconds to first probe, the duration and number of seconds before the long probe and the duration of the long probe were statistically similar on all plant types. The average residency time of green peach aphids on each plant type was also statistically the same. WingedMyzus persicae (Sulzer) (green peach aphid) landed in greater numbers on older than on younger plants regardless of plant type. This suggests that the level of aphid colonization is determined by a combination of factors such as height, surface and age of plants but not by plant origin. Where virus vectors are present early in the season, the greater number of aphids landing on the relatively older and larger transplants may contribute to higher infection levels, but later in the season plant physiological factors favorable to virus multiplication or translocation are more likely factors     

A modified enzymic browning assay for potato glandular trichomes

Tetralobulate (Type A) glandular trichomes are present on foliage of many wild as well as cultivated species of potatoes. These trichomes confer resistance to insect pests if their density, gland size and biochemical products are similar as those ofSolanum berthaultti Hawkes andS. polyadenium Greenm. Hybrids ofS. tuberosum L. ×S. berthaultii have been screened for resistance by measurement of phenolic oxidation activity in their Type A trichomes by use of an enzymic browning assay. However, variability in the procedure has prompted the development of an improved assay.     

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     

Distribution of the 241 bp deletion of chloroplast DNA in wild potato species

The common potato (Solanum tuberosum L. ssp.tuberosum) has T-type chloroplast DNA characterized by a 241 bp deletion. To explore the maternal ancestry of the common potato, a total of 566 accessions of 35 wild species, collected mostly from central Bolivia to northern Argentina, were determined for presence or absence of the deletion by a simple PCR assay using primers flanking the deleted region of chloroplast DNA. Sixteen out of 80 accessions ofS. tarijense, S. berthaultii, andS. neorossii showed a shorter PCR amplified fragment. Sequencing of these fragments revealed that the same 241 bp was deleted at the same position in these accessions. This strongly suggests that the deletion event had occurred in wild species.     

The Inter-genebank potato database and the dimensions of available wild potato germplasm

The Association of Potato Inter-genebank Collaborators (APIC) constructed a database of all wild potato holdings of the most important potato genebanks in Europe, the United States, Peru, and Argentina. The Inter-genebank Potato Database (IPD) now contains data of 11,819 wild potato accessions conserved in seven potato genebanks. The collector’s number is the key identifier used to merge all databases into the IPD. A total of 7,112 different wild potato accessions were identified, which comprise 5,306 accessions with known collector’s numbers. The IPD passport database showed that almost 30% of accessions held in APIC genebanks are from Argentina, a country that comprises less species diversity than Peru and Bolivia. These latter countries are represented by 24% and 20% of accessions, respectively. APIC genebanks maintain 188Solarium taxa out of more than 230 recognized by the latest comprehensive treatment of potatoes(Solarium sect.Petota) by Hawkes (1990). About 60% of the accessions comprise only 20 taxa represented by 785 to 92 accessions each. Conversely, 2% of the accessions comprise 72 taxa and are represented by five or fewer accessions each. About 70 taxa are not available in any genebank. The IPD evaluation database comprises 5,603 records with data from more than 33,000 evaluations of wild potato accessions. Fifty-five traits are summarized, including the reactions of the accessions to 12 pathotypes or races or strains of 12 fungi, four bacteria, 12 viruses, one viroid, 13 nematodes, and seven insects; response to heat and cold stress; and content of dry matter, starch, vitamin C, amylase, reducing sugars, and glycoalkaloids. About 30% of the wild potato populations screened showed various levels of resistance to most of the diseases and pests evaluated. The IPD database is available on the Internet at www.potgenebank.org.