Infection of Pratylenchus penetrans by Nematode-pathogenic fungi

Eleven fungal isolates were tested in agar dishes for pathogenicity to Pratylenchus penetrans. Of the fungi that produce adhesive conidia, Hirsutella rhossiliensis was a virulent pathogen; Verticillium balanoides, Drechmeria coniospora, and Nematoctonus sp. were weak or nonpathogens. The trapping fungi, Arthrobotrys dactyloides, A. oligospora, Monacrosporium dlipsosporum, and M. cionopagum, killed most of the P. penetrans adults and juveniles added to the fungus cultures. An isolate of Nematoctonus that forms adhesive knobs trapped only a small proportion of the nematodes. In 17-cm³ vials, soil moisture influenced survival of P. penetrans in the presence of H. rhossiliensis; nematode survival decreased with diminishing soil moisture. Hirsutella rhossiliensis and M. ellipsosporum were equally effective in reducing numbers of P. penetrans by 24-25% after 4 days in sand. After 25 days in soil artificially infested with H. rhossiliensis, numbers of P. penetrans were reduced by 28-53%.     

Effect of Single and Interplantings on Pathogenicity of Pratyenchus penetrans and P. neglectus to Alfalfa and Crested Wheatgrass

Alfalfa is a host of Pratylenchus penetrans and P. neglectus, whereas crested wheatgrass is a host of P. neglectus but not of P. penetrans. In a 120-day greenhouse experiment at 24 ñ 3 C, P. neglectus inhibited the growth of ''Lahontan'' alfalfa and ''Fairway'' crested wheatgrass. There were no differences in persistence and plant growth of alfalfa and crested wheatgrass, or reproduction of P. neglectus, in single plantings of alfalfa (AO) or crested wheatgrass (CWO), or in interplanted alfalfa and crested wheatgrass (ACW) treatments. On alfalfa, P. penetrans inhibited growth and reproduced more than did P. neglectus. Inhibition of plant growth and reproduction of P. penetrans was greater on alfalfa in AO than in ACW treatments. Pratylenchus penetrans did not reproduce on crested wheatgrass, but inhibited growth of crested wheatgrass in interplanted treatments and was avirulent in single planted treatments. Results were similar in a controlled growth chamber experiment at 15, 20, 25, and 30 C. Both nematode species inhibited alfalfa growth at all temperatures, and P. penetrans was more virulent than was P. neglectus to alfalfa at all temperatures and treatments. Plant growth inhibition and reproduction of P. penetrans on alfalfa in single and interplanted treatments were similar at 15-20 C, but were greater in single than in interplanted treatments at 25-30 C. Pratylenchus penetrans was avirulent to crested wheatgrass in the single planted treatments at all temperatures, but inhibited growth of crested wheatgrass in interplanted treatments at 20-30 C. Plant growth and reproduction of P. neglectus on crested wheatgrass was similar in single and interplanted treatments at 20-30 C and 15-30 C, respectively.     

Modeling of Yield Loss in Potato Early Dying Caused by Pratylenchus penetrans and Verticillium dahliae

Yield-loss models were developed for potato early dying, caused by an interaction between Verticillium dahliae and Pratylenchus penetrans. Yield data were collected over 5 years (1985-1989) from potato plants grown in microplots infested with V. dahliae and (or) P. penetrans. The model y = b₀ + (1 - b₀)/(1 + [VD/36.7]), where y was the relative yield (with uninfested controls = 1.0) and VD was the preplant density of V. dahliae microsclerotia per cm³ soil, was fitted to the data set. When P. penetrans = 0, b₀ = 0.55 (SE = 0.099), and when P. penetrans > 0, b₀ = 0.23 (SE = 0.035). This model assumed that yield loss was proportional to the concentration of preplant microsclerotia of V. dahliae, and only qualitatively related to presence or absence of P. penetrans. This study contrasts with previous reports that predict yield loss being proportional to preplant population densities of both P. penetrans and V. dahliae.     

Pathogenicity of Pratylenchus penetrans to Navy Bean (Phaseolus vulgaris L.)

The pathogenicity of Pratylenchus penetrans (root-lesion nematode) to Phaseolus vulgaris (navy bean) was evaluated in greenhouse experiments. Shoot and root fresh weight of cv. Sanilac plants were increased 4 and 21%, respectively, by an initial population density (Pi) of 25 P. penetrans per 100 cm³ soil. Leaf area and shoot fresh and dry weights were decreased by a Pi of 50 or more P. penetrans per 100 cm³ soil. A significant positive linear relationship existed between initial soil population densities of P. penetrans and final soil and root population densities of this nematode. Three dry bean cultivars, Sanilac, Seafarer, and Tuscola, were susceptible to P. penetrans, and yields were reduced by 43-76% when plants were exposed to a Pi of 150 P. penetrans per 100 cm³ soil. P. penetrans also reproduced on bean cultivars Saginaw, Gratiot, and Kentwood, but did not decrease bean yields, suggesting that these cultivars were tolerant to this nematode.     

Effect of Initial Nematode Population Density on the Interaction of Pratylenchus penetrans and Verticillium dahliae on 'Russet Burbank' Potato

Four similar growth chamber experiments were conducted to test the hypothesis that the initial population density (Pi) of Pratylenchus penetrans influences the severity of interactive effects of P. penetrans and Verticillium dahliae on shoot growth, photosynthesis, and tuber yield of Russet Burbank potato. In each experiment, three population densities of P. penetrans with and without concomitant inoculation with V. dahliae were compared with nematode-free controls. The three specific Pi of JR penetrans tested varied from experiment to experiment but fell in the ranges 0.8-2.5, 1.8-3.9, 2.1-8.8, and 7.5-32.4 nematodes/cm³ soil. Inoculum of V. dahliaewas mixed into soil, and the assayed density was 5.4 propagules/gram dry soil. Plants were grown 60 to 80 days in a controlled environment. Plant growth parameters in two experiments indicated significant interactions between P. penetrans and V. dahliae. In the absence of V. dahliae, P. penetrans did not reduce plant growth and tuber yield below that of the nematode-free control or did so only at the highest one or two population densities tested. In the presence of K dahliae, the lowest population density significantly reduced shoot weight and photosynthesis in three and four experiments, respectively. Higher densities had no additional effect on shoot weight and caused additional reductions in photosynthesis in only one experiment. Population densities of 0.8 and 7.5 nematodes/cm³ soil reduced tuber yield by 51% and 45%, whereas higher densities had no effect or a 15% additional effect, respectively. These data indicate that interactive effects between P. penetrans and V. dahliae on Russet Burbank potato are manifested at P. penetrans population densities less than 1 nematode/cm³ soil and that the nematode population density must be substantially higher before additional effects are apparent.     

Suppression of Meloidogyne incognita and M. javanica by Pasteuria penetrans in Field Soil

The role of Pasteuria penetrans in suppressing numbers of root-knot nematodes was investigated in a 7-year monocuhure of tobacco in a field naturally infested with a mixed population of Meloidogyne incognita race 1 and M. javanica. The suppressiveness of the soil was tested using four treatments: autoclaving (AC), microwaving (MW), air drying (DR), and untreated. The treated soil bioassays consisted of tobacco cv. Northrup King 326 (resistant to M. incognita but susceptible to M. javanica) and cv. Coker 371 Gold (susceptible to M. incognita and M. javanica) in pots inoculated with 0 or 2,000 second-stage juveniles of M. incognita race 1. Endospores of P. penetrans were killed by AC but were only slightly affected by MW, whereas most fungal propagules were destroyed or inhibited in both treatments. Root galls, egg masses, and numbers of eggs were fewer on Coker 371 Gold in MW, DR, and untreated soil than in AC-treated soil. There were fewer egg masses than root galls on both tobacco cultivars in MW, DR, and untreated soil than in the AC treatment. Because both Meloidogyne spp. were suppressed in MW soil (with few fungi present) as well as in DR and untreated soil, the reduction in root galling, as well as numbers of egg masses and eggs appeared to have resulted from infection of both nematode species by P. penetrans.     

Influence of Pratylenchus penetrans on Plant Growth and Water Relations in Potato

Plants of potato (Solanum tuberosum) cultivars Katahdin and Superior were inoculated with 0, 1,500, or 15,000 Pratylenchus penetrans. Transpiration, measured in the greenhouse with a porometer after 56 days of growth, was not significantly different among nematode inoculum levels or between cultivars. The rate of xylem exudation from decapitated root systems of Katahdin plants inoculated with 1,500 or 15,000 P. penetrans and Superior plants inoculated with 15,000 P. penetrans was lower than from noninoculated plants. Root weight of Katahdin and Superior was not affected by P. penetrans inoculum level. Transpiration of plants inoculated with 0, 500, 5,000 or 50,000 P. penetrans was recorded weekly from 14 to 56 days after planting. No consistent effects of nematode inoculum density on transpiration rate were observed. Root hydraulic conductivity was lower in Katahdin plants inoculated with 266 P. penetrans per plant and in Chippewa with 5,081 per plant than in noninoculated plants. Nematodes reduced leaf area of Superior, Chippewa, and Katahdin and root dry weight of Chippewa but had no effect on growth of Hudson, Onaway, or Russet Burbank plants. Assessing nematode effects on root hydraulic conductivity may provide a measure of the tolerance of potato cultivars to nematodes.     

Growth Response of Three Vegetables to Smooth- and Crenate-Tailed Females of Three Species of Pratylenchus

The effect of morphological variants of females of Pratylenchus penetrans, P. neglectus, and P. crenatus on the growth of three vegetables was studied. Variants were characterized by having either a smooth or crenate tail terminus. Pea was inoculated with variants of P. penetrans, one female per seedling, and grown at light intensities ranging from 1,350 to 21,600 lux in a series of five experiments. Only crenate-tailed females of P. penetrans suppressed the growth of pea and only when pea was grown at 3,900 lux. Radish was inoculated with morphological variants of P. penetrans, P. neglectus, and P. crenatus, four females per seedling, and grown at 3,900 lux in two experiments. Again, truly creuate-tailed females of P. penetrans inhibited growth. The two variants of P. penetrans had a similar infectivity, greater than that of the other two species of Pratylenchus. Only crenate-tailed P. penetrans reproduced on radish. Onion was inoculated with variants of P. penetrans and P. crenatus, four females per seedling, and grown at 14 C at 12,900 lux. Again, only crenate-tailed P. penetrans inhibited growth. The variants of P. penetrans had a similar infectivity, greater than that of P. crenatus. Neither species reproduced on onion at low temperatures.     

Population Growth of Pratylenchus penetrans on Winter Cover Crops Grown in the Pacific Northwest

Population growth of Pratylenchus penetrans on 13 fall and winter cover crops was studied in the greenhouse and field. All crops except oat cv. Saia supported population growth of P. penetrans in greenhouse experiments, although the response of P. penetrans to oat cv. Saia varied considerably between experiments. The mean ratio of the final population density/initial population density (Pf/Pi) after 16 weeks for P. penetrans added to a greenhouse soil mix was 0.09, whereas Pf/Pi values after 10 weeks for two experiments with naturally infested soil were 0.95 and 2.3. Although P. penetrans increased on sudangrass cv. Trudan 8 and sudangrass × sorghum hybrid cv. SS 222, subsequent incorporation of sudangrass vegetation into soil reduced P. penetrans populations to preplant levels. Field experiments were inconclusive but suggested that oat cv. Saia or rye cv. Wheeler may be better choices for winter cover than weed-contaminated fallow or other crops on P. penetrans-infested sites in the Pacific Northwest.     

Population Development of Pasteuria penetrans on Meloidogyne arenaria

A microplot study on the influence of cropping sequences with peanut in summer and bare fallowed or cover crops of rye or vetch in winter on the population development of Pasteuria penetrans was initiated in the spring of 1987. The number of spores of P. penetrans attached per second-stage juvenile of Meloidogyne arenaria race 1 increased from 0.11 in the fall of 1987 to 7.6, 8.6, and 3.6 in the fall of 1989 in the rye, vetch, and fallowed plots, respectively. Higher (P ≤ 0.05) levels of P. penetrans occurred in the rye and vetch plots than in fallowed plots. No influence of P. penetrans on peanut, rye, or vetch yield was observed in 1987 and 1988, but in 1989 peanut yield was 64% higher (P ≤ 0.05) in plots infested with P. penetrans than in plots without P. penetrans. Numbers of M. arenaria in plots without P. penetrans were influenced by the cropping sequences in the spring of 1988 and 1989 but not in the fall following the peanut crop. In the spring the plots with rye had the lowest nematode numbers in either year (P ≤ 0.05). Nematode numbers were lower (P ≤ 0.05) in plots with P. penetrans than in plots without P. penetrans in the spring of 1989 (vetch) and the fall of 1989 (rye, vetch, and fallowed).     

Pathogenicity of Pratylenchus penetrans,Heterodera glycines,and Meloidogyne incognita on Soybean Genotypes

The pathogenicity of Heterodera glycines, Meloidogyne incognita, and Pratylenchus penetrans on H. glycines-resistant ''Bryan,'' tolerant-susceptible ''G88-20092,'' and intolerant-susceptible ''Tracy M'' soybean cultivars was tested using plants grown in 800 cm³ of soil in 15-cm-diam. clay pots in three greenhouse experiments. Plants were inoculated with 0, 1,000, 3,000, or 9,000 H. glycines race 3 or M. incognita eggs, or vermiform stages of P. penetrans/pot. Forty days after inoculation, nmnbers of all three nematodes, except H. glycines on Bryan, generally increased with increasing inoculum levels in Experiment I. Heterodera glycines and M. incognita significantly decreased growth only of Tracy M. At 45 and 57 days after inoculation with 6,000 individuals/pot in experiments II and III, respectively, significantly more P. penetrans and M. incognita than H. glycines were found on Bryan. However, H. glycines and M. incognita population densities were greater than P. penetrans on G88-20092 and Tracy M. Growth of Tracy M infected by H. glycines and M. incognita and growth of G88-20092 infected by M. incognita decreased in Experiment III. Pratylenchus penetrans did not affect plant growth. Reduction in plant growth differed according to the particular nematode species and cultivar, indicating that nematodes other than the species for which resistance is targeted can have different effects on cultivars of the same crop species.     

Effects of Hydroclytic Enzymes on Plant-parasitic Nematodes

Proteases, lipase, and chitinase killed Tylenchorhynchus dubius in vitro and in soil. Tylenchorhynchus dubius was more susceptible to the enzymes than Pratylenchus penetrans. Papain was the most effective protease, and other enzymes were less effective. Heating enzymes to 80 C for 10 min greatly reduced nematicidal effectiveness. Scanning electron micrographs showed that papain and chitinase produced structural changes in the cuticle of T. dubius. Lipase removed a thin outer layer. Papain removed material filling the striata, or furrow, between the horizontal bands. When added to soil, chitinase, lipase, collagenase, and proteases (papain and bromelain) decreased motility of T. dubius populations up to 75%. Bromelain was the most active in soil against T. dubius, and collagenase was the most active in soil against P. penetrans.     

Interactions Among Pratylenchus penetrans,P. scribneri,and Verticillium dahliae in the Potato Early Dying Disease Complex

Microplots were infested with combinations of the fungus Verticillium dahliae and Pratylenchus penetrans and P. scribneri to test for individual and combined effects of these organisms on potato yield and nematode reproduction. Verticillium dahliae alone caused yield losses in all 3 years of the experiment, and the interaction between P. penetrans and V. dahliae was significant (P ≤ 0.05) in 2 years. Pratylenchus penetrans alone caused yield losses in 2 years and P. scribneri alone caused yield losses in 1 year. No two-way or three-way interaction was found involving P. scribneri. In 1987, reproduction for low densities of P. penetrans was 5 times higher when P. scribneri was also present than when it was absent, and 3.5 times higher in 1988. In nematode species mixtures, reproduction of P. scribneri was decreased by V. dahliae in 1987-88. The final population density of P. scribneri was negatively affected by V. dahliae and positively related to the initial proportion of P. scribneri to P. penetrans. In species mixtures with proportions of P. penetrans ranging from 0.1 to 0.5, reproduction of P. penetrans was negatively affected by V. dahliae and decreased linearly in relation to the increase in the initial proportion of P. penetrans in both years. The final population density of P. penetrans was affected only by V. dahliae.     

Temperature-Dependent Development of Pasteuria penetrans in Meloidogyne arenaria

Pasteuria penetrans is a promising biological control agent of plant-parasitic nematodes. This study was conducted to determine effects of temperature on the bacterium''s development in Meloidogyne arenaria. Developmental stages of P. penetrans were viewed with a compound microscope and verified with scanning electron microscopy within each nematode at 100 accumulated degree-day intervals by tracking accumulated degree-days at three temperatures (21, 28, and 35 °C). Five predominant developmental stages of P. penetrans were identified with light microscopy: endospore germination, vegetative growth, differentiation, sporulation, and maturation. Mature endospores were detected at 28, 35, and >90 calendar days at 35, 28, and 21 °C, respectively. The number of accumulated degree-days required for P. penetrans to reach a specific developmental stage was different for each temperature. Differences were observed in the development of P. penetrans at 21, 28, and 35 °C based on regression values fitted for data from 100 to 600 accumulated degree-days. A linear response was observed between 100 to 600 accumulated degree-days; however, after 600 accumulated degree-days the rate of development of P. penetrans leveled off at 21 and 28 °C, whereas at 35 °C the rate decreased. Results suggest that accumulated degree-days may be useful only in predicting early-developmental stages of P. penetrans.     

Temporal Changes in the Vertical Distribution of Pratylenchus penetrans under Raspberry

Population densities of Pratylenchus penetrans and the biomass of fine roots of raspberry at depths of 0-5, 5-10, 10-20, and 20-30 cm were determined every 2 weeks for 2 years. The vertical distribution of P. penetrans varied from season to season, but the seasonal changes were not similar for the 2 years. In most seasons, the greatest population density was in the 5 to 10-cm-depth interval. Population densities of P. penetrans were not consistently correlated with the vertical distribution of raspberry roots in any season.     

Interactions of Pratylenchus penetrans and Meloidogyne incognita as Coinhabitants in Tomato

Greenhouse experiments on the interactions of Pratylenchus penetrans and Meloidogyne incognita showed that the population densities of both nematode species were depressed whenthey coinhabited tomato roots. Fifty days after inoculation, the population level of a P. penetrans monoculture was about four times higher than when M. incognita was present. Conversely, M. incognita reproduced twice as fast alone as in combination. There were no significant differences in the numbers of P. penetrans when they were inoculated either 10 days prior to or after introduction of M. incognita. Root entry by P. penetrans was significantly inhibited by the presence of M. incognita. Split-root experiments showed that the inhibitory effects of M. incognita upon reproduction of P. penetrans involved factors other than the availability of feeding sites. On the other hand, the inhibitory effects of P. penetrans on M. incognita appeared to be primarily due to the quantity of available roots.     

Effects of Pratylenchus penetrans and Rhizoctonia fragariae on Vigor and Yield of Strawberry

Microplot and small field-plot experiments were conducted to determine the effects of Pratylenchus penetrans on strawberry yield over several seasons and to evaluate the effects of nematode control on strawberry vigor and yield. Pratylenchus penetrans alone or in combination with the black root rot pathogen, Rhizoctonia fragariae, reduced strawberry yield in microplots over time. There were no differences in effects on yield among R. fragariae anastomosis groups A, G, or I. The interaction of the two pathogens appeared to be additive rather than synergistic. In field plots infested with P. penetrans alone, plant vigor and yield were increased by the application of carbofuran and fenamiphos nematicides. Nematode control was transitory, as P. penetrans populations were initially suppressed but were not different in samples taken 10 months after treatment. These data highlight the error in associating causality between plant damage and nematode populations based on a correlation of root disease with nematode diagnostic assays from severely diseased plants. These findings may help to explain how nematode numbers can sometimes be higher in healthy plants than in severely diseased plants that lack sufficient roots to maintain nematode populations. Because nematode populations from up to a year before harvest are better correlated with berry yield, preplant nematode diagnostic assays taken a year in advance of harvest may be superior in predicting damage to perennial strawberry yield.     

Anhydrobiosis in Pratylenchus penetrans and Tylenchorhynchus n. sp. in Cultivated Soils Cropped to Winter Rye

Anhydrobiotes of Pratylenchus penetrans were found in two cultivated soils sown to rye in southern Ontario during the growing season. Anhydrobiotes at the 0-2.5-cm depth were recovered from 9 and 6 of 11 samplings, respectively, of a Vineland silt loam and a Fox loamy sand during the dry summer of 1983. At the 2.5-15.0-cm depth, anhydrobiotes were recovered less frequently. In the summer of 1984, anhydrobiotes of P. penetrans were recovered once and anhydrobiotes of Tylenchorhynchus n. sp. twice in 11 samplings. The percentages of P. penetrans populations that were anhydrobiotes in 1983 and 1984 were closely related to soil moisture content and corresponding moisture tensions. Populations of P. penetrans were greatest in October in the lower soil depth, 2.5-15.0 cm; those of Tylenchorhynchus n. sp. were greater in the surface layer of soil, 0-2.5 cm, and peaked in August.     

Effect of Meloidogyne incognita on Reproduction of Pratylenchus penetrans in Red Clover and Alfalfa

Roots of seedlings of red clover and alfalfa growing on 10⁻¹ Hoagland and Arnon solution agar were inoculated with various combinations of Meloidogyne incognita and Pratylenchus penetrans. Egg-laying by P. penetrans decreased as the number of nematodes, the ratio of entrant M. incognita to entrant P. penetrans, and the priority of invasion of roots by M. incognita increased. Embryogeny and hatching of eggs of P. penetrans, and development of larvae of M. incognita, were not affected. In red clover, the greatest red uction occurred when there were 65 entrant nematodes, the ratio of M. incognita:P. penetrans was 4:1 and M. incognita was inoculated four days prior to P. penetrans. In alfalfa, the less-favorable host for both nematodes, the greatest reduction occurred when there were 45 entrant nematodes, the ratio of M. incognita:P. penetrans was 2:1, and M. incognita was inoculated 4 days prior to P. penetrans.     

Suppression of Pratylenchus penetrans Populations in Potato and Tomato using African Marigolds

Current strategies for management of Pratylenchus penetrans in both white potato and tomato consist of the use of fumigant or non-fumigant nematicides or crop rotation. The objective of this study was to determine if double-cropping African marigolds (Tagetes erecta) with potatoes or tomatoes could reduce P. penetrans populations. Plots were 10 m × 3 m arranged in a randomized complete block design with four replications. Treatments included marigolds, potatoes or tomatoes, and natural weedy fallow followed by either potatoes or tomatoes. Nematode populations were sampled before spring planting, between crops in August and after harvest in November. During the 3 years of the study, P. penetrans soil population density declined by an average of 93% from the pre-plant level when marigold was grown in rotation with potato and by 98% when marigold was grown.in rotation with tomato. Weedy fallow preceding potato resulted in an average decline in P. penetrans soil population density of 38%, and a similar decrease (37%) was seen when fallow preceded tomato. There was a significant reduction in the number of P. penetrans found in both potato and tomato roots when the crops followed marigolds. These results suggest that P. penetrans population density may be significantly reduced when marigolds are double-cropped with potatoes or tomatoes.