Effects of Potassium Fertilization and Pratylenchus penetrans on Yield and Potassium Content of Red Clover and Alfalfa

Red clover and alfalfa were inoculated with Pratylenchus penetrans and grown in an Alberry sandy loam soil to which potassium (K⁺) was added at seeding at 0, 41.5, 83, and 166 μg/g. In one experiment with alfalfa, additional K⁺ was added after each forage cut to replace that which was removed. Nematode populations were not consistently affected by K⁺ fertilization. Nematode infection stunted red clover and alfalfa and resulted in lower yields at all K⁺ levels, except for alfalfa at the lowest K⁺ level. Nematode infection had no effect on taproot yields. However, it resulted in lower rootlet yields from red clover at all K⁺ levels, lower rootlet yields from alfalfa only at the highest K⁺ level in one experiment, and lower rootlet yields at all but the lowest K⁺ level in a second experiment, potassium fertilization enhanced yield of red clover and alfalfa. Yield increases were smaller from increased K⁺ fertilization in nematode-infested soil than in noninfested soil. Pralylenchus penetrans had little effect on the K⁺ content of red clover or alfalfa. The stunting of plants from nematode infection resulted in less K⁺ being removed from the soil.     

The Influence of Pratylenchus penetrans on the Incidence and Severity of Verticillium Wilt of Potato

The influence of Pratylenchus penetrans on the incidence and severity of Verticillium wilt was examined in the potato cultivars ''Kennebec'', ''Katahdin'', and ''Abnaki''. Single-stem plants were grown in soil maintained at a temperature of 22 ± 1 C. Axenically cultured nematodes were suspended in water and introduced to the soil, at a rate of ca 5,000/25.4-cm pot, through holes made around each stem. Ten days after infestation with nematodes, conidial suspensions of Verticillium albo-atrum were introduced into the soil at a rate of ca 1,000,000/pot. Among Katahdin plants, the severity of foliar symptoms was increased in the presence of both pathogens 2 and 3 weeks after soil intestation. During the remaining 5 weeks, severity of foliar symptoms was not different between plants infected by both pathogens and those infected by Verticillium alone. Within the wilt-susceptible cultivar Kennebec and the resistant eultivar Abnaki, no effects on foliar symptom severity were observed. When plant heights, shoot weights, and tuber yields were analyzed, a Pratylenchus-Verticillium interaction was not evident within any of the cultivars tested. Nematode populations in roots and rhizosphere were suppressed in Kennebec and Katahdin plants in the presence of Verticillium.     

Varietal Response of Tomato to the Interaction of Salinity and Meloidogyne incognita Infection

Response of tomato (Lycopersicon esculentum) cultivars to a range of conductivity levels was tested in the presence and absence of Meloidogyne incognita. The conductivity levels were produced by appropriate adjustment of a 1:1 solution of sodium chloride and calcium chloride. The growth of M. incognita resistant (''Beefmaster'' and ''Atkinson'') and susceptible (''Hunts 2580'' and ''Ronita'') tomato plants was inversely related to soil salinity between EC_e 0 and 5 mmhos/cm. Nematode inoculation of salt-stressed plants significantly reduced plant height, fresh and dry weight, number of flowers, and fruit weight in most cultivars. In Hunts 2580, flower number and fruit weight increased; apparently flower production shifted from determinate to indeterminate, with negative implications for mechanical harvesting. Nematode reproduction on susceptible varieties also decreased with increase in salinity.     

Factors Affecting Population Trends of Plant-Parasitic Nematodes on Rangeland Grasses

The effects of environmental conditions on population trends of plant-parasitic nematodes were studied in experimental plots of five wheatgrasses in the western Utah desert. In a 3-year (1984-86) field study, soil water and temperature affected the population trends of the ectoparasites, Tylenchorhynchus acutoides and Xiphinema americanum, and the migratory endoparasite, Pratylenchus neglectus, on Fairway crested wheatgrass, Agropyron cristatum; ''Hycrest'' crested wheatgrass, A. cristatum X A. desertorura; ''Rosana'' western wheatgrass, Pascopyrum smithii; ''Oahe'' intermediate wheatgrass, Thinopyrum intermedium; and RS-1 hybrid (Elytrigia repens X Pseudoroegneria spicata). The largest soil populations of these nematode species were collected in 1984 under good plant-growth conditions. A reduction in nematode populations occurred in 1985 and 1986, possibly because of low soil-water conditions. There was a positive relationship between high soil water and maximum population densities of T. acutoides in the spring and fall of 1984, and between low soil water and minimum population densities of the nematode in 1985 and 1986. Pratylenchus neglectus populations were affected by soil water, although to a lesser degree than the ectoparasitic nematodes. Population densities of the three nematode species were significantly lower in the drier years of 1985 and 1986 than in 1984. Nematode populations were greater at the lower soil depths in the fall than in the spring or summer.     

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.     

Survival of Rhizobium in Acid Soils

A Rhizobium strain nodulating cowpeas did not decline in abundance after it was added to sterile soils at pH 6.9 and 4.4, and the numbers fell slowly in nonsterile soils at pH 5.5 and 4.1. A strain of R. phaseoli grew when added to sterile soils at pH 6.7 and 6.9; it maintained large, stable populations in soils of pH 4.4, 5.5, and 6.0, but the numbers fell markedly and then reached a stable population size in sterile soils at pH 4.3 and 4.4. The abundance of R. phaseoli added to nonsterile soils with pH values of 4.3 to 6.7 decreased similarly with time regardless of soil acidity, and the final numbers were less than in the comparable sterile soils. The minimum pH values for the growth of strains of R. meliloti in liquid media ranged from 5.3 to 5.9. Two R. meliloti strains, which differed in acid tolerance for growth in culture, did not differ in numbers or decline when added to sterile soils at pH 4.8, 5.2, and 6.3. The population size of these two strains was reduced after they were introduced into nonsterile soils at pH 4.8, 5.4, and 6.4, and the number of survivors was related to the soil pH. The R. meliloti strain that was more acid sensitive in culture declined more readily in sterile soil at pH 4.6 than did the less sensitive strain, and only the former strain was eliminated from nonsterile soil at pH 4.8; however, the less sensitive strain also survived better in limed soil. The cell density of the two R. meliloti strains was increased in pH 6.4 soil in the presence of growing alfalfa. The decline and elimination of the tolerant, but not the sensitive, strain was delayed in soil at pH 4.6 by roots of growing alfalfa.     

Effect of Ditylenchus dipsaci and Pratylenchus penetrans on Verticillium Wilt of Alfalfa

Verticillium albo-atrum wilt symptoms appeared faster and were significantly more severe in the presence of Ditylenchus dipsaci in Vernal, a wilt-susceptible cultivar, than in Marls Kabul, a wilt-resistant cultivar. Winter kill in the field was not affected by the nematode during the first winter, but 50% of plants were killed in the second winter. Forage yield from nematode-infected plants was significantly reduced the second year. Interaction with V. albo-atrum did not significantly reduce forage yields below that of D. dipsaci alone. Pratylenchus penetrans did not increase the severity of wilt symptoms in the presence of V. albo-atrum, nor did it affect forage yield in the greenhouse. It did, however, reduce alfalfa yields in presence of V. albo-atrum under field conditions. D. dipsaci and P. penetrans reproduced faster in Vernal than in Maris Kabul when the fungus was present.     

Effect of Soil Temperature and pH on Resistance of Soybean to Heterodera glycines

Soybean cyst nematode (SCN), Heterodera glycines Ichinohe, is a major pest of soybean, Glycine max L. Merr. Soybean cultivars resistant to SCN are commonly grown in nematode-infested fields. The objective of this study was to examine the stability of SCN resistance in soybean genotypes at different soil temperatures and pH levels. Reactions of five SCN-resistant genotypes, Peking, Plant Introduction (PI) 88788, Custer, Bedford, and Forrest, to SCN races 3, 5, and 14 were studied at 20, 26, and 32 C, and at soil pH''s 5.5, 6.5, and 7.5. Soybean cultivar Essex was included as a susceptible check. Temperature, SCN race, soybean genotype, and their interactions significantly affected SCN reproduction. The effect of temperature on reproduction was quadratic with the three races producing significantly greater numbers of cysts at 26 C; however, reproduction on resistant genotypes remained at a low level. Higher numbers of females matured at the soil pH levels of 6.5 and 7.5 than at pH 5.5. Across the ranges of temperature and soil pH studied, resistance to SCN in the soybean genotypes remained stable.     

Meloidogyne incognita Resistance Characteristics in Tomato Genotypes Developed for Processing

Nine resistant processing tomato (Lycopersicon esculentum) cultivars and advanced lines were compared with four susceptible cultivars in 1,3-dichloropropene-fumigated and nontreated plots on Meloidogyne incognita-infested sites over 3 years. Yield of all resistant genotypes grown in nontreated and nematicide-treated plots did not differ and was greater than yield of susceptible genotypes. M. incognita initial soil population densities caused 39.3-56.5% significant (P = 0.05) yield suppressions of susceptible genotypes. Nematode injury to susceptible plants usually caused both fruit soluble solids content and pH to increase significantly (P = 0.05). Only trace nematode reproduction occurred on resistant genotypes in nontreated plots, whereas large population density increases occurred on susceptible genotypes. Slightly greater nematode reproduction occurred on resistant genotypes at the southern desert location, where soil temperature exceeded 30 C, than at other locations. At two locations resistant MOX 3076 supported greater reproduction than other resistant genotypes.     

Effect of Temperature on Infection and Survival of Rotylenchulus reniformis

From infestation of lettuce with preinfective females to egg deposition, populations of Rotylenchulus reniformis from Baton Rouge, Louisiana; Lubbock and Weslaco, Texas; and Mayaguez, Puerto Rico, required 41, 13, 7, and 7 days at 15, 20, 25, and 34 C, respectively. No nematode infection occurred at 10 C with any R. reniformis population, and the population from Puerto Rico did not reproduce at 15 C. Nematode survival was not influenced by temperature, since populations from Texas and Louisiana survived for 6 months without a host at - 5 , - 1 , 4, and 25 C. Survival of R. reniformis was substantially influenced by soil moisture. Soil moistures greater than 7% (< 1 bar) aided nematode survival at storage temperature of 25 C, whereas moisture adversely affected nematode survival below freezing. Soil moisture below 4% (> 15 bars) favored nematode survival below freezing but adversely affected nematodes in soils stored at 25 C. Soil moisture effects on nematode survival were less accentuated at 4 and 0 C.     

Influence of Population Densities of Heterodera schachtii on Sugar Beets Grown in Microplots

High initial population densities of Heterodera schachtii larvae (36 and 108/gm of soil) greatly retarded the seedling emergence of sugar beet ''Monogerm CSF 1971'' in Vineland fine sandy loam. In comparison with controls, initial population densities (P_i''s) of 1.7, 3.0, 6.2, and 14.4 larvae/gm of soil respectively reduced the weight of storage roots by 38, 56, 64, and 92%. Weights of tops also decreased with increases in P_i; weights of tap and small feeder roots tended to be higher at all P_i''s except the highest. Sucrose percentage was not affected by any initial nematode density. The populations were lower at midseason than at seeding, and at harvest had increased greatly, with respective populations of 339, 402, 222, and 140 larvae/gm of soil. At harvest, cysts/gm of soil and cysts/gm of root were respectively 4.4 and 72, 6.1 and 99, 6.1 and 191, and 5.8 and 140. The maximum rate of multiplication was 150-200. and maximum density was 400 larvae/gm of soil. The high pathogenicity and multiplication rate of the nematode was attributed to optimum temperature conditions and soil type.     

Analysis of Crop Losses in Tomato Due to Pratylenchus penetrans

The effects of Pratylenchus penetrans upon yields of ''Veebrite'' tomato were studied at initial soil population densities (P_i) of 360, 2,010, 4,580, and 14,360 nematodes/kg of soil in 20-cm (i.d.) clay-tile microplots. The lowest P_i appeared to stimulate fruit production. Higher P_i''s suppressed fruit production (total weight of marketable tomatoes and numbers of intermediate- and large-sized fruits), in comparison to control yields, the highest P_i resulted in 38% fewer fruits which weighed 44% less. These losses were at least partly due to a delay in fruit ripening, caused by the nematodes, which did not become apparent until the fourth week. Nematode populations in the soil increased at all but the highest P_i; final populations were around 7,000/kg of soil. Nematode populations in roots ranged from 230-590/gm of root at the completion of the experiment. Nematode control by fumigation would definitely be warranted at soil population densities of 2,000/kg or higher; with 500-2,000/kg, the decision to fumigate would depend on soil type and economic and hiological factors.     

Early Growth of Soybean as Altered by Heterodera glycines,Phenamiphos and/or Alachlor

Greenhouse and field experiments were conducted to determine the effects of phenamiphos and/or alachlor on early growth of soybean, root morphology, and infection and resurgence of Heterodera glycines (race 1). All tests were planted to ''Ransom'' soybeans. In greenhouse experiments without nematodes, root growth was inhibited at 5 days by alachlor treatments and at 10 days by phenamiphos treatments; with nematodes, phenamiphos treatments enhanced root growth. Phenamiphos also suppressed early penetration of soybean roots by H. glycines in the greenhouse. Early soybean growth parameters among treatments were generally similar in the field. Nematode penetration was limited with treatments containing phenamiphos at one location. Plants treated with only alachlor had less nematode infection than did the control; however, plants treated with herbicide/nematicide combinations had more nematode penetration than did plants treated with phenamiphos alone. Alterations of root growth and interference with the efficacy of phenamiphos are two processes by which alachlor may enhance soybean susceptibility or suitability to H. glycines.     

Influence of Meloidogyne chitwoodi and M. hapla on Wheat Growth

Meloidogyne chitwoodi reduced the growth of winter wheat ''Nugaines'' directly in relation to nematode density in the greenhouse, The relationship between top dry weight and initial nematode density suggests a tolerance limit of Nugaines wheat to M. chitwoodi of between 0.03 and 0.18 eggs/cm³ of soil; the value for relative minimum plant top weight was 0.45 g and 0.75 g, respectively. Growth of wheat in field microplots containing four population densities (0.003, 0.05, 0.75 and 9 eggs/cm³ soil) was not affected significantly at any inoculum level compared to controls during September to July, However, suppression of head weights of ''Fielder'' spring wheat grown May-July occurred in microplots initially infested with 0.75 and 9 eggs/cm³ soil. Reproduction (Pf/Pi) was poorer at these two inoculum levels as compared to the lower densities. In another greenhouse experiment, roots of wheat cultivars Fielder, ''Fieldwin,'' ''Gaines,'' ''Hyslop,'' and Nugaines became infected by M. chitwoodi, but not by M. hapla. Reproduction of M. chitwoodi was less on Gaines and Nugaines than on Fielder, Fieldwin, or Hyslop.     

Differential Effects of Pratylenchus neglectus Populations on Single and Interplantings of Alfalfa Grasses

The invasion by three different Utah populations of Pratylenchus neglectus (UTI, UT2, UT3) was similar in single and interplantings of ''Lahontan'' alfalfa and ''Fairway'' crested wheatgrass at 24 ñ 3 °C. Population UT3 was more pathogenic than UT1 and UT2 on both alfalfa and crested wheatgrass. Inoculum density was positively correlated with an invasion by P. neglectus. Invasions by UT3 at all initial populations (Pi) exceeded that of UT1 and UT2 for both single and interplanted treatments. The greatest reductions in shoot and root weights of alfalfa and crested wheatgrass were at a Pi of 8 P. neglectus/cm³ soil. Pi was negatively correlated with alfalfa and crested wheatgrass shoot and root growth and nematode reproduction. The reproductive factor (Rf) for UT3 exceeded that of UT1 and UT2 in single and interplantings at all inoculum levels. There were no differences in Rfin the Utah populations in single or interplantings. A nematode invasion increased with temperature and was greatest at 30 °C. Population UT3 was more pathogenic than UT1 and UT2 and reduced shoot and root growth at all soil temperatures. Populations UT1 and UT2 reduced shoot and root growth at 20-30 °C. Soil temperature was negatively correlated with shoot and root growth and positively correlated with nematode reproduction. Reproduction of UT3 exceeded that of UT1 and UT2 at all soil temperatures.     

A Nematicide Seed Treatment to Control Ditylenchus dipsaci on Seedling Alfalfa

Three nematicides were evaluated as seed treatments to control the alfalfa stem nematode (Ditylenchus dipsaci) on seedling alfalfa. Alfalfa seeds were soaked for 10 hours in a 0.5% (formulated by weight) concentration of either carbofuran, phenamiphos or oxamyl in acetone with no adverse effect on seed germination. All three treatments decreased nematode damage and increased survival of ''Ranger'' (susceptible) and ''Lahontan'' (resistant) alfalfa plants, when seeds were planted in soil infested with D. dipsaci. Mean live plant counts after 6 weeks in the untreated control, acetone alone, carbofuran, phenamiphos, and oxamyl treatments, respectively, were 4.3, 6.3, 19.0, 19.8, and 19.0 for Lahontan and 4.5, 1.5, 18.5, 19.3, and 18.0 for Ranger from 20 seeds/pot. Nematicide seed treatments resulted in significantly healthier Ranger alfalfa plants 4 months after planting. The combination of seed treatment and host resistance may provide a means of establishing alfalfa in an alfalfa monocropped system where soil populations of D. dipsaci are high.     

Population Development of Meloidogyne incognita on Soybean Defoliated by Pseudoplusia includens

Greenhouse studies examined population densities of Meloidogyne incognita race 4 on soybean (Glycine max ''Davis'') defoliated by larvae of soybean looper (Pseudoplusia indudens (Walker)). Plants were defoliated over a 2-week period beginning 5 weeks after seedlings were transplanted. Four groups of plants were infested with nematodes (5,000 eggs/pot) at 2-week intervals to allow harvesting of plants at 0, 2, 4, and 6 weeks postdefoliation (WPD). Plants in each group were harvested 4 weeks after nematode infestation. Root and nodule weights of defoliated plants were suppressed at 0 WPD, but differences were not detectable at 2, 4, and 6 WPD. Population densities of M. incognita were similar on defoliated and control plants at 0 WPD but were greater on defoliated plants at 4 and 6 WPD. Percentage hatching of eggs produced on the latter plants also was higher. Effects of insect-induced defoliation on development of M. incognita remained detectable even after soybean plant growth apparently returned to normal.     

Interactions between Nematodes and Earthworms: Enhanced Dispersal of Steinernema carpocapsae

Dispersal of the nematode Steinernema carpocapsae (All strain), applied on the top or the bottom of soil columns, was tested in the presence or absence of two earthworm species, Lumbricus terrestris or Aporrectodea trapezoides. Nematode dispersal was estimated after a 2-week period with a bioassay against the greater wax moth, Galleria mellonella. Vertical dispersal of nematodes was increased in the presence of earthworms. When nematodes were placed on the surface of soil columns, significantly more nematodes dispersed to the lower half of the columns when either earthworm species was present than when earthworms were not present. When nematodes were placed on the bottom of soil columns, significantly more nematodes dispersed to the upper half of the columns when L. terrestris was present than when A. trapezoides was present or in the absence of earthworms. Because nematodes were found on the exterior and in the interior of earthworms, nematode dispersal may be enhanced by direct contact with the earthworms.     

Early Root Response to Meloidogyne incognita in Resistant and Susceptible Alfalfa Cultivars

The early events of Meloidogyne incognita behavior and associated host responses following root penetration were studied in resistant (cv. Moapa 69) and susceptible (cv. Lahontan) alfalfa. Ten-day-old seedlings of alfalfa cultivars were inoculated with second-stage juveniles (J2) and harvested 12, 24, 48, and 72 hours and 7, 14, and 21 days later. Both cultivars supported similar root penetration and initial J2 migration. By 72 hours after inoculation the majority of J2 were amassed inside the vascular cylinder in roots of susceptible Lahontan, while J2 had not entered the vascular cylinder of resistant Moapa 69 and remained clumped at the root apex. Nematode development progressed normally in Lahontan, but J2 were not observed in Moapa 69 after day 7. The greatest differences between RNA translation products isolated from inoculated and uninoculated roots of Lahanton occurred 72 hours after inoculation. Only minor differences in gene expression were observed between inoculated and uninoculated Moapa 69 roots at 72 hours. Comparison of translation products from inoculated versus mechanically wounded Lahontan roots revealed products that were specific to or enhanced in nematode-infected plants. Moapa 69 appears to possess a type of resistance to M. incognita that does not depend on a conventional hypersensitive response.     

Activity and Differential Induction of Chitinase Isozymes in Soybean Cultivars Resistant or Susceptible to Root-knot Nematodes

Host physiological events in relation to infestation by parasitic nematodes are not well documented. Soybean plant responses to Meloidogyne incognita infestation were compared to resistant (Bryan) and susceptible (Brim) cultivars at 0, 1, 3, 10, 20, and 34 days after infestation (DAI). The resistant cultivar had higher chitinase activity than the susceptible cultivar at every sample time beginning at 3 DAI. Results from isoelectric focusing gel electrophoresis analyses indicated that three acidic chitinase isozymes with isoelectric points (pIs) of 4.8, 4.4, and 4.2 accumulated to a greater extent in the resistant compared to the susceptible cultivar following challenge. SDS-PAGE analysis of root proteins revealed that two proteins with molecular weights of approximately 31 and 46 kD accumulated more rapidly and to a higher level in the resistant than in the susceptible cultivar. Additionally, three major protein bands (33, 22, and 20 kD) with chitinase activity were detected with a modified SDS-PAGE analysis in which glycolchitin was added into the gel matrix. These results indicate that higher chitinase activity and early induction of specific chitinase isozymes may be associated with resistance to root-knot nematode in soybean.