Effects of Interactions among Heterodera glycines,Meloidogyne incognita,and Host Genotype on Soybean Yield and Nematode Population Densities

The effects of host genotype and initial nematode population densities (Pi) on yield of soybean and soil population densities of Heterodera glycines (Hg) race 3 and Meloidogyne incognita (Mi) race 3 were studied in a greenhouse and field microplots in 1983 and 1984. Centennial (resistant to Hg and Mi), Braxton (resistant to Mi, susceptible to Hg), and Coker 237 (susceptible to Hg and Mi) were planted in soil infested with 0, 31, or 124 eggs of Hg and Mi, individually and in all combinations, per 100 cm³ soil. Yield responses of the soybean cultivars to individual and combined infestations of Hg and Mi were primarily dependent on soybean resistance or susceptibility to each species separately. Yield of Centennial was stimulated or unaffected by nematode treatments, yield of Braxton was suppressed by Hg only, and yield suppressions caused by Hg and Mi were additive and dependent on Pi for Coker 237. Other plant responses to nematodes were also dependent on host resistance or susceptibility. Population densities of Mi second-stage juveniles (J2) in soil were related to Mi Pi and remained constant in the presence of Hg for all three cultivars. Population densities of Hg J2 on the two Hg-susceptible Cultivars, Braxton and Coker 237, were suppressed in the presence of Mi at low Hg Pi.     

Parasitic Variability of Meloidogyne incognita Populations on Susceptible and Resistant Cotton

Root gall induction and egg production by the four recognized host races and two cytological races of Meloidogyne incognita were compared on cotton Gossypium hirsutum cvs. Deltapine 16 (root-knot susceptible) and Auburn 634 (highly resistant). The 12 nematode populations included in the study were from various parts of the world. No population increases occurred on the highly resistant cultivar. After 45 days, populations of host races 1 and 2 induced slight root galling on both cuhivars with only limited reproduction. Host race 4 populations induced moderate root galling with higher reproduction on Deltapine 16 than that of race 1 or race 2 populations. Host race 3 populations induced severe root galling with population density increases of 7-30-fold. In a complementary study, 24 cotton cultivars or breeding lines were compared for suitability as hosts for a typical population of M. incognita race 3. The poorest hosts, ''Aubnru 623,'' ''Auburn 634,'' and ''McNair 220,'' yielded fewer eggs after 45 days than were added initially. The best hosts - ''M-8.'' ''DES 24-8,'' ''McNair 235,'' and ''Coker 20l'' - yielded > 5 times as many eggs as were added initially.     

Host Suitability and Susceptibility of Glycine max cv. Bedford to Race 1 of Heterodera glycines

Populations of Heterodera glycines identifiable as race 1 reproduced on the race 1 resistant ''Bedford'' soybean. A Beaufort County, North Carolina, population had an index of parasitism of 112% on Bedford in greenhouse tests. Indices of parasitism for this population on race 1 resistant cultivars Pickett 71, Centennial, and Forrest were less than 10%. The Beaufort County population had significantly greater reproduction on Bedford in microplots than did populations of race 3 or race 4. In field tests, a race 1 population suppressed yields of Bedford but not yields of Centennial. Based on these data, Bedford is no longer recommended in North Carolina as a race 1 resistant cultivar.     

An Alternative Method for Evaluating Soybean Tolerance to Heterodera glycines in Field Plots

Alternate planting dates and periodic destruction of the previous year''s soybean crop as well as 1-year bare fallow were used to establish a range of population densities ofHeterodera glycines for the subsequent year. Soybean cultivar Coker 156 (susceptible, moderately tolerant) was compared to cultivars Essex (susceptible, intolerant) and Bedford (resistant) to evaluate tolerance at different H. glycines population densities established through the previous year''s treatments. Yield of Coker 156 was consistently intermediate between yields of Bedford and Essex in 1986 and 1987. Yield of Essex was negatively correlated (P = 0.05) with preplant egg numbers of H. glycines in 1987, whereas yield of Bedford and Coker 156 were not related to nematode density. Reproduction of H. glycines was greater (P = 0.05) on the moderately tolerant Coker 156 than on either of the other cultivars.     

Susceptibility of Soybean Introductions to Races 1, 2, 3, and 4 of Heterodera glycines

Thirteen soybean plant introduction (PI) lines, selected for their apparent susceptibility to Heterodera glycines, were compared with cultivar Lee 74 as hosts of H. glycines races 1, 2, 3, and 4. Race 3 produced the highest average number of females of the four races. Compared to Lee 74, more (P = 0.05) females of H. glycines race 1 were extracted from eI 274420, PI 274423, and PI 317333; PI 86457 had more females of H. glycines race 2; and PI 86443, PI 86457, PI 261467, PI 274420, PI 274421, and PI 274423 had more females of H. glycines race 3. Similar numbers of females of H. glycines race 4 developed on all of the soybean lines and Lee 74. PI 274421, PI 274420, or PI 196159 could provide a more or equally susceptible host for H. glycines races 1, 2, 3, and 4 than Lee 74. One of these three lines could be substituted for Lee as the standard susceptible cultivar in the race determination test.     

Effects of Continuous Cropping of Resistant and Susceptible Cultivars on Reproduction Potentials of Heterodera glycines and Globodera tabacum solanacearum

The reproductive potentials of Heterodera glycines (mixture of races 3 and 4 and unidentified races) and a tobacco cyst nematode Globodera tabacum solanacearum were studied in the field. The experiments involved four cultivars of soybean Glycine max and four cultivars of Nicotiana tabacum. The reproductive potential of the H. glycines population was high on Essex and Lee 74 soybean but low on Forrest and Bedford over the 3 years (1982-84) of continuous cropping. The reproductive potential of H. glycines was 12% on Forrest and 6% on Bedford in 1982 but increased to 37 and 35% in 1983 and to 71 and 41% in 1984, respectively, on these two cultivars. The reproductive potential of G. tabacum solanacearum was high on McNair 944 and Coker 319 tobacco cultivars and low on VA 81 and PD 4 over the 3 years of cropping. The reproductive potential of G. tabacum solanacearum on VA 81 and PD 4 was 18 and 17% in 1982, 7 and 16% in 1983, and 5 and 5% in 1984, respectively. The changes in reproductive potentials of H. glycines and G. tabacum solanacearum may be related to inherent genetic variability in the systems that control reproduction of the two cyst nematodes and nature of resistance incorporated in the soybean and tobacco cultivars.     

Variation in Resistance of Soybean Lines to Races of Heterodera glycines

The objective of this study was to determine the interrelationships of Heterodera glycines races based on their resistance to soybean (Glycine max) cultivars and lines against which they were tested. Greenhouse tests determined the numbers of females of each of eight races of H. glycines that developed on 277 to 522 soybean cultivars and lines. A Female Index (number of females on a test cultivar as a percentage of the number on ''Lee 74'') was calculated and used in frequency distributions, correlations, and duster analyses of the resistance reactions to the different races in an attempt to determine relationships among cultivars. Frequency distribution patterns of all cultivars and lines tested against each race were skewed in favor of resistance, and in some cases bimodality was observed. The majority of correlations between pairs of races were highly significant. Cluster analyses based on the correlations divided eight races into four clusters that explained 73% of the variation in resistance. Cluster 1 was comprised of races 2, 4, and 14; Cluster 2 was comprised of races 6 and 9; Cluster 3 was comprised of races 1 and 3; and Cluster 4 was comprised of race 5. The information obtained in this study could increase the efficiency of testing resistant soybean breeding lines for resistance to H. glycines.     

Effects of Heterodera glycines and Meloidogyne incognita on Early Growth of Soybean

Greenhouse and field microplot studies were conducted to compare soybean shoot and root growth responses to root penetration by Heterodera glycines (Hg) and Meloidogyne incognita (Mi) individually and in combination. Soybean cultivars Centennial (resistant to Hg and Mi), Braxton (resistant to Mi, susceptible to Hg), and Coker 237 (susceptible to Hg and Mi) were selected for study. In the greenhouse, pot size and number of plants per pot had no effect on Hg or Mi penetration of Coker 237 roots; root weight was higher in the presence of either nematode species compared with the noninoculated controls. In greenhouse studies using a sand or soil medium, and in field microplot studies, each cultivar was grown with increasing initial population densities (Pi) of Hg or Mi. Interactions between Hg and Mi did not affect early plant growth or number of nematodes penetrating roots. Root penetration was the only response related to Pi. Mi penetration was higher in sand than in soil, and higher in the greenhouse than in the field, whereas Hg penetration was similar under all conditions. At 14 days after planting, more second-stage juveniles were present in roots of susceptible than in roots of resistant plants. Roots continued to lengthen in the greenhouse in the presence of either Mi or Hg regardless of host genotype, but only in the presence of Mi in microplots; otherwise, responses in field and greenhouse studies were similar and differed only in magnitude and variability.     

Effects of Environments,Meloidogyne incognita Inoculum Levels,and Glycine max Genotype on Root-knot Nematode-Soybean Interactions in Field Microplots

Five soybean cultivars (Braxton, Gordon, Jeff, Bragg, and Wright) resistant to Meloidogyne incognita (Mi) and three susceptible cultivars (Coker 156, GaSoy 17, and Coker 237) were grown at two locations for four seasons in microplots with increasing initial soil population densities (Pi) of Mi. The resistant cultivars and Coker 156 yielded better than GaSoy 17 and Coker 237 at all Pi. Yield response was dependent on environmental conditions and at one location was stimulated on Braxton, Gordon, Jeff, and Bragg by low Pi. Although Mi reproduced well on all cultivars, the pattern of reproduction differed. Population densities of Mi leveled off after 90 days on GaSoy 17 and Coker 237 but were still increasing after 120 days on the resistant cultivars; population densities were lower on resistant than on the susceptible cultivars. The population density of Mi on Coker 156 after 120 days was intermediate between those on the other susceptible and on the resistant cultivars. Mi population densities followed the same pattern under varying environmental conditions.     

Effects of Peanut-Tobacco Rotations on Population Dynamics of Meloidogyne arenaria in Mixed Race Populations

A 3-year microplot study was initiated to characterize the population dynamics, reproduction potential, and survivorship of single or mixed populations of Meloidogyne arenaria race 1 (Ma1) and race 2 (Ma2), as affected by crop rotations of peanut ''Florigiant'' and M. incognita races 1 and 3-resistant ''McNair 373'' and susceptible ''Coker 371-Gold'' tobacco. Infection, reproduction, and root damage by Ma2 on peanut and by Ma1 on resistant tobacco were limited in the first year. Infection, reproduction, and root-damage potentials on susceptible tobacco were similar for Ma1 and Ma2. In the mixed (1:1) population, Ma1 was dominant on peanut and Ma2 was dominant on both tobacco cultivars. Crop rotation affected the population dynamics of different nematode races. For years 2 and 3, the low numbers of Ma1 and Ma2 from a previous-year poor host increased rapidly on suitable hosts. Ma1 had greater reproduction factors ([RF] = population density at harvest/population density at preplandng) than did Ma2 and Ma1 + Ma2 in second-year peanut plots following first-year resistant tobacco, and in third-year peanut plots following second-year tobacco. In mixed infestations, Ma1 predominated over Ma2 in previous-year peanut plots, whereas Ma2 predominated over Ma1 in previous-year tobacco plots. Moderate damage on resistant tobacco was induced by Ma1 in the second year. In the third year, moderate damage on peanut was associated with ''Ma2'' from previous-year peanut plots. The resistant tobacco supported sufficient reproduction of Ma1 over 2 years to effect moderate damage and yield suppression to peanut in year 3.     

Effects of Meloidogyne incognita on Nitrogen Fixation in Soybean

The effects of a North Carolina population of Meloidogyne incognita on N₂ fixation on root-knot-susceptible ''Lee 68'' and moderately resistant ''Forrest'' soybean were evaluated 50, 75, I00, and 135 days after inoculation with nematodes. Nematodes stimulated N₂ fixation in Lee 68 by 50 days and in Forrest by 75 days. At all other intervals, N₂ fixation was either depressed or unaffected by nematodes. Additional observations indicate that the susceptibility of Lee 68 is associated with greater rates of penetration by larvae and more favorable responses of host tissues to nematodes than occur in Forrest. With time, however, the histological reactions of both hosts became less favorable for nematode development. Resistant or hypersensitive responses became common in Forrest by 75 days but not in Lee 68 until 90 days after inoculation. This population of M. incognita may stimulate N₂ fixation at a specific time interval and depress it at others; therefore, disease of susceptible soybeans caused by this nematode is probably not primarily due to a net loss of fixed nitrogen but to pathogenicity similar to that which occurs on nonlegume hosts.     

Influence of Calonectria crotalariae on Reproduction of Heterodera glycines on Soybean

Calonectria crotalariae enhanced root penetration of Lee 74 (susceptible) and Centennial (resistant) soybeans by juveniles of race 3 of Heterodera glycines. Numbers of cysts in and on the roots of Lee 74 increased during the first 30 days in the presence of the fungus. Percentage of root infection by the fungus increased at 40 days in Lee 74 in the presence of the nematode. Numbers of cysts in soil at 80 and 120 days after inoculation with both organisms accounted for the significantly increased nematode population levels on Lee 74. In the presence of the fungus on the resistant cultivar, significantly increased levels of cysts were recovered from soil at 120 days. Fungus infection of Centennial roots also infected with the nematode increased from 58 to 86% at 120 days. An inoculum timing study in which Lee 74 was infested with the nematode and fungus individually, sequentially, and in combination at days 0 and 35 indicated that enhanced nematode reproduction was related more to early plant-fungus than to early plant-fungus-nematode interaction(s).     

Galling and Yields of Soybean Cultivars Grown in Meloidogyne arenana-Infested Soil

Field trials with 39 soybean cultivars and five breeding lines from public and private sources were conducted from 1982 through 1985 at sites infested with Meloidogyne arenaria. Nematode population densities and root-knot galling were measured for each soybean entry. All were efficient hosts for the nematode, and average juvenile numbers in the soil increased 5-50 × from planting to harvest. Differences (P < 0.05) in galling were found among entries in each year. Centennial, Cobb, Coker 368, Hutton, and Jeff cultivars, recognized for their resistance to M. incognita, were severely galled and yielded poorly. Bedford, Forrest, A7372, Bragg, Braxton, Gordon, and Kirby, also recognized for their resistance to M. incognita, were among the least galled cultivars. Yields of all entries, however, were too low to justify their planting in sites heavily infested with M. arenaria.     

Host Suitability of Selected Hybrids,Varieties and Inbreds of Corn to Populations of Meloidogyne spp.

Rates of reproduction of root-knot nematodes on corn varied with Meloidogyne species, with different populations of certain species, and with corn cultivars. M. arenaria, M. incognita and M. javanica reproduced at varying rates on all corn cultivars tested. None of the three selections of M. hapla reproduced on corn. Most of the Meloidogyne populations increased more rapidly on ''Coker'' and ''Pioneer'' hybrids than on ''McNair'' hybrids or on open-pollinated varieties or inbreds. Nematodes often reduced root growth, but the differences within given nematode-cultivar treatments were not usually significant. Root growth of ''Coker 911,'' which supported a high rate of reproduction, was affected less than ''Pioneer 309B'' which supported a low rate of nematode reproduction.     

Damage and Reproductive Potentials of Pratylenchus brachyurus and P. penetrans on soybean

Damage and reproductive potentials of Pratylenchus brachyurus and P. penetrans on soybean, Glycine max, cvs. Essex, Forrest, and Lee 68, were determined in microplot tests. Cultivar Essex was generally tolerant to P. brachyurus. Yield of Forrest was suppressed linearly with increasing P_i''s in the sandy soil (r = -0.92) and loamy sand soil (r = -0.99). Low to moderate P_i''s in the sandy clay loam gave an increase in yields as compared to plants without nematodes. Yield was not affected by this nematode in muck. Lee 68 was very sensitive to P. penetrans in microplots. Yield vs. P_i was fitted by a quadratic model (r = 0.82) with yield decreasing sharply as P_i''s were increased. The reproduction of both species decreased with increases in P_i. Lee 68 was a good host for P. penetrans, whereas Essex and Forrest were fair to poor hosts for P. brachyurus.     

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.     

Resistance to Meloidogyne spp. in Allohexaploid Wheat Derived from Triticum turgidum and Aegilops squarrosa

Expression of resistance to Meloidogyne incognita and M. javanica from Aegilops squarrosa was studied in a synthetic allohexaploid produced from Triticum turgidum var. durum cv. Produra and Ae. squarrosa G 3489. The reproductive rate of different races of M. incognita and M. javanica, expressed in eggs per gram of fresh root, was low (P < 0.05) on the synthetic allohexaploid and the resistant parent, Ae. squarrosa G 3489, compared with different bread and durum wheat cultivars. Reproduction of race 2 and race 3 of M. incognita and an isolate of M. javanica was studied on the synthetic allohexaploid and seven cultivars of T. aestivum: Anza, Coker 747, Coker 68-15, Delta Queen, Double Crop, McNair 1813, and Southern Bell. The latter six cultivars are grown in the southeastern United States and reportedly were resistant to M. incognita. Significant differences (P < 0.05) were detected in nematode reproduction on the seven bread wheat cultivars. Reproduction of M. incognita race 3 and M. javanica was highest on Anza. Reproductive rates on the six southeastern United States bread wheat cultivars varied both within and among nematode isolates. The lowest reproductive rates of the three root-knot isolates were detected in the synthetic allohexaploid.     

Variability in Host Preference Among Field Populations of Heterodera glycines

Eighteen soybean fields, six each with race 3, race 4, or Bedford population of Heterodera glycines, were selected for testing of host variability. Each field was divided into three sections, and a bulk soil sample was taken from each section. The 54 bulk soil populations (BSP) and 270 single cyst populations (SCP) were subjected to race determination tests. Tests of the 18 BSP and 90 SCP from the race 3 fields revealed that race 3 was the predominant race; however, 68 of the populations tested were other races. Tests of the 18-BSP and 90 SCP from race 4 fields demonstrated that races 2 and 4 were predominant, with 38 and 39 populations; respectively. Tests of the 18 BSP and 90 SCP from the Bedford population fields revealed tremendous variability. Races 2, 4, and 6 were the predominant races, with 32, 31, and 28 populations, respectively. These results indicate that of the three races studied, the Bedford population is the most variable, race 3 shows considerable variability, and race 4 shows very little.     

Relationship between Meloidogyne incognita and Rotylenchulus reniformis as Influenced by Soybean Genotype

The effect of soybean genotype on competition between Meloidogyne incognita race 2 (Mi) and Rotylenchulus reniformis (Rr) was evaluated in greenhouse and microplot replacement series experiments. Soil in pots containing seedlings of ''Davis'' (susceptible to Mi) or ''Buckshot 66'' (resistant to Mi) was infested with 1,000 vermiform individuals in the following Mi:Rr ratios: 0:0, 100:0, 75:25, 50:50, 25:75, or 0:100. After 91 days, the relative nematode yields (number of nematodes in mixed culture divided by the number in nonmixed culture) of each species were calculated based on soil and root nematode populations expressed as nematodes per gram of dry root tissue. To define the relationship between the two species, calculated relative nematode yields were compared with a theoretical noncompetition model using lack-of-fit regression. In the greenhouse, Mi populations on ''Davis'' were stimulated in the presence of Rr. In microplots, low Mi and Rr population densities likely resulted from severe galling and destruction of feeder roots that probably occurred early in the season. Enhanced susceptibility to Mi was not observed on ''Buckshot 66'', which remained resistant to Mi even when colonized by Rr. Host resistance is a key factor in determining the nature of the relationship between Mi and Rr.     

Reproduction of Meloidogyne incognita Race 3 on Flue-cured Tobacco Homozygous for Rk1 and/or Rk2 Resistance Genes

Most commercial tobacco cultivars possess the Rk1 resistance gene to races 1 and 3 of Meloidogyne incognita and race 1 of Meloidogyne arenaria, which has caused a shift in population prevalence in Virginia tobacco fields toward other species and races. A number of cultivars now also possess the Rk2 gene for root-knot resistance. Experiments were conducted in 2013 to 2014 to examine whether possessing both Rk1 and Rk2 increases resistance to a variant of M. incognita race 3 compared to either gene alone. Greenhouse trials were arranged in a completely randomized design with Coker 371-Gold (C371G; susceptible), NC 95 and SC 72 (Rk1Rk1), T-15-1-1 (Rk2Rk2), and STNCB-2-28 and NOD 8 (Rk1Rk1 and Rk2Rk2). Each plant was inoculated with 5,000 root-knot nematode eggs; data were collected 60 d postinoculation. Percent galling and numbers of egg masses and eggs were counted, the latter being used to calculate the reproductive index on each host. Despite variability, entries with both Rk1 and Rk2 conferred greater resistance to a variant of M. incognita race 3 than plants with Rk1 or Rk2 alone. Entries with Rk1 alone were successful in reducing root galling and nematode reproduction compared to the susceptible control. Entry T-15-1-1 did not reduce galling compared to the susceptible control but often suppressed reproduction.