Interrelationship of Heterodera schachtii and Meloidogyne hapla on Tomato

Invasion of tomato (Lycopersicon esculentum L.) roots by combined and sequential inoculations of Meloidogyne hapla and a tomato population of Heterodera schachtii was affected more by soil temperature than by nematode competition. Maximum invasion of tomato roots, by M. hapla and H. schachtii occurred at 30 and 26 C, respectively. Female development and nematode reproduction (eggs per plant) of M. hapla was adversely affected by H. schachtii in combined inoculations of the two nematode species. Inhibition of M. hapla development and reproduction on tomato roots from combined nematode inoculations was more pronounced as soil temperature was increased over a range of 18-30 C and with prior inoculation of tomato with H. schachtii. M. hapla minimally affected H. schachtii female development, but there was significant reduction in the buildup of H. schachtii when M. hapla inoculation preceded that of H. schachtii by 20 days.     

Self-interactions of Meloidogyne hapla and of Heterodera schachtii on Beta Vulgaris

Double inoculations of sugar beet with larvae of Meloidogyne hapla resulted in a higher galling incidence in only one treatment than did a single inoculation using the same number of larvae. Double inoculations with larvae of Heterodera schachtii, however, resulted in three- to five-fold more cysts in most cases than did single inoculations using the same number of larvae. In general, plants died more quickly after double inoculations than after single inoculations of the same total number of either nematode. Ratios of total soluble carbohydrates to reducing carbohydrates were lower in multiple inoculated treatments than in other treatments. Plants infected with M. hapla had lower quantities of B, K, and P in leaf tissue than noninoculated plants, but no differences were correlated with type of inoculation. Plants inoculated with H. schachtii had lower quantities of B, K, and Mg than noninoculated plants. Also, quantities of Mn, Cu, and Zn were much lower in plants inoculated twice with H. schachtii larvae than in plants inoculated with the same total number of larvae in a single dose.     

Influence of Meloidogyne hapla Chitwood, 1949 on Development and Establishment of Heterodera schachtii Schmidt, 1871 on Beta vulgaris L

Influence of Meloidogyne hapla on estahlishnrent and maturity of Heterodera schachtii in sugarbeet was studied. Results indicated that when the majority of M. hapla were in second, third, or fourth larval stages within plants prior to H. schachtii inoculation, growth and development of the latter was retarded. However, when M. hapla reached the young female stage prior to inoculation of H. schachtii, establishment and development of the latter was greatly enhanced. As M. hapla reached maturity before and after egg production prior to H. schachtii inoculation, establishment and growth of the latter was progressively decreased. In each instance, M. hapla developed independently and matured at the same rate as in plants inoculated with only M. hapla. Usually ratios of total soluble carbohydrates to reducing carbohydrates were lower, but not significantly different, in plants receiving both nematodes as compared to other treatments.     

Pathological Interaction of a Combination of Heterodera schachtii and Meloidogyne hapla on Tomato

Increased culturing of a tomato population of Heterodera schachtii (UT1C) on tomato for 480 days (eight inoculation periods of 60 days each) significantly increased virulence to ''Stone Improved'' tomato. A synergistic relationship existed between Meloidogyne hapla and H. schaehtii on tomato. A combination of H. schachtii (UTIC) and M. hapla significantly reduced tomato root weights by 65, 64, and 61% below root weights of untreated controls, and single inoculations of M. hapla and H. schachtii, respectively. This corresponded to root reductions of 42, 44, and 46% from a combination of H. schachtii (UT1B) and M. hapla. Antagonism existed between H. schachtii and M. hapla with regard to infection courts and feeding sites. The root-knot galling index dropped from 6.0 with a single inoculation of M. hapla to 4.3 and 3.3 with combined inoculations of M. hapla plus UT1B and M. hapla plus UTIC cyst nematode populations. The pathological virulence of H. schachtii to sugarbeet was not lost by extended culturing on tomato; there were no differences in penetration, maturation, and reproduction between sugarbeet populations continually cultured on sugarbeet and the population continually cultured on tomato.     

Relationship Between Heterodera schachtii,Meloidogyne hapla,and Nacobbus aberrans on Sugarbeet

Heterodera schachtii, Meloidogyne hapla, and Nacobbus aberrans either alone, or in various combinations with each other, can, when inoculated at a concentration of 12 second-stage juveniles/ cm³ of soil, cause a significant (P = 0.01) suppression of growth of sugarbeet (cv. Tasco AH14) seedlings. M. hapla and H. schachtii decreased growth of sugarbeet more than N. aberrans over a 60-day period. The adverse effect of N. aberrans on the final population/initial population (Pf/Pi) ratio for either M. hapla or H. schachtii was dependent on time, and was more accentuated on that of M. hapla than on that of H. schachtii. Neither M. hapla nor H. schachtii had an adverse effect on the Pf/ Pi ratio of N. aberrans. N. aberrans is considered to be less aggressive on sugarbeet than either H. schachtii or M. hapla. Sections of sugarbeet roots infected simultaneously with H. schachtii and N. aberrans showed scattered vascular elements between the N. aberrans syncytium located in the central part of the root and that of H. schachtii in the peripheral position.     

Induced Resistance to Meloidogyne hapla by other Meloidogyne species on Tomato and Pyrethrum Plants

Advance inoculation of the tomato cv. Celebrity or the pyrethrum clone 223 with host-incompatible Meloidogyne incognita or M. javanica elicited induced resistance to host-compatible M. hapla in pot and field experiments. Induced resistance increased with the length of the time between inoculations and with the population density of the induction inoculum. Optimum interval before challenge inoculation, or population density of inoculum for inducing resistance, was 10 days, or 5,000 infective nematodes per 500-cm³ pot. The induced resistance suppressed population increase of M. hapla by 84% on potted tomato, 72% on potted pyrethrum, and 55% on field-grown pyrethrum seedlings, relative to unprotected treatments. Pyrethrum seedlings inoculated with M. javanica 10 days before infection with M. hapla were not stunted, whereas those that did not receive the advance inoculum were stunted 33% in pots and 36% in field plots. The results indicated that advance infection of plants with incompatible or mildly virulent nematode species induced resistance to normally compatible nematodes and that the induced resistance response may have potential as a biological control method for plant nematodes.     

The Interrelationship of Heterodera schachtii and Ditylenchus dipsaci on Sugarbeet

Heterodera schachtii significantly (P = 0.05) reduced sugarbeet root growth below that of uninoculated controls at 20, 24, and 28 C, and Ditylenchus dipsaci significantly (P = 0.05) reduced root growth below that of uninoculated controls at 16, 20, 24, and 28 C. A combination of H. schachtii and D. dipsaci significantly (P = 0.05) reduced root growth below that of single inoculations of H. schachtii at all temperatures and D. dipsaci at 20, 24, and 28 C. Single inoculations of H. schachtii and D. dipsaci significantly (P = 0.05) reduced top growth of sugarbeet below that of uninoculated controls at 20, 24, and 28 C, and 16, 20, 24, and 28 C, respectively. A combination of the two nematodes significantly (P = 0.05) reduced top growth below that of single inoculations of H. schachtii at all temperatures. However, a combination of the two nematodes failed to significantly (P = 0.05) reduce top growth below that of single inoculations of D. dipsaci at any temperature. Inoculations of either H. schachtii or D. dipsaci did not affect penetration of the other nematode, and D. dipsaci did not affect development and reproduction of H. schachtii. D. dipsaci did not reproduce on sugarbeet.     

Effects of Oxime Carbamate Nematicides on Development of Heterodera schachtii on Sugarbeet

Treatment of sugarbeet, Beta vulgaris L., with aldicarb, aldicarb sulfoxide, or aldicarb sulfone 10 days after plants were inoculated with Heterodera schachtii prevented development of the nematode, but second-stage larvae penetrated the roots. These chemicals had no measurable effects on nematodes in plants treated 15 days after inoculation. The tests established that soil treatments of aldicarb are directly or indirectly lethal to larvae developing within roots of sugarbeet. Heterodera schachtii failed to develop on root slices of red table beet grown in soil treated with aldicarb or aldicarb sulfoxide. Similar treatment of plants with aldicarb sulfone or oxamyl did not affect subsequent development of H. schachtii on root slices of treated plants.     

Antioxidant Enzymes in Phytoparasitic Nematodes

Presence of different antioxidant enzymes, such as superoxide dismutase (SOD), catalase, and ascorbate, p-phenilendiamine-pyrocathecol (PPD-PC), o-dianisidine, and guaiacol isoperoxidases, was shown in the phytoparasific nematode species Meloidogyne incognita, M. hapla, Globodera rostochiensis, G. pallida, Heterodera schachtii, H. carotae, and Xiphinema index. The activity of the enzymes tested differed among the life stages examined. SOD was present in cysts but was not detected in Meloidogyne egg masses. Catalase activity of Meloidogyne females was higher than that of preparasitic stages and cyst-nematode females. For the first time, ascorbate peroxidase was found to occur commonly in phytoparasitic nematodes, with the highest activity in the invading life-stages. In all the life stages examined, the antioxidant enzyme activities of M. hapla were markedly higher than those of M. incognita. Glutathione peroxidase was not found in the species examined.     

Effect of Soil Temperature on Reproduction of Meloidogyne chitwoodi and M. hapla Alone and in Combination on Potato and M. chitwoodi on Rotation Plants

Meloidogyne chitwoodi developed and reproduced more rapidly than M. hapla in potato roots at 15, 20, or 25 C when both species of nematodes were inoculated simultaneously at 250 or 1,000 juveniles of each. At 30 C significantly more M. hapla than M. chitwoodi females were found at the lower inoculum level after 41 days. More M. chitwoodi than M. hapla juveniles were extracted from soil at 15, 20, and 25 C, but only at the lower inoculum level at 30 C. Potato was considered a more suitable host for M. chitwoodi than M. hapla because of M. chitwoodi''s greater reproduction at 15, 20, and 25 C. Corn and wheat cultivars tested supported M. chitwoodi reproduction at temperatures of 10, 15, 20, and 25 C, but fewest eggs were produced on these plants at 20 C. Temperatures of 10 to 25 C had little influence on the low reproduction of M. chitwoodi on four alfalfa cultivars. M. chitwoodi reproduced on the alfalfa entry Mn PL9HF.     

Interaction of Ditylenchus dipsaci and Meloidogyne hapla on Resistant and Susceptible Plant Species

Numbers ofDitylenchus dipsaci or Meloidogyne hapla invading Ranger alfalfa, Tender crop bean, Stone Improved tomato, AH-14 sugarbeet, Yellow sweet clover, and Wasatch wheat from single inoculations were not significantly different from numbers by invasion of combined inoculations. D. dipsaci was recovered only from shoot and M. hapla only from root tissue. Combined inoculations did not affect reproduction of either D. dipsaci or M. hapla. D. dipsaci suppressed shoot growth of all species at 15-30 C, and M. hapla suppressed shoot growth of tomato, sugarbeet, and sweet clover at 20, 25, and 30 C. There was a positive correlation (P < 0.05) between shoot and root growth suppression by D. dipsaci on all cultivars except wheat at 20 C and tomato at 30 C. M. hapla suppressed (P < 0.05) root growth of sugarbeet at 20-50 C and wheat at 30 C. Growth suppression was synergistic in combined inoculations of sweet clover shoot growth at 15 C and root growth at 20-30 C, wheat root growth at 15 and 20 C, and tomato root growth at 15-30 C (P < 0.05) D. dipsaci invasions caused mortality of alfalfa and sweet clover at 15-30 C and sugarbeet at 20-30 C. Mortality rates of alfalfa and sweet clover increased synergistically (P < 0.05) from combined inoculations.     

Suppression of Cyst Nematode by Natural Infestation of a Nematophagous Fungus

Penetration of cabbage roots by Heterodera schachtii was suppressed 50-77% in loamy sand naturally infested with the nematophagous fungus Hirsutella rhossiliensis. When Heterodera schachtii was incubated in the suppressive soil without plants for 2 days, 40-63% of the juveniles had Hirsutella rhossiliensis spores adhering to their cuticles. Of those with spores, 82-92% were infected. Infected nematodes were killed and filled with hyphae within 2-3 days. Addition of KCl to soil did not increase infection of Heterodera schachtii by Hirsutella rhossiliensis. The percentage of infection was lower when nematodes were touched to two spores and incubated in KCl solution than when nematodes naturally acquired two spores in soil.     

In Vitro Parasitism of Rotylenchus robustus by Isolates of Hirsutella rhossiliensis

We tested the hypothesis that isolates of Hirsutella rhossiliensis from host nematodes in the family Hoplolaimidae (Rotylenchus robustus and Hoplolaimus galeatus) would be more virulent to R. robustus than would isolates from host nematodes not in the Hoplolaimidae (Heterodera schachtii and Criconemella xenoplax). Nematodes were touched to 10-20 spores of different isolates and incubated at 20 C in 4.5 mM KC1; the percentage of nematodes colonized (filled with hyphae) was determined after 2, 5, 10, 20, and 30 days. The hypothesis was rejected because isolates from H. schachtii and C. xenoplax were equivalent or better at parasitizing R. robustus than were isolates from R. robustus and H. galeatus. In addition, the R. robustus and H. galeatus isolates were as pathogenic to C. curvata as they were to R. robustus, but produced fewer spores per colonized nematode (H. schachtii) than did the other isolates.     

Predisposition of Broadleaf Tobacco to Fusarium Wilt by Early Infection with Globodera tabacum tabacum or Meloidogyne hapla

In greenhouse experiments, broadleaf tobacco plants were inoculated with tobacco cyst (Globodera tabacum tabacum) or root-knot (Meloidogyne hapla) nematodes 3, 2, or 1 week before or at the same time as Fusarium oxysporum. Plants infected with nematodes prior to fungal inoculation had greater Fusarium wilt incidence and severity than those simultaneously inoculated. G. t. tabacum increased wilt incidence and severity more than did M. hapla. Mechanical root wounding within 1 week of F. oxysporum inoculation increased wilt severity. In field experiments, early-season G. t. tabacum control by preplant soil application of oxamyl indirectly limited the incidence and severity of wilt. Wilt incidence was 48%, 23%, and 8% in 1989 and 64%, 60%, and 19% in 1990 for 0.0, 2.2, and 6.7 kg oxamyl/ha, respectively. Early infection of tobacco by G. t. tabacum predisposed broadleaf tobacco to wilt by F. oxysporum.     

Influence of Glomus fasciculatum on Meloidogyne hapla Infecting Allium cepa

The impact of Glomus fasciculatum on Meloidogyne hapla associated with Allium cepa was evaluated in two experiments. Nematode density was not different in mycorrhizal and nonmycorrhizal plants 10 weeks after the joint inoculation of M. hapla and G. fasciculatum. Differences in the age structure of M. hapla populations reared on mycorrhizal and nonmycorrhizat plants were noted. G. fasciculatum enhanced leaf and bulb growth of A. cepa in the absence of M. hapla, but did not affect plant weight when nematodes were present. Survival and reproduction of M. hapla were not affected by G. fasciculatum or phosphorus (P). The estimated time required for inoculated second-stage juveniles (J2) to mature to the adult stage was 1,000 degree hours (base = 9 C) greater in mycorrhizal than in nonmycorrhizal plants supplemented with P. Although the infectivity of J2 was not measured directly, colonization of A. cepa by G. fasciculatum appeared to alter the ability of M. hapla to penetrate roots.     

Histopathology of Selected cultivars of Tobacco infected with Meloidogyne species

Rates of nematode penetration and the histopathology of root infections in fluecured tobacco cultivars ''McNair-944,'' ''Speight G-28,'' and ''NC-89'' with either Meloidogyne arenaria, M. incognita, M. hapla, or M. javanica were investigated. Penetration of root tips by juveniles of all species into the M. incognita-resistant NC-89 and G-28 was much less than that on the susceptible McNair-944. Few juveniles of M. incognita were detected in resistant cultivars 7 and 14 days after inoculation. Infection sites exhibited some cavities and extensive necrotic tissue at 14 days; less necrotic tissue and no intact nematodes were observed 35 days after inoculation. Although some females of M. arenaria reached maturity and produced eggs, considerable necrosis was induced in the resistant cultivars. Meloidogyne hapla and M. javanica developed on all cultivars, but there was necrotic tissue at some infection sites in the resistant cultivars. The occurrence of single multistructured nuclei in the syncytia of most M. hapla infections differed from the numerous small nuclei found in syncytia caused by the other three species.     

Resistance of Trisomic and Diploid Hybrids of Beta vulgaris and B. procumbens to the Sugarbeet Nematode,Heterodera schachtiis Arnold

Trisomic and diploid hybrids of sugarbeet (Beta vulgaris L.) X wild beet (B. procumbens Chr. Sin.) inherited the gene for resistance to Heterodera schachtii Schm. from B. procumbens. The hybrids showed partial resistance to H. schachtii, manifested in failure of larvae to reach maturity. Although significantly greater numbers of female nematodes developed on plants inoculated with populations from the Netherlands or Italy than on plants inoculated with a population from the Salinas Valley, California. the totals for all populations on resistant plants were small. Greater numbers of males than females developed on root-slice cultures of resistant hyhrids when compared to a susceptible cultivar.     

Effect of Plant Age and Transplanting Damage on Sugar Beets infected by Heterodera schachtii

Sugar beet (Beta vulgaris L. cv. Monogerm C.S.F. 1971) seeds sown into Vineland fine sandy loam, infested with 15,500 H. schachtii juveniles/pot, showed little growth during an 11-week test in the greenhouse. Seedlings transplanted at 2, 4, and 6 weeks of age had 32, 30, and 31% less top weight and 71, 68, and 59% less root weight, respectively, compared to controls grown in nematode-free soil. Nematode reproduction in both direct-seeded and transplanted sugar beets was limited and related to root weight. Shoot/root ratios were increased by the nematodes in all nematode-infected beets compared to those grown in soil without nematodes. In contrast to seeding or transplanting sugar beets into nematode-infested Vineland fine sandy loam, an inoculation of Beverly fine sandy loam supporting 0 (seeds), 2-, 4-, and 6-week-old sugar beet seedlings with 7,400 juveniles/pot, followed by 11 weeks of growth in the growth-room, resulted in top weight losses of only 13, 3, 18, and 15% and losses in root weight of 44, 38, 36, and 38%, respectively. Nematode reproduction was high and all shoot/root ratios were increased by the nematode compared to the noninoculated controls. These experiments have shown that sugar beets sown into nematode-infested soil are damaged much more heavily by H. schachtii juveniles than seeds inoculated with the nematode immediately following sowing. Results indicate that an increase in tolerance of sugar beets to attack by H. schachtii does not occur beyond the first 2 weeks of growth and that transplanting damage lowers the tolerance of seedlings to nematode attack.     

Interaction of Vesicular-Arbuscular Mycorrhizae and Cultivars of Alfalfa Susceptible and Resistant to Meloidogyne hapla

The interaction between vesicular-arbuscular mycorrhizal (VAM) fungi and the root-knot nematode (Meloidogyne hapla) was investigated using both nematode-susceptible (Grasslands Wairau) and nematode-resistant (Nevada Synthetic XX) cultivars of alfalfa (Medicago sativa) at four levels of applied phosphate. Mycorrhizal inoculation improved plant growth and reduced nematode numbers and adult development in roots in dually infected cultures of the susceptible cultivar. The tolerance of plants to nematode infection and development when preinfected with mycorrhizal fungi was no greater than when they were inoculated with nematodes and mycorrhizal fungi simultaneously. Growth of plants of the resistant cultivar was unaffected by nematode inoculation but was improved by mycorrhizal inoculation. Numbers of nematode juveniles were lower in the roots of the resistant than of the susceptible cultivar and were further reduced by mycorrhizal inoculation, although no adult nematodes developed in any resistant cultivar treatment. Inoculation of alfalfa with VAM fungi increased the tolerance and resistance of a cultivar susceptible to M. hapla and improved the resistance of a resistant cultivar.     

Response of Pinus ponderosa Seedlings to Stylet-Bearing Nematodes

Of 12 stylet-bearing nematodes used for inoculations, Pratylenchus penetrans, P. brachyurus, P. vulnus, Ditylenchus destructor, Meloidogyne incognita, M. javanica, and M. hapla reproduced on Pinus ponderosa, while Xiphinema index, Aphelenchus avenae, Paratylenehus neoamblycephalus, Tylenchulus semipenetrans, and Macroposthonia xenoplax did not. P. vulnus, P. brachyurus, P. penetrans, A. avenae, D. destructor, T. semipenetrans, and P. neoamblycephalus significantly suppressed both the shoot and root wet weights of ponderosa pine seedlings obtained from stands in five different locations. X. index significantly suppressed root wet weights, M. xenoplax siguificantly suppressed shoot wet weight, and M. incognita, M. javanica, and M. hapla suppressed neither at the inoculation levels used. Injurious nematodes tended to suppress root growth more than shoot growth. Seedlings from two locations produced greater shoot growth wet weight than did seedlings from the other three locations. The more injurious nematodes tended to cause an increase in the water content of shoots. Frequency analyses of seedling population shoot-root ratios indicated that ponderosa pine seedlings could be selected for better shoot-root ratios as well as for resistance to several pathogenic nematodes.