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Effects of Earthworms on the Dispersal of Steinernema Spp. D Plant Pathology and Microbiology Publications Plant Pathology and Microbiology 3-1995 Effects of Earthworms on the Dispersal of Steinernema spp. D. I. Shapiro Iowa State University G. L. Tylka Iowa State University, [email protected] E. C. Berry United States Department of Agriculture L. C. Lewis United States Department of Agriculture Follow this and additional works at: http://lib.dr.iastate.edu/plantpath_pubs Part of the Agricultural Science Commons, Agriculture Commons, Entomology Commons, and the Plant Pathology Commons The ompc lete bibliographic information for this item can be found at http://lib.dr.iastate.edu/ plantpath_pubs/122. For information on how to cite this item, please visit http://lib.dr.iastate.edu/ howtocite.html. This Article is brought to you for free and open access by the Plant Pathology and Microbiology at Iowa State University Digital Repository. It has been accepted for inclusion in Plant Pathology and Microbiology Publications by an authorized administrator of Iowa State University Digital Repository. For more information, please contact [email protected]. Journal of Nematology 27(1):21-28. 1995. © The Society of Nematologists 1995. Effects of Earthworms on the Dispersal of Steinernema spp.1 D. I. SHAPIRO, 2 G. L. TYLKA, 3 E. C. BERRY, 4 AND L. C. LEWIS5 Abstract: Previous studies indicated that dispersal of S. carpocapsae may be enhanced in soil with earthworms. The objective of this research was to determine and compare the effects of earthworms on dispersal of other Steinernema spp. Vertical dispersal of Steinernema carpocapsae, S. feltiae, and S. glaseri was tested in soil columns in the presence and absence of earthworms (Lumbricus terrestris). Dispersal was evaluated by a bioassay and by direct extraction of nematodes from soil. Upward dispersal of S. carpocapsae and S. feltiae increased in the presence of earthworms, whereas upward dispersal of S. glaseri was not affected by earthworms. No significant differences were detected in downward dispersal of S. carpocapsae and S. feltiae in soil with earthworms compared to soil without earthworms. Downward dispersal of S. gla~eri, however, was greater in soil without earthworms relative to soil with earthworms. In soil void of earthworms, dispersal of S. glaseri was greatest followed by dispersal of S. carpocapsae. The presence of earthworm burrows in soil did not influence nematode dispersal. Nematodes were recovered from the surface, interior, and casts of earthworms. Therefore, nematodes may have a phoretic association with earthworms. Key words: earthworm, entomophyllic nematode, nematode, nematode dispersal, Steinernema spp. Entomopathogenic nematodes in the ge- host presence, temperature, soil texture, nus Steinernema have many attributes as bi- and soil moisture (14). ological control agents, including wide Dispersal characteristics vary among host ranges, safety to nontarget organisms, Steinernema spp. Steinernema carpocapsae and vectoring a bacterium, Xenorhabdus (Weiser) moves little from the site of appli- sp., which rapidly kills insect hosts (15). cation, whereas S. glaseri (Steiner) is a rel- Despite their great potential as biological atively active disperser (18). Steinernema control agents, results of nematode appli- carpocapsae applied to the surface of verti- cations have been inconsistent (11). Biotic cal soil columns remained within the up- and abiotic factors that influence the effi- per 15 cm after 30 days, but S. glaseri was cacy of nematode applications must be in- recovered 90 cm below the application vestigated (8). point (18). Because of these differences in Increased dispersal may increase the ef- behavior, S. carpocapsae is classified as an ficacy of nematodes against insect pests (9). ambusher and S. glaseri is classified as a Adverse environmental conditions can de- cruise forager (15). The foraging behavior crease nematode mobility, thereby de- of S. feltiae (Filipjev) is considered interme- creasing their effectiveness (8). Factors diate between S. carpocapsae and S. glaseri that may affect nematode dispersal include (1). Differences in dispersal behavior may be important in selecting nematode species for use against pests occupying different Received for publication 8 August 1994. niches in soil habitats (16). I Journal Paper No. 15954 of the Iowa Agricultural and Home Economics Experiment Station, Ames; Project No. Interactions with soil invertebrates may 3130. increase nematode dispersal (14). For ex- 2 Department of Entomology, Iowa State University, Ames, IA 50011. ample, a phoretic relationship was ob- Department of Plant Pathology, Iowa State University, served between nematophagous mites and Ames, IA 50011. 4 USDA ARS, National Soil Tilth Laboratory, Ames, IA S. carpocapsae (6). Shapiro et al. (19) re- 50011. ported increased dispersal of S. carpocapsae USDA ARS, Corn Insects Research Unit, Ames, IA 50011. in the presence of earthworms. Earth- We thank M. Abbas, R. D. Gunnarson, D. Soh, and S. Sou- worms are not adversely affected by ento- hrada for assistance in the laboratory and Drs, I. Glazer, E. E. Lewis, and J. J. Obrycki for reviewing an earlier draft of this mopathogenic nematodes (4,17). There- manuscript. We also thank Dr. G. O. Poinar for nematode fore, it may be possible to exploit the asso- identification. This research was supported in part by the Leopold Center of Sustainable Agriculture, Ames, IA, Grant ciation between S. carpocapsae and No. 92-18. earthworms in the soil to increase the effi- 21 22 Journal of Nematology, Volume 27, No. 1, March 1995 cacy of the nematode as a biological con- earthworms, Lumbricus terrestris (L.), ob- trol agent. The influence of earthworms tained from the USDA ARS National Soil on the dispersal of other Steinernema spe- Tilth Laboratory (Ames, IA), were added cies and the mechanism(s) of how earth- to the selected columns and allowed to worms affect nematode dispersal have not burrow for 1 week before nematode appli- been examined. Because of differences in cation. Subsequently, 10,000 infective ju- dispersal behavior, the effects of earth- veniles of each species were applied in 0.3- worms on dispersal may vary among 0.4 ml of distilled water to the bottom sur- nematode species. Shapiro et al. (19) sug- face of the columns (section 6) to test gested two hypotheses relating to the upward movement, and to the top surface earthworm-nematode relationship: the of the columns (section 1) to evaluate nematodes are dispersed by direct contact downward movement. Soil columns were with earthworms and (or) nematodes dis- then incubated for 14 days at approxi- perse at different rates in soil with earth- mately 21 C and 54% relative humidity, worm burrows than in soil without bur- after which they were dismantled to eval- rows. The objectives of this research were uate nematode dispersal. to determine and compare the effects of Nematode dispersal was evaluated using earthworms on dispersal of S. carpocapsae, two methods: a bioassay against G. mel- S. glaseri, and S. feltiae and to investigate lonella larvae, and direct extraction and the mechanisms involved in how earth- counting of nematodes. Four replicate soil worms influence nematode dispersal. columns per treatment were used for each method of evaluation. For the bioassay, MATERIALS AND METHODS soil from each column section was re- moved and placed in plastic petri dishes The effects of earthworms on dispersal (150 x 25 ram) with 20 last instar G. mel- of three nematode species were deter- loneUa (19). The petri dishes were incu- mined in vertical soil columns. Steinernema bated at approximately 27 C and 80% rel- carpocapsae, All strain, S. glaseri, NC strain, ative humidity for 72 hours, at which time and S. feltiae, strain 27, were originally ob- nematode-induced larval mortality was re- tained from biosys (Palo Alto, CA) and corded. Mortality was considered to be were reared in last instars of the greater nematode induced if cadavers exhibited wax moth, Galleria meUonella (L.). Experi- flaccidity and coloration characteristic of mental units were polyvinyl chloride Steinernema infections. In the second (PVC) pipe consisting of six 4-cm-long sec- method, nematodes were extracted from tions and 5-cm-d pipe joined with duct the soil in each column section using su- tape and filled with soil (34% sand, 28% crose centrifugation (13), and Steinernema silt, and 38% clay, pH 7.0). Soil moisture in spp. were quantified by observation with columns was adjusted to approximately an inverted compound microscope at a field capacity. The column sections were magnification of x 40. numbered vertically; the top section was Analysis of nematode movement was ac- one and the bottom section was six. complished by determining the number of The experiments were organized as ran- nematodes reaching the half of the column domized block designs with depth as a opposite where the nematodes were placed repeated measure. Experiments testing (sections 1-3 for upward movement and upward and downward movement of 4-6 for downward movement). Bioassay nematodes were run simultaneously. Six data were analyzed through ANOVA of treatments (three nematode species, with mean numbers of dead G. meUonella larvae and without earthworms) were each ap- obtained from soil in the three sections op- plied to eight soil columns for upward posite to where the nematodes were movement and eight columns for down- placed. Because differences in virulence ward movement (96 columns total). Two against G. melloneUa among the three Dispersal of Steinernema spp.: Shapiro et al. 23 nematode species were not determined, in- with S. carpocapsae) were dissected and ex- terspecific comparisons of dispersal could posed to G. mellonella as previously de- not be made using the bioassay data. For scribed. the direct extraction method, ANOVA was An additional experiment was con- performed on mean proportions of nema- ducted to determine whether nematode todes recovered. Proportions were calcu- dispersal is greater in soil with earthworm lated based on the number of nematodes burrows than in soil without burrows. Up- recovered from the half of the column op- ward dispersal of S.
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