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Hatch and Reproduction of Globodera Tabacum Tabacum in Response to Tobacco, Tomato, Or Black Nightshade J

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Hatch and Reproduction of Globodera Tabacum Tabacum in Response to Tobacco, Tomato, Or Black Nightshade J Journal of Nematology 27(3):382-386. 1995. © The Society of Nematologists 1995. Hatch and Reproduction of Globodera tabacum tabacum in Response to Tobacco, Tomato, or Black Nightshade J. A. LAMONDIA 1 Abstract: The effects of broadleaf tobacco, tomato, and black nightshade on juvenile hatch and reproduction of Globodera tabacum tabacum were determined in laboratory and greenhouse experi- ments. Root exudates from nightshade stimulated greater egg hatch than those from either 'Rutgers' tomato or '86-4' tobacco. Hatch was greater at higher proportions of root exudates for all three plant species. Root exudates from plants greater than 3 weeks old stimulated more hatch than younger plants. No regression relationships existed between plant age and nematode batch. In other exper- iments, hatch from eggs in cysts was higher for tomato and nightshade after 10 weeks in greenhouse pots compared to tobacco and bare soil. Numbers of second-stage juveniles in eggs in cysts produced from a previous generation on the same host were highest on nightshade and less on tomato and tobacco. Cysts of variable age recovered from field soil had increased hatch in both root exudates or water compared to recently produced cysts from plants in growth chambers. Globodera t. tabacum may be subject to both host and environmentally mediated diapause. Key words: hatch stimulation, Nicotiana tabacum, Lycopersicon esculentum, nematode, root exudates, Solanum nigrum, tobacco cyst nematode. The tobacco cyst nematode, Globodera experiments. G. t. tabacum, however, can tabacum tabacum (Lownsbery and Lowns- persist in soil for many years in the absence bery) Behrens, is an important parasite of of a host (1), suggesting some form of dor- shade (9,13) and broadleaf (11) tobacco in mancy with survival value. the Connecticut River Valley of Connecti- The objectives of this research were to cut and Massachusetts. While most impor- determine the i) effects of tobacco, tomato, tant as a pathogen of tobacco, the host and black nightshade on G. t. tabacum ju- range of G. t. tabacum also includes tomato, venile hatch stimulation; ii) effect of exu- eggplant, and solanaceous weeds such as date dilution and plant age on hatch; iii) black nightshade (7,12). and relation of hatch stimulation to host Hatch of many cyst nematodes may be efficiency for these plant species. stimulated to various degrees by root exu- dates of host plants (2,3,14,16,19) or, more MATERIALS AND METHODS uncommonly, by nonhost plants (19). Al- Root exudates were collected from '86-4' though Globodera tabacum solanacearum broadleaf tobacco (Nicotiana tabacum L.), hatch was increased after exposure to to- 'Rutgers' tomato (Lycopersicon esculentum bacco roots in combination with soil micro- Mill.), and Solanum nigrum L. collected in organisms (6), the extent of G. t. tabacum Windsor, CT. Specimens of S. nigrum have hatch stimulation by root exudates of to- been deposited at the Herbarium, Ento- bacco or other weed or crop plants is cur- mology Dept., CAES, P.O. Box 1106, New rently unknown. Approximately 80% of Haven, CT, 06504-1106, C. Lemmon, Di- encysted second-stage juveniles in eggs rector. Seeds were planted in 1:1 sand:pot- hatched in response to resistant or suscep- ting mix (Sunshine mix no. 3, Fisons West- tible tobacco in greenhouse pot experi- ern Corp., Downers Grove, IL) in 96-cavity ments (8), but the influence of root exu- seedling trays. Three weeks after emer- dates on hatch was not examined in these gence and weekly thereafter, roots were gently washed free of soil, and 2 g fresh- weight roots from three plants were Received for publication 25 August 1994. soaked for 2 hours in 100 ml distilled wa- a Associate Plant Pathologist, Department of Plant Pathol- ter. Root exudate solutions were filtered ogy and Ecology, Connecticut Agricultural Experiment Sta- tion, Valley Laboratory, P.O. Box 248, Windsor, CT 06095. through medium-porosity, slow flow-rate The author thanks F. J. Ferrandino for assistance with sta- filter paper and frozen until experimental tistical analyses, J. F. Ahrens for identification of Solarium nigmm, and Jane Canepa-Morrison for technical assistance. use at room temperature. 382 Tobacco Cyst Nematode Hatch: LaMondia 383 Hatching tests were performed in 96- ing soil and cysts in bags that were not well trays using standardized cysts of planted served as bare soil or fallow con- known size and age (10). Ten uniformly trols. Pots were placed in the greenhouse sized (0.35-0.45 mm) Globodera t. tabacum on propagation mats at 22 C at 15-hour cysts produced in pots in the greenhouse, daylength. Cysts were recovered from the with a mean viable egg content of 288, bags in pots after 10 weeks, crushed in wa- were placed in 10 replicate cells of 0.3 ml ter, and the number of unhatched J2 in of the appropriate root exudate. Hatching eggs was counted. All of the soil in the pots trays were incubated in the dark at 20 C. was dried and washed using a modified Nematode hatch was counted weekly for 4 Fenwick can to recover new cysts produced weeks, and the exudate solution was re- on the roots (18). Cysts were counted and placed with fresh exudate each week. Dis- crushed to determine the number of j2 in tilled water served as controls. Root exu- eggs per cyst. The experiments were per- dates from each host source were added to formed twice. separate hatching trays. Hatch from cysts recovered from field The effect of plant host on G. t. tabacum soil planted to tobacco for 6 consecutive hatch was examined using root exudate so- years was compared to hatch of cysts re- lutions collected 3 weeks after emergence. covered from tobacco plants grown in Root exudate solution was used at full growth chambers at 24 C. Cysts were re- strength or serially diluted with distilled covered from soil 2 weeks after tobacco water to result in 1:1, 1:10, 1:100, or harvest and after 8 weeks of tobacco 1:1000 parts exudate to distilled water. growth. Five cysts of each type were ex- The effect of plant age on juvenile hatch posed to tobacco root exudate or distilled from eggs was examined using exudates water for 4 weeks in hatch chambers. Field collected 3, 4, 5, 6, 7, or 8 weeks after seed- cysts were of unknown age and contained ling emergence. Root exudate solution was fewer and more variable numbers of juve- used at full strength. niles per cyst than growth chamber pro- Root exudate dilution and plant age ef- duced cysts. Cysts were crushed in water fects on juvenile hatch from eggs in cysts after 4 weeks, and hatch was expressed as were analyzed by stepwise polynomial re- percentage of total juveniles in cysts. gression up to a fourth-order polynomial There were five replicates of each treat- with alpha 0.05 to enter and 0.10 to re- ment, and the data were analyzed after move (15). square-ro0t transformation to standardize The influence of 86-4 tobacco, Rutgers the variance. tomato, or S. nigrum on G. t. tabacum hatch and reproduction in soil in greenhouse RESULTS pots was also examined. Nylon mesh bags (8) (2 cm 2, mesh opening 210 Ixm) were Globodera t. tabacum juvenile hatch was placed in the center of 450-cm 3 pots con- stimulated by root exudates of black night- taining a pasteurized Merrimac fine loamy shade, tomato, and tobacco (Figs. 1,2). sand field soil free of cyst nematodes There were no significant differences be- (72.7% sand, 21.7% silt, 5.6% clay, pH = tween constants (intercepts) in the nonlin- 6.8). Each nylon bag contained 25 G. t. ear regression equations generated by tabacum cysts of uniform size and age (av- stepwise polynomial regression. Paired erage 285 viable J2 in eggs per cyst). One t-tests between means indicated that root bag containing cysts was placed in the cen- exudates from nightshade stimulated ter of each pot. Five-week-old transplants greater hatch than tomato or tobacco, of 86-4 tobacco, Rutgers tomato, or S. ni- which were not different. Hatch was grum were each transplanted to eight rep- greater at higher proportions of root exu- licate pots so that the root ball was 1 cm dates for all three plant species. While ju- above the nylon mesh bags. Pots contain- venile hatch was much lower in the second 384 Journal of Nematology, Volume 27, No. 3, September 1995 1200 TABLE i. Hatch stimulation of Globodera tabacum tabacum by root exudates of '86-4' tobacco, 'Rutgers' .~1000 tomato, and black nightshade of different plant ages after 4 weeks. 8oo n~ightshade, 600 Experiment l itomato Hatched J2 per 10 cysts'~ 400 Ltobacco Plant age weeks Nightshade Tomato Tobacco 200 "1- 3 839.7 289.2 108.1 i ~ t I i 4 1,515.8 330.1 340.9 0.2 0.4 0.6 0.8 5 1,628.4 406.0 444.2 Proportion of Root Exudate in Solution 6 1,394.4 504,7 571.7 7 1,555.6 971.1 454.2 F~G. 1. Relationship between the proportion of 8 1,666.6 157.6 245.0 nightshade, tobacco, or tomato root exudate in solu- LSD 259,0 151.9 174.4 tion and Globodera tabacum tabacum hatch, experiment 1. Nightshade: Y = 49.0 + 5513.1X - 9370.4X 2 + Source of variation df Mean square P 4822.3XZ; R 2 = 0.998, P = 0.001. Tomato: Y = 33.6 + 137.6X + 181.3X2; R 2 = 0.998, P = 0.001. To- Host 2 21361836 0.001 bacco: Y = 34.8 + 440.7X - 282.5X2; R e = 0.992, Age 5 1131605 0.001 P = 0.001.
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