Influence of aldicarb, carbofuran, propoxur and fenamiphos on activity of Pratylenchus penelrans (Cobb) and P. crenatus Eoof (’)

Joe KIMPINSKIand Salah EL ERAKI Agriculture Canada, Research Station, P. O. Box 1210, Charlottetown, Prince Edward Island, Canada ClA 7M8 and Nematology Department, Institute of Plant Pathology, Agriculfural Research Centre, Giza, Egypt.

SUMMARY Aldicarb was the most effective nematicide in reducing the activity of Prafylenchus penetrans and P. crenatus, followed by fenamiphos, carbofuran and propoxur. Nematode populations which moved down through vertical tubes in 31 hours or less did not show any significant increase or decrease in passage time when nematicide concen- trations were increased to 24 pg/ml. A significant interaction existed between temperature, aldicarb concentra- tion, and nematode species. This was interpreted to mean thata dose response to aldicarb was observed for P. cre- natus at 100 and 200, but not at 300, and for P. penetrans at 200 and 300, but not at 100. Activity of fourth-stage juvenile and adult P. crenafus at 220 was inhibited more than the activity of second- or third- stage juveniles when aldicarb concentrations were 6 or 12 pg/ml. Propoxur did not have this effect. There was no significant difference in activity between the various stagesof P. penetrans when treated with aldicarb or propoxur. Aldicarb treatments of 6 and 12 pglg dry soi1 yeight reduced numbers of P. penetrans and P. crenatus in red clover and timothy in the greenhouse moreso than equivalent treatmentsof fenamiphos or propoxur. Propoxur only reduced numbers of P. crenafus to a significant degree in red clover. Yield increases were associated with aldicarb treat- ments on red clover and timothy harbouring P. penetrans and P. crenafus respectively. Aldicarb at 12 pg/g on red clover harbouring P. crenatus also was associated with yield increases. Aldicarb and fenamiphos caused changes in body shape and activity of female nematodes. Pratylenchus penetrans recovered more quicltly from the effects of fenamiphos than from aldicarb after exposure to solutions of 12 or 24 pg/ml for 24 hours. Pratylenchus crenafus recovered more quickly than P. penetrans after similar exposure to aldicarb.

RÉSUMÉ Influence de l’aldicarbe, du carbofuran, du propoxur et du fenamiphos sur l’activiti de Pratylenchus penetrans (Cobb) et P. crenatus Loof L’aldicarbe a été le nématicide le plus actif pour réduire l’activité de Pratylenchuspenetrans et P. crenatus, suivi par le fenamiphos, le carbofuran et le propoxur. Le passage de populations de nématodes se déplaçant vers le bas dans des tubes verticaux, en 31 heures, n’était pas significativement ralenti ou accéléré quand les concen- trations de nématicide étaient augmentées jusqu’à 24 pg/ml. Une interaction significative a été observée entre latempérature, la concentrationd’aldicarbe et l’espèce dunématode. Onconsidére qu’uneréponse à la dose d’aldicarbe est observée pour P. crenafus à 100 et 200 mais pas à 300 et pour P. penetrans à 200 et 300 mais pas zi 100. Aux doses d’aldicarbe de6 à 12 pg/ml, l’activité des juvéniles de4e stade et des adultes deP. crenatus, à 220, était plus inhibée que celle des juvéniles de 2e et 3e stade. Cet effet n’était pas observé avec le propoxur. Aucune différence significative n’a étéobservée entre les activités des différents stades P.de penetrans traités avec l’aldicarbe ou le propoxur. En serre, des traitements à raison de 6 à 12 pg par gramme de sol sec ont réduit le nombre de P. penetrans et P. crenatus dans le LrBfle et lafléole des prés plus que des traitements équivalents avecle fenamiphos ou le propoxur. Ce dernier produit n’a réduit significativementle nombre de P. crenatus que dans letrèfle. L’aldicarbe a provoqué des augmentations de rendementchez le trkfle et lafléole des prés hébergeant respectivementP. penefrans et P. crenafus.

(l) Contribution no 492 of the Charlottetown Research Station.

Revue Nématol. 6 (1) : 103-110 (1983) 103 J. Kimpinski & S. El Eraki

A la dose de 12 pg/g de sol sec, I’aldicarbe a causé des augmentations de rendement chez le trèfle attaqué par P. crenatus. L’aldicarbe et le fenamiphos ont causé des modifications dans la forme du corpset l’activité des néma- todes. Chez P. penetrans, les effets du fenamiphos se dissipent plus vite que ceux de l’aldicarbe après exposition à des solutions de 12 ou 24 pg/ml pendant 24 heures. Après une exposition similaire avec l’aldicarbe, les effets se sont dissipés plus vite chez P. crenatus que chez P. penetrans.

Althoughmany of thecarbamate and organo- rinsedthree times in sterile tap water to reduce phosphatepesticides are systemics, their major microbial contaminants. effect is often as potent contact nematicides against Thenematicides included in this study were plant-parasiticnematodes inthe soil (Bromilow aldicarb10% granular, carbofuran 1.1% liyuid, & Lord, 1979 ; Hague, 1979 ; Hague & Pain, 1973 ; fenamiphos 1.66% liquid, and propoxur 2% wettable Jorgenson,1978). These compounds usually do not powder.Fresh stock solutions of 100 pg/mlwere kill plantparasitic nematodes at concentrations prepared for eachexperiment. The nematicides used in the field. Instead, they depress populations were agitated in steriletap water for several hours in a byinterfering witcarbamate and organophosphate pesticides is through to a depth of 2.5 cm with dry sterileSand of particle theirability to inhibit esterases in the nervous size 150-250pm. Thetubes wereplaced vertically system of vertebratesand insects (Corbett, 1974 ; inwatch glasses so that the meshinterfaced witth O’Brien et al., 1974).Acet>ylcholinesterase has been nematicidesolutions containing O, 3,6, 12, or detectedinplant-parasitic nematodes (Dickson, 24 pg/ml. It was assumed that most of the nematicide Huisingh & Sasser, 1971; Rohde, 1960), and the t,oxic stayedin solution and did not adsorb to the rela- modeis believed to besimilar to that in insects tively large particles of sterile washed Sand (Hough, (Bunt, 1975 ; Homeyer & Wagner,1981). However Thomason & Farmer,1975). Aliquots of 0.1ml there is no direct evidence of an operational cholin- suspensions of 36.5 f 1.9 SZ P. penetrans or esterasesystem nematodes in (VanGundy 30.6 f 1.2 SX P. crenatus werepipetted ont0 the & McKenry, 1977). surface of the Sand in each tube. Tubes were covered The objective of this study was to determine the with plastic to reduce evaporation and held at 220. effect of threecarbamates : aldicarb,carbofuran, Thenumber of nematodes that moveddown the and propoxur, and the organophosphate, fenamiphm, tube into the solution after 31 hours was expressed on the activity and population size of two common asa percentage of the total. In addition, the tinle species of rootlesionnematode in eastern Canada, which allowed 50% of the mobilepopulation to Pratylenchus penetrans (Cobb) and P. crenatus Loof. passthrough the tubes was calculated from the grouped data by the algorithm of Sampford (1952) and with the assumptionof a log normal distribution. Asecond experiment recorded theactivity of Materials and methods nematodesin the Sand columns at 100, 200, and 300, at aldicarb concentrations of O, 3, and 6 pg/ml. Populations of Pratylenchuspenetrans and P. The exposure time was twenty hours and the inocu- crenatus weremaintained on Ottawared clover lation levels were58.3 f 1.2 SX P. penetrans and (Trifolium pratense L.) and Climax timothy (Phleum 42.6 f 1.3 SZ P. crenatus. pratense L.), respectively. Nematodes were extracted In the third experiment data were obtained on the by pIacing up to- 10 g of rootin a mist chamber number of nematodes that moveddown through (Seinhorst, 1950) or 50 g of soil in a Baermann pan vertical Sand columns in 24 hours at concentrations (Townshend,1963). Samples then were dried at of O, 3, 6, and 12 pg/ml of aldicarb and propoxur. 1000 for24 hours and counts wereexpressed as Theactivity of second- andt>hird-stage juveniles numbers of nematodes per g of dry root or per kg was compared to the activity of fourth- stage juve- of dry soil. Nematodes usedfor inoculumwere niles and adults. The distinction between categories

104 Revue Nématol. 6 (1) : 103-110 (1983) Influence of nematicides on Pratylenchus penetrans and P. crenatus was based primarily on nematode size and the pro- The statistical analyses showed that, in addition portion of each was similar for P. penetrans and P. to the main effects of temperatureand nematicide crenatus. Theinoculation levels wereequivalent to concentration, a significant second-order interaction the previousexperiment and the temperature was existed between temperature, aldicarb concentration 220. andnematode species. Thisindicated that adose Agreenhouse experiment examined the effect response toaldicarb was observed for P. crenatus of aldicarb, fenamiphos, and propoxur on population at 100 and 200 but not at 300, and for P. penetrans size of P. penetrans and P. crenatus. Climax timothy at 200 and 300 but not at 100 (Fig. 1). or Ottawa red clover were seeded in pots containing Aldicarb at concentrations of 6 and 12 g/ml 1.0 kg of sterilizedCharlottetown fine sandyloam reduced the activityof P. penetrans, while the move- (70% sand,20% silt, 10% Clay). Afterfour days ment of P. crenatzrs was lessened at 12pg/ml (Tab. 2). 10 O00 nematodes in 50 ml of water were distributed Fourth-stagejuveniles and adults of P. crenafus over the surface of each pot and covered with about wereaffected more than second- or third-stage 1 cm of soil. Seven days later nematicide solutions juveniles.Numerically the trend was opposite for wereadded atrates of O, 6, or 12 pg/g dry soil. P. penetrans but was notstatistically significant. Plants wereheld in the greenhousein a light-dark Propoxur did not significantly affect the activity of regime of 14 and 10 hours, respectively, at a mean P. penetrans at any of the concentrations. However, temperature of220. Nematodecounts and foliage the mobility of P. crenatus was reduced as propoxur weights were recorded eleven weeks after seeding. concentrations were increased, butthere was no The finaltest consisted of placing five active significant difference between the stages. P. penetrans or P. crenatus females in 1 ml solutions Aldicarb reduced the numbers of both nematode of aldicarb or fenamiphos at O, 1,3,6,12,or 24 pg/ml. species in the greenhouse pots except for the 6 pg/g After 24 hours the nematodes were examined with level in timothy pots inoculated with P. peizetrarzs astereomicroscope at 70 x for changesin activity (Tab. 3). The same trend in red clover occurred for or body shape. They were then transferred to ster- fenamiphos,though the reductions were notas ilized tap water for another 24 hours and re-exam- marked as in the aldicarb treatments. Propoxur at ined.The procedure used toestimate the amount 12 pg/g reduced the population size of P. crenatus of coiling or body shrinkage was a modified version in redclover but to a lesser degree thanaldicarb. of themethods employed by Keetch (1974) and The population size of P. penefrans was not reduced Bunt(1975). The relative scores were assigned on significantly by propoxur in red clover and timothy. a three-pointa scale (Snedecor & Cochran1980). The categoriesused here, with assigned scores in brackets, were normalsinusoidal motion (l),mod- eratecoiling and decreased movement (2), and severe coiling with immobility andbody shrinkage(3). The temperature was 200.

Results

Aldicarbcaused the greatest reduction in the numbers of nematodes passing through the columns within 31 .hours followed byfenamiphos, carbo- furan, and propoxur (Tab. 1). This trend was similar at 4, 8, and 21 hours and these data were not shown for the sake of brevity. Most of the population was still in the sand columns after31 hours when aldicarb concentrations were 24 pg/ml. For the active popu- lationwhich included only those nematodes which ALDICARB pglml moved down the columns in 31 hours or less, there was no definite trend in the amount of passage time Fig. 1. Arcsin dpercent of populations of Praly- asnematicide concentrations increased. In some Ienchuspenefrans and P. crenatus moving within cases the passage time increased as the nematicide 20 hours at 100,200, or 300 down vertical sand columns concentrationsincreased, while in other instances treatedwith aldicarb. Vertical bar represents the therewas no change or the oppositeoccurred. standard error at P = 0.05 and n = 4.

Revue Nématol. 6 (1) : 103-110 (1983) 105 J. Iiimpinski 6: S. El Eraki

Aldicarbtreatments were associated with both an Both aldicarband fenamiphos caused significant increase in yield and a decrease innumbers of P. changesinnematode activity and body shape penetrans and P. crenatus in red clover and timothy, (Table 4).Pralylenchus penetrans appeared to recover respectively.The aldicarb treatment of 12 pg/g fromthe effects of fenamiphosearlier thanfrom also resultedin an increasein yield of redclover aldicarbafter being exposed to concentrations of when P. crenatus numberswere reduced. The only 12and 24 pg/mlfor 24 hours. Also, P. crenatus other significant yield increases were for fenamiphos recoveredmore fully t>han P. penetrans from the at 12 pg/g inred clover and timothy potJs that effects of aldicarb at concentrations of 12and contained P. penetrans and P. crenalus, respectively. 24 p g/ml.

Table 1 Percentage of the total population at 31 hours which moved down through 2.5 cm vertical soi1 columns treated with nematicides, and the number of hours (1) required for 50 % of the active population tomove through the same columns

Nematicide concentrationsNematicide(pglml)Nematode species ( 2, O 3 6 12 24

Total Population (3) (percent)

Aldicarb PP 95.1 f 2.3 (4) 80.7 f 3.7 41.1 f 6.0 30.6 & 2.6 13.0 f 2.1 PC 89.1 f 4.2 88.1 f 7.2 58.8 & 8.2 50.3 f 3.6 14.3 f 2.8 Fenamiphos PP 94.2 f 2.4 80.2 f 4.0 61.6 f 5.0 68.0 f 7.4 17.5 f 5.1 PC 88.0 f 5.7 89.8 f 4.9 85.8 f 5.8 50.3 f 3.0 36.1 f 8.0 Carbofuran PP 92.0 f 1.8 83.6 f 4.5 66.3 & 10.4 59.5 f 7.4 42.7 f 7.7 PC 89.6 f 1.7 84.6 f 3.0 71.5 & 3.1 70.2 f 8.0 39.8 f 9.3 Propoxur PP 93.1 f 2.8 88.1 f 3.3 71.3 f 4.1 80.6 f 3.2 61.8 f 4.0 PC 89.8 f 4.4 83.8 & 4.5 76.5 f 4.1 68.8 f 3.5 68.0 f 9.4

Active Population (3) (no. of hours) Aldicarb PP 4.8 f 0.4 (*) 4.7 f 0.2 5.1 f 0.3 6.3 f 0.4 6.1 f 0.3 PC 6.9 f 0.4 7.0 f 0.6 5.9 f 0.3 5.7 f 0.3 -(? Fenamiphos PP 4.9 f 0.4 4.6 f 0.2 5.0 f 0.4 4.9 f 0.3 4.9 f 0.4 PC 6.3 f 0.3 6.1 f 0.2 5.7 f 0.2 5.2 f 0.5 4.4 f 0.2 Carbofuran PP 4.4 f 0.3 4.1 f 0.2 4.1 f 0.7 4.4 f 0.2 4.5 f 0.3 PC 6.2 & 0.3 6.1 f 0.5 6.0 f 0.4 7.6 f 1.0 7.7 f 1.1 Propoxur PP 4.4 f 0.3 4.7 f 0.2 4.5 f 0.2 3.7 f 0.4 3.0.f 0.8 PC 6.7 f 0.3 7.8 f 0.5 7.2 f 0.7 7.5 f 0.7 8.3 f 0.6

(l) Observation times were 4, 8, 21, and 31 hours after nematodes were added to columns. (2) PP = Pratylenchus penetrans, PC = P. crenatus. (3) Total population includes those nematodes left in the columns after 31 hours ; active population represents only those nematodes which passed through the columns in 31 hours or less. (4) Mean of four replicates ; standard errors may be compared within each population. (5) Not enough specimens.

106 Revue Nématol. 6 ,(l): 103-110 (1983) Influence of nematicides on Pratylenchus penetrans and P. crenatus

Table 2

Percentages (l) of populations of second- and third-stage juveniles (2 -/- 3) and fourth-stage juveniles and adults (4 + A) moving down 2.5 cm vertical sand columns treated with aldicarbor propoxur

Nem aticide Nematode Category Nematicide concentration (pglml)concentration Nematicide Category NematodeNematicide species O 3 6 12

Aldicarb P. penetrans 2+3 68.7 (2) 66.232.1 41.3 4tA 75.4 69.2 51.3 41.2 P. crenatus 2+3 46.0 68.3 61.9 66.4 4+A 70.344.9 51.8 31.2 Propoxur P. penetrans 2+3 77.6 72.174.7 72.5 4+A 75.4 78.7 67.0 73.8 P. crenatus 2+3 72.3 74.2 38.4 54.9 4+A 62.4 76.3 35.6 49.0

(l) Arcsin transformation ; counts were done24 hours after nematodes were addedto the topsof the sandcolumns; tcmpcrature was 220. (2) Mean of four replicates ; standard error of a mean derived from the error mean square = 5.2.

Table3 Effect of nematicide treatments on populations of Pratylenchus penetrans (PP) and P. crenatus (PC), and on foliage yield’s (1)

plant Nematode Nematicide treatmentsNematicide Nematode Host plant ( 2, species O A6 42 F6 FI, p,. Pl,

No. of nematodeslpot (x 100) Red clover Red PP 72a (3) 8c 7c 23bc 15bc 32ab 29ab PC 83a 1lbc 7c 16bc 17bc 19bc 23 b Timothy PP 28a 13ab 10b 17ab 12ab 27a 17ab PC 41a 14bc 8bc 17ab 20ab 20ab 16ab Foliage yield in gramslpot

Red clover Red PP 1.63 (4) 2.47 2.47 2.20 2.30 1.93 2.17 PG 1.63 2.17 2.37 1.50 1.87 1.63 2.13 Timothy PP 5.07 5.03 5.52 4.97 5.55 5.01 4.88 PG 4.1 1 4.93 5.08 4.51 4.83 4.63 4.38

(l) 10 O00 nematodes added to each pot 4 days after seeding ; nematicides applied seven days later ; nematode and yield data obtained eleven weeks after seeding. (2) Aldicarb (A), fenamiphos (F),and propoxur (P) applied at 6 and 12 pg/g in relation to the soi1 weight; O is the check. (3) Geometric mean of three replicates ; numbers in horizontal rom followed by the same letter are not signifi- cantly different according to Duncan’s multiple range test atP = 0.05. (4) Arithmetic mean of the dryweight of three replicates ; standard error of a mean equals& 0.21.

Revue Nèmabol. Revue 6 (1) : 103-110 (1983) 107 J. Kimpinski & S. EL Eraki

Table 4

Effect. of aldicarb and fenamiphos on activity and body shape (l) of female nematodes after 24 hours of exPosure, and degree of recovery after 24 hours in water

Nematode Treatment Nematicide. concentrationNematicide. Treatment Nematode (pglml) species O 1 3 6 12 24

Nematicide

P. penetrans Aldicarb 1.00 (2) 2.40 2.532.93 2.73 2.87 Fenamiphos 1.O0 1.53 2.20 2.933.00 3.00 P. crenatus Aldicarb 1.27 1.87 2.53 2.733.00 2.80 Fenamiphos 1.13 1.403.00 2.132.87 2.60 Water P. penetrans Aldicarb 1.O0 1.27 1.20 1.20 2.47 2.53 Fenamiphos 1.O0 1.53 1.27 1.53 2.13 1.93 P. crenatus2.07 1.87Aldicarb 1.60 1.131.67 1.13 Fenamiphos 1.07 1.40 1.53 2.07 1.80 1.80

Difference (3) P. penetrans Aldicarb 0.00 1.20 1.26 1.530.34 0.46 Fenamiphos 0.000.67 0.26 1.401.07 0.87 P. crenatus Aldicarb 1.13 O. 140.86 0.74 0.93 0.93 Fenamiphos 0.061.07 0.000.53 0.60 1.20

(I) Assigned relative scores (Snedecor & Cochran, 1980) ; normal sinusoidal motion = 1.00, modest body coiling with some decrease in motion = 2.00, severe body coiling, immobility, and shrinkage 5 3.00. Five females in each replicate. (2) Mean of three replicates ; standard error of a mean for Nematicide, Water, and Difference categories are & 0.08, & 0.12, and f 0.14, respectively. (3) Indicates the relative change or reduction in body coiling and shrinkage when nematodes were removed from the nematicide solution, and washed in water for24 hours.

Discussion Previous work showed aldicarb to be more effective than fenamiphos in the control of Globodera rosto- Aldicarb caused the greatest reduction in nematode chiensis (Woll.) under field conditions(Whitehead activity,fenamiphos and carbofuran were some- et al., 1973 ; MOSS, Crump & Whitehead,1975). whatintermediate, while propoxurhad the least Brodie and Good (1973) observed that less aldicarb effect (Tab. 1, 3, 4). This trend agreed closely with thancarbofuran or fenamiphoswas required for the acute oralLD5Os for rsts reported by Martin and control of Pleloidogyne incognita (Kofoid & White), Worthing (1977) at 0.9, 8.0-14.0, 15.3-19.4, and thelatter two pesticides having about the same 90.0-128.0 mg of pesticide per kg of rat weight for effect. However,Marban-Mendoza and Viglierchio aldicarb,carbofuran, fenamiphos, and propoxur, (1980)needed carbofuran solutions which were respectively.The similarity is interestingsince 100times thestrength of fenamiphossolutions nematodes do notpossess a respiratory or circulatory to immobilize Pratylenchusvulnus Allen & Jensen. system and would not be subject to the same dysfunc-Propoxur was shown previously to be somewhat tion as in rats. innocuous as a nematicide (Bunt, 1975).

108 Revue Nématol. 6 (1) : 103-110 (1983) Influence of nematicides on Pratylenchus penetrans and P. crenatus

There was little clear indication that some nema- onlya few nematodespecies, often from different todeswere lessaffected bynematicides since the families. timerequired for 50% of theactive population to movethrough the columns in 31 hours or less increased in some instances and decreased in others ACKNOWLEDGEMENTS as the nematicide concentrations increased (Tab. 1). The increase in nematode activity as concentrations We thank D. C. Read,Senior Research Scientist, of nematicides increased was due probably to rapid CharlottetownResearch Station, for reviewing the bodycontractions. For example propoxur has a manuscript,K. B. McRae,Regional Statistician, rapidknock-down effect onflying insects caused Kentville,Nova Scotia, for statistical suggestions, by inhibition of acetylcholinesterase (Martin & Wor- and C. E.Gallant, Charlottetown Research Station, thing, 1977), and a similar action causing rapid body for technical assistance. contractionsin P. penetrans couldbe responsible for the increase in activity for a short time. The influence of temperature on aldicarb (Fig. 1) was notunexpected since this phenomenon is commonfor many pesticides (Bunt, 1975). The REFERENCES differing dose response to aldicarb is related to the temperature of the twospecies, in whichP. penefrans BATTERBY,S. (1979).Toxic effects of aldicarband generallylives at highertemperatures than P. its metabolites on second stage larvae of Heterodera crenatus (Kimpinski & Willis, 1981). Aldicarb there- schachtii. Nematologica, 25 : 377-384. fore wasnot as effective on P. penetrans at 100, BATTERBY,S., LE PATOUREL,G.N.J. & WRIGIIT, D.J. or on P. crenatus at 300. (1977).Accumulation and metabolism of aldicarb Adultsand fourth-stage juveniles of P. crenatus bythe free-livingnematodes Aphelenchusauenae wereless activethan the younger stages when and Panagrellusredivivus. Ann. appl.Biol., 86 : aldicarbwas present. The situation was reversed 69-76. for P. penetrarzs thoughthe differenceswere only BRODIE,B.B. & GOOD,J.M. (1973). Relative efflcacy significant at P = 0.1(Tab. 2). Previousreports of selectedvolatile and nonvolatilenematicides indicated that immature stageswere more susceptible forcontrol of Meloidogyneincognita ontobacco. thanadults to nematicides (Evans & Thomason, J. Nematol., 5 : 14-18. 1971 ; Marban-Mendoza & Viglierchio, 1980). Fourth- BROMILOW,R.H. & LORD,K.A. (1979).Distribution stagejuveniles and adults of P. penetrarzs invaded of nematicides in soi1 and its influence on control alfalfamore readily than second- andthird-stage of cyst-nematodes ( Globodera and Heterodera spp.). juveniles(Sontirat & Chapman,1970 ; Townshend, Ann. appl.Biol., 92 : 93-104. 1978),which suggests that olderstages are more BUNT, J.A. (1975).Effect and mode of action of some active.Active nematode species metabolizeand systemicnematicides. Meded.Landbouwhogesch. eliminate nematicides more rapidly than less active Wageningen, 75-10 : 1-128. species (Batterby, Le Patourel & Wright,1977 ; CORBETT, J.R. (1974). The biochemical mode of action Marks,Thomason & Castro,1968), andthe same of pesticides. London & New York, Academic process mayapply to differentstages within a Press, 330 p. species. Unfortunately,information is lackingon DICKSON,D.W., HUISINGH,D. & SASSER, J.N. (1971). the activity of the different stages of P. crenatus. Dehydrogenases,acid and alkalinephosphatases, andesterases forchemotaxonomy of selected The reason for the better recovery of P. crenatus Meloidogyne,Ditylenchus, Heterodera and Aphe- ascompared to P. penetrans fromthe effects of lenchus spp. J. Nematol., 3 : 1-16. aldicarb(Tab. maybe related to cholinesterase 4) EVANS,A.A.F. & THOMASON,I.J. (1971). Ethylene concentrationsin the nematodes. However, more dibromide toxicity to adults, larvae and moulting information is needed on the biochemistry of plant- stages of Aphelenchusavenue. Nematologica, 17 : parasiticnematodes before majoradvances can be 243-254. madein understanding the action and selectivity HAGUE,N.G.M. (1979). A techniqueto assess the of nematicides(Marks, 1971 ; Wright, 1981). Voss effkacy of non-volatilenematicides against the (1981) reasoned that the use of acetylcholinesterase potatocyst nematode (Globoderarostochiensis). inhibiting insecticides with high specificity to closely Ann. appl.Biol., 93 : 205-211. relatedspecies was limited, though families and HAGUE,N.G.M. & PAIN, B.F.(1973). The effect of orderscould be separated. There may be a better organophosphoruscompounds and oxime carba- chance to developnematicides with appropriate -mateson thepotato cyst nematode Heterodera selectivity since agricultural practices usually favour rostochiensis Woll. Pestic. Sci., 4 : 459-465.

Revue Nêmatol. 6 (1) :103-110 (2983) 109 J. Kimpinski & S. El Eraki

HOMEYER,B. & WAGNER,K. (1981). Mode of action cholinesteraseinhibition. In : Kohn, G. K. (Ed.) of fenamiphos and its behavioursoil. in Nematologica, Mechanism of pesticideaction. Washington, USA. 27 : 215-219. American Chemical Society : 1-13. HOUGH,A. & THOMASON,I.J. (1975). Effects of ROHDE,R.A. (1960).Acetylcholinesterase inplant- aldicarb on the behavior of Heterodera schachtii and parasiticnematodes and ananticholinesterase Meloidogyne javanica. J. Nematol., 7 : 221-229. fromasparagus. Proc.helminth. Soc. Wash., 27 : HOUGH,A., THOMASON,I.J. & FARMER,W.J. (1975). 121-123. Behavior of aldicarb in soil relat.ive tocontrol of SAMPFORD,M.R. (1952,). The estimation of response- Heteroderaschaclztii. J. Nematol., 7 : 214-221. timedistributions 1. Fundamental c0ncept.s and JORGENSON,E.C. (1978). Effects of aldicarb on Fusa- general methods. Biometrics, 8 : 13-32. rium wilt-root-knotnematode disease of Cotton. SEINHORST, J.W. (1950). De betekenis van de toestand J. Nematol., 10 : 372-374. vande grond voor het optreden van aanstasting KEETCEI,D.P. (1974). The effect of nematicideson door het stengelaaltje (Ditylenchus dipsaci, (Kühn.) feeding, postureand dispersa1 of Aphelenchus Filipjev). Tijdschr.Plziekt., 56 : 291-349. avenue. Nematologica, 20 : 107-118. SNEDECOR,G.W. & COCHRAN, W.G.(1980). Sta- KIMPINSKI,J. & WILLIS, C.B. (1981).Influence of tisticalmethods. 7 th Ed., lowa StateUniversity soil temperature and pH on Pratylenchus penetrans Press, 507 p. and P. crenatus in alfalfa and timothy. J. Nematol., SONTIRAT,S. & CHAPMAN, R.A. (1970).Penetration 13 : 333-338. of alfalfa roots by different stages of Pratylenchus MARBAN-MENDOZA,N. & VIGLIERCIIIO,D.R. (1980). penetrans (Cobb). J. Nematol., 2 : 270-271. Behavioral effects of carbofuran and phenamiphos STEELE,A.E. & HODGES,L.R. (1975). In-vitro and on Pratylenchusvulnus. 1. Motility and dispersion. in-vivo effects of aldicarb on survival and develop- J. Nematol., 12 : 102-114. ment of Heteroderaschachtii. J. Nematol., 7 : MARKS, C.F. (1971).Respiration responses of a 305-312. Caenorhabditis sp.and Aphelenchusavenue to TOWNSHEND, J.L.(1963). A modification and evalua- the nematicide 1,2-dibromoethane (EDB). J. Nema- tion of theapparatus for the Oostenbrinkdirect fol., 3 : 113-118. cot.tonwoo1 filterextraction method. Nematologica MARKS, C.F., THOMASON, I.J.& CASTRO, C.E. (1968). 9 : 106-110. Dynamics of thepermeation of nematodesby TOWNSHEND, J.L.(1978). Infectivity of Pratylenchus water,nematicides and other substances. Expl penetrans on alfalfa. J. Nematol., 10 : 318-323. Parasit., 22 : 321-337. VANGUNDY, S.D. & MCKENRY,M.V. (1977).Action MARTIN, H. & WORTHING,C.R. (1977). Pesticide of nematicides. In : Horsfall, J. G. & Cowling, manzzal. 5thEd. British Crop Protection Council, E. B. (Eds). Plantdisease an advancedtreatise. 593 p. Vol. 1. Howdisease is managed. London & New MCLEOD,R.W. & KHAIR, G.T. (1975).Effects of York, AcademicPress. : 263-283. oximecarbamate,organophosphate and benzimi- Voss, G. (1981).Taxonomy-related cholinesterase dazolenematicides on life cycle stages of root- inhibitionpatterns in insects. J. econ.Entom., knotnematodes, Meloidogyne spp. Ann. appl. 74 : 555-557. Biol., 79 : 329-341. WHTIEIIEAD,A.G., TITE,D.J., FRASER, J.E. & Moss, S.R., CRUMP, D. & WIIITEIIEAD,A.G. (1975). FRENCH,E. M. (1973).Control of potato cyst- Control of potato cyst-nematodes, Heterodera rosto- nematode, Heteroderarostochiensis, inthree soils chiensis and H.pallida, insandy, peaty and silt by smallamounts of aldicarb,Dupont 1,410 or loam soils byoximecarbamate and organophos- nemacurapplied tothe soil atplanting time. phate nematicides. Ann. appl. Biol., 81 : 359-365. Ann. appl.Biol., 74 : 113-118. NELMES, A.J.(1970). Behavioral responses of Hete- WRIGHT,D.J. (1981).Nematicides : mode of action roderarostochiensis larvaeto aldicarb and its sul- and newapproaches to chemicalcontrol. In : foxide and sulfone. J. Nematol. 2 : 223-227. Zuckerman, B. M. & Rohde,R. A. (Eds). Plant O’BRIEN,R.D., HETNARSKI, B., TRIPATHI,R.R. parasiticnematodes, New York, AcademicPress, & HART,G.J. (1974). Recent studies onacetyl- Vol. 3 : 421-449.

Accepté pour publication le 2 août 1982.

110 Revue NEmatol. 6 (1) : 103-110 (1983)