Entomologia Experimentalis et Applicata 88: 73–80, 1998. © 1998 Kluwer Academic Publishers. Printed in the Netherlands.

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Probing and feeding characteristics of the greenhouse whitefly in association with host-plant acceptance and whitefly strains
H. Lei1,2,∗ , W. F. Tjallingii1 & J. C. van Lenteren1
of Entomology, Wageningen Agricultural University, 6700 EH Wageningen, The Netherlands; of Biology, Beijing Normal University, Beijing 100875, P.R. China; ∗ Current address: Department of Ecology, Lund University, S-223 62 Lund, Sweden
2 Department 1 Department

Accepted: April 16, 1998

Key words: Trialeurodes vaporariorum, rearing history, host plants, acceptance ranks, probing, phloem feeding, electrical penetration graph, DC-EPG

Abstract Host-plant and whitefly strain effects and their interactions on the probing and sap feeding of the greenhouse whitefly, Trialeurodes vaporariorum (Westwood), have been investigated in this study using the DC-EPG (Electrical Penetration Graph) technique. Whiteflies generally displayed fewer but longer probes on highly acceptable cucumber than on less acceptable tomato. Both whitefly strains, the T(omato)-strain and the C(ucumber)-strain, showed a significantly lower number of phloem phases on cucumber than on tomato. However, the duration of total phloem phases achieved by either of the whitefly strains on these two host plants was not significantly different. These data indicate that a more continuous phloem feeding has occurred on cucumber plants. Indeed, the percentage of phloem feeding time after the first sustained phloem phase (longer than 15 min) was higher on cucumber for the C-strain whiteflies. When comparing these two whitefly strains, the T-strain whiteflies probed less frequently but longer than the C-strain whiteflies did on both host plants. Also, the T-strain whiteflies displayed a longer duration of total phloem phases on tomato. An interaction between the whitefly strain and plant effects was detected on a parameter, which showed that whiteflies probed significantly longer before reaching the first phloem phase on the host plants that had been previously experienced. In conclusion, both plant species and whitefly strains affect whitefly’s probing and feeding behaviour, though plant effects are much stronger.

Introduction The greenhouse whitefly, Trialeurodes vaporariorum (Westwood) is a serious pest insect in temperate regions, causing damage to plants in greenhouses and open fields by feeding on plant assimilates and transmitting viruses. Sooty mould fungi which develop on whitefly’s excreted honeydew also cause economic loss by reducing the commercial values of products. Trialeurodes vaporariorum is a polyphagous insect, but large differences exist in its host-plant utilisation pattern, which can be expressed as a performance or acceptance ranking. The acceptance ranks of several host plants, based on the life history parameters of whiteflies such as mortality and fecundity, has been established as follows: eggplant > gherkin > cucum-

ber > gerbera > melon > tomato > sweet pepper (van Lenteren & Noldus, 1990). The quality of the plant sap, the presence of secondary metabolites and certain leaf surface features are thought to contribute to this ranking (Noldus et al., 1986; van Vianen et al., 1988; van Lenteren & de Ponti, 1990). The acceptance ranks are not only influenced by plant factors, but also by whitefly strains (van Boxtel et al., 1978; Thomas, 1993), i.e., a whitefly population that has been reared on a certain plant species for more than 50 generations. Transferring insects from a plant on which they have been reared for a long time to a new host plant species can result in changed performance. The cotton whitefly has been observed to have a high mortality when transferred from cotton or sweet potato to cassava, although cassava is also an

as this diameter has least influence on whitefly’s probing behaviour (Lei et al. However. They were used as indicators of phloem finding and phloem acceptance. Tables 1 and 2). In adult EPGs. but one plant was used for two EPG recordings. referred to as C-strain. The sequence of probing activities by each individual insect was divided into two or three periods on the basis of the following key events: the first phloem phase and the first phloem phase longer than 15 min. 1988). The electrical penetration graph technique (EPG.. About 20 females were tested for each combination of the host plant and whitefly strain. to a 2 cm length of gold wire of diameter 10 µm. has also been used to investigate the probing activities by greenhouse whiteflies (Janssen et al. three main phases are distinct: pathway phase (C and F waveform). EPG parameters. Seedlings with 3–5 true leaves were used in the experiments. the summed non-probing periods (%) before the 1st phloem phase could reflect to what degree the pathway probing was interrupted either by mesophyll factors or by surface factors (Parameter 9. Tables 1 and 2). Lange Groene Giganta) for about 5 years. Then. including salivation and ingestion. to reflect the pathway factors affecting the phloem finding process.74 acceptable host plant for certain strains of this species (Legg. the time needed to reach the 1st phloem phase and the 1st sustained phloem phase were measured. the waveforms from whitefly larvae appeared different than that from adults (Lei et al. Moneymaker) for about 8 years.. However. reflecting phloem feeding. 1988). The main advantage of this technique is that it allows visualisation and quantification of the invisible probing activities within the plant tissue. 1992). xylem phase (G waveform). developed to monitor the probing activities by aphids (Tjallingii. whitefly. in association with host plants of different acceptance levels and with different whitefly strains.. and another on cucumber (cv. EPG recording. we used a proportional phloem phase in the total time after the Materials and methods Plants and insects. referred to as Tstrain. DC system). Numbers and mean duration of total probes and phloem phases were used as general parameters (Parameters 1–4. reflecting water ingestion. Moneymaker were grown in a glasshouse at 18–22 ◦ C and L16:D8 photoperiod. The whitefly stock cultures were kept in separate cages in glasshouse compartments with the same climate conditions as above. and phloem phase (E waveform). 1996a). Cucumber. The whitefly was attached. respectively. Recording was conducted for a total of 12 h for each treatment. The first EPG application to whitefly adults yielded waveforms similar to aphids (Janssen et al. whiteflies need to probe through the epidermis to the vascular bundles using their stylets before they can start feeding on the phloem. The output EPG waveforms are determined by the insect’s behaviour and by the stylet tip positions in leaf tissue. As the absolute duration of the total phloem phases (Parameter 4. 1989). and amplifier were placed in a Faraday cage to shield the set-up from external electric noise sources. Lange Groene Giganta and tomato. As phloem feeders. a droplet of conductive water-based silver paint was deposited on the thorax. one reared on tomato (cv. the whitefly was immobilised using a vacuum device. This paper concerns the investigations into the probing behaviour of the greenhouse whitefly. It is uncertain whether this ‘strain effect’ is reflected in the probing behaviour of whitefly. and the white wax powder on thorax was cleaned off with a fine brush.0 software (Tjallingii & Mayoral. 1989). either from the start of an experiment or from the start of a probe during which the event occurred (Parameters 5–8. Within probing. The EPG signals were acquired (A/D conversion) by a PC and subsequently analysed with STYLET 2. . Whitefly individuals were used once. Three-day-old females were used in the experiments. Before starting the EPG recording. Tables 1 and 2). probing and non-probing can be distinguished. Cucumis sativus cv.. Two whitefly strains were used. it is unknown whether this pattern would be maintained for the host plants of less distinguishable acceptance levels and would be affected by the host-plant origin of whitefly. with silver paint. Plant.. Tables 1 and 2) could be limited by the experimental time. 1997). The recording of EPGs was conducted using a DC system with an input resistance of 109 Ohm (Tjallingii. Therefore. Moreover. Various invisible probing activities take place when the stylets are penetrating into the leaf tissue. 1996b). Preliminary studies regarding whitefly probing behaviour on plants of different acceptance levels have shown that whiteflies probe and feed less on the lowest-ranking host plant (sweet pepper) than on the highest-ranking host plant (cucumber) (Lei et al. Lycopersicon esculentum cv. reflecting intercellular stylet penetration. 1996).

duration. Also. The mean duration of total phloem phases (Parameter 4. To test the efficiency of these transformations. the host plants had a much stronger influence on the probing and feeding activities of whiteflies . Statistical tests were carried out using the software package STATGRAPHICS 7. the percentage of time spent in the phloem after the first sustained phloem phase in the experiment (Parameter 10. only the time to the 1st phloem phase in a probe (Parameter 6) showed a significant interaction between plant and whitefly strain effects in the ANOVA. Before running the ANOVA. the whiteflies had significantly fewer but longer probes and a lower number of phloem phases (Parameters 1–3. Host-plant effects. respectively.75 1st sustained phloem phase to reflect the continuity of the whitefly’s phloem feeding (Parameter 10. but a longer time was needed to reach the first phloem phase. than the whitefly strains and the interactions between the two.025. P=0. P<0.061.001.). a significant interaction between whitefly strain and plant effects was also detected (P=0. Table 2) was not significantly different between cucumber and tomato for both whitefly strains. Table 1). ANOVA analysis was conducted for Parameters 1–3 and 5–9. Statistics. Similarly.77E-6 and 0. respectively. followed by the Tukey– Kramer multiple comparison of means. Table 2). and their interactions. Therefore.05 significance level and thus concluded that the variates of this parameter have heterogeneous variances. Table 2). Finally.001) but not on cucumber (Parameter 10. Results Two-way ANOVA. P<0. logarithmic and arcsine transformations were applied to the data concerning the number. Moreover. a significantly lower percentage of non-probing time before the first phloem phase (Parameter 9. The values of the general parameters (1–4) were calculated using all replicates.001. In contrast. A two-way analysis of variance (ANOVA). Mann-Whitney U. P=0. Whitefly strain effects. but the phloem phase related parameters (5–10) were based on the individuals that showed any phloem phase. Table 2) was calculated. P=0. Table 2). and Statistical Graphics Corp.005) and the mean duration of total probes (P=0.047) but not on cucumber (Parameter 4.042). the plant effect. the T-strain whiteflies probed fewer times but for longer periods than the C-strain whiteflies did (Table 2). which are important assumptions for ANOVA. Table 2).59) was bigger than the critical value (F = 5. Table 2) was found on cucumber than on tomato. Note that for the time to the first phloem phase in a probe (Parameter 6). Interactions between plant and whitefly strain effects. Table 2). the T-strain whiteflies showed a significantly higher percentage of phloem feeding time after the first sustained phloem phase than the Cstrain whiteflies on tomato (Kruskal–Wallis. Table 2) was higher on cucumber than on tomato for the C-strain whiteflies. All other parameters showed only significant plant effects. In general. Significant whitefly strain effects were detected in the number of probes (P=0. Among the 10 parameters. Also. and thus concluded that the variates of these parameters do not fit to a normal distribution.362). Mann–Whitney U. and the results generated from this analysis are presented in Table 1. but not significantly so (Kruskal– Wallis. both from the start of an experiment as well as from the beginning of the probe concerned (Parameters 5–6. P=0. On cucumber. Table 2) was lower on cucumber than on tomato. in the further analysis. the percentage of whiteflies showing phloem phases (Parameter 11. On tomato. Tables 1 and 2). and percentage of the probing activities. Both whitefly strain and plant effects were significant for the number and duration of total probes (Parameters 1 and 2). square root. the percentage of whiteflies that showed any phloem phase within one EPG recording period (12 h) (Parameter 11. was adopted to test the whitefly strain effect. Among the EPG parameters shown in Table 1. The T-strain whiteflies showed a significantly longer duration of total phloem phases than the C-strain whiteflies on tomato (Kruskal–Wallis. non-parametric Kruskal–Wallis and Mann–Whitney U tests were applied to these data. the SchefféBox and Kolmogorov-Smirnov methods were used to examine the variance homogeneity and the data normality. Mann–Whitney U.95) at 0. in addition to a strong plant effect. The P values of both Parameters 4 and 10 in Table 2 calculated from the Kolmogorov-Smirnov method were 3. Scheffé-Box method revealed that the F value for Parameter 10 in Table 2 (F = 8. When tested either on tomato or on cucumber. the time needed to achieve a sustained phloem feeding was longer on cucumber than on tomato (Parameters 7–8.0 (Manugistics Inc.008) in the ANOVA (Parameters 1–2.

f. some other parameters showed a similar trend on average.492 61 68.003 0.507 – – – Residual 78 305. Duration of total probes 3.211 <0.056 0. by phloem sap feeding.301 0. before feeding on the phloem sap. and cessation of each subsequent event in this process and thus contribute to .076 0. significance level. cucumber).f.065 0.452 1 0. P.481 75 9.001 – – – 1 2.0431 0. Number of phloem phases (E) 4. Continuous phloem sap feeding can be regarded as acceptance of a host plant.018 0. However.001 – – – Strain × plant 1 2. Time to the 1st sustained E in probe 9. Number of probes 2. ultimately. SS.331 0. the C-strain whiteflies showed a longer pathway phase before reaching the phloem phase when tested on cucumber than the T-strain whiteflies (Figure 1).874 0.f. E. many factors can play different roles in the initiation.601 <0. followed by the stylet penetration through successive tissue layers between the epidermis and the phloem sieve elements and. SS P 1 32. The process of host-plant selection by whiteflies consists of a series of consecutive events starting with the first labial contact with the plant surface. such as the time to the 1st sustained phloem phase in an experiment and in a probe (Parameters 7–8.8 – – – 1 0.f.408 1 0.982 78 0.05 are marked in bold face. Time to the 1st E in probe 7.f.891 – – – 1 0.005 1 0.012 1 2.042 1 0. sum of squares.001 0. SS P d.941 <0.558 1 0. degree of freedom.047 0.151 1 0.002 1 67.f. and their interactions. SS P d.76 Table 1. Abbreviations: d. phloem phase. maintenance.459 1 0.f. dash line. Although no significant differences were found.f. Discussion and conclusions Effects of host-plant tissue factors on acceptance ranks.f.022 0.074 0. SS P d. sustained E. SS P d.305 – – – Plant 1 85. Time to the 1st E in experiment 6. SS P d.112 0. Percentage of non-probe before the 1st E 76 3.001 1 1.382 1 0. SS P d. Results of two-way ANOVA for EPG parameters concerning whitefly strains (T-.825 1 0.621 65 4. Time to the 1st sustained E in experiment 8.001 1 0.f.f. data violating ANOVA assumptions not shown Whitefly strain d. P values smaller than 0. Similarly.201 <0.691 0.002 <0. SS P d. Percentage of phloem period after the 1st sustained E the T-strain whiteflies spent more time in the pathway probing which led to phloem feeding than the C-strain whiteflies did.001 1 0.852 1 0.392 1 0. C-strain).933 60 1.051 – – – 10. host-plants (tomato.001 1 0.008 1 0. Duration of total phloem phases (E) 5. SS P d.442 – – – 76 9. Table 2) and the proportional phloem feeding time after the 1st sustained phloem phase (Parameter 10.091 0.048 0.122 <0. SS P d.098 0. Table 2). phloem phase longer than 15 min.

A host plant is considered to be of higher acceptance rank when the phloem feeding is more continuous (Parameter 10.7 ± 1. specific phloem properties play a predominant role which may lead to acceptance of the sap as a food source. On suitable host plants.3 ± 27.4 ± 27.0 248.4 58. phloem phase longer than 15 min. in other words phloem acceptance.2 ± 3. The significance test for Parameter 4 and 10 was carried out using Kruskal-Wallis and Mann–Whitney U methods T-whitefly strain Cucumber N = 20 1. and nontransport tissue in the vascular bundle are involved.1 292.7 ± 20.4 168. 1990).8 ± 22. pathway activities are predominant. Percentage of non-probe before the 1st E 10. i.9 ± 1. we took a period longer than 15 min as a criterion for the sustained ingestion. as reflected by the Parameters 5–9 in Table 2. Time to the 1st E in experiment (min) 6. phloem phase.1 539.6 35.9 ± 6.5 162. 1989). we found that whitefly did not excrete any honeydew droplets when the phloem phase was in periods shorter than 15 min. Number of phloem phases (E) 4.5 16. de- pending on whether the first phloem phase is longer than 15 min.4 ± 33.3 1.3 ± 6. with 95% confidence interval. the first phloem phase is usually sustained already.4 ± 52. (1) the first phloem phase and (2) the first phloem phase that is sustained for more than 15 min.2 ± 4.8 ± 52.0 ± 18.8 95 c a b b a a b a c b Tomato N = 19 50. In a few observations where concurrent EPG and honeydew recordings were conducted. which can also be regarded as the first sign of host-plant acceptance. As some of the apical sensilla on a whitefly’s labium.0 454. are chemosensilla (Walker & Gordth.3 ± 4.0 24. sustained E.1 65 ab bc a ab b b a b ab ab 79. These two key events divide the probing process into two or three natural parts.3 ± 9.2 389.0 23.2 9. in contrast to aphids.0 603. Percentage of phloem period after the 1st sustained E 11.2 ± 55. Duration of total phloem phases (E) (min) 5.8 18.. the non-probing duration may be influenced by the surface semiochemicals (van Lenteren & Noldus. Graphic expression of an interaction between the plant effect and the whitefly strain effect.7 ± 9.5 ± 9.4 ± 1. the first phloem phase.3 ± 0. For the duration of the stylet path- .0 ± 2.6 ± 20. Time to the 1st sustained E in experiment (min) 8.6 478.0 34.0 ± 3. Different letters in the same row indicate significance derived from Tukey–Kramer method.5 ± 3. Before the first key event. In our selection of parameters (numbered 1–11 in Tables 1 and 2).0 ± 19.8 185.1 443.9 ± 55.5 1.5 ± 6. A critical selection of parameters to evaluate the acceptance ranks is therefore important.7 49.6 9.9 ± 5. Thus.9 ± 4.8 ± 49.2 ± 50.1 35. the total acceptance rank.8 258. These parameters have been thought to reflect the host-plant recognition as well as the phloem finding process in which factors from epidermis.0 ± 4.3 364.1 32. we used two key events.4 ± 0. E. Percentage of whiteflies showing sustained phloem phase 29.8 43.6 ± 39.1 ± 3.8 ± 3. Time to the 1st sustained E in probe (min) 9. Number of probes 2.4 41.2 31.0 ± 6.4 95 Figure 1.77 Table 2.3 184. Duration of total probes (min) 3. Means and standard errors of EPG parameters in association with two host-plant species (cucumber and tomato) and two whitefly strains (T-strain and C-strain). After the first key event.5 528.0 ± 3. Tables 1 and 2). Chemical and physical properties of plant tissue can greatly affect each element of the probing process.2 46.5 70 a c a ab ab b a b a ab C-Whitefly strain Cucumber Tomato N = 23 N = 20 b b b a a a ab a bc a 43.9 43.2 226.e.6 15.3 ± 29. mesophyll. The data points are the means of log (time to 1st phloem phase (E) in a probe (seconds). Time to the 1st E in probe (min) 7.

in contrast to aphids. 1997). the time to the first or the first sustained phloem phase in an experiment or in a probe (Parameters 5–8. Table 2). Whiteflies may use these chemoreceptors to taste intercellular plant fluids. 1988. Cole. Differences in performance between strains of whitefly have been shown by other authors. Caillaud et al. only the figures of the total phloem phases (Parameter 4. The question arises whether physical factors could have caused this prolonged pathway phase. However. on tomato plants. These parameters include: fewer probing times (Parameter 1. Different pathway probing time may be caused by stimulating or deterring effects of some chemicals in the tissue. than a Dutch whitefly strain. These differences were also observed in host-plant ranking for aphids (Webster et al. more comparative studies are greatly needed to confirm this point. In fact. Table 2) can be enhanced by prolonging the duration of the experiment (Lei et al. (1989) reported that a Hungarian strain of the greenhouse whitefly performed much better on sweet pepper. Table 2) showed the same tendency. Within the total probing time. factors in the superficial tissue layers seem very important. especially the distance to the phloem. very few potential drops. This postponement may decrease the number of whiteflies that reached the phloem phase in the experimental time (Parameter 11. Table 2). Therefore. 1994.78 way phase. 1996). Usually. Van Lenteren et al. Therefore. Table 2). Table 2). 1969. When the pathway phase exerts an analogous influence on the whitefly’s probing. based on a number of EPG parameters. the T-strain whiteflies had a significantly longer phloem phase than the C-strain whiteflies. 1989. Table 2) but almost equal total duration on two host plants. We can speculate that some stimulants present in cucumber tissue have been detected during the occurrence of the C waveform of a whitefly and result in a prolonged intercellular exploration which may give the whitefly a better chance to contact a sieve element. 1995). as indicated by the insects’ performance.. the differences in distance cannot explain this phenomenon completely. complicates this point.. the phloem phase. Apparently. However. Table 2). 1996a). becomes important for ranking a host-plant acceptance level. In our study. such a negative effect can be overcome by using the same plant species or varieties as control. The results of this study demonstrate that cucumber is more readily accepted as a host plant than tomato. No further experiment was carried out to clarify what factors were involved. longer total probing duration (Parameter 2. fewer number of phloem phases (Parameter 3. 1995. due to the paradox that whiteflies actually spent longer time in the pathway probing and thus caused a postponement of subsequent phloem phases on cucumber. does not necessarily reflect the acceptance at all tissue levels in the plant. the higher the acceptance ranks the longer the phloem ingestion and vice versa. one may consider that the overall acceptance rank of a host plant. which is generally a poor host plant for whiteflies. Table 2).. the proportional increase in distance is much lower than that in the pathway probing time prior to the first phloem phase in a probe (Table 3). Janssen et al.. An involvement of chemical factors can certainly not be excluded. Usually. This might be related to a whitefly’s characteristic pathway probing (e. Table 1) showed significant whitefly strain effects. Whitefly strain effects and their interactive role in host-plant acceptance.g. shorter non-probing time before the first phloem phase (Parameter 9. The percentage of whiteflies that reached the phloem phase (Parameter 11. This result could relate to the presence of a large number of pre-cibarial and cibarial chemosensilla in whiteflies (Wensler & Filshie. Hunter et al. The fact of a longer total pathway in high-ranking cucumber is contrary to the results of some aphid studies where more acceptable host-plants were shown to cause shorter pathway probing (Givovich & Niemeyer. and more persistent phloem feeding after the 1st phloem phase in the C-strain whiteflies (Parameter 10. could account for this. 1993. Consequently. This impediment may be more serious on cucumber leaves than tomato leaves due to different morphological leaf features. thus forming the principal contribution. together with other possible factors such as the impediment of whitefly’s free movement caused by wiring. clearly distinguishable subpatterns of C waveform). Anatomical differences between the two host plants. two parameters (Parameters 1–2. but no significance was observed on cu- . most likely do not sample cells intracellularly (Tjallingii.. this overall acceptance may be inconsistent with some phases of the probing process.. and references therein). The T-strain whiteflies exhibited fewer probes and longer total probing time than the C-strain ones on cucumber as well as on tomato. we measured the distance to the phloem in the tertiary veins (the preferred probing sites) and it is indeed larger in cucumber than in tomato (Table 3). as these homopterans. especially the sustained phloem ingestion.. Alternatively. Lei et al.

the only parameter that showed significant interactive effects in the ANOVA was the time to the first phloem phase in a probe (Parameter 6. such as those related to the two key events (Parameters 5–10. Host-plant induced changes on the whitefly’s performance have been previously reported. a certain selection ‘pressure’ has been effective on the insects. P. in comparison with the increased whitefly pathway time before the 1st phloem phase and 1st sustained phloem phase in the probe concerned (Parameters 6 and 8 in Table 2) Tomato Phloem distance (µm) Time to 1st E in probe (min) T-strain C-strain Time to 1st sustained e in probe T-strain C-strain 97 23 16 18 15 Cucumber 108 35 42 35 44 % increment in cucumber 11 52 163 94 193 cumber plants. the resulting effect could be partly genetic and partly phenotypic. 1996. Acknowledgements This study was part of the co-operative research program between Wageningen Agricultural University (WAU). Other parameters. Caillaud. the Dutch Ministry of Agriculture & Fisheries and WAU. van Lenteren. some other parameters (Parameters 7. previous experience also resulted in more persistent phloem feeding after the 1st phloem phase (Parameter 10. C.). P. In conclusion. van. Medeldelingen van de Faculteit Landbouwwetenschappen Rijksuniversiteit Gent. Determination of host plant quality of eggplant (Solanum melongena L.) for the greenhouse whitefly (Trialeurodes vaporariorum (Westwood)) (Homoptera: Aleyrodidae). B. S. Pierre. known as ‘hostplant induction’ (Bernays & Weiss.79 Table 3. C. So. a long-time association with a host-plant species may result in behavioural modifications. but this effect was not independent. J. & M. Long-time rearing of whiteflies on a single host-plant species can form a certain strain with its own probing characteristics. 43/2. References Bernays. Instead. Table 2) showed the same non-significant tendency. Woets & J. Entomologia Experimentalis et Applicata 78: 1–8. the Netherlands and Beijing Normal University. the whitefly strains interacted so that the ranking of the strains on tomato. 8 and 10. cucumber (Cucumis sativus L. The adults tested in our experiments were reared on their strain host plants and had no previous experience on any other plants. This interactive effect indicates that a previous experience can cause the insects to spend a longer time on pathway probing. The host-plant origin of whiteflies should be taken into account when performing EPG tests on different host plants. Besides the prolonged pathway probing. Table 1).R. Boxtel. Our experiments do not allow the separation of these two components. Table 2) showed no clear strain differences. In the course of their rearing history. the probing and feeding profile of whiteflies is mainly determined by plant factors at all tissue levels. 1978. Table 2). Di Pietro. 397–408. tomato (Lycopersicon esculentum L.). Enkegaard (1993) examined the performance of two cotton whitefly (Bemisia tabaci (Gennadius)) strains (poinsettia strain and tobacco strain) on poinsettia plants and discovered that the poinsettia-strain whiteflies had longer longevity and higher fecundity than the tobacco-strain whiteflies. Induced food preferences in caterpillars: the need to identify mechanisms. according to the values of this parameter. J. M. This time was longer on cucumber than on tomato. Actually. Proportional increment (%) in cucumber (Cucumis sativus) phloem distance from lower epidermis. Analysis of wheat resistance to the cereal aphid Sitobion avenae using electrical penetration graphs and flow charts combined .) and paprika (Capsicum annuum L.. The senior author was financially supported by the Dutch Royal Academy of Sciences (KNAW). Chaubet & J. E. was the reverse of that on cucumber (Figure 1). 1995. This might be a favourable strategy for whiteflies to find the phloem vessel by exploring the tissue more extensively. A. On the other hand. R. In our experiments. W. as shown in plant effects. China. 1996). Weiss.

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