HOST ALTERNATION IN APHIS FABAE SCOP.

Experiments on the feeding preferences and relative fecundity of Aphis fabae among leaves of different ages and kinds were extended to the comparison of different forms of the aphid: apterous and alate virginoparae and gynoparae.
All three forms showed a preference for the primary host, Euonymus , over a secondary host, sugar beet, but this preference was strongest in the gynoparae, weaker in the alate virginoparae and weakest in the apterae. The relative fecundity of the aphids on the two kinds of leaf paralleled their feeding preferences most closely in the gynoparae, less so in the alate virginoparae and least in the apterae.
All three forms also showed some preference for growing over mature leaves of the same kind, but this preference was strongest in the apterous virginoparae, weaker in the alate virginoparae and weakest in the gynoparae. The relative fecundity of the aphids on the two ages of leaf paralleled their feeding preferences most closely in the apterae, less so in the alate virginoparae and least in the gynoparae.
The physiological, ecological and evolutionary significance of the results is discussed in the light of the dual discrimination theory of aphid host selection.     

METHODS FOR MASS REARING AND INVESTIGATING THE HOST RELATIONS OF APHIS FABAE SCOP.

A method is described for producing some hundreds of apterous and thousands of alate virginoparous Aphis fabae every day continuously throughout the year. Broad beans are used as the host plants, in a 'production-line' arrangement of eleven ventilated breeding cages, with additional cages for starting subcultures and rearing alatae, all contained in a portion of a small greenhouse with supplementary lighting in winter. When running smoothly, the culture requires about half an hour's work by one man per day.
An 'aphid gun', being a modified type of aspirator catcher, is described, with which large numbers of aphids could be handled and counted rapidly without damaging them.
Amodified technique for host-transference experiments is described, which permits individual leaves to be compared one with another as host-units for the aphids. Two types of small metal cages are used, both easily attached to leaves in their natural positions on growing plants. One type confines the aphids to a part of a single leaf and is used mainly for determining relative reproduction rates on different leaves. The other type encloses equal areas of two different leaves between which the aphids can move freely, and is used for determining their behavioural preferences. The principles underlying the use of such cages, the experimental procedures and the interpretation of the results obtained are discussed.     

HOST FINDING BY APHIDS IN THE FIELD: APHIS FABAE SCOP. (GYNOPARAE) AND BREVICORYNE BRASSICAE L.; WITH A RE-APPRAISAL OF THE ROLE OF HOST-FINDING BEHAVIOUR IN VIRUS SPREAD

Aphis fabae gynoparae occurred in the same large proportion in simultaneous collections of all aphids alighting and probing on, and taking off from, a host plant (spindle) and a non-host (peach), and behaved similarly when approaching and leaving them in the same conditions. Most alighters took off again from leaves of both kinds within a few minutes, staying longer and probing more times on the host. In atmospheric conditions favouring local 'hovering' instead of dispersal, flying and alighting aphids became concentrated around host plants, not through any specific attraction to them, but apparently because more aphids had accumulated upon them and were now taking off.
Brevicoryne brassicae occurred in the same large proportion in simultaneous collections of aphids alighting on a host plant (cabbage) and a non-host (sugar beet). No satisfactory evidence was found of preferential alightment on cabbage and there were indications of preferential alightment on the non-host. A very small minority of the A. fabae and B. brassicae that alighted on their hosts stayed there long enough to larviposit. This minority was rather larger among alighters late in the day, but in the absolute sense, more colonization occurred during earlier periods when more aphids arrived.
The intensely dispersive type of host-finding behaviour in Myzus persicae, A. fabae and B. brassicae may be common among Aphididae. It seems ideal for the dissemination of non-persistent plant viruses, more particularly among the less-favoured host plants of each aphid. The tendency to commensal association between virus and vector provides an ecological framework which may govern the incidence of virus-vector specificity and symbiosis.     

蚜虫寄主植物与取食部位的多样性

在中国科学院动物研究所已有标本采集记录和国内外文献资料的基础上,分别从蚜虫类Aphidina科级和属级阶元系统研究世界范围内蚜虫寄主植物与取食部位的多样性.蚜虫的寄主植物种类繁多,涉及267科2120属,尤其以菊科、禾本科、豆科、伞形科、唇形科、蔷薇科、茜草科、兰科、壳斗科、杨柳科、胡桃科等植物为主.13科蚜虫的寄主植物差别很大:在科级水平,球蚜科Adelgidae、纩蚜科Mindaridae和平翅绵蚜科Phloeomyzidae的寄主植物类群比较单一,蚜科Aphididae和瘿绵蚜科Pemphigidae的寄主植物范围最为广泛;在属级水平,各属蚜虫间寄主植物也有明显差异,有143属蚜虫的寄主植物多于2科,其中蚜科的属占多数,蚜属Aphis、瘤蚜属Myzus、长管蚜属Macrosiphum、粗额蚜属Aulacorthum和声蚜属Toxoptera的寄主植物最多,各超过100科290属.在不同的寄主植物上寄生着不同种类的蚜虫,其中16科47属寄主植物上寄生的蚜虫多于14属.蚜虫在寄主植物上的取食部位丰富多样,可分为7类,分别为叶片、嫩梢、嫩枝、茎、花、根部、果实等.以叶片、茎、嫩枝和嫩梢为蚜虫主要取食部位.平翅绵蚜科主要取食嫩枝、茎和根部,短痣蚜科Anoeciidae主要取食叶片、嫩梢和根部,球蚜科、群蚜科Thelaxidae和毛管蚜科Greenideidae取食叶片、嫩梢、嫩枝和茎等部位,纩蚜科取食叶片、嫩梢、茎和花等部位,大蚜科Lachnidae和斑蚜科Drepanosiphidae除取食叶片、嫩梢、嫩枝、茎等部位外,前者还在根部取食,后者还寄生在果实上,根瘤蚜科Phylloxeridae在除花以外的其它6个部位取食,蚜科、瘿绵蚜科和毛蚜科Chaitophoridae的取食部位最为多样,它们在上述7个部位均可取食.还初步讨论了在不同寄主植物上蚜虫物种的分化,以及蚜虫与寄主植物之间的对应关系.     

THE DISTRIBUTION OF APHID INFESTATION IN RELATION TO LEAF AGE

Infestations of apterous Aphis fabae Scop, on potted sugar beets have been followed in detail for several weeks. The plants were somewhat stunted and their crowns presented an unusually complete series of leaf ages. Records were kept of the changing number and size of the leaves and of their stage of growth. Parallel records were kept of the changing population of aphids on every leaf, and the figures are analysed in various ways to show how suitability for the aphids varied through the life cycle of the leaves.
The leaves were very suitable when young, became unsuitable as they matured, became suitable again just after maturity and then unsuitable again as they senesced. But among leaves at any given stage, those which were growing or senescing rapidly were more suitable than those changing slowly, unless the rate of senescence was very high. The differences of population density on different-aged leaves were due largely to the preferences exercised by the apterous adults. The added effect of differences in the fecundity of these mothers while feeding on different leaves was not excluded, but could not be assessed. It is concluded that the physiological development of the plant as a whole determines, through the growth and senescence among its total complement of leaves, the progress and pattern of its aphid infestation.     

Climate effects on life cycle variation and population genetic architecture of the black bean aphid, Aphis fabae

Aphid species may exhibit different reproductive modes ranging from cyclical to obligate parthenogenesis. The distribution of life cycle variation in aphids is generally determined by ecological forces, mainly climate, because only sexually produced diapausing eggs can survive harsh winters or periods of absence of suitable host plants. Aphids are thus interesting models to investigate intrinsic and environmental factors shaping the competition among sexual and asexual lineages. We conducted a Europe-wide sampling of black bean aphids, Aphis fabae, and combined population genetic analyses based on microsatellite data with an experimental determination of life cycle strategies. Aphids were collected from broad beans (Vicia faba) as well as some Chenopodiaceae, but we detected no genetic differentiation between aphids from different host plants. Consistent with model predictions, life cycle variation was related to climate, with aphids from areas with cold winters investing more in sexual reproduction than aphids from areas with mild winters. Accordingly, only populations from mild areas exhibited a clear genetic signature of clonal reproduction. These differences arise despite substantial gene flow over large distances, which was evident from a very low geographic population structure and a lack of isolation-by-distance among 18 sites across distances of more than 1000 km. There was virtually no genetic differentiation between aphids with different reproductive modes, suggesting that new asexual lineages are formed continuously. Indeed, a surprising number of A. fabae genotypes even from colder climates produced some parthenogenetic offspring under simulated winter conditions. From this we predict that a shift to predominantly asexual reproduction could take place rapidly under climate warming.     

Leaf toughness changes the effectiveness of larval aggregation in the butterfly Byasa alcinous bradanus (Lepidoptera: Papilionidae)

Two subspecies of the papilionid butterfly Byasa alcinous , B. a. bradanus and B. a. alcinous , have varying degrees of larval aggregation. Early instar larvae of ssp. bradanus always occur in aggregations. To determine the functions of larval aggregation in this subspecies, we examined the effects of leaf toughness on larval performance when caterpillars were reared alone and in aggregations. Newly hatched larvae were reared either individually or in groups of 10 and were fed either tough or tender leaves of Aristolochia debilis . When fed tough leaves, more gregarious larvae survived the first instar. This difference between solitary and aggregated larvae did not occur when caterpillars were fed soft leaves. The effects of aggregation on larval weight and duration were not significant between leaf-toughness treatments. Larval aggregation of B. a. bradanus improves larval survivorship in early instars that use host plants with tough leaves.     

3种寄主上桃蚜的选择性及形态分化

桃蚜是一种重要的农业害虫,寄主广泛,种下分化复杂。以采自黄土高原旱作区桃树、烟草、甘蓝上的桃蚜为研究对象,通过叶片选择法、传统比较形态测定法研究了3种寄主上桃蚜的选择性及形态分化。结果表明:在3种寄主同时存在的情况下,烟草上的桃蚜嗜食烟草,表现为63.5%的桃蚜选择烟草叶,13.8%选择甘蓝叶,8.2%选择桃叶,而甘蓝和桃树上的桃蚜对原寄主没有表现出明显的嗜好性;从形态指标来看,3种寄主上的桃蚜在体长、触角末节长度、后足腿节长度、触角与体长的比例方面存在显著差异(P0.05),说明这几个特征可以作为区分这3种寄主上桃蚜的依据。综合分析可以初步认为黄土高原旱作区烟草上的桃蚜可能形成了寄主专化型-烟草型。     

On the processes shaping reproductive isolation in aphids of the Aphis fabae (Scop.) complex (Aphididae: Homoptera)

Models of sympatric speciation for phytophagous insects are based on the premise that ecological specialization on different host plants can indirectly result in significant reproductive isolation. A candidate example of host plant shift is provided by the partially reproductively-isolated aphids Aphis fabae fabae and A. f. mordwilkoi , which utilize different hosts for sexual reproduction. However, unexpected isolation based on mating behaviour was found for these two aphids. In olfactometer trials, males responded to the pheromones of females of both subspecies when presented a subspecies in isolation but preferentially to females of the same subspecies in choice tests. On contact, the incidence or duration of between-subspecies copulations was significantly lower than same-subspecies copulations, and spermatozoa were transferred during 88% of the same-subspecies copulations, but only 19% of between-subspecies copulations. These data are indicative of strong pre-zygotic isolation between the subspecies. Complementary restriction analysis of mitochondrial DNA and a plasmid (pAFE/eu) in an obligate symbiotic bacterium (Buchnera) revealed a paraphyletic relationship among four subspecies of A. fabae (A. f. fabae, A. f mordwilkoi, A. f cirsiiacanthoidis and A. f. solanella). We discuss how the lack of clear genetic differentiation between A. f. fabae and A. f. mordwilkoi , despite substantial pre-zygotic isolation, may be a consequence of reinforcement.     

APHID BEHAVIOUR ON HEALTHY AND ON YELLOWS-VIRUS-INFECTED SUGAR BEET

Myzus persicae Sulz., M. ascalonicus Doncaster, Aphis fabae Scop, and Aulacorthum solani Kalt., when caged on sugar-beet leaves in the glasshouse preferred yellows-virus-infected leaves to healthy ones; they chose those with the most severe symptoms on which they bred more rapidly and lived longer than on green leaves. M. persicae behaved similarly on whole plants in the glasshouse. The previous host influenced the aphid's preference. Differences between the multiplication rates of aphids on healthy plants of inbred sugar-beet varieties were eliminated or reversed by infection with yellows virus.
Spraying healthy plants with either sugar solutions or hydrolysate of casein increased the multiplication rate of M. persicae , but much less than did virus infection.     

Variation in Eastern cottonwood (Populus deltoides Bartr.) phloem sap content caused by leaf development may affect feeding site selection behavior of the aphid, Chaitophorous populicola Thomas (Homoptera: Aphididae)

Apterous populations of Chaitophorous populicola Thomas (Homoptera: Aphididae) appear to track Eastern cottonwood (Populus deltoides Bartr.) leaf development. Few aphids occur on mature leaves. Marked individual aphids on leaves of different developmental stages were observed through a period of new leaf initiation. Nymph and adult C. populicola frequently track leaf development by moving up to younger leaves. A comparison of phloem sap constituents and leaf toughness among leaf developmental stages revealed some differences that could be used by C. populicola to determine leaf age. Phloem sap exudates, collected from P. deltoides leaves of different developmental stages, were analyzed by high-performance liquid chromatography for free amino acids and the phenolic glycoside salicin. Sucrose concentration in exudates, indicative of phloem sap exudation rate, was uniform among leaf stages. Of 20 amino acids examined, only aspartic acid and gamma-amino-n-butyric acid (GABA) concentrations differed significantly between leaf stages. Forward stepwise discriminant function analysis showed that seven of the amino acids analyzed are useful for classifying leaf maturity groupings. Aphid-infested cottonwoods had lower cystine concentrations in phloem sap than aphid-free plants. Salicin concentration was significantly higher in new leaves. Leaf toughness was assessed by lignin density and distance measurements in petiole cross-sections. Rapidly expanding leaves had significantly less lignification and new leaves had shorter distances to the vascular bundles than senescent leaves. These physiological and phytochemical differences among P. deltoides leaf developmental stages may contribute to the leaf stage selection patterns exhibited by the aphid, C. populicola.     

Aphids do not attend to leaf colour as visual signal, but to the handicap of reproductive investment

The evolution of visual warning signals is well known in animals but has received scant attention in plants. The coevolutionary hypothesis is the most influential hypothesis on warning signals in plants proposing that red and yellow leaf colours in autumn signal defensive strength to herbivores. So far, evidence in support of the hypothesis, which assumes a coevolutionary origin of autumnal leaf colours, is correlative and open to alternative explanations. We therefore tested the coevolutionary hypothesis experimentally by colouring the leaves either red or green of same-aged mountain ash (Sorbus aucuparia) individuals. We monitored the response of winged aphids to leaf colour using insect glue on branches with natural and artificial leaf colours in each individual. In contrast to the prediction of the coevolutionary hypothesis, aphid numbers did not differ between the individuals with artificial green or artificial red leaves. Likewise, at the within-plant level, aphids did not colonize branches with natural green leaves preferentially. However, we suggest that plants emitted warning signals because aphids colonized the hosts non-randomly. We found a strong positive correlation between aphid numbers and fruit production, suggesting an allocation trade-off between investment in plant defence and reproduction. Our study demonstrates that aphids use warning signals or cues in host selection, probably volatiles, but that they did not use leaf colour.     

Evaluation of induced responses, insect population growth, and host-plant fitness may change the outcome of tests of the preference-performance hypothesis: a case study

The preference‐performance hypothesis predicts that insect preference should correspond to host suitability for offspring development. We studied the pattern of within‐plant preference in the aphid Sipha flava and its consequences for offspring performance on the host‐plant Sorghum halepense, regarding the role of induced responses of plants to aphid feeding. The consequences of within‐plant preference on aphid population growth and host‐plant traits were also evaluated. Our results showed that winged and wingless aphids preferred to settle on mature rather than young leaves. In contrast, aphid individual growth rate was higher on young leaves when compared with mature leaves, suggesting that the outcome of this test rejected the preference‐performance hypothesis. However, the inclusion of the factor ‘previous aphid infestation’ changed the outcome from a maladaptive choice to a neutral one. Thus, individual growth rates of S. flava increased when aphids developed on leaves that had been previously infested. Interestingly, aphid growth rate on previously infested leaves did not differ between young and mature leaves. On the other hand, aphid population reproductive rate was higher and the percentage of winged aphids lower when infestation occurred on mature rather than young leaves. Aphid infestation reduced plant and shoot biomass, and increased leaf mortality. These negative effects on plant traits related to plant fitness were greater when aphid infestation occurred on young leaves. Likewise, whereas infestation on mature leaves did not cause a significant reduction in the number of flowering plants compared with control plants, aphid infestation on young leaves did reduce the number of plants at the flowering stage. Consequently, if both the reproductive rate of aphids in the mid‐term, and host‐plant fitness are taken into account, the results indicate that aphid preference for mature leaves may be an adaptive choice, thus supporting the preference‐performance hypothesis.     

Nutritional enhancement of host plants by aphids - a comparison of three aphid species on grasses

Three aphid species were compared with respect to ability of enhancing the nutritional quality of their host plants. Rhopalosiphum padi, which does not induce macroscopic changes in its host plants, was compared with Schizaphis graminum and Diuraphis noxia, both of which induce distinctive types of chlorotic lesions. Phloem sap samples were collected from severed stylets of feeding aphids and from exudates of cut leaves of plants uninfested or infested with each aphid species. Samples were analyzed for concentrations of individual amino acids.Compared to R. padi, S. graminum ingested phloem sap with a two-fold higher concentration of amino acids and a much higher proportion of essential amino acids. Similar differences between these two aphid species were observed on both wheat and barley. For each aphid species, the absolute concentrations of amino acids and the relative proportions of essential amino acids were similar between the two host plants. Effects of D. noxia were similar to those of S. graminum, though less dramatic. Exudates from leaves infested with each aphid species showed relative concentrations of individual amino acids that were similar to those in the corresponding stylet exudates. Based on comparison of stylet exudates and cut leaf exudates from infested and uninfested plants, R. padi seems to have little effect on amino acid composition of phloem. Changes in the phloem induced by both S. graminum and D. noxia appear to be systemic, affecting at least the whole leaf they are feeding on. The changes observed for D. noxia and for S. graminum are likely to be nutritionally advantageous for the aphids and are expected to affect the aphids' dependence on nutritional supplementation by intracellular symbionts (Buchnera).     

General Gregariousness and Specific Social Preferences among Wild Chimpanzees

Wild chimpanzees form temporary parties that vary in size and composition. Previous studies have revealed considerable intraspecific variation in party compositions. We examined patterns of association among age, sex, and reproductive classes of chimpanzees at Ngogo in the Kibale National Park, Uganda. We employed a class-based association index and a randomization procedure to control for confounding factors and to test for differences between classes. Results indicate that males associated with other males significantly more than expected if all classes behaved equivalently, while females generally associated with individuals of the same sex less than expected. To interpret these patterns we used two additional indices that separate associations into two components: general gregariousness and preference for particular classes of associates. Males and estrous females were more gregarious than other classes, while anestrous females were less so. After controlling for general gregariousness, adult males as a class showed no specific preference for associating with each other. Anestrous females preferred each other as party members, and estrous females avoided each other. These results are consistent with previous findings that adult males are more gregarious than females. They diverge from the standard picture of chimpanzee society, however, by suggesting a mutual affinity among anestrous females, but not among adult males as a class.     

MORPHOLOGICAL STUDIES ON MYZUS PERSICAE IN CHINA

Abstract  Experiments were carried out to study the morphological characters of Myzus persicae (Sulzer) on tobacco, rape and peach from different localities of China. The results showed that the number of accessory hairs on ultimate rostral segment had no difference among all samples in 14 morphological characters used by Blackman in this study. Cluster analysis indicated that the morphological differences between samples from peach and samples from other host plants was the biggest, the next was the samples from Changchun on tobacco. Compared with peach, the aphid from rape was more similar to that from tobacco in morphology. For the samples on tobacco from the central and southern China, they can be clustered into yellowish-green group and reddish-brown group. It shows unlikely to separate successfully the aphids on tobacco from those on non-tobacco host plants only by morphological characters.     

Flight behaviour of the black bean aphid, Aphis fabae, and the cabbage aphid, Brevicoryne brassicae, in host and non-host plant odour

Abstract. Walking alate virginoparae of Brevicoryne brassicae (L.) and Aphis fabae Scopoli were presented with odours of steam-distilled extracts of the non-host plants summer savoury ( Satureja hortensis L.) and tansy ( Tanacetum vulgare L.) in an olfactometer. No effects of the extracts were observed on B. brassicae. However, A.fabae were repelled by summer savoury and tansy odour; both extracts also masked an attractant response to bean (host plant) odour. In a flight chamber, air permeated with odour from host or non-host plants was blown over flying alates of both species, with a green, plant-mimicking target presented once a minute. The incidence of targeted (host-responsive) flight of B.brassicae was not affected by odour from a growing host plant ( Brassica oleoracea ) or a non-host plant tansy. Host plant ( Vicia faba ) odour did not affect the initial rate of climb or the incidence of targeted flight of A.fabae , but when the bean odour was alternated with odour from non-host tansy plants a greater number of targeted flights occurred in the host plant odour. The volatile extracts of tansy and summer savoury were also presented to flying A.fabae. Aphids flying in air permeated with tansy odour at 450g plant equivalents initiated fewer targeted flights than when flying in clean air. However, no differences in flight behaviour were observed with summer savoury extract. In a horizontal wind tunnel the tansy extract at 90 g plant equivalents blown across the surface of yellow targets reduced the numbers of alate A.fabae landing. The results indicate that plant odours can affect flight and landing of aphids.     

中国桃蚜形态学研究(英文)

对来自中国不同地区的烟草、油菜和桃树的桃蚜形态特征研究结果表明：在所研究的14个形态特征中,喙末节端部的次生毛的数目这一形态特征在所有的样本中均无显著差异。聚类分析表明采自桃树上的桃蚜的形态特征与采自其它寄生植物上的桃蚜的形态特征差异最大;其次是采自长春烟草上的桃蚜。油菜上的桃蚜在形态上比桃树上的桃蚜更接近烟草上的桃蚜。我国中南部烟草上的桃蚜又可区分为黄绿和红褐两类。在本研究中所用的14个形态特征不能明确地将我国烟草上的桃蚜与其它寄生植物上的桃蚜区分开。     

How does the host‐specialized aphid deal with food deficiency?

Aphis gossypii Glover shows obvious host specialization, with cucurbit‐ and cotton‐specialized biotypes or host races in many regions. Because its annual natal host crops senesce earlier the cucurbit‐specialized biotype may suffer food deficiency. The method this biotype uses to overcome this challenge is still poorly understood. In order to understand the potential of the cucurbit‐specialized biotype aphids in host shift and usage, the performance of this biotype on cotton (Gossypium hirsutum), a common but poor quality host plant, was explored in this study. The cucurbit‐specialized aphids could establish populations on cotton only when these plants had at least nine leaves, and subsequent populations developed rather slowly. The presence of whitefly populations on cotton improved the success rate of cucurbit‐specialized aphids. The cucurbit‐specialized aphids were mainly distributed on the older leaves of cotton, with only a few settling on the upper leaves. The cucurbit‐specialized aphids reared on cotton for 40, 54 and 61 days still maintained strong preference for their natal host plant, cucumber (Cucumis sativus), rather than cotton, and their net reproductive rates and intrinsic rates of natural increase were dramatically lower when they were transferred onto new six‐leaf cotton plants or detached leaves. Therefore, we concluded that the cucurbit‐specialized aphids have the potential to utilize mature or whitefly‐stressed cotton plants, but that this feeding experience on cotton did not alter their specialization for cucurbits. Some cotton plants could act as a temporary host for the cucurbit‐specialized aphids to overcome food deficiency arising from senescing cucurbits.     

Direct and Indirect Impacts of Infestation of Tomato Plant by Myzus persicae (Hemiptera: Aphididae) on Bemisia tabaci (Hemiptera: Aleyrodidae)

The impacts of infestation by the green peach aphid (Myzus persicae) on sweetpotato whitefly (Bemisia tabaci) settling on tomato were determined in seven separate experiments with whole plants and with detached leaves through manipulation of four factors: durations of aphid infestation, density of aphids, intervals between aphid removal after different durations of infestation and the time of whitefly release, and leaf positions on the plants. The results demonstrated that B. tabaci preferred to settle on the plant leaves that had not been infested by aphids when they had a choice. The plant leaves on which aphids were still present (direct effect) had fewer whiteflies than those previously infested by aphids (indirect effect). The whiteflies were able to settle on the plant which aphids had previously infested, and also could settle on leaves with aphids if no uninfested plants were available. Tests of direct factors revealed that duration of aphid infestation had a stronger effect on whitefly landing preference than aphid density; whitefly preference was the least when 20 aphids fed on the leaves for 72 h. Tests of indirect effects revealed that the major factor that affected whitefly preference for a host plant was the interval between the time of aphid removal after infestation and the time of whitefly release. The importance of the four factors that affected the induced plant defense against whiteflies can be arranged in the following order: time intervals between aphid removal and whitefly release > durations of aphid infestation > density of aphids > leaf positions on the plants. In conclusion, the density of aphid infestation and time for which they were feeding influenced the production of induced compounds by tomatoes, the whitefly responses to the plants, and reduced interspecific competition.