Presence of trehalulose and other oligosaccharides in hemipteran honeydew, particularly Aleyrodidae

Abstract. Production by a bacterium of the disaccharide trehalulose was reported 30 years ago. The association between the fabrication of trehalulose and Insecta was found more recently. It was initially discovered in the honeydew, excreta, of the whitefly Bemisia tabaci feeding on Euphorbia pulcherrima where it was the predominant sugar. In the present study, B. tabaci -produced trehalulose was again found in honeydew at significant levels, this time when the whitefly fed on 10 different plants. In seven of 10 Bemisia /host combinations, trehalulose accounted for more than 30% of the total carbohydrates found in their honeydew and was the principal oligosaccharide. Trehalulose constituents, glucose and fructose, were also present at lower levels, as were other oligosaccharides such as melezitose. Feeding by B. tabaci on the three other plant hosts also resulted in the production of trehalulose at relatively high levels (6.1–16.5%). Other whitefly species examined had little (e.g. B. afer and Aleurothrixus spp.), or no trehalulose (e.g. Trialeurodes spp. and Siphoninus phillyreae ), in their honeydew. Trehalulose was also found in the honeydew of two aphid and one scale insect species. In insects whose honeydew had low levels or no trehalulose, sucrose, its constituents, or larger sugars predominated. The trisaccharide bemisiose was also found in the honeydew of half the whitefly species examined. Bemisiose was discovered here for the first time in the honeydew of three aphid species and three species of scale insects. Reasons for the production of trehalulose are often linked to high levels of dietary sucrose in whiteflies. This is probably true in our case because cotton and cucurbits, at least, are known to contain a great deal of sucrose. Although other functional possibilities were explored, the data suggest that trehalulose, being less susceptible to hydrolysis than sucrose, is involved in osmoregulation.     

The longevity effects of trehalulose feeding by parasitoids

Abstract.  Insects, particularly phloem-feeding Sternorrhyncha, are known to produce sugars in their honeydew (excreta) that are not found in their host plants. Of these, Bemisia tabaci , the sweet potato whitefly, is the only insect known to produce trehalulose [α- d -glucose (1,1) d -fructose] as a major component of its honeydew. The present study aims to determine whether trehalulose is comparable to sucrose as a nutrient source for three whitefly parasitoids ( Encarsia formosa , Encarsia pergandiella and Eretmocerus eremicus ). In addition, the study also examines trehalulose feeding effects on longevity for a parasitoid of muscoid Diptera, Nasonia vitripennis . Parasitoids are provided diets of either sucrose or trehalulose in varying concentrations (from 0.1% to 70%) or a water control. Sucrose and trehalulose are not significantly different in affecting survival when compared at the same concentration. This was true for all Bemisia parasitoids and N. vitripennis. Certain specific diets are significantly different in pairwise combination tests. There is a significant effect of species, diet type and the interaction of these two factors on the longevity of the three different Bemisia parasitoid species; however, within species, there is no significant increase in longevity observed for either carbohydrate diet. This result contrasts with expectations for the effects of host-modified carbohydrates on longevity. The implications are that, although carbohydrate feeding is essential for these parasitoids, these host-provided sources of carbohydrates are equally capable of extending longevity.     

Distinct sucrose isomerases catalyze trehalulose synthesis in whiteflies,Bemisia argentifolii,and Erwinia rhapontici

Isomaltulose [alpha-D-glucopyranosyl-(1,6)-D-fructofuranose] and trehalulose [alpha-D-glucopyranosyl-(1,1)-D-fructofuranose] are commercially valuable sucrose-substitutes that are produced in several microorganisms by the palI gene product, a sucrose isomerase. Trehalulose also occurs in the silverleaf whitefly, Bemisia argentifoli, as the major carbohydrate in the insect's honeydew. To determine if the enzyme that synthesizes trehalulose in whiteflies was similar to the well-characterized sucrose isomerase from microbial sources, the properties of the enzymes from whiteflies and the bacterium, Erwinia rhapontici, were compared. Partial purification of both enzymes showed that the enzyme from whiteflies was a 116 kD membrane-associated polypeptide, in contrast to the enzyme from E. rhapontici, which was soluble and 66 kD. The enzyme from E. rhapontici converted sucrose to isomaltulose and trehalulose in a 5:1 ratio, whereas the enzyme from whiteflies produced only trehalulose. Unlike the E. rhapontici enzyme, the whitefly enzyme did not convert isomaltulose to trehalulose, but both enzymes catalyzed the transfer of fructose to trehalulose using sucrose as the glucosyl donor. The results indicate that trehalulose synthase from whiteflies is structurally and functionally distinct from the sucrose isomerases described in bacteria. The whitefly enzyme is the first reported case of an enzyme that converts sucrose to exclusively trehalulose.     

Nutritional ecology of a parasitic wasp: food source affects gustatory response, metabolic utilization, and survivorship

The success of biological control is partly mediated by the longevity and reproductive success of beneficial insects. Availability of nectar and honeydew can improve the nutrition of parasitic insects, and thereby increase their longevity and realized fecundity. The egg parasitoid, Anaphes iole, showed strong gustatory perception of trehalulose, a carbohydrate found in homopteran honeydew. Chromatographic analysis demonstrated that enzymatic hydrolysis of sucrose, a common nectar sugar, proceeded at a faster rate than that of melezitose, a sugar common in aphid honeydew. A long-term bioassay showed that longevity was greater at 20 °C than at 27 °C, and at both temperatures survival was generally greatest for wasps provisioned with the three major nectar sugars, sucrose, glucose, and fructose. Patterns of food acceptance and utilization showed that A. iole accepted and utilized a broad range of sugars found in nature, including those found in nectar as well as honeydew. Glucose, fructose, and several oligosaccharides composed of these monosaccharide units appear to be more suitable for A. iole than other sugars tested. Evidence suggests that individual fitness benefits afforded by food sources are important for a time-limited parasitoid, and that continued investigations on the interface between nutrition and biological control are warranted for A. iole.     

Effect of six carbohydrate sources on the longevity of a whitefly parasitoid Eretmocerus hayati (Hymenoptera: Aphelinidae)

Parasitoid adults often acquire carbohydrates by feeding on floral nectar and honeydew which provides them with energy and prolongs their life span. The concentration and type of saccharide in nectar and honeydew are variable by species of plant and insect. To explore the effects of various sugar type and concentrations on parasitoid fitness, we compared 5%, 10% and 20% (w/v) solutions of six different sugar resources (glucose, fructose, sucrose, trehalose, melezitose and honey) on the longevity of Eretmocerus hayati, a larval parasitoid of the whitefly Bemisia tabaci in China. Male and female longevity was increased by all of the sugar diets, but female wasps survived longer than the males when the same sugar diet was supplied. Female parasitoids feeding on 10% glucose and 10% honey increased longevity, respectively up to 6.2- and 5.9-fold longer than distil water; 5% honey and 10% fructose had the greatest effects on male longevity, up to 3.5- and 3.3-fold. All six sugar diets, no matter which concentration, significantly changed the survival curves. Glucose, sucrose and honey were optimal sugar diets for this wasp, and 10% was the optimal concentration. Our results could provide an insight into the nutritional requirements of E. hayati under laboratory conditions. Such information can be a basis to improve the longevity of this biological control agent by sugar feeding during the indoor mass-rearing process.     

寄主植物对双斑恩蚜小蜂的发育、存活和繁殖的影响

研究了茄子和扶桑两种不同寄主植物对烟粉虱寄生性天敌双斑恩蚜小蜂Encarsia bimaculata发育、存活和繁殖等特性的影响。在26±0.5℃的条件下,在扶桑上,双斑恩蚜小蜂卵至成虫的发育历期为13.6天,2龄幼虫至成虫的存活率为93.2%;而在茄子上双斑恩蚜小蜂卵至成虫的发育历期和2龄幼虫至成虫的存活率分别为12.1天和91.1％。扶桑上双斑恩蚜小蜂雌性成虫的平均寿命为8.0天,平均单雌产卵量为35.9粒;而在茄子上雌性成虫的平均寿命为6.6天,平均单雌产卵量为27.6粒。双斑恩蚜小蜂在扶桑和茄子植物上的种群内禀增长率分别为0.2081和0.892。两种不同寄主植物对双斑恩蚜小蜂在若虫发育历期和存活率、成虫寿命和平均产卵量以及内禀增长率等方面均存在着显著差异。研究结果表明,寄主植物对双斑恩蚜小蜂的发育、存活和繁殖有较大的影响。     

烟粉虱B生物型若虫、皮蜕及成虫提取物对双斑蚜小蜂行为的影响

烟粉虱B生物型的若虫、皮蜕及其成虫的提取物作为一种利它素信息源,在室内对其在双斑蚜小蜂寻找寄主取食、寄生行为的影响进行了生物测定。烟粉虱的若虫、皮蜕及其成虫分别用正己烷、乙醇和无菌水进行粗提。研究结果发现,双斑蚜小蜂在处理区寻找寄主停留的时间高于对照区。在处理区,双斑蚜小蜂行动活泼,对利它素源表现出高的正趋向性和选择性。对于同一利它素源、同一提取介质的两种不同浓度,双斑蚜小蜂在若虫 水提取物的高浓度区停留的时间（111.23s）最长,与在低浓度区的停留时间差异显著;而在烟粉虱皮蜕及其成虫的水、正己烷和乙醇提取物处理区,不同浓度的提取物对蚜小蜂停留的时间影响差异不显著。本研究的结果表明,利它素可以增加蚜小蜂寻找寄主的效率,有利于蚜小蜂寻找到适宜的寄主。     

Effect of six sugars on the longevity,oviposition performance and nutrition accumulation in an endoparasitoid,Meteorus pulchricornis (Hymenoptera: Braconidae)

The effects of six sugar resources (fructose, glucose, sucrose, trehalose, raffinose and honey) on the longevity, oviposition performance and nutrition levels of Meteorus pulchricornis, a thelytokous larval endoparasitoid of the common cutworm Spodoptera litura were examined under laboratory conditions. Female adults of M. pulchricornis fed 1 M fructose, glucose, trehalose or sucrose solutions survived longer than those fed on other sugar solutions or water. When provided with honey or sucrose solutions, the female parasitoids laid more offspring than those fed other sugar diets or the control. The body size of offspring driven from honey-, fructose-, sucrose-, and glucose-fed females, along with water-fed group, were larger than the trehalose- and raffinose-fed females. However, the emergence rates of all offspring generated from different sugars- and water-fed females were similar. When separately given honey, sucrose or fructose, M. pulchricornis females accumulated fructose at a higher level than the other groups. Parasitoid wasps fed trehalose solution accumulated the highest level of total sugar. Glycogen levels and lipid content were highest at emergence and then decreased across all diets. In addition, females fed on trehalose had the highest level of glycogen compared to other sugar diets and water control regardless of emergency level. Females fed trehalsoe, fructose, and glucose solutions had a higher level of lipid than those fed other sugar solutions and water at life end. The outcome of this study can benefit both laboratory rearing and management interventions that improve sugar sources for the parasitoid in the field.     

Effects of aphid honeydew sugars on the longevity and fecundity of the aphidophagous gall midge Aphidoletes aphidimyza

The predatory gall midge Aphidoletes aphidimyza, which is used for augmentative biological control of aphids in greenhouses, uses aphid honeydew as an energy source. Currently, Rhopalosiphum padi on barley plants and Melanaphis sacchari on sorghum plants are used as the banker plant systems for A. aphidimyza to control Aphis gossypii on eggplant in Japan. We analyzed the sugar components of the honeydew of these three species by HPLC. The major components of honeydew were sucrose, fructose, and melezitose for A. gossypii; glucose and fructose for R. padi; and glucose, fructose, and melezitose for M. sacchari. Maltose and trehalose were minor components of the honeydew of these species. We investigated the influence of sugars, including three artificial aphid honeydews, six sugar components, and distilled water, on the longevity of unmated A. aphidimyza. Both females and males attained the greatest longevity on sucrose and artificial honeydew of A. gossypii. Mean longevities of both females and males were shortest when they were provided only with water. We conducted another experiment in which a mated female was released in a cage with an eggplant seedling infested with A. gossypii and fed with sucrose or only water. Females fed with sucrose lived significantly longer and had higher lifetime fecundity than those fed only water.     

Searching behaviour of Encarsia formosa as mediated by colour and honeydew

The habitat- and host-searching behaviour of female Encarsia formosa Gahan (Hymenoptera: Aphelinidae) was assessed using an airflow olfactometer and a filter paper test. Responses to different odour cues, colours, host-produced honeydew, non-host honeydew and single carbohydrates were determined. The parasitoid was not attracted to or arrested by odours emanating from clean tobacco leaves, tobacco leaves heavily infested with the host Trialeurodes vaporariorum (Westwood) (Homoptera: Aleyrodidae) and covered with honeydew, or honeydew alone. However, E. formosa females showed a significant response to green light transmitted through a tobacco leaf. The yellow part of the spectrum was partly responsible for this response. Thus, the long-range orientation is random with respect to the presence of hosts. Filter paper tests showed that the short-range searching behaviour is influenced by water soluble, non-volatile contact-kairomones contained in the host-produced honeydew. Contact with honeydew excreted by L3/L4 T. vaporariorum resulted in longer searching times than honeydew from adult T. vaporariorum or L3/L4 Bemisia tabaci Gennadius (Homoptera: Aleyrodidae). No difference was found between the response to honeydew excreted by adult and L3/L4 B. tabaci. The parasitoids' response to honeydew was unaffected by the host plant on which the whiteflies had fed. Non-host honeydew and single carbohydrates also affected the searching behaviour of E. formosa but to a lower extent than host honeydew. The possible differences in the carbohydrate and amino acid composition of the honeydew excreted by different life-stages of T. vaporariorum and B. tabaci are discussed.     

两种蚜小蜂对烟粉虱寄主的产卵选择性（英文）

双斑恩蚜小蜂和桨角蚜小蜂是华南地区烟粉虱的两种优势种寄生蜂。本文研究了两种寄生蜂对不同龄期烟粉虱寄主的产卵选择特性。结果表明: 两种寄生蜂均可寄生烟粉虱的1～4龄若虫。当只有1个龄期的烟粉虱若虫存在时,双斑恩蚜小蜂更多地寄生3龄和4龄若虫,较少寄生1龄和2龄若虫;而桨角蚜小蜂则更多地寄生3龄和2龄若虫,较少寄生1龄和4龄若虫。在4个龄期若虫同时存在时,双斑恩蚜小蜂会明显增加对3龄和4龄若虫的产卵寄生,降低对1龄和2龄若虫的寄生;而桨角蚜小蜂则增加对2龄和3龄若虫的寄生,减少对1龄和4龄若虫的寄生。寄主植物的差异不影响这二种蚜小蜂对各龄期烟粉虱若虫的产卵选择倾向。结果提示,烟粉虱若虫3龄和4龄是双斑恩蚜小蜂最适宜的寄主,而桨角蚜小蜂最适宜的寄主是烟粉虱2龄和3龄若虫。     

两种蚜小蜂对烟粉虱寄主的产卵选择性(英文)

双斑恩蚜小蜂和桨角蚜小蜂是华南地区烟粉虱的两种优势种寄生蜂。本文研究了两种寄生蜂对不同龄期烟粉虱寄主的产卵选择特性。结果表明:两种寄生蜂均可寄生烟粉虱的1~4龄若虫。当只有1个龄期的烟粉虱若虫存在时,双斑恩蚜小蜂更多地寄生3龄和4龄若虫,较少寄生1龄和2龄若虫;而桨角蚜小蜂则更多地寄生3龄和2龄若虫,较少寄生1龄和4龄若虫。在4个龄期若虫同时存在时,双斑恩蚜小蜂会明显增加对3龄和4龄若虫的产卵寄生,降低对1龄和2龄若虫的寄生;而桨角蚜小蜂则增加对2龄和3龄若虫的寄生,减少对1龄和4龄若虫的寄生。寄主植物的差异不影响这二种蚜小蜂对各龄期烟粉虱若虫的产卵选择倾向。结果提示,烟粉虱若虫3龄和4龄是双斑恩蚜小蜂最适宜的寄主,而桨角蚜小蜂最适宜的寄主是烟粉虱2龄和3龄若虫。     

Sugar, organic acid, phenolic acid and plant growth regulator content of extracts of honeydew of the aphid Myzus persicae and of its host plant, Raphanus sativus

The honeydew of Myzus persicae feeding on radish seedlings contains glucose, fructose, trehalose, melezitose and sucrose, of which glucose and fructose are present in radish seedlings. In addition, the honeydew contains eight organic acids, of which seven are present in uninfested seedlings and six are present in infested radish seedlings: of the eighteen phenolic acids in the honeydew, five are present in uninfested radish seedlings. Auxins, gibberellins, growth inhibitors and cytokinins are present in the honeydew. The presence of carbonate and bicarbonate previously recorded in honeydew of M. persicae could not be confirmed, but ammonia is present in fresh honeydew and accounts for its high pH.     

Plutella xylostella (diamondback moth) and its parasitoid Diadegma semiclausum show different gustatory and longevity responses to a range of nectar and honeydew sugars

Parasitoids as well as many of their herbivorous hosts, depend on carbohydrate‐rich food during the adult stage. Different types of nectar and honeydew vary with regard to their sugar composition. In order to successfully exploit a food source, the insect must show a positive gustatory response to its component sugars and be able to digest and metabolise them. Here we tested the herbivore Plutella xylostella L. (Lepidoptera: Plutellidae) and its parasitoid Diadegma semiclausum (Hellén) (Ichneumonidae: Campopleginae) with respect to their feeding response and longevity when provided with one of nine sugars (fructose, glucose, lactose, maltose, melibiose, melezitose, raffinose, sucrose, or trehalose). Both species responded to and showed a prolonged life span on a broad range of sugars. The impact of food supply on life span was about sixfold higher for the parasitoid than for the herbivore. In general, there was a good fit between gustatory response and achieved longevity, with some outliers. Both species showed only weak responses to melibiose, which significantly prolonged life span. The parasitoid showed a gustatory response to melezitose, which did not prolong its life span. The parasitoid and its herbivorous host responded differently to trehalose. These differences in gustatory response and longevity show the potential for application of selective sugar sources in conservation biological control. At the same time it also reveals a risk that the indiscriminate application of sugar sources may stimulate herbivory.     

Effect of high temperature on the metabolic processes affecting sorbitol synthesis in the silverleaf whitefly,Bemisia argentifolii

Whiteflies accumulate the polyhydric alcohol, sorbitol, when exposed to temperatures greater than about 30 degrees C. Feeding experiments using artificial diets containing labeled sucrose showed that more of the label was incorporated into whitefly bodies and less was excreted in the honeydew when feeding was conducted at 41 compared with 25 degrees C. Analysis of the components of the honeydew showed that more of the excreted label was in glucose and fructose and less in trehalulose at 41 degrees C than at 25 degrees C. A similar effect of temperature on honeydew composition occurred for whiteflies feeding on cotton leaves. Measurement of the activities of glycolytic, pentose-phosphate and polyol pathway enzymes at 30 and 42 degrees C showed that NADPH-dependent ketose reductase/sorbitol dehydrogenase (NADPH-KR/SDH), sucrase, glucokinase and glucose-6-phosphate dehydrogenase activities were stimulated to a greater extent at 42 degrees C than trehalulose synthase and fructokinase. NAD(+)-sorbitol dehydrogenase (NAD(+)-SDH) activity was inhibited at 42 degrees C. We propose that high temperature alters metabolic activity in a way that increases the availability of fructose and stimulates pentose-phosphate pathway activity, providing both the substrate and coenzyme for sorbitol synthesis. High temperature also increases the activity of NADPH-KR/SDH, the enzyme in whiteflies that synthesizes sorbitol, but inhibits the activity of NAD(+)-SDH, the enzyme that degrades sorbitol.     

甘蓝与黄瓜寄主上B型烟粉虱和温室白粉虱蜜露糖分、氨基酸和挥发物组分的比较分析

粉虱蜜露是粉虱寄生性天敌搜索寄主的主要利它素源。应用离子色谱分别对甘蓝与黄瓜上B型烟粉虱(Bemisia tabaci B-biotype)蜜露以及黄瓜上温室白粉虱Trialeurodes vaporariorum蜜露的接触性利它素糖和氨基酸组分和含量进行了比较研究。结果表明：2种粉虱在不同寄主植物上的蜜露均富含糖和氨基酸,其中糖含量占绝对优势,甘蓝上B型烟粉虱蜜露、黄瓜上B型烟粉虱蜜露和黄瓜上温室白粉虱蜜露中的糖含量分别是相应氨基酸含量的42.5、2.6和5.4倍,其中糖类物质中又以寡糖含量占绝对优势,分别占89.3%、81.7%和88.2%。不同寄主植物和粉虱种类显著影响蜜露中糖和氨基酸的组成和含量。其中,甘蓝上B型烟粉虱蜜露中的寡糖以二糖占优势,占97.3%;二糖中又以蔗糖异构糖和松二糖占优势,分别占52.7%和35.4%。黄瓜上B型烟粉虱蜜露和温室白粉虱蜜露寡糖中以三糖和四糖占优势,分别占73.1%和85.4%;优势糖水苏（四）糖和松三糖分别占40.3%和 26.2%及49.9%和27.0%。甘蓝上B型烟粉虱蜜露中氨基酸以丙氨酸占优势,含量为66.5%;而黄瓜上B型烟粉虱及温室白粉虱蜜露中氨基酸以甘氨酸含量最高,分别占氨基酸总量的38.2%和51.7%。应用GC-MS对甘蓝上B型烟粉虱蜜露和黄瓜上温室白粉虱蜜露挥发物组分的鉴定结果显示,两种粉虱蜜露中共同含有的主要挥发物为邻苯二甲酸二（2-乙基）己酯。     

High Performance Liquid Chromatography Analysis of Carbohydrates of Cotton-Phloem Sap and of Honeydew Produced by Bemisia tabaci Feeding on Cotton

Phloem sap from cotton (Gossypium hirsutum L.) was collected from young and mature leaves by the aphid-stylet technique. Exudate was analyzed for carbohydrates by HPLC using sensitive pulsed amperometric detection. The predominant carbohydrate present (>90%) was identified as sucrose. A second, unidentified compound that was not one of the more commonly translocated sugars was detected in mature leaves. Carbohydrates in honeydew produced by the sweet-potato whitefly (Bemisia tabaci [Genn.]) feeding on cotton were sucrose, glucose, fructose, trehalulose, and a series of oligosaccharides.     

Oligosaccharide synthesis regulates gut osmolality in the ant-attended aphid Metopeurum fuscoviride but not in the unattended aphid Macrosiphoniella tanacetaria

Abstract:  An artificial phloem sap (APS) for Metopeurum fuscoviride and Macrosiphoniella tanacetaria , based on analysis of their host plant, Tanacetum vulgare , phloem sap, contained 308  μ g/ μ l (900  μ mol/ μ l) sucrose (and no other sugars) and a mixture of 14 amino acids with a total concentration of 82.5 nmol/ μ l. There was no significant difference in the total amino acid concentration of the honeydew of adult M. fuscoviride fed on the host plant and aphids fed on APS. Incubation of isolated guts in APS indicated no role of gut bacteria or gut enzymes on the amino acid pattern in the gut. The sugar composition of the honeydew of the ant-attended M. fuscoviride indicated a rapid digestion of sucrose into glucose and fructose, and the simultaneous synthesis of considerable amounts of melezitose and some trehalose. The sugar composition of the honeydew of the unattended M. tanacetaria in contrast showed only traces of trehalose and melezitose, but up to 20% erlose in plant-fed aphids. Incubation of isolated guts of M. fuscoviride in APS demonstrated a steady high rate of melezitose synthesis by gut enzymes over an 8-h period. Incubation of isolated guts of M. tanacetaria on the other hand demonstrated only a moderate rate of erlose synthesis and no detectable melezitose or trehalose. Melezitose in the aphid M. fuscoviride is a signal sugar for ants (ecological function), indicating the presence of abundant sugar rich honeydew [ Woodring et al. (2004) Physiol. Entomol. , vol. 29, pp. 311–319]. It was estimated that melezitose reduces the gut osmolality of M. fuscoviride to approximately 25–35% of what it would be without the synthetases (physiological function). M. tanacetaria on the other hand produces very little honeydew, is not attended by ants, and thus there is little need to synthesize large amounts of oligosaccharides to attract ants or for osmoregulation.     

Comparing the effects of five naturally occurring monosaccharide and oligosaccharide sugars on longevity and carbohydrate nutrient levels of a parasitic phorid fly, Pseudacteon tricuspis

Abstract.  The longevity and nutrient levels of Pseudacteon tricuspis provided with 1  m solutions of five naturally occurring sugars, fructose, glucose, sucrose, trehalose and melezitose, are compared. All but melezitose, result in significant increases in the longevity of P. tricuspis in comparison with sugar-starved flies (flies provided with water only). Sugar-starved female and male P. tricuspis have an average longevity of 3.3 and 4.1 days, respectively. Provision of free water in addition to sugar solution is necessary for optimum longevity by female and male flies. Longevity is increased by 2.4–2.6-fold by the two monosaccharides, fructose and glucose, and by 2.6–2.8-fold by the disaccharides, sucrose and trehalose. Phorid flies provided with the trisaccharide sugar, melezitose, had a marginal increase in lifespan (approximately 1 day), but this is not significantly different from the longevity of sugar-starved flies. Significantly greater levels of total sugars are detected in P. tricuspis fed the disaccharide sugars (sucrose, trehalose) or the monosaccharide sugars (fructose, glucose), compared with flies provided with melezitose (trisaccharide), or to sugar-starved flies. Fructose is not detected in sugar-starved flies, or in flies fed glucose or trehalose. However, high levels of fructose are detected in flies fed sucrose or fructose, whereas levels of fructose in melezitose-fed flies are intermediate. In general, significantly greater glycogen levels are detected in P. tricuspis fed sucrose, glucose, trehalose or fructose, compared with melezitose-fed or sugar-starved flies. Levels of total sugars and glycogen in sugar-fed flies are positively correlated with wing length, possibly indicating a higher accumulation of storage sugars by larger flies. These results are discussed in relation to the nutritional ecology of the phorid fly.     

Behavioral responses of the parasitoid Psyllaephagus pistaciae (Hymenoptera: Encyrtidae) to host plant volatiles and honeydew

The behavioral responses of the parasitoid Psyllaephagus pistaciae, the major biocontrol agent of the common pistachio psylla, Agonoscena pistaciae, to volatiles emanating from its host plant and host honeydew, were examined using a four‐arm airflow olfactometer. In addition, the arrestment behavior of this parasitoid on clean and honeydew‐treated leaves of the pistachio, Pistacia vera, was monitored. The infested pistachio leaves were the most favored source of the volatile attracting the parasitoids. The parasitoid clearly distinguished and responded to infochemicals emitted by psyllid honeydew but at a lower level than to the volatiles from infested host plants. However, the searching time, locomotory behavior, antennal drumming and ovipositor probing were all affected when they encountered honeydew‐contaminated zones on pistachio leaves. These findings suggest that the psyllid honeydew releases kairomones that stimulate the parasitoids to greater searching activity, as well as providing a directional cue. The intensive searching activities in the presence of the volatiles tested were very similar to responses by the parasitoid females when encountering patches treated with psyllid honeydew. Such behavior could retain the parasitoid in a favorable area, thereby increasing the probability of additional host encounters.