A seed parasitoid wasp prevents berries from changing their colour,reducing their attractiveness to frugivorous birds

1. Frugivorous and seed‐feeding insects may alter the traits of fruits, such as shape and size, which may influence fruit attractiveness to frugivorous birds. Consequently, trait‐mediated interactions may occur in systems where plants, seed‐dispersing frugivorous vertebrates, and frugivorous or seed‐feeding insects interact. We investigated colour manipulation in Ilex integra Thunb. berries caused by the seed parasitoid wasp Macrodasyceras hirsutum Kamijo and how that manipulation relates to fruit attractiveness for frugivorous birds. 2. In winter, the colour of I. integra berries varied from green to red, but most berries were greenish, indicating that the berries were immature. Berry dissection indicated that the number of live parasitoid larvae present within each berry was closely related to berry colour – the greater the number of live larvae, more intense is the green colour of the berry. However, the wasp larvae did not modify the shape or size of the berries. More than 98% of berries that were protected from the insects by gauze bags ripened and turned red. In the present study, berries with unfertilised seeds alone turned red. Field‐feeding preference tests showed that the brown‐eared bulbul Hypsipetes amaurotis Temminck preferred red berries to green berries. 3. We demonstrated that the seed parasitoid wasp manipulates the berry colour, but not its shape or size, in a density‐dependent manner. Because green berries suffered less from bird foraging, we believe that this colour manipulation helps the wasps to avoid being killed by the birds. The present study indicates that manipulation by wasps may reduce the level of mutualism between the tree and seed‐dispersing birds.     

The Importance of Behavioral Plasticity for Maximizing Foraging Efficiency in Frugivorous Lepidopteran Larvae

In this study, we present evidence that the larvae of Acrobasis vaccinii (Lepidoptera: Pyralidae), a species that feeds on cranberry fruit, are able to take into account at least three cues (fruit size, fruit color, and the distance between fruits) when searching for food. In laboratory experiments, the relative impact of each cue depended on which cues were presented in a given bioassay. Fruit color was the dominant cue in some contexts (e.g., larvae showed a significant perference for green fruit over red ones regardless of fruit size when fruits were equidistant from the larvae) but not in others (e.g., when given a choice between a large red fruit placed close to the larvae and a small green fruit placed farther away, a significantly higher proportion of larvae chose the former). This plasticity in foraging behavior allows larvae to maximize foraging efficiency under conditions of differing fruit and larval densities, which this species experiences in nature.     

Signals of profitability? Food colour preferences in migrating juvenile blackcaps differ for fruits and insects

Red is a common colour signal in both aposematic warning displays, and in fruit displays. One common feature is that red is conspicuous against the natural background of the prey and fruits. However, there is a potential conflict between fruits and aposematic prey in how a bird predator should react to red colours, where fruits aim to attract birds and aposematic insects aim to ward off, often the same bird individuals. Here we investigate possible differences in red/green colour preferences of frugivorous, wild-caught, young blackcaps (Sylvia atricapilla), when food is either a fruit or an insect. Birds in two groups were presented with a series of pairs of food items that had been artificially painted red and green, in the order of (I) fruits, crickets and maggots, or (II) crickets, fruits, and maggots. Birds first presented with crickets or fruits differed in first attacks directed at the two colours: They showed no colour preference between fruits, but showed a clear preference for green over red crickets. Also, birds in both experimental groups clearly preferred green to red maggots. These results provide evidence that wild, frugivorous birds are able to differentiate between prey types, and show different colour preferences depending on whether food is insect or fruit. We conclude that blackcaps show an attack bias against red insects, and that one important function of the signal in insects, is to inhibit attack after discovery. However, the lack of preference for red fruits suggests other functions to red fruit displays, such as facilitating discovery per se, rather than directly stimulating attack after discovery.     

Red foliage color reliably indicates low host quality and increased metabolic load for development of an herbivorous insect

Plant chemical defense and coevolved detoxification mechanisms in specialized herbivorous insects are fundamental in determining many insect–plant interactions. For example, Brassicale plants protect themselves from herbivory by producing glucosinolates, but these secondary metabolites are effectively detoxified by larvae of Pierid butterflies. Nevertheless, not all Brassicales are equally preferred by these specialist herbivores. Female Pieris butterflies avoid laying eggs on anthocyanin-rich red foliage, suggesting red color is a visual cue affecting oviposition behavior. In this study, we reared P. brassicae larvae on green and red cabbage leaves, to determine whether foliage color reliably indicates host plant quality. We did not find a difference in survival rates or maximal larval body mass in the two food treatments. However, larvae feeding on red cabbage leaves exhibited significantly lower growth rates and longer durations of larval development. Interestingly, this longer development was coupled with a higher consumption rate of dry food matter. The lower ratio of body mass gain to food consumption in larvae feeding on red cabbage leaves was coupled with significantly higher (ca. 10 %) larval metabolic rates. This suggests that development on red foliage may incur an increased metabolic load associated with detoxification of secondary plant metabolites. Energy and oxygen allocation to detoxification could come at the expense of growth and thus compromise larval fitness as a result of extended development. From an evolutionary perspective, red foliage color may serve as an honest defensive cue, as it reliably indicates the plant’s low quality as a substrate for larval development.     

Is fruit size important in the selection of oviposition sites by cranberry fruitworm,Acrobasis vaccinii?

Under laboratory conditions, females of Acrobasis vaccinii (Riley) (Lepidoptera: Pyralidae) oviposit significantly more often on larger than on smaller cranberry fruit. This behaviour ensures that neonate larvae have access to maximum resources and are larger when moving to the next fruit. However, while there is some evidence that on a large spatial scale females may lay eggs in patches of larger fruit, there was no evidence of such an oviposition preference within patches in natural bogs. These results may be explained by the fact that the oviposition period for A. vaccinii begins early in the cranberry season, before fructification is completed, leading to potential competition among females for the first fruit present in the environment. Laying eggs on the first fruit available may be advantageous as this behaviour increases the probability that females select unexploited fruit. This strategy would be particularly effective for maximizing female fitness in grazer frugivorous species (such as the cranberry fruitworm) that live in environments where severe climatic conditions induce limited flowering/fruit formation and where hosts thus are rare.     

Exploring host‐associated differentiation in the North American native cranberry fruitworm,Acrobasis vaccinii,from blueberries and cranberries

The factors explaining host‐associated differentiation (HAD) have not yet been fully characterized, especially in agricultural systems. It is thought that certain characteristics within a system may increase the probability for HAD to occur. These characteristics include relatively long‐standing evolutionary relationships between insects and their host plants, endophagy, and allochrony in host‐plant phenologies. We assessed the status of these characteristics as well as the presence of HAD in the cranberry fruitworm, Acrobasis vaccinii Riley (Lepidoptera: Pyralidae), a pest associated with blueberry and cranberry in eastern North America. We reveal the occurrence of two distinct populations of A. vaccinii that are allochronically isolated by the phenological stage of their respective host plants (cranberries or blueberries). Laboratory‐reared A. vaccinii adults collected from blueberries emerge at least 1 week earlier than adults from cranberries and the antennal sensitivity of adults to host‐plant volatiles differs between A. vaccinii collected from blueberry and cranberry. Despite finding characteristics indicative of HAD, we did not detect a genetic signature of HAD in A. vaccinii. These findings suggest that HAD may occur through behavioral and phenological mechanisms before there is sufficient genetic variation to be detected.     

Red light stimulates flowering and anthocyanin biosynthesis in American cranberry

Morphological responses of American cranberry (Vacciniummacrocarpon Ait, Ericaceae) to different light conditions (red,far-red, white light and sunlight) were examined. Root growth and development,stem elongation, leaf enlargement, de-etiolation of stem and leaf, flower budformation, and flowering of American cranberry were measured under each lightcondition and in the dark. It was found that red light promotes development ofroots and leaves, flowering, and de-etiolation of stem and leaf of Americancranberry. Stem elongation and etiolation of stem and leaf were shown infar-redlight and dark. Anthocyanin biosynthesis as phytochemical response in cranberryplants was most sensitive to red light. Estimation of anthocyanin levels indifferent parts of cranberry plant suggested that anthocyanins were presentonlyin red fruit skins, and not in peeled fruits, green fruits, green leaves, greenstems, roots and seeds.     

Differences in larval emergence chronotypes for sympatric Rhagoletis brncici Frías and Rhagoletis conversa (Bréthes) (Diptera,Tephritidae)

In central Chile, Rhagoletis brncici and R. conversa, can be found in sympatry, associated with the fruit of their native host plants: Solanum tomatillo and S. nigrum (Solanaceae), respectively. Third-stage larvae must emerge from its host in search of pupation sites, and during this period larvae must find an appropriate pupation microhabitat while avoiding predation and adverse abiotic factors. In this study, we explored whether these sympatric species differ in terms of the timing of their larval exit from the host fruit in search of pupation sites. Field-collected fruits from host plants were checked daily for larval emergence, within 24 h, under laboratory conditions, in order to determine the time of the event. We found that these species differed significantly in their diel larval emergence. For R. brncici, most larvae left the host fruit between late evening and past midnight, meanwhile larvae from R. conversa concentrated their peak of emergence near midnight and early morning. We discuss these findings in terms of the ecological and evolutionary implications of the temporal separation of larval emergence regarding the use of pupation sites, abiotic stress and risk of predation for these sympatric species.     

Conspicuousness, not colour as foraging cue in plant–animal signalling

The global prevalence of red and black fruits has still not been explained. Hypotheses based on innate consumer preferences have been tested and rejected. Though colour itself plays an important role in animal foraging, it is only one component of signals. Another major component are colour contrasts against background achieving the conspicuousness of signals. In order to evaluate which signal component determines consumers behaviour, we measured fruit colour and colour contrasts of 43 species against their natural background under ambient light conditions. Red and black fruits exhibit stronger contrasts and are therefore more conspicuous to consumers than fruits of other colours. Subsequently, trials were carried out to determine whether colour or conspicuousness influences avian food choice. Four bird species strongly preferred contrasting red–green or black–green over uni-coloured red, green, or black fruit displays, while no preference for particular hues was found. We therefore hypothesize that conspicuousness determines avian food selection and define the contrast hypothesis: Diurnal dispersers select fruit colours based on their conspicuousness and not their colour itself.
Because colour vision is an ancient trait, the entire heterogeneous group of frugivorous birds most likely perceives conspicuousness uniformly over evolutionary time spans. Conspicuousness has thus the potential to explain the global prevalence of red and black fruits.     

Developmental regulation of phospholipase D in tomato fruits

The catabolism of phospholipids initiated by phospholipase D (PLD, EC 3.1.4.4) is an inherent feature of developmental processes that include fruit growth and ripening. In cherry tomatoes (Lycopersicon esculentum Mill.), soluble and membrane-associated PLD activities increased during fruit development, which peaked at the mature green and orange stages. The increase in PLD activity was associated with a similar increase in the intensity of a 92 kDa band as demonstrated by western blot analysis. A full-length cDNA having 2430 bp and encoding a putative polypeptide with 809 amino acids, was isolated using tomato RNA, RT-PCR and 5' and 3' rapid amplification of cloned ends (RACE). Analysis of the primary and secondary structures showed the presence of the C2 domain, the PLD domain and several other features characteristic of PLD alpha. Microtom tomato plants transformed with antisense PLD alpha cDNA, were similar to untransformed plants and showed normal fruit set and development. The ethylene climacteric was delayed by over 7 d in the antisense PLD fruits, indicative of a slower ripening process. The leaves and unripened fruits of antisense PLD microtom plants possessed lowered PLD activity and PLD protein, as demonstrated by western blotting. However, during ripening, PLD activity in the transgenic fruits was maintained at a higher level than that in the untransformed control. Immunolocalization of PLD in microtom tomato fruits revealed the cytosol-membrane translocation of PLD during fruit development. The ripe fruits of antisense PLD celebrity plants possessed lowered PLD expression and activity and showed increased firmness and red colour. These results suggest that the expression of antisense PLD cDNA could be variable in different tomato varieties. The potential role of PLD in ethylene signal transduction events is discussed.     

Oviposition behaviour of two tephritid fruit flies,Dacus tryoni and Dacus jarvisi,as influenced by the presence of larvae in the host fruit

Summary When offered a choice, females of the fruit flies Dacus tryoni (Frogg.) and D. jarvisi (Tryon) strongly preferred to lay in fruits without larvae rather than fruits which already contained larvae. Fruits which contained even low densities of larvae, including newly hatched ones, received many fewer eggs than control fruits. This preference was not influenced by the species of larvae present in the fruits nor by the distance to uninfested fruits. Discrimination occurred when fruits with and without larvae were close together (10 cm apart) and also when they were separated by distances of about one metre. Laboratory assays suggested that the flies detect chemical changes in the fruit associated with the decomposition which accompanies larval feeding, but they do not seem to detect the larvae Perse. This behaviour may be significant when these two species utilise the same host since the species which is able to infest fruits first will reduce the availability of hosts for the other species. In contrast to many other Tephritids (e.g. Rhagoletis, Anastrepha and Ceratitis) female Dacus don't discriminate against fruits which contain eggs nor do they deposit a pheromone to deter oviposition by females that subsequently visit the fruit. An hypothesis is proposed to explain the absence of oviposition-deterring pheromones in Dacus, and their presence in many other species of Tephritidae, on the basis of differences in life history and population structure.     

Tomato fruit size, maturity and alpha-tomatine content influence the performance of larvae of potato tuber moth Phthorimaea operculella (Lepidoptera: Gelechiidae)

Various physical and chemical properties of host plants influence insect larval performance and subsequent adult fitness. Tomato plants are relatively new hosts to the potato tuber moth, Phthorimaea operculella (Zeller), with the fruit being its preferred feeding site. However, it is unclear how the biochemical and physical properties of tomato fruits relate to potato tuber moth performance. Significant amounts of alpha-tomatine were detected in maturing green and ripening fruits of cherry (cv. Ceres) and processing (cv. Serio) types of tomatoes whereas none was detected in a fresh market variety (cv. Marglobe), at comparable stages. alpha-Tomatine is negatively and significantly correlated with development rate (head capsule size) of larvae reared in the fruits of the cherry and processing type tomatoes. Generally, survival, growth and development were significantly superior for larvae reared in the ripening fruits of the fresh market cultivar. At this stage, the fruits of this cultivar are also the largest. Based on these results it is concluded that fruit alpha-tomatine content, as well as fruit size and maturity, all affect performance of P. operculella larvae in the fruits of cultivated tomatoes.     

Leaf Colour as a Signal of Chemical Defence to Insect Herbivores in Wild Cabbage (Brassica oleracea)

Leaf colour has been proposed to signal levels of host defence to insect herbivores, but we lack data on herbivory, leaf colour and levels of defence for wild host populations necessary to test this hypothesis. Such a test requires measurements of leaf spectra as they would be sensed by herbivore visual systems, as well as simultaneous measurements of chemical defences and herbivore responses to leaf colour in natural host-herbivore populations. In a large-scale field survey of wild cabbage (Brassica oleracea) populations, we show that variation in leaf colour and brightness, measured according to herbivore spectral sensitivities, predicts both levels of chemical defences (glucosinolates) and abundance of specialist lepidopteran (Pieris rapae) and hemipteran (Brevicoryne brassicae) herbivores. In subsequent experiments, P. rapae larvae achieved faster growth and greater pupal mass when feeding on plants with bluer leaves, which contained lower levels of aliphatic glucosinolates. Glucosinolate-mediated effects on larval performance may thus contribute to the association between P. rapae herbivory and leaf colour observed in the field. However, preference tests found no evidence that adult butterflies selected host plants based on leaf coloration. In the field, B. brassicae abundance varied with leaf brightness but greenhouse experiments were unable to identify any effects of brightness on aphid preference or performance. Our findings suggest that although leaf colour reflects both levels of host defences and herbivore abundance in the field, the ability of herbivores to respond to colour signals may be limited, even in species where performance is correlated with leaf colour.     

桔小实蝇和番石榴实蝇对6种寄主果实的产卵选择适应性

植食性昆虫对寄主植物的选择适应性是研究昆虫和植物协同进化关系的核心内容之一。评估寄主植物对植食性昆虫种群的适合度,需要综合分析昆虫对寄主的产卵选择性和寄主对昆虫的取食适合性。以桔小实蝇和番石榴实蝇为研究对象,分别测定了这两种实蝇对6种寄主果实:番石榴、香蕉、杨桃、木瓜、甜橙、番茄的产卵选择性以及幼虫取食后对其生长发育的影响。寄主产卵选择性实验分别采用完整寄主果实直接供试产卵和块状寄主果实气味引诱产卵两种处理方式;在生长发育适应性实验中,以幼虫和蛹的存活和生长发育等相关参数作为评价指标。实验结果表明,寄主的供试方式不同,两种实蝇的选择性均有明显差异;对寄主气味选择性强的寄主更适合于两种实蝇后代的生长发育。两种实蝇对6种寄主果实的产卵选择性和后代发育适合性两者相关性不显著,与许多文献报道单一地采用发育适合性(如发育历期、存活率或蛹重等)作为评价寄主选择性的结果不一致。两种实蝇之间对6种寄主果实的产卵选择和幼虫取食适合性既具相似性也具差异性,表明这两种实蝇在寄主生态位上既有重叠性又有分化性。     

Studies on host preference and its biological effects on the red palm weevil,Rhynchophorus ferrugineus Olivier in Egypt The fruit trees

A comparative bioassay (choice or non-choice) was developed to evaluate the efficacy of different species of fruit trees as alternative host plants on the morphological and biological aspects of the red palm weevil, Rhynchophorus ferrugineus and also the latent effect of feeding the larvae on pupal and adult stages. Highly significant differences exist between averages of larval body weight of R. ferrugineus larvae fed on fruit trees. The lowest average of larval weight occurred in the larvae fed on the pieces of sour orange, mandarin followed by olive and lime trees. A small increase in body weight appeared in larvae fed on mango, fig and guava in comparison with those fed on date palm trees (control). The larval mortality reached the highest percentages (100%) by obligatory feeding of R. ferrugineus insects on olive, lime, sour orange and mandarin followed by mango (97%) and guava (96%) compared with control (0%), respectively. The latent effect of obligatory feeding on the larvae of R. ferrugineus was highly decreased during pupation (4%, 4% and 8%) when the insect fed on mango, guava and fig, compared with 100% pupation in the case of larvae fed on ornamental palm or date palm, respectively. Most pupae resulting from the treated larvae by R. ferrugineus failed in adult emergence especially those insects fed on olive, lime, sour orange and mandarin trees compared to untreated insects (95.6–96.2%), respectively. Data also proved that if an insect is forced to feed on a non-preferred host plant, this results in death of the insect often during the larval stage, a decrease of pupation or failure of adult emergence. These results led to the conclusion that mango, fig, and guava trees are believed to be vulnerable hosts for red weevil attack. The lowest percentage of food consumption appeared in those larvae fed on mandarin, sour orange, lime and olive trees compared with those fed on palm. The present results emphasise the need to test the efficiency of the compounds extracted from these host plants if it is considered as an insecticide or an alternative host protectant for management of the red palm weevil R. ferrugineus.     

Accumulation of health promoting phytochemicals in wild relatives of tomato and their contribution to in vitro antioxidant activity

Harnessing natural variation is an important aspect of modern marker assisted breeding. Traditionally breeding programmes have focused on increased yield and resistance to biotic and abiotic pressures. However, consumer demands for improved quality have lead to increased effort into the breeding of nutritional quality traits in crop plants. In the present study, health-related phytochemicals (carotenoids, tocopherols and phenolics) present in green, yellow and red wild relatives of tomato have been analyzed during fruit development and ripening. This study shows that the differences in the final colour of the fruits were due to a distinct accumulation of carotenoids mainly related to the expression of the phytoene synthase-1 gene (Psy-1). In ripe red-fruited tomatoes, the different deposition of pigments gave rise in some cases to colour differences visually discernible by the consumer. Important quantitative differences between and across taxa were noticed for the in vitro antioxidant activity (AA) of the samples.     

Olfactory responses of Grapholita molesta to different developmental stages of immature fruit of peach,plum, and apple

The Oriental fruit moth, Grapholita molesta (Busck) (Lepidoptera: Tortricidae), is known to shift its host use from stone fruits (the primary hosts) to pome fruits (secondary hosts) as the season progresses. Grapholita molesta recognizes potential hosts by semiochemicals from fruits, but few studies have addressed the olfactory responses of G. molesta to host fruits during the seasonal transition of the moth from stone to pome fruits. So, we investigated the olfactory responses of larvae and gravid females of G. molesta to different fruit stages (collected on May 31, June 11, and June 25) of immature peach, plum, and apple fruits, using both multiple and two choice tests. In the multiple choice tests, for fruits collected after June 11, larvae showed a significant preference for peach as their first choice, compared to plum or apple. The final choice rate (when larvae had stayed on the fruit for more than 1 h) of larvae was also highest for peach fruits collected on June 11 and June 25. In two choice tests (Y-tube test), the response rate and overall choice rate of larvae and mated ovipositing adult females were also highest on peaches collected on May 31 (larvae) and June 25 (adults), respectively. In conclusion, we found that both the adults and larvae of G. molesta were highly attracted to immature peach fruits in laboratory choice tests.     

Geographic variation in fruit colour is associated with contrasting seed disperser assemblages in a south‐Andean mistletoe

Fruit colour influences fruit choice by seed dispersers. The mistletoe Tristerix corymbosus (Loranthaceae) produces mature fruits of two different colours in two different biomes: yellow in the Chilean matorral and green in the temperate forest of southern South America. We conducted field surveys to establish the association between fruit colour and disperser identity throughout the entire geographical range of T. corymbosus. We selected 22 populations, eight of which were located in the Chilean matorral and 14 in the temperate forest south of the matorral. To identify the seed dispersers of the mistletoe we used direct observation, camera traps, and live‐trapping of small mammals. We also report experiments to assess fruit selection by seed dispersers based on differences in colour. The assemblages of dispersers of T. corymbosus differ between the two biomes: yellow fruits in Chilean matorral are exclusively dispersed by three bird species while green fruits in the temperate forest are exclusively dispersed by a marsupial. The differences in the assemblages of seed dispersers can be explained by differences in food‐finding strategies between the two assemblages. Green fruits in temperate forest are not easily detected by birds, while colour might not be an important cue for the marsupial because it is nocturnal and uses other senses to locate food. We propose that the association between the marsupial and the green‐fruited mistletoe constitutes an ecological fitting rather than the outcome of a co‐evolutionary process. The marsupial might have allowed the mistletoe T. corymbosus to retain green coloration in mature fruit, a condition to which it is preadapted by a slower ripening process in temperate forest populations.     

Studies on host preference and its biological effects on the Red Palm weevil,Rhynchophorus ferrugineus Olivier in Egypt B. vegetable crops

The investigation on some Egyptian vegetables preference by the Red Palm weevil (RPW), Rhynchophorus ferrugineus has been carried out under laboratory conditions to explore a suitability of alternative host plants. The choice test of insect larvae showed that Taro, Squash, Eggplant, common potato, yellow carrot, tomato and sweet pepper were accepted food, while the Table beet (Vegetable) was rejected. Non-choice feeding of R. ferrugineus larvae on vegetables affected larval body weight and increased larval mortality. Highest rate of pupation was recorded from larvae fed on C. pepo (92%) and B. vulgaris (97%). Adult emergence reached 80% and 92% on C. pepo and Sugar beet (field crop) (field crop). The external skin of sugar cane was not acceptable food for insect larvae, while injured sites and internal soft fibres were preferred as food. Therefore, vegetable fruits, stem of sugarcane are believed to be the most vulnerable hosts for red weevil attack. The alternative host plant examined can be used to monitor the dynamics of the population of RPW in order to take effective preventive measures and most powerful tool available for use in pest management.