Life history evolution in a globally invading tephritid: patterns of survival and reproduction in medflies from six world regions

Comparisons among populations from different localities represent an important tool in the study of evolution. Medflies have colonized many temperate and tropical areas all over the world during the last few centuries. In a common garden environment, we examined whether medfly populations obtained from six global regions [Africa (Kenya), Pacific (Hawaii), Central America (Guatemala), South America (Brazil), Extra-Mediterranean (Portugal) and Mediterranean (Greece)] have evolved different survival and reproductive schedules. Whereas females were either short-lived [life expectancy at eclosion (e₀) 48–58 days; Kenya, Hawaii and Guatemala] or long-lived (e₀ 72–76 days; Greece, Portugal and Brazil], males with one exception (Guatemala) were generally long-lived (e₀ 106–122 days). Although males universally outlived females in all populations, the longevity gender gap was highly variable (20–58 days). Lifetime fecundity rates were similar among populations. However, large differences were observed in their age-specific reproductive patterns. Short-lived populations mature at earlier ages and allocate more of their resources to reproduction early in life compared with long-lived ones. In all populations, females experienced a post-reproductive lifespan, with this segment being significantly longer in Kenyan flies. Therefore, it seems plausible that medfly populations, inhabiting ecologically diverse habitats, have evolved different life history strategies to cope with local environmental conditions.  © 2009 The Linnean Society of London, Biological Journal of the Linnean Society , 2009, 97 , 106–117.     

Hosts of Mediterranean fruit fly Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) from Broome and the Broome Peninsula, Western Australia

Abstract  In the Kimberley region of Western Australia, Mediterranean fruit fly (medfly) Ceratitis capitata (Wiedemann) (Diptera: Tephritidae) is only established at Broome. From 1998 to 2001 extensive fruit sampling was carried out at Broome during a trial investigating the eradication of medfly from the Kimberley using the sterile insect technique. Fruit was collected from 82 native and introduced plants growing in and surrounding the town of Broome and held for emergence of medfly. Eighteen plants, including the native tree Terminalia petiolaris , were found to be medfly hosts . On the basis of abundance, fruiting phenology and host suitability, eight species (kumquat Fortunella japonica , mango Magnifera indica , Barbados cherry Malpighia glabra , orange jessamine Murraya paniculata , guava Psidium guajava , Pacific almond Terminalia catapa , blackberry tree Terminalia petiolaris and yellow oleander Thevetia peruviana ) were the most important hosts ensuring medfly survival and population growth. Despite medfly being reared from T. petiolaris , it is likely that medfly can only maintain populations in areas close to human habitation, and eradication from these areas would lead to eradication from the whole Kimberley region.     

Stifled sex in sympatry: patterns of sexual incompatibility among desmognathine salamanders

Sexual incompatibility due to differences in mate-recognition systems can be an important factor in preventing hybridization and gene flow between animal populations in sympatry. We tested in the laboratory for sexual incompatibility between populations of two species of desmognathine salamanders that occur in sympatry in south-eastern Kentucky, North America, Desmognathus monticola and D. welteri (Caudata: Plethodontidae: Desmognathinae). Spermatophore deposition and insemination (two related measures of sexual success) were frequent in intraspecific encounters, but interspecific encounters never resulted in the deposition of spermatophores by males (Experiment 1). The sexual behaviour patterns of these two species are similar qualitatively, but all encounters staged between them failed to progress beyond the most preliminary of courtship interactions (Experiment 2), perhaps due to species differences in chemosensory communication. Our data on sexual incompatibility between D. monticola and D. welteri are similar to those available for other desmognathine taxa that occur in sympatry. Review of a broader data-set on patterns of sexual incompatibility both within and between species, and in allopatry and in sympatry, provides evidence for both divergence of mate-recognition systems in allopatry and enhancement of sexual incompatibility in sympatry for desmognathine salamanders. We hypothesize that diversification in allopatry may be a consequence of natural and/or sexual selection acting to promote sexual success within populations. Enhancement in sympatry may be a consequence of subsequent selection for accurate species recognition under the threat of hybridization or gene flow between species.  © 2004 The Linnean Society of London, Biological Journal of the Linnean Society , 2004, 82 , 367–375.     

Short- and long-range dispersal of medfly, Ceratitis capitata (Dipt., Tephritidae), and its invasive potential

Abstract:  Data were obtained from mark recapture trials pertaining to the dispersal of medfly, Ceratitis capitata (Dipt., Tephritidae), over both short (10–160 m) and very long distances (0.5–9.5 km) within the surveillance trapping array in Adelaide, Australia. They could be related to previously reported data sets by expressing the capture rates of each set in common terms that corrected for differences in recapture rate resulting from type of trap, season or climate. The mean capture rate at each distance from the point of release in each data set was expressed as a percentage of the real or inferred rate of that set at a distance of 100 m. The resulting distribution of dispersal distances conformed to both an inverse power model and a modified Cauchy model regardless of whether the present and previous data were combined or not. The modified Cauchy model inferred that the median distance flown was extremely short and 90% of flies displaced only 400–700 m despite the fact that a consistent trend in declining catch rates was obtained up to 9.5 km. The spread of invading propagules in quarantined zones in the first generation is likely to be limited by a decline to non-viable density within 1 km or less of the incursion point and the spread of larger infestations could be limited by the longevity of the dispersers. The results also have significance to the ability of surveillance trapping arrays to detect infestations and also to methods of distributing insects for the 'sterile insect technique'.     

The impact of nitrogen fertilization of tomato on feeding site selection and oviposition by Trialeurodes vaporariorum

The sexual calling incidence of wild and mass-reared Mediterranean fruit fly males, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae), was studied at 25 °C and a photophase of 14 h starting at 0600 h. Wild males fed on yeast hydrolysate and sugar attained sexual maturity earlier and exhibited higher calling activity than males fed on sugar alone. In both sugar and yeast hydrolysate-fed wild males, calling levels were high during the first hours of the photophase, declined at 0900–1000 h, then increased again reaching high levels between 1200 and 1500 h, and finally declined and ceased after 1600 h. Food deprivation for 8 h or more caused a significantly lower frequency of calling. Irradiated (sterile) and unirradiated mass-reared males fed on either sugar and yeast or sugar alone did not differ in the age of attaining sexual maturity (maximum levels of calling occurred about 8 days earlier than in the wild males). However, irradiated males fed on sugar called less frequently than males in the other treatments of mass-reared males. The daily rhythm of calling in protein and sugar-fed irradiated males differed somewhat from that of wild males. Calling was recorded at high levels from the beginning of the photophase until 1300 h without any decrease at 0900–1000 h. It then decreased to very low levels from 1500–1800 h and increased slightly at 1900 h. These findings are discussed in relation to the mating behavior of the fly and to the effectiveness of released sterile males in Sterile Insect Technique programs.     

The influence of urea feeding on the bacterial and archaeal community in the forestomach of collared peccary (Artiodactyla, Tayassuidae)

Aims:  This study was carried out to test whether bacterial and archaeal populations, and products of fermentation in each compartment of collared peccary stomach, vary significantly with urea feeding. Bacteria and archaeal population variation among the four stomach compartments were also compared.
Methods and Results:  Archaeal and bacterial communities in the forestomach of four individuals per treatment – peccaries fed diets with and without urea – were analysed at molecular level using PCR followed by denaturing gradient gel electrophoresis. Volatile fatty acids profiles in the three different compartments of the forestomach were also compared. The bacterial community composition varied considerably among each compartment and with urea provision, but no variation was observed between archaeal populations. Differences in bacterial communities between treatments – with and without urea – were greater than amongst stomach compartments. The acetate: propionate proportion decreased with urea provision in diet. Some differences in bacterial but not archaeal community composition were observed in each compartment of the collared peccary forestomach.
Conclusions:  There are some differences in bacterial but not archaeal populations in each compartment of collared peccary stomach. Use of urea in the diet of peccary can substantially modify the profile of volatile fatty acids released in its forestomach, but does not influence the archaeal community composition. Urea has an important effect on bacterial population DGGE profile present in the peccary's forestomach.
Significance and Impact of the Study:  These results demonstrate the ability of the collared peccary to use urea as source of nonprotein nitrogen, and confirm a hypothesis that the collared peccary has a digestive physiology more similar to ruminant than nonruminant animals.     

Ontogeny of gonadal sex development relative to growth in fathead minnow

Ovarian differentiation of fathead minnow Pimephales promelas occurred at between 10 and 25 days post‐hatch (dph)(8–11 mm fork length, L _F, and 7–12 mg), and was characterized by the presence of meiotic cells in the centre of the gonad, location of the somatic cells at the periphery of the gonad and the formation of an ovarian cavity. In contrast with the developing ovary, in the presumptive testis somatic cells were scattered throughout the gonads and this was evident from 25 dph (fish >10 mm and >11 mg). In males, at 60 dph (15–26 mm and 39–220 mg) the efferent ducts (sperm ducts) were apparent and the testis lobules started to form, but germ cells (spermatogonia) did not enter meiosis until between 90 and 120 dph. Fish of both sexes reached full sexual maturity at between 120 and 150 dph (males: 33–59 mm and 400–2895 mg; females: 24–48 mm and 160–1464 mg). Differences in body size ( L _F and mass) between males and females were only apparent when the fish were approaching full sexual maturity (120 dph).     

Comparison of the mitotic karyotypes of Ceratitis capitata, Ceratitis rosa, and Trirhithrum coffeae (Diptera: Tephritidae) by C-banding and FISH.

The sex chromosomes of the tephritid fruit fly Ceratitis capitata (Wiedemann) are heteromorphic. The male-determining region was located on the Y chromosome by deletion mapping using unbalanced offspring from several translocation strains. In addition, we showed that only 15% of the Y chromosome is required for male determination and male fertility. Based on this result, we expected to find Y-chromosomal length polymorphism in natural populations. Using fluorescence in situ hybridization with two repetitive DNA probes that label the Y chromosome, no obvious size differences were detected in seven wild-type strains and three mutant strains. As the medfly is probably of East African origin, we also analyzed two wild-type strains established recently from pupae sampled in Kenya. The Y chromosomes show a polymorphism in the hybridization pattern of a repetitive Y-specific medfly clone. However, the overall size of the Y chromosome is similar to that of the other strains. Besides C. capitata, the tephritid fruit flies Ceratitis (Pterandrus) rosa Karsch and Trirhithrum coffeae Bezzi also emerged from pupae sampled in Kenya. Their karyotype was analyzed by C-banding. Furthermore, the ribosomal genes were mapped to the sex chromosomes in these two species. Key words : Ceratitis capitata, Tephritidae, C-Banding, FISH, rDNA.     

Estimates of gene flow from rare alleles in natural populations of medfly Ceratitis capitata (Diptera: Tephritidae)

Gene flow based on the spatial distribution of rare alleles at 25 gene loci was estimated in 15 populations of Ceratitis capitata (Wiedemann) from different parts of the world. Estimates of Nm, the number of migrants exchanged per generation among populations in different regions of the world, appeared to be quite similar, ranging from 3.36 in tropical Africa to 2.94 in the New World and 2.72 in Mediterranean basin populations. This suggests that gene flow among neighbouring populations of medfly is quite extensive. The genetic differentiation in American, Mediterranean and African populations was related to major climatic differences between North and South. These differences arise mainly from five loci that showed gene frequency patterns suggestive of latitudinal clines in allele frequencies. The clinal variation was such that tropical-subtropical populations were more heterozygous than temperate populations. It was concluded that gene flow, counteracting the forces of natural selection and genetic drift, determines the extent to which geographical populations of C. capitata are differentiated.     

Effects of gamma-irradiation on midgut proteolytic activity of the mediterranean fruit fly, Ceratitis capitata (Diptera: Tephritidae)

The Mediterranean fruit fly (medfly), Ceratitis capitata (Wiedemann), is a key pest of citrus in Spain because of significant yield losses and to quarantine restrictions. Biologically based control methods, such as the Sterile Insect Technique (SIT), which relies on the sterilization by irradiation of large numbers of insects, is gaining an increasing role in the control of medfly in Mediterranean areas. However, gamma-irradiation might damage the midgut epithelium cells, causing a lowering of nutritive assimilation that can negatively affect adult performance. Irradiation effects on digestive physiology are well established for a number of insect pests, but there is no information on medfly. Our aim was to determine the effects of gamma-irradiation on C. capitata digestive protease activity. Both larvae and adults were found to use a similar proteolytic system based on aspartyl-, trypsin-, chymotrypsin-, amino peptidase-, and carboxypeptidase A- and B-like activities. Pupae of the Vienna-7 (tsl) strain were irradiated at 70 or 140 Gy, two days before emergence, and the adults fed during 5 days on sugar-protein (4:1) diets. Protease activity was measured in midgut extracts and compared with males non-irradiated reared in the same conditions. The results showed that the irradiation doses tested had no effect on the digestive proteolytic activities of medfly adults. Moreover, the longevity of irradiated medflies at the highest dose (140 Gy) was similar to that of controls.     

Parasitism on medfly by Diachasmimorpha longicaudata (Ashmead) (Hymenoptera: Braconidae) in different guava cultivars

The parasitism efficiency of the Braconidae wasp, Diachasmimorpha longicaudata (Ashmead), was checked on four guava cultivars (Paluma, Sassaoca, Pedro Sato and Kumagai) infested with larvae of medfly, Ceratitis capitata (Wiedemann) (Diptera: Tephritidae). Five blocks of eight fruits, each with two fruits of each cultivar, were put inside C. capitata adult cages, during 2h for oviposition, and a week later, when the larvae inside guavas were developed, the fruits were exposed to parasitoids for 24h. The mean fruit weight, larvae mortality, number of pupae, percentage of medfly and parasitoid emergence were evaluated. There was not statistical difference among cultivars to weight, larvae mortality, number of pupae e emergence of medfly. The percentage of parasitism was higher in Pedro Sato cultivar (19.8%) compared with Kumagai cultivar (2.9%), but it was statistically similar to the other cultivars.     

Intron size polymorphism of the Adh1 gene parallels the worldwide colonization history of the Mediterranean fruit fly, Ceratitis capitata

The exon-primed intron-crossing (EPIC) PCR technique was used to analyse the size variation at the first intron of the Ceratitis capitata Adh ₁ gene. A total of 27 samples from 16 natural populations was analysed from five geographical regions in the species range: Africa, Mediterranean Basin, Latin America, Hawaii and Australia. The Adh ₁ first intron varies extensively in length with at least 18 size variants ranging from 1400 bp to 3450 bp. These variants can be grouped into four distinct size categories: short, medium, long and very long. The majority of these variants are present only in the African populations. Only a subset of the ancestral variants appear to have succeeded in migrating from Africa during the medfly colonization process. The medfly population structure inferred from the intron size polymorphism is congruent with that observed from the analysis of allozyme variation. The geographical dispersal of the medfly from its source area is associated with a gradual and great reduction in intron variability which parallels the trend of decreasing variability evaluated at 26 biochemical loci. The intron phylogenetic tree is in agreement with allozyme data in portraying the dynamic population history of the medfly. Stochastic evolutionary forces such as drift, bottleneck effects and migration seem to have played the major roles in the dispersion pattern of Adh ₁ intron variation during the colonization of the medfly.     

Camouflage and colour change: antipredator responses to bird and snake predators across multiple populations in a dwarf chameleon

Potential prey are often exposed to multiple predators that vary in their foraging tactics and ability to detect prey. For animals that rely on crypsis to avoid predators, one solution is to alter their behaviour or appearance to maximize crypsis in ways that are specific to different types of predator. We tested whether dwarf chameleons ( Bradypodion transvaalense ) showed different behavioural responses, including colour change, towards multiple predators (bird and snake models) that detect and capture prey in different ways, and whether these antipredator responses varied geographically. Chameleons consistently used the same body postures (lateral compression and flipping to the opposite side of the branch) and displayed similar chromatic (colour) contrast against the natural background in response to both predator types. However, they became significantly more achromatically contrasting (brighter) in the presence of the snake compared to the bird. This relative difference in achromatic contrast towards the two types of predator was consistent among populations. There were also significant differences in both absolute achromatic and chromatic contrast among populations despite very similar light environment, background coloration and habitat structure. Our results highlight facultative crypsis as one type of flexible antipredator tactic and emphasize the importance of visual ecology in understanding prey–predator interactions.  © 2006 The Linnean Society of London, Biological Journal of the Linnean Society , 2006, 88 , 437–446.     

Sex-specific niche partitioning and sexual size dimorphism in Australian pythons (Morelia spilota imbricata)

Sexual dimorphism is usually interpreted in terms of reproductive adaptations, but the degree of sex divergence also may be affected by sex-based niche partitioning. In gape-limited animals like snakes, the degree of sexual dimorphism in body size (SSD) or relative head size can determine the size spectrum of ingestible prey for each sex. Our studies of one mainland and four insular Western Australian populations of carpet pythons ( Morelia spilota ) reveal remarkable geographical variation in SSD, associated with differences in prey resources available to the snakes. In all five populations, females grew larger than males and had larger heads relative to body length. However, the populations differed in mean body sizes and relative head sizes, as well as in the degree of sexual dimorphism in these traits. Adult males and females also diverged strongly in dietary composition: males consumed small prey (lizards, mice and small birds), while females took larger mammals such as possums and wallabies. Geographic differences in the availability of large mammalian prey were linked to differences in mean adult body sizes of females (the larger sex) and thus contributed to sex-based resource partitioning. For example, in one population adult male snakes ate mice and adult females ate wallabies; in another, birds and lizards were important prey types for both sexes. Thus, the high degree of geographical variation among python populations in sexually dimorphic aspects of body size and shape plausibly results from geographical variation in prey availability.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 77 , 113–125.     

Adhesive powder uptake and transfer by Mediterranean fruit flies, Ceratitis capitata (Dipt., Tephritidae)

Abstract:  Entostat^TM is an electrostatically charged wax powder that is used as a carrier particle in novel delivery systems for contaminating target insect pests with insecticides, biologicals or pheromones. Here, the adhesion of two forms of Entostat to the Mediterranean fruit fly (medfly) Ceratitis capitata (Wiedemann) was examined, and the adhesion of Entostat to live and dead medflies was compared. From controlled contaminations of medflies, it was shown that live medflies acquired larger quantities of Entostat than dead medflies, which could be due to the electrostatic charge shown to be carried by live insects. Air-milled Entostat (7.59  μ m mean diameter) adhered in larger quantities to medflies than pestle and mortar-ground Entostat (9.17  μ m mean diameter). Exposing medflies to different quantities of Entostat affected the initially adhering quantity but did not alter the proportion of powder retained over time. Medfly males contaminated with air-milled Entostat were shown to transfer small quantities to females during mating. This documentation of secondary powder transfer underscores the potential for using slow-acting killing agents on the basis of this delivery system.     

High sexual signalling rates of young individuals predict extended life span in male Mediterranean fruit flies

In a laboratory study, we monitored the lifetime sexual signalling (advertisement) of wild male Mediterranean fruit flies, and we tested the hypothesis that high lifetime intensity of sexual signalling indicates high survival probabilities. Almost all males exhibited signalling and individual signalling rates were highly variable from the beginning of the adults maturity and throughout their life span (average life span 62.3 days). Sexual signalling rates after day 10 (peak maturity) were consistently high until about 1 week before death. There was a positive relationship between daily signalling rates and life span, and an increase in signalling level by one unit over all times was associated with an approximately 50% decrease in mortality rate. Signalling rates early in adult life (day 6–20) were higher in the longest-lived than in the shortest-lived flies. These results support the hypothesis that intense sexual signalling indicates longer life span. We discuss the importance of age-specific behavioural studies for understanding the evolution of male life histories.     

Differential Migration by Sex in the Great Bustard: Possible Consequences of an Extreme Sexual Size Dimorphism

We explored migration patterns in Great bustards ( Otis tarda ), a species that shows strong sexual selection and the most extreme sexual size dimorphism among birds. The aim was to explain differential migration, examining whether Great bustards fulfil the main predictions of bird migration theory hypotheses and sexual segregation theory hypotheses. We radio-tracked the seasonal movements of 65 males and 68 females in central Spain. We found four main sexual differences. First, the proportion of migratory males was higher than that of migratory females (86% vs. 51%). Second, males abandoned the leks immediately after the mating season (late May to early Jun.), whereas females remained there for another 3–7 mo. Third, 54% of the migratory males used two different post-breeding areas, the first located northwards at 82 km from the breeding sites in summer, and the second southwards at 50 km in autumn–winter. Migratory females used only one area in autumn–winter which coincided geographically with that of males. And fourth, males returned to the breeding areas earlier (between Sep. and Mar.) than females (between Jan. and Apr.). These results show that the Great bustard is a differential migrant by sex in central Spain and support the weather sensitivity hypothesis (males were less tolerant to summer heat) and the specialization hypothesis (exclusive maternal care of the brood by females). Sexual differences in migratory behaviour are probably ultimately determined by the strong sexual selection in this species.     

Population-specific demography and invasion potential in medfly

Biological invasions are constantly gaining recognition as a significant component of global change. The Mediterranean fruit fly (medfly) constitutes an ideal model species for the study of biological invasions due to its (1) almost cosmopolitan geographic distribution, (2) huge economic importance, and (3) well-documented invasion history. Under a common garden experimental set up, we tested the hypothesis that medfly populations obtained from six global regions [Africa (Kenya), Pacific (Hawaii), Central America (Guatemala), South America (Brazil), Extra-Mediterranean (Portugal), and Mediterranean (Greece)] have diverged in important immature life-history traits such as preadult survival and developmental times. We also tested the hypothesis that medfly populations from the above regions exhibit different population growth rates. For this purpose, data on the life history of immatures were combined with adult survival and reproduction data derived from an earlier study in order to calculate population parameters for the above six populations. Our results clearly show that medfly populations worldwide exhibit significant differences in preadult survival, developmental rates of immatures and important population parameters such as the intrinsic rate of increase. Therefore, geographically isolated medfly populations may share different invasion potential, since population growth rates could influence basic population processes that operate mostly during the last two stages of an invasion event, such as establishment and spread. Our findings provide valuable information for designing population suppression measures and managing invasiveness of medfly populations worldwide.     

Bioinvasions of the medfly Ceratitis capitata: source estimation using DNA sequences at multiple intron loci.

The Mediterranean fruit fly, Ceratitis capitata, is a devastating agricultural pest that threatens to become established in vulnerable areas such as California and Florida. Considerable controversy surrounds the status of Californian medfly infestations: Do they represent repeated introductions or the persistence of a resident population? Attempts to resolve this question using traditional population genetic markers and statistical methods are problematic because the most likely source populations in Latin America were themselves only recently colonized and are genetically very similar. Here, significant population structure among several New World medfly populations is demonstrated through the analysis of DNA sequence variation at four intron loci. Surprisingly, in these newly founded populations, estimates of population structure increase when measures of subdivision take into account the relatedness of alleles as well as their frequency. A nonequilibrium, likelihood-based statistical test that utilizes multilocus genotypes suggests that the sole medfly captured in California during 1996 was introduced from Latin America and was less likely to be a remnant of an ancestral Californian population. Many bioinvasions are hierarchical in nature, consisting of several sequential or overlapping invasion events, the totality of which can be termed a metainvasion. Phylogenetic data from multilocus DNA sequences will be vital to understanding the evolutionary and ecological processes that underlie metainvasions and to resolving their constituent levels.     

Gustatory adaptation affects sexual maturation in male German cockroaches,Blattella germanica

Adaptations to hazardous environmental factors are essential for survival, although they may be maladaptive in conditions where the hazard is absent. In German cockroach (Blattella germanica L.) populations, glucose aversion has evolved rapidly in response to glucose‐containing insecticidal baits, but little is known about the consequences of this behaviour in the absence of bait. In the present study, glucose‐averse (GA) and wild‐type (WT) male German cockroaches are restricted to a range of nutritionally defined diets containing either glucose or fructose as the sole carbohydrate source, and time to first expression of courtship is measured by stimulating the male antennae daily with isolated antennae from receptive, 6‐day‐old females. Glucose‐averse males that are restricted to glucose‐containing diets mature their courtship responses significantly later than GA males restricted to fructose‐containing diets, whereas there is no difference in maturation of courtship responses between GA males restricted to fructose‐containing diets and WT males restricted to diets containing either sugar type. Glucose‐averse males furthermore respond later to GA female antennae than to WT female antennae, all from 6‐day‐old females. This suggests that GA females are less sexually stimulating, and the results are also consistent with earlier findings showing that GA females contain less developed oocytes than WT females at this age. These findings demonstrate that an adaptive gustatory mutation conferring protection from a toxin may have comparatively detrimental effects under conditions where the toxin has vanished, both by delaying female sexual maturation and signalling and by delaying male sexual maturation and courtship under conditions where glucose is a major energy source.