Alcohol dehydrogenase polymorphism in the seaweed fly,Coelopa frigida

Seaweed flies (Coelopa frigida) inhabit piles of decaying seaweed on the seashore. All populations so far studied have been found to be polymorphic at the alcohol dehydrogenase locus (Adh). This article reports an attempt to identify some of the forces of natural selection that may be maintaining this polymorphism. First, the genetic determination of the rather complex isozyme system is described. Several inbred lines homozygous at the Adh locus were derived and the biochemical properties of their allozymes compared. Significant differences in both specific activities and thermal stabilities were found between ADH allozymes. A simple experiment is reported in which individuals with different Adh genotypes were cultured in competition with each other in the presence of elevated levels of ethanol. Although the presence of ethanol resulted in greater mortality, there is no evidence that it was selective with respect to the Adh genotypes. The possible relevance of these results to the maintenance of the Adh polymorphism is discussed.This work was supported by a grant to T. H. D. from the Science Research Council.     

Asymmetrical costs of sexual conflict in the seaweed fly, Coelopa frigida

Abstract.  1. Sexual conflict can play an important role in the evolution of animal life-history characteristics, including lifespan. Seaweed flies show an increase in mortality rates when exposed to brown algae. The seaweed stimulates females to oviposit and males to mount females. Females typically respond to male mounts by performing a violent rejection response.
2. Here the contribution of sexual conflict to the increase in mortality seen in the presence of seaweed was determined. The survival of single and mixed sex pairs of flies was followed in the presence and absence of seaweed.
3. The two sexes showed differential survival rates, with females living longer in the absence of seaweed. The presence of seaweed reduced survival in both sexes. In the presence of seaweed, female survival was lower when paired with a male. Over 40% of the reduction in survival in females in the presence of seaweed appears to be attributable to sexual conflict.
4. The presence of a female did not significantly affect male survival. Thus the mortality cost of being in the presence of the opposite sex and seaweed appears highly asymmetric.
5. In the presence of seaweed, female survival was lower when females were paired with small males. Small males exhibit higher levels of harassment of females, thus it is argued that pre-copulatory sexual conflict is the probable cause of the increased mortality cost to females of being in the presence of both males and seaweed.     

The effect of an inversion system and the time interval between matings on postcopulatory sexual selection in the seaweed fly, Coelopa frigida

The seaweed fly, Coelopa frigida, exhibits LMSP. A large chromosomal inversion system affects many traits including egg-to-adult viability via heterosis. Consequently, there is also considerable potential for cryptic female mate choice to operate on the basis of sperm karyotype. Here, we investigated the effect of time interval and chromosomal inversion karyotype on postcopulatory sexual selection. Homokaryotypic females were mated with a male of the same and a male of the opposite homokaryotype. The order of the matings was varied so cryptic female mate choice could operate either in concert or antagonistically with LMSP. LMSP was found when there was a 24 h time interval between matings, irrespective of the order in which the males were mated. However, when the males were mated in quick succession the order of mating was important. When LMSP and cryptic female mate choice work in concert a high level of LMSP was found. However, when the male of opposite homokaryotype mated first, then first male sperm precedence was observed. This suggests that polyandrous females might be able to bias paternity but only when matings occur in quick succession. Consequently, population density is likely to affect the operation of postcopulatory sexual selection.     

Genetic divergence and phenotypic plasticity contribute to variation in cuticular hydrocarbons in the seaweed fly Coelopa frigida

Cuticular hydrocarbons (CHCs) form the boundary between insects and their environments and often act as essential cues for species, mate, and kin recognition. This complex polygenic trait can be highly variable both among and within species, but the causes of this variation, especially the genetic basis, are largely unknown. In this study, we investigated phenotypic and genetic variation of CHCs in the seaweed fly, Coelopa frigida, and found that composition was affected by both genetic (sex and population) and environmental (larval diet) factors. We subsequently conducted behavioral trials that show CHCs are likely used as a sexual signal. We identified general shifts in CHC chemistry as well as individual compounds and found that the methylated compounds, mean chain length, proportion of alkenes, and normalized total CHCs differed between sexes and populations. We combined these data with whole genome resequencing data to examine the genetic underpinnings of these differences. We identified 11 genes related to CHC synthesis and found population‐level outlier SNPs in 5 that are concordant with phenotypic differences. Together these results reveal that the CHC composition of C. frigida is dynamic, strongly affected by the larval environment, and likely under natural and sexual selection.     

Intraspecific molecular variation in the seaweed fly Coelopa frigida consistent with behavioural distinctness of British and Swedish populations

The major aim of this study was to compare the intraspecific variation and genetic structure of the behaviourally distinct British and Swedish populations of the seaweed fly Coelopa frigida. C. frigida has been the subject of intense study into the basis of female choice. The behaviour of British females is consistent with a 'good genes' model, whereas that of the Swedish flies suggests a Fisher process, in which the difference between the former and the latter is defined by female choice increasing offspring viability in 'good genes' models. Through a study of variability in the mitochondrial cytochrome oxidase II gene from more than 600 flies, we show that there is clear differentiation at the molecular level between the two countries' populations, with an FST of > 75% and no shared haplotypes. Tajima's test reveals an excess of rare variants relative to expectation, which, if not the result of selective sweep, indicates either a population expansion or purifying selection against weakly deleterious variants. Within the two populations, substantial subpopulation differentiation is observed in the UK, where there is also evidence of isolation by distance. Swedish populations exhibit lower variability, and no evidence of isolation by distance, with the latter result possibly being related to the continuous distribution of suitable habitat. The pattern of intraspecific variation is explainable by a combination of contemporary and also historical factors. British and Swedish populations may have been descended from at least two separate founding populations during the recolonization of these areas following Pleistocene glaciations.     

Early development of the kelp fly,Coelopa frigida (Diptera). II. Morphology of cleavage and blastoderm formation

Cleavage and blastoderm formation in Coelopa frigida are extremely rapid developmental processes. In short (6–7 minutes) successive cell cycles, nuclei multiply and spread out through the egg. The movement seems to be aided by endoplasmic vesicles and cisternae which are in direct contact with the nuclear membrane. The first cells to separate from the egg plasmodium in early superficial cleavage stages are the pole cells. Precursor material from multivesicular bodies forms the pole cell membranes. The primary nuclei from the posterior pole region are removed from the blastoderm by the pole cell segregation. Blastoderm nuclei from the regions adjacent to the posterior pole migrate into the residual periplasm after pole cell segregation has been completed and constitute the blastoderm nuclei in that region of the egg. Nucleoli are not revealed during internal cleavage. They appear in pole cells shortly after their segregation. The generation time of the blastoderm nuclei increases after the twelfth cleavage. Concurrently, nucleoli form in the blastoderm nuclei and permanent cell membranes separate individual blastoderm cells. After blastoderm cells have been separated from each other, they remain in contact with the interior yolk sac by means of cytoplasmic canals. This contact is maintained at least during the early phases of blastokinesis. Observations on nuclear migration and rapid membrane formation are discussed as examples of protein assembly from subunits as an alternative to de novo protein synthesis in early stages of development.     

Quantification of ribonucleotides and uridine metabolism in ovaries of the kelp fly (Coelopa frigida,Diptera) during final stages of oogenesis

The concentrations of twelve 5′-ribonucleotides (AMP, ADP, ATP, CMP, CDP, CTP, GMP, GDP, GTP, UMP, UDP, UTP) and four ribonucleotide sugars: CDP-glucose (CDPG), UDP-glucose (UDPG), UDP-N-acetyl-glucosamine (UDPGlcNAc), and UDP-glucoronic acid (UDPGlcUA) in homogenates from Coelopa frigida ovarioles were determined by high pressure liquid chromotography. It was possible to separate the ribonucleotides from each other and from the ribonucleotide sugars in the same analysis.The amount of mono- and di-nucleotides per ovariole remains low during the final stages of oogenesis (stages 10–14). Of the mono-, di- and tri-phosphates the amounts of triphosphates per ovariole are the highest in all cases and double during this phase of follicle development. Of all nucleoside triphosphates, ATP reaches the highest concentration with an increase from 30 pmoles per ovariole to 58 pmoles between stage 10 and 14.     

Transgenerational effects of parental larval diet on offspring development time, adult body size and pathogen resistance in Drosophila melanogaster

Environmental conditions experienced by parents are increasingly recognized to affect offspring performance. We set out to investigate the effect of parental larval diet on offspring development time, adult body size and adult resistance to the bacterium Serratia marcescens in Drosophila melanogaster. Flies for the parental generation were raised on either poor or standard diet and then mated in the four possible sex-by-parental diet crosses. Females that were raised on poor food produced larger offspring than females that were raised on standard food. Furthermore, male progeny sired by fathers that were raised on poor food were larger than male progeny sired by males raised on standard food. Development times were shortest for offspring whose one parent (mother or the father) was raised on standard and the other parent on poor food and longest for offspring whose parents both were raised on poor food. No evidence for transgenerational effects of parental diet on offspring disease resistance was found. Although paternal effects have been previously demonstrated in D. melanogaster, no earlier studies have investigated male-mediated transgenerational effects of diet in this species. The results highlight the importance of not only considering the relative contribution each parental sex has on progeny performance but also the combined effects that the two sexes may have on offspring performance.     

The effects of food shortage during larval development on adult body size, body mass, physiology and developmental time in a tropical damselfly

Few studies have looked jointly at the effects of larval stressors on life history and physiology across metamorphosis, especially in tropical insects. Here we investigated how the variation of food availability during the larval stage of the tropical and territorial American rubyspot damselfly (Hetaerina americana) affects adult body size and body mass, and two physiological indicators of condition--phenoloxidase activity (an indicator of immune ability) and protein concentration. We also investigated whether larval developmental time is prolonged when food is scarce, an expected situation for tropical species whose larval time is less constrained, compared to temperate species. Second instar larvae were collected from their natural environments and reared in one of two diet regimes: (i) "rich" provided with five Artemia salina prey every day, and (ii) "poor" provided with two A. salina prey every day. In order to compare how distinct our treatments were from natural conditions, a second set of last-instar larvae were also collected and allowed to emerge. Only body size and phenoloxidase increased in the rich regime, possibly to prioritize investment on sexually selected traits (which increase mating opportunities), and immune ability, given pathogen pressure. The sexes did not differ in body size in relation to food regimes but they did differ in body mass and protein concentration; this can be explained on the basis of the energetically demanding territorial activities by males (for the case of body mass), and female allocation to egg production (for the case of protein). Finally, animals delayed larval development when food was scarce, which is coherent for tropical environments. These findings provide key insights in the role of food availability in a tropical species.     

Investigating dietary preferences in two competing dipterans, Coelopa frigida and Coelopa pilipes, using stable isotope ratios of carbon and nitrogen

Two species of seaweed fly, Coelopa frigida (Fabricius) and Coelopa pilipes (Halliday) (both Diptera: Coelopidae: Coelopini), compete for resources within deposits of marine algae washed ashore on British beaches. Previous studies report that adult flies exhibit algal‐specific behaviour that may influence interspecific interactions. It is predicted that coelopid larvae may also demonstrate algal‐specific dietary preferences. Larval dietary preferences are investigated by comparing the ratios of ¹³C/¹²C and ¹⁵N/¹⁴N in both wild flies and macroalgae to those of laboratory‐reared flies. Results showed only a small difference between the stable isotope ratios of the most abundant algae, Laminaria spp. (Laminariaceae) and Fucus spp. (Fucaceae), although there were significant differences between wild adult coelopids. This result illustrates different metabolic processes in two closely related species. The stable isotope ratios of wild‐caught coelopids were found to differ significantly from laboratory‐reared coelopids. This is either the result of red algae in the diet of natural populations or a difference in bacterial communities. We suggest that experiments with laboratory‐reared flies/specimens can greatly increase the utility of stable isotope analysis in the investigation of animal food webs, even where potential diets are isotopically similar. However, this approach is dependent on re‐creations that accurately mimic natural conditions.     

Rapid evolution of larval development time during mass-rearing in the melon fly, Bactrocera cucurbitae

A quantitative genetic analysis of rapid evolution of a life history trait has been conducted on the first 24 generations of mass-rearing in the melon fly Bactrocera cucurbitae (Coquillett) (Diptera: Tephritidae). The phenotypic variance of larval development time in each generation was divided into genetic and residual components. Mean and phenotypic coefficients of variation of larval development time decreased gradually as generations proceeded as a result of artificial selection for shorter larval period in the mass-rearing procedure. There was a trend that additive genetic coefficients of variation in larval development time decreased with generations. These changes are entirely attributed to genetic responses to laboratory selection under the mass-rearing environment because the population was maintained at a very large size so as to exclude random genetic drift and inbreeding depression, which would be other factors responsible for the observed genetic changes. The residual coefficients of variation in larval development time did not change with generations. Realized heritability of larval development time was low. The heritabilities for larval development time estimated from parent–offspring regression at generations 60 and 70, when the evolutionary plateau was asymptotically reached, were not significantly larger than 0. Received: April 22, 1999 / Accepted: September 20, 1999     

Effects of intraspecific larval competition on the development of the African buffalo fly Haematobia thirouxi potans

The effects of larval crowding on development, survival and size of the African buffalo fly, Haematobia thirouxi potans (Bezzi), were examined in small (10 or 20 g) and 1 000 g masses of cattle dung in the laboratory at 25°C. Dung was infested with fly eggs at densities which ranged from 0.05 to 32.0 eggs per g dung; in most cases 90–100% of eggs hatched. The duration of larval developmentin 10 g dung masses was extended by larval crowding at densities above 0.8 eggs per g dung and the development period with 32 eggs per g dung (11 days) was double that observed in uncrowded dung masses. Both size and survival decreased with increasing larval density, but stunting was observed at lower densities than those which reduced survival and so size was the more sensitive index of the adequacy of the larval environment. The effects of larval crowding became evident at lower larval densities in the 1 000 g dung masses than in the 10 g dung masses. There was a minimum size below which third instar larvae failed to pupariate. The probability of survival was related to pupal size and a high proportion of the smaller individuals failed to eclose.

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Effets de la competition intraspécifique sur le développement des larves, la taille des pupes et la survie d'Haematobia thirouxi potans

Résumé Les effets du surpeuplement larvaire sur le développement, la survie et la taille d'Haematobia thirouxi potans, ont été examinés sur de petites quantités de bouse (10 et 20 g) et de plus importantes (1 000 g), au laboratoire à 25°C. Les bouses étaient contaminées avec des oeufs de la mouche à des densités de 0,05 à 32 oeufs par gramme de bouse. Dans la plupart des cas le taux d'éclosion était de 90–100%. La durée du dévelopement larvaire dans 10 g de bouse a été prolongée pour les densités supérieures à 0,8 oeuf per g, et la durée développement pour 32 oeufs per g (11 jours) était le double de celle obtenue grand il n'y avait pas surpeuplement. La taille et la survie diminuent toutes les deux avec la densité larvaire, mais la miniaturisation était observée à des densités où la survie n'était pas modifiée; ainsi la taille est un indice plus sensible de l'adéquation de l'habitat larvaire. Les effets du surpeuplement larvaires deviennent clairs à des densités plus faibles avec 1 000 g de la bouse qu'avec 10 g. Il y a eu une taille minimale endessous de laquelle les larves étaient incapables de former une pupe. La probabilité d'éclosion était liée à la taille de la pupe, et une plus forte proportion de petits individus ne sont pas parvenu à éclore.

     

Effect of four chlorophytes on larval survival, development and adult body size of the mosquito Aedes aegypti

The effect of four microalgal chlorophytes, Chlorella vulgaris,Scenedesmus quadricauda, Chlorococcum sp. and Ankistrodesmus convolutus, on the survival, larval development and adultbody size of the mosquito Aedes aegypti was investigated. The percentage mortality of larvae fed with C. vulgaris, Chlorococcum sp.or S. quadricauda after six days was 100%, 84% and 88%,respectively. Delayed pupation and body size reduction of the mosquitoesfed with C. vulgaris, Chlorococcum sp. and S. quadricaudawere observed. In contrast, larvae fed with A. convolutus werebigger than those fed with normal insectory feed. The study showed thatC. vulgaris, S. quadricauda and Chlorococcum sp. have potentialto be used as larvicidal agents.     

The influence of competition between foragers on clutch size decisions in an insect parasitoid with scramble larval competition

The effect of competition between ovipositing females on theirclutch size decisions is studied in animals that lay their eggsin discrete units of larval food (hosts). In such species theeffect of competition depends on the form of the larval competitionwithin such units. In insect parasitoids, there might eitherbe contest (solitary parasitoids) or scramble competition (gregariousparasitoids) between larvae within a host For gregarious parasitoids,a decreasing clutch size with increasing competition betweenforagers is predicted. This prediction is tested in experimentsusing the parasitoid Aphaertta minuta. Parasitoids were eitherkept alone or in groups of four before the experiment, in whichthey were introduced singly in a patch containing unparasitizedhosts. Animals kept together laid on average clutches of 0.74eggs smaller than females kept alone (average clutch is 5.3),thereby confirming the prediction. Clutch size decreased withencounter number, which might be due to the adjustment of thefemale's estimate of the encounter rate with hosts. Finally,the results are compared with those reported for solitary parasitoids(that have scramble larval competition), for which it is predictedthat the clutch size will increase with increasing levels ofcompetition between females.     

The effect of temperature on the development time and brood size of diaptomus pallidus herrick

Diaptomus pallidus individuals were raised in the laboratory at three temperatures (15, 20, and 25°C) and fed an alfalfa and trout-food diet ad libitum. Data were taken on the development times of the egg, naupliar, and each copepodid stage and the brood sizes of field animals acclimated to the test conditions.The results indicated D. pallidus does not have a temperature range over which its development rate is nearly constant as earlier reported. Rather, the development rate is temperature dependent within the experimental range. Broods produced at 20°C and 25°C were significantly smaller than those produced at 15°C but not significantly different from each other.     

The genetic effects of competition in seaweed flies

In spite of abundant evidence that intra- and inter-specific competition occurs in natural communities, there is surprisingly little to suggest it is a major force promoting genetic change. This report assesses the genetic effects of competition in two species of seaweed fly, Coelopa frigida and C. pilipes. In laboratory cultures of C. frigida the relative survival of heterozygotes at the Adh locus, which was being used as a marker for the large αβ chromosomal inversion, was greater than that of homozygotes. In monocultures of C. frigida this competitive superiority was dependent on larval density. At low densities facilitation was seen, whereas at high larval densities there was competition. In mixed cultures of the two species, interspecific competition contributed to the differential mortality of C. frigida , and observations of natural populations suggested that competition may have similar effects to those described in laboratory culture. A possible mechanism involving the supply of nutritive microorganisms is proposed to underly both intra- and inter-specific competition. In seaweed flies, competition and the consequent differential mortality appear to be forces maintaining rather than reducing genetic variation.     

Covariance of adult size and development time in the parasitoid waspAphidius ervi in relation to the size of its host,Acyrthosiphon pisum

Summary Adult size (in terms of dry weight; DW) and development time (T _p ) of the solitary parasitoidAphidius ervi varied when reared in different nymphal instars of its host, apterous virginoparae of the pea aphid (Acyrthosiphon pisum). Parasitoid DW increased with an increase in the DW of the host at parasitization, from the first to the third aphid instar. Female wasps gained 1.1 times more in DW than their male counterparts in all four host classes, butT _p did not significantly differ between the sexes. Parasitoid DW was consistently more variable thanT _p . The two traits covaried positively with an increase in host size from the first to the third instar, but they varied independently in parasitoids from fourth-instar hosts. The host size (and stage) at the time of parasitization imposes constraints on the growth and development of immatureA. ervi that are reflected in the pattern of covariation between DW andT _p . When growing in aphids below a certain size threshold, parasitoids can maximize fitness by a trade-off between DW andT _p . Consequently, the assumption implicit in host-size models of parasitoid oviposition decisions — that females incur a relatively greater reduction in size (used as an index of fecundity) than males when developing in poor quality hosts — can be falsified.