Accumulation of B-chromosomes by preferential segregation in females of the grasshopper Melanoplus femur-rubrum

In a population of Melanoplus femur-rubrum 13.9% of the males and 9.1% of the females sampled possessed a metacentric B chromosome (B). In crosses of females with one B (1 B females) and 0 B males 0.82 ± 0.05 of the offspring received the B. The value expected from Mendelian segregation is 0.5. In crosses of 1 B males and 0 B females the frequency of offspring receiving the B was 0.53 ± 0.02. The B is heterochromatic during prophase I of spermatogenesis but is euchromatic during prophase I of oogenesis. The observation that in 1 B females only one B was present in metaphase I of oogenesis suggested strongly that the high rate of transmission of the B by the females resulted from preferential segregation of the B into the secondary oocyte. The maintenance of the B in the species in discussed.Supported by Grant GB 23665 from the National Science Foundation, Washington, D.C.     

Mitotic instability leading to an accumulation of B-chromosomes in grasshoppers

The study of mitotically unstable B-chromosomes (supernumeraries) of two grasshopper species confirmed a suggestion made earlier (Nur, 1963) that the instability should always be associated with a tendency of the B's to increase in frequency. Among 780 Camnula pellucida (Scudder) males from California, 105 had B's. In the testes of these males the number of B's varied from follicle to follicle and ranged between 0 and 4. Because of this variation, the number with which each male started to develop could not be determined. However, the relatively low frequency of males with B's and the regular meiotic behavior of the latter suggested that most of the 105 males started with a single B. Cytological analysis of the cells of the gastric caeca of 31 males whose testes contained B's confirmed this suggestion by showing that only one male had two B's in these cells; all the rest had one. In the testes of the 74 other males the mean number of B's ranged from 0.89–2.50, but only two males had means higher than 2.00. The observed ratio of one male with two B's to 30 with one, suggested that only the two males with the highest means started to develop with two B's and that the other 72 males all started with one. Since the mean for the 72 males was 1.37 B's per male, it was concluded that during the development of the testes of these males the mean increased by 37%. The males with B's had fewer follicles in their testes and apparently had also a lower frequency of normal sperm. — The analysis of the testes of Locusta migratoria L. males from Japan gave results which agreed with those from C. pellucida.Supported by grants GB 1585 and GB 6745 from the National Science Foundation, Washington, D.C.     

Cytogenetic studies on natural populations of grasshoppers with special reference to B-chromosomes

In Atractomorpha bedeli the frequencies of males with B-chromosomes in the males from the Tofuro-ato population were 21, 30, 32, 40, and 26%, respectively, over five years. Numbers of B's (0 to 7) in the primary spermatocytes were usually constant within the follicles but were variable among the follicles within the individuals. In males from the Okazyo-ato population, 35% contained B's in their primary spermatocytes. The cells of the gastric caeca were observed in 8 males with B's in the primary spermatocytes but no cells of the gastric caeca contained B's. Cells of ovariole walls and those of gastric caeca were analysed in 80 females from the same population; none of them contained B's. These facts indicate that B's are present in the germ line but absent in the somatic line. The difference between the germ line and the somatic one was explained on the basis of elimination of B's from the somatic cells. The inter-follicular variation in number of B's was ascribed to mitotic non-disjunction in the germ line prior to differentiation of the follicles.Contribution No. 86 from the Institute of Biology, Oita University.     

Offspring sex ratio produced by female guppies in the wild correlates with sexual ornaments of their sons

The attractiveness hypothesis predicts that females produce broods with male-biased sex ratios when they mate with attractive males. This hypothesis presumes that sons in broods with male-biased sex ratios sired by attractive males have high reproductive success, whereas the reproductive success of daughters is relatively constant, regardless of the attractiveness of their sires. However, there is little direct evidence for this assumption. We have examined the relationships between offspring sex ratios and (1) sexual ornamentation of sons and (2) body size of daughters in broods from wild female guppies Poecilia reticulata. Wild pregnant females were collected and allowed to give birth in the laboratory. Body size and sexual ornamentation of offspring were measured at maturity. Our analysis revealed a significant positive correlation between offspring sex ratios (the proportion of sons per brood) and the total length as well as the area of orange spots of sons, two attributes that influence female mating preferences in guppies. The sex ratio was not associated with the body size of daughters. These results suggest that by performing adaptive sex allocation according to the expected reproductive success of sons and daughters, female guppies can enhance the overall fitness of their offspring.     

Intralocus sexual conflict,adaptive sex allocation,and the heritability of fitness

Intralocus sexual conflict arises when selection favours alternative fitness optima in males and females. Unresolved conflict can create negative between‐sex genetic correlations for fitness, such that high‐fitness parents produce high‐fitness progeny of their same sex, but low‐fitness progeny of the opposite sex. This cost of sexual conflict could be mitigated if high‐fitness parents bias sex allocation to produce more offspring of their same sex. Previous studies of the brown anole lizard (Anolis sagrei) show that viability selection on body size is sexually antagonistic, favouring large males and smaller females. However, sexual conflict over body size may be partially mitigated by adaptive sex allocation: large males sire more sons than daughters, whereas small males sire more daughters than sons. We explored the evolutionary implications of these phenomena by assessing the additive genetic (co)variance of fitness within and between sexes in a wild population. We measured two components of fitness: viability of adults over the breeding season, and the number of their progeny that survived to sexual maturity, which includes components of parental reproductive success and offspring viability (RS^V). Viability of parents was not correlated with adult viability of their sons or daughters. RS^V was positively correlated between sires and their offspring, but not between dams and their offspring. Neither component of fitness was significantly heritable, and neither exhibited negative between‐sex genetic correlations that would indicate unresolved sexual conflict. Rather, our results are more consistent with predictions regarding adaptive sex allocation in that, as the number of sons produced by a sire increased, the adult viability of his male progeny increased.     

Replication of euchromatin and heterochromatin in a mealybug

Asynchronous DNA replication of euchromatic (E) and heterochromatic (H) chromosomes and heterochromatic B chromosomes (B) were studied in the mealybug, Pseudococcus obscurus Essig (Homoptera: Coccoidea). The study was carried out on mycetocytes of adult females and on spermatocytes of mid-second instar males by employing tritiated thymidine labeling and autoradiography. In the mycetocytes the incorporation of the labeled thymidine began and ended later in the B's than in the E chromosomes. The S period was found to be about 21 hours. The DNA replication of the E chromosomes occupied about 86% of the S period and that of the B's 33%; during 18% of the mid-S period the replication of the two types of chromosomes overlapped. In the meiotic S period of the spermatocytes, the DNA of the E chromosomes started to replicate earlier than that of the H chromosomes and the B's, but the replication of the E chromosomes, the H chromosomes, and the B's overlapped. The H chromosomes completed their replication much later than the E chromosomes and slightly later than the B's.Supported by grants GB 1585 and GB 6745 to Dr. Uzi Nur from the National Science Foundation, Washington, D. C.Part of a thesis submitted to the University of Rochester in partial fulfillment of the requirements for the degree of Doctor of Philosophy.     

Related male Drosophila melanogaster reared together as larvae fight less and sire longer lived daughters

Competition over access to reproductive opportunities can lead males to harm females. However, recent work has shown that, in Drosophila melanogaster, male competition and male harm of females are both reduced under conditions simulating male-specific population viscosity (i.e., in groups where males are related and reared with each other as larvae). Here, we seek to replicate these findings and investigate whether male population viscosity can have repercussions for the fitness of offspring in the next generation. We show that groups of unrelated-unfamiliar (i.e., unrelated individuals raised apart) males fight more intensely than groups of related-familiar males (i.e., full siblings raised together as larvae), supporting previous findings, and that exposure to a female is required to trigger these differential patterns of male–male competition. Importantly, we show that differences in male–male competition can be associated with transgenerational effects: the daughters of females exposed to unrelated-unfamiliar males suffered higher mortality than the daughters of females exposed to related-familiar males. Collectively, these results suggest that population structure (i.e., variation in the relatedness and/or larval familiarity of local male groups) can modulate male–male competition with important transgenerational consequences.     

Does egg carotenoid improve larval quality in Arctic charr (Salvelinus alpinus)?

Females in mutually ornamented species are often less conspicuously ornamented than their male conspecifics. It has been hypothesized that offspring quality may decrease if females invest more resources into ornaments at the expense of resources in eggs. An experiment was carried out to test whether natural variation in carotenoid in the eggs from a wild population of Arctic charr (Salvelinus alpinus) was associated with survival and growth of their offspring until hatching. Wild Arctic charr were caught at a spawning ground during the spawning period. Eggs from two different females, one female with yellowish carotenoid‐rich eggs and one with paler eggs, were fertilized by sperm from the same male. This was repeated until gametes were collected from 42 females and 21 males, giving a total of 21 groups. After fertilization, the zygotes from each of the two females were reared in four replicated groups. These 168 groups were reared separately until hatching when the surviving larvae were counted and their body length measured. For the two response variables survival and body length at hatching, no effect was demonstrated of any of the predictors (i) amount of carotenoid in the unfertilized eggs, (ii) the mothers'' body condition, or (iii) ornament intensity of their red carotenoid‐based abdominal ornament. Thus, this study gives no support for the hypothesis that females investing less carotenoid into their eggs suffer from decreased offspring quality until hatching. This lack of association between female ornament intensity and their fitness is not as expected if female ornaments evolved due to direct sexual selection from males on the more ornamented females (“direct selection hypothesis”).     

POLYANDRY INCREASES OFFSPRING FECUNDITY IN THE BULB MITE

Abstract The common occurrence of polyandry continues to puzzle evolutionary biologists, as female reproductive success is thought to be limited mostly by her fecundity. Here we test whether females of the bulb mite, a species in which the females are highly promiscuous, benefit from polyandry in terms of increased fitness of their progeny. Females were given opportunity to mate with either one or six males, but the experiment was designed to allow the same number of matings per female in both groups, that is, irrespective of the number of males. We found that daughters of females mated to six males had significantly higher fecundity than daughters of females mated to one male, whereas other fitness components of progeny (male virility and longevity of both sexes) were not affected. These findings appear to support hypotheses proposing that multi-male mating enables females to exercise postcopulatory mate-choice (direct or indirect, via sperm competition) and thus accrue genetic benefits.     

Fitness effects of a selfish gene (the Mus t complex) are revealed in an ecological context

In wild house mice, genes linked to the t transmission distortion complex cause meiotic drive by sabotaging wild-type gametes. The t complex is consequently inherited at frequencies higher than 90%. Yet, for unclear reasons, in wild mouse populations this selfish DNA is found at frequencies much lower than expected. Here, we examine selection on the t complex in 10 seminatural populations of wild mice based on data from 234 founders and nearly 2000 progeny. Eight of the 10 populations decreased in t frequency over one generation, and the overall frequency of t haplotypes across all 10 populations was 48.5% below expectations based on transmission distortion and 34.3% below Mendelian (or Hardy-Weinberg) expectations. Behavioral and reproductive data were collected for 10 months for each population, and microsatellite genotyping was performed on seven of the populations to determine parentage. These combined data show t-associated fitness declines in both males and females. This is the first study to show evidence for a reduction in the ability of +/t males to maintain territories. Because females tend to mate with dominant males, impairment of territorial success can explain much of the selection against t observed in our populations. In nature, selection against heterozygote carriers of the t complex helps solve the puzzlingly low t frequencies found in wild populations. This ecological approach for determining fitness consequences of genetic variants has broad application for the discovery of gene function in general.     

Do assortative mating and immigrant inviability help maintain population genetic structuring of an herbivore on a crop and a wild relative?

Population genetic structuring is common among herbivorous insects and frequently is associated with divergent host plants, such as crops and their wild relatives. Previous studies showed population genetic structuring in corn leafhopper Dulbulus maidis in Mexico, such that the species consists of two sympatric, host plant-associated populations: an abundant and widespread "pestiferous” population on maize (Zea mays mays), and a small and localized "wild" population on perennial teosinte (Zea diploperennis). a maize wild relative with a limited distribution. This study addressed whether assortative mating and immigrant inviability mediate genetic structuring of corn leafliopper by comparing the mating and reproductive successes of pestiferous and wild females that colonize their nonassociated host plants against the successes of females colonizing their associated host plants. Assortative mating was assessed by comparing mating frequencies and premating and mating times among females of each population on each host plant: immigrant inviability was assessed by comparing, across two generations, the fecundity, survival, development time, sex ratio, and population growth rate among leafhopper populations and host plants. Our results showed that on maize, and compared to resident, pestiferous females, wild females were more likely to mate, and greater proportions of their offspring survived to adult stage and were daughters;consequently, the per-generation population growth rate on maize was greater for immigrant, wild leafhoppers compared to resident, pestiferous leafhoppers. Our results suggested that wild leafhoppers emigrating to maize have a fitness advantage over resident, pestiferous leafhoppers, while immigrant pestiferous and resident wild leafhoppers on teosinte have similar fitnesses.     

ASSOCIATION BETWEEN SEX RATIO DISTORTION AND SEXUALLY ANTAGONISTIC FITNESS CONSEQUENCES OF FEMALE CHOICE

Genetic variation can be beneficial to one sex yet harmful when expressed in the other—a condition referred to as sexual antagonism. Because X chromosomes are transmitted from fathers to daughters, and sexually antagonistic fitness variation is predicted to often be X-linked, mates of relatively low-fitness males might produce high-fitness daughters whereas mates of high-fitness males produce low-fitness daughters. Such fitness consequences have been predicted to influence the evolution of female mating biases and the offspring sex ratio. Females might evolve to prefer mates that provide good genes for daughters or might adjust offspring sex ratios in favor of the sex with the highest relative fitness. We test these possibilities in a laboratory-adapted population of Drosophila melanogaster , and find that females preferentially mate with males carrying genes that are deleterious for daughters. Preferred males produce equal numbers of sons and daughters, whereas unpreferred males produce female-biased sex ratios. As a consequence, mean offspring fitness of unpreferred males is higher than offspring fitness of preferred males. This observation has several interesting implications for sexual selection and the maintenance of population genetic variation for fitness.     

Host instar, body size and fitness in the koinobiotic parasitoid Aphidius nigripes

In aphidiine parasitoids, resources for growth and adult body size increase with host instar used by ovipositing females, but the fitness consequences of body size on fitness are poorly documented. We compared the fitness of male and female A. nigripesadults that varied in size as a consequence of developing in different instars of their host Macrosiphum euphorbiae. When reproductive fitness was measured without considering time, female wasps from small and large hosts performed similarly, contributing 125–175 foundresses plus 100–180 sons to the next generation. However, when expressed as the innate capacity for increase (r _m), female fitness correlated with host-induced variation of wasp size, indicating that micropopulations initiated by large wasps would increase faster. In a wind-tunnel, a sex pheromone plume from large female wasps induced more males to fly upwind when released at a distance of 50 cm downwind than small females, indicating that large females were sexually more attractive. With respect to male body size effects on fitness, large individuals performed similar to small ones, whether fitness was measured by lifetime mating frequency, fertile inseminations, or proportion of daughters among progeny born from their mates. When young naive males of unequal size were directly competing for mating with a virgin female, small and large males had equal mating success, and large individuals were no more successful than small ones at displacing a competitor already positioned on a receptive female. In a wind-tunnel test where males were scored on their ability to reach a female pheromone source, small and large males were equally affected by wind speed but reached the source located 50 cm downwind in equal proportions, suggesting similar capacity for finding mates by flying upwind. Our results indicate that despite host resources not being fixed at the time of attack for the koinobiont A. nigripes, fitness consequences of resource limitation by the mother may be perceived to be greater for daughters than sons, which would explain male-biased sex ratio in early-instar hosts.     

Pest control by the release of insects carrying a female-killing allele on multiple loci

With recent advances in genetics, many new strategies for pest control have become feasible. This is the second article in which we model new techniques for pest control based on the mass release of genetically modified insects. In this article we model the release of insects carrying a dominant and redundant female killing or sterilizing (FK) allele on multiple genetic loci. If such insects are released into a target population, the FK allele can become widely spread in the population through the males while reducing the population each generation by killing females. We allow the number of loci used to vary from 1 to 20. We also allow the FK allele to carry a fitness cost in males due to the gene insertions. Using a model, we explore the effectiveness and optimal strategies for such releases. In the most ideal circumstances (no density-dependence and released insects equal in fitness to wild ones), FK releases are several orders of magnitude more effective than equal sized sterile male releases. For example, a single release of 19 FK-bearing males for every two wild males, with the released males carrying the FK allele on 10 loci, reduces the target population to 0.002% of no-release size. An equal sized sterile release reduces the target population to 5% of no-release size. We also show how the effectiveness of the technique decreases as the fitness cost of the FK alleles in males increases. For example, the above mentioned release reduces the target population to 0.7% of no-release size if each FK allele carries a fitness cost in males of 5%. Adding a simple model for density-dependence and assuming that each of the released males carries the FK allele on six loci, we show that the release size necessary to reduce the target population to 1/100 of no-release size in 10 generations of releases varies from 0.44:1 to 4:1 (depending on parameter values). We also calculate the optimal number of loci on which to put the FK allele under various circumstances.     

Male-biased sex ratio in litters of Alpine marmots supports the helper repayment hypothesis

In a French population of Alpine marmots (Marmota marmota),the sex ratio at weaning was biased in favor of males. Thisbias also seemed to exist at birth. Under Fisher's equal allocationprinciple, this means that daughters should be more costlyto produce than sons. Because the Alpine marmot can be considereda cooperative breeding species, we investigated whether thedifferential cost between sons and daughters may be explainedby the helper repayment hypothesis. The Alpine marmot usessocial thermoregulation during hibernation, allowing juvenilesto better survive over winter. In the study population, juvenilesurvival during winter increased with group size. More precisely,juvenile survival during winter increased with the number andwith the proportion of subordinate males in the hibernatinggroup, but juvenile survival did not depend on the number of subordinate females. As our results did not support alternativehypotheses to explain the observed bias in sex ratio amongoffspring at emergence, we conclude that the helper repaymenthypothesis is the best candidate to explain the observed offspringsex ratio bias in Alpine marmots. By participating in socialthermoregulation, subordinate males may repay part of the investment they received from their parents and thus become less costlyto produce. We suggest that only subordinate males helped becausethey may gain direct fitness benefits, whereas subordinatefemales may only expect indirect fitness benefits from helping.Finally, the offspring sex ratio per individual parent wasmale biased, but mothers adjusted the size and the sex compositionof their litters according to their phenotypic condition asexpected from the Trivers-Willard hypothesis.     

A mitotically unstable supernumerary chromosome with an accumulation mechanism in a grasshopper

Summary A population of the grasshopper Calliptamus palaestinensis Bdhr. from near Jerusalem, Israel was analyzed cytologically. Twenty-two percent of the males had extra heterochromatic chromosome segments or supernumerary chromosomes. The extra segments were present in either a heterozygous or a homozygous condition and in frequencies similar to those expected according to the Hardy-Weinberg law.The supernumeraries were heteropycnotic in the first meiotic division but paired and segregated regularly. In almost all the males with supernumeraries the number of supernumeraries varied between follicles in the testes but was constant within each follicle. Only 12.3% of the males had supernumeraries, but among those with supernumeraries the majority had means higher than 1.5. A calculation based on the frequency of the supernumeraries in the zygotes which are expected in the next generation indicated that of the 31 males with supernumeraries, probably not more than 3 males originated from zygotes with more than a single supernumerary. The variation in the number of supernumeraries and the high means probably resulted from preferential non-disjunction of the supernumeraries in the early mitoses prior to the differentiation of the follicles.Mitotic non-disjunction of supernumeraries as the cause of intra-individual variation is considered. It is suggested that variation due to non-disjunction would be maintained only in association with accumulation mechanisms.Postdoctoral Trainee in Biology, U. S. Public Health Service, 1962–1963.     

Maintenance of a "Parasitic" B Chromosome in the Grasshopper MELANOPLUS FEMUR-RUBRUM

About 10-15% of the males and females of the grasshopper Melanoplus femur-rubrum collected near Rochester, New York, possessed a supernumerary B chromosome. The frequency of the B chromosome remained fairly constant during the years 1971-1974. The B chromosome was shown previously to be transmitted at a rate of about 0.5 and 0.8 by 1B males and females, respectively. This study was designed to determine the forces preventing the B chromosome from increasing in frequency due to the high rate of transmission by the females. Eighty inseminated females collected in the wild were analyzed cytologically together with their embryos (10-20 per female). Ten of the 80 females had a B chromosome, and they transmitted it at a rate of about 0.75. Among the 983 embryos analyzed, 0.141 had one B, 0.007 had two, and the mean number of B chromosomes per embryo was 0.155. The frequency of the B chromosome in the sperm pool (0.061) was consistent with a 0.5 rate of transmission. Individuals with two B chromosomes apparently have low viability, because about six were expected, but none was found among 851 adult males and females examined. The data suggest that the viability of the 1B individuals was only about 0.86 that of the OB individuals. There was no evidence that the B chromosome increased the fecundity of either the 1B males or females. It was concluded, therefore, that the B chromosome reduced the fitness of all the individuals carrying it and was thus "parasitic," and that it was maintained in the population only because of its high transmission rate. The maintenance of other B chromosomes with high transmission rates is reviewed.     

Natural variation in female mating frequency in a polyandrous butterfly: effects of size and age

In the polyandrous gift-giving butterfly Pieris napi, females mature at a smaller size than males under poor food conditions, so it has been suggested that females can compensate for their smaller size through nuptial feeding. We tested this hypothesis by assessing female polyandry in relation to female size in a study of a wild bivoltine population and in a laboratory experiment. Contrary to expectation, larger females had a higher mating frequency. In the wild population female polyandry was positively correlated with size and in the laboratory experiment larger females were also more polyandrous. Hence, smaller females cannot compensate for their size by increasing their mating frequency, perhaps because the rate of spermatophore breakdown sets the limit for polyandry and larger females are able to break down the spermatophore faster. In addition, the lifetime number of matings for wild females varied between one and five. As previous studies indicate that female fitness in P. napi appears to increase monotonically with number of mates as a result of the increase in spermatophore material received, it is surprising that 12% of the wild females in the oldest age classes had mated only once and that another 35-40% had mated only twice. This apparently maladaptive behaviour is discussed in terms of sexually antagonistic coevolution and whether environmental conditions influence optimal mating frequency. Copyright 2002 The Association for the Study of Animal Behaviour. Published by Elsevier Science Ltd. All rights reserved.     

Association of polyandry and sex-ratio drive prevalence in natural populations of Drosophila neotestacea

Selfish genetic elements bias their own transmission to the next generation, even at the expense of the fitness of their carrier. Sex-ratio (SR) meiotic drive occurs when an X-chromosome causes Y-bearing sperm to die during male spermatogenesis, so that it is passed on to all of the male''s offspring, which are all daughters. How SR is maintained as a stable polymorphism in the absence of genetic suppressors of drive is unknown. Here, we investigate the potential for the female remating rate to affect SR dynamics in natural populations, using the fly Drosophila neotestacea. In controlled laboratory conditions, females from populations where SR is rare mate more often than females from populations where SR is common. Furthermore, only when males mate multiply does the average fertility of SR males relative to wild-type males decrease to a level that can prevent SR from spreading. Our results suggest that differences in the female mating rate among populations may contribute to SR dynamics in the wild, and thus also affect the outcome of this intragenomic conflict. In line with this, we also present evidence of a localized population crash due to SR that may have resulted from habitat fragmentation along with a reduced mating rate.     

Reproductive strategies of Aphidius ervi Haliday (Hymenoptera: Aphidiidae)

Hymenopteran parasitoids are usually arrhenotokous parthenogenetic, where females arise from fertilized and males from unfertilized eggs. Therefore, the reproductive fitness of females is a function of egg production and furthermore affected by mating, whereas that of males is mainly determined by the number of daughters they father. Aphidius ervi Haliday is a quasi-gregarious parasitoid of a number of aphid pests on economically important crops such as legumes and cereals. Females are monandrous whereas males are polygynous. Here, we tested how parental age at mating and male mating history affected mating success, fecundity and daughter production in this species. Once-mated males perform significantly better than naïve males with regard to mating success, suggesting that males learn from previous matings. The fecundity of virgin females is not significantly different from that of mated females regardless of parental age at mating and male mating history, indicating that mating does not stimulate egg production or contribute to female nutrient supply. Males can replenish sperm supply after mating, implying that they are at least moderately synspermatogenic. Preference for young over old mates for mating by both sexes may be explained by the fact that aging of both sexes contributes to the reduction of daughter production. Rather than sperm depletion, the reduced daughter production may be attributed to diminishing sperm viability and mobility in aging males and increasing constraints in fertilization process in aging females. Our results also show that female age has a stronger impact on the production of daughters, suggesting that fertilization process in females is more sensitive to aging than sperm vigor in males.