Differences in tomato spotted wilt virus vector competency between males and females of Frankliniella occidentalis

Possible differences in tomato spotted wilt virus (TSWV) transmission vector competency between Frankliniella occidentalis (Pergande) (Thysanoptera: Thripidae) males and females were investigated. The males of the Dutch reference population NL3 transmitted TSWV at a notably higher rate (57%) than the females (32%). The viruliferous males also transmitted more frequently within the first six days after adult emergence than the females. For both sexes, the transmission efficiency dropped with age, simultaneously with the consumption rate. The higher vector efficiency for males appeared to be a general phenomenon as this feature was also found for thirteen other F. occidentalis populations, which originated from distinct geographic regions.     

Effects of tomato spotted wilt virus (TSWV) isolates, host plants, and temperature on survival, size, and development time of Frankliniella fusca

The effects of different isolates of the tomato spotted wilt tospovirus (TSWV), host plants, and temperatures on Frankliniella fusca (Hinds) (Thysanoptera: Thripidae), the most important vector of TSWV in North Carolina, were measured in the laboratory. Thrips were reared at either 18.3, 23.9, or 29.4 °C until adult eclosion on excised leaves of Datura stramonium L. or Emilia sonchifolia (L.). Plants were either infected with the TSWV isolates CFL or RG2, or left uninfected (control). The results revealed a positive relationship between larval survival and temperature, regardless of host plant or TSWV isolate. Both survival to adult and percentage transmission of TSWV by F. fusca were significantly affected by the interaction between host plant and TSWV isolate. The consequence of this interaction was that the cohort‐based percentage transmission from infected E. sonchifolia plants for CFL was 1.3‐fold greater than that of RG2, whereas the percentage transmission from infected D. stramonium plants for RG2 was twice that of CFL. Both host plant and TSWV isolates showed significant effects on thrips development time to adult and head capsule width of adult thrips, as well as on the incidence of thrips infection with TSWV. The infection status of these thrips was determined by ELISA for the NSs viral protein. Infected thrips reared on infected host foliage took longer to develop to adult and were smaller than non‐infected thrips which had also been reared on infected host foliage, demonstrating a direct effect of the TSWV on thrips. However, non‐infected thrips reared on non‐infected leaves took longer to develop than non‐infected thrips reared on infected leaves, suggesting an effect of the plant tissue on thrips. In addition, adult thrips reared on TSWV‐infected D. stramonium at 29.4 °C developed smaller head capsules than thrips developing on infected foliage at lower temperatures and on non‐infected leaves of D. stramonium or E. sonchifolia. Both TSWV isolates and host plants differentially affected females more than males. In conclusion, both the infection of thrips by TSWV and TSWV‐mediated changes in host plant quality were found to have significant biological effects on F. fusca.     

Female-biased infection and transmission of the gastrointestinal nematode Trichuris arvicolae infecting the common vole, Microtus arvalis

Previous studies addressing the importance of host gender in parasite transmission have shed light on males as the more important hosts, with the higher transmission potential of males being explained by the fact that they often harbour higher parasite loads than females. However, in some systems females are more heavily infected than males and may be responsible for driving infection under such circumstances. Using a wild population of common voles (Microtus arvalis), we showed that females were more frequently infected by the intestinal nematode Trichuris arvicolae than males (i.e. prevalence based on the presence of eggs in the faeces) and that females were shedding greater numbers of parasite eggs per gram of faeces (EPG) than males. By applying an anthelmintic treatment to either male or female voles, we demonstrated that treating females significantly reduced parasite burdens (i.e. prevalence and EPG) of both male and female hosts, while treating males only reduced parasite burden in males. These findings indicate that in this female-biased infection system females play a more important role than males in driving the dynamics of parasite transmission.     

Specific Insect-Virus Interactions Are Responsible for Variation in Competency of Different Thrips tabaci Isolines to Transmit Different Tomato Spotted Wilt Virus Isolates

Local adaptation between sympatric host and parasite populations driven by vector genetics appears to be a factor that influences dynamics of disease epidemics and evolution of insect-vectored viruses. Although T. tabaci is the primary vector of Tomato spotted wilt virus (TSWV) in some areas of the world, it is not an important vector of this economically important plant virus in many areas where it occurs. Previous studies suggest that genetic variation of thrips populations, virus isolates, or both are important factors underlying the localized importance of this species as a vector of TSWV. This study was undertaken to quantify variation in transmissibility of TSWV isolates by T. tabaci, in the ability of T. tabaci to transmit isolates of TSWV, and to examine the possibility that genetic interactions and local adaptation contribute to the localized nature of this species as a vector of TSWV. Isofemale lines of Thrips tabaci from multiple locations were tested for their ability to transmit multiple TSWV isolates collected at the same and different locations as the thrips. Results revealed that the probability of an isofemale line transmitting TSWV varied among virus isolates, and the probability of an isolate being transmitted varied among isofemale lines. These results indicate that the interaction of T. tabaci and TSWV isolate genetic determinants underlie successful transmission of TSWV by T. tabaci. Further analysis revealed sympatric vector-virus pairing resulted in higher transmission than allopatric pairing, which suggests that local adaptation is occurring between T. tabaci and TSWV isolates.     

Do sexual dimorphisms in reproductive allocation and new shoot biomass increase with an increase of altitude? A case of the shrub willow Salix reinii (Salicaceae)

Based on the general tendency for females of dioecious plants to pay higher reproductive cost than males, it has been predicted that females should have much more reduced reproductive outputs and diminished vegetative production than males in energy-limited habitats. Nevertheless, this prediction has rarely been directly investigated. We investigated altitudinal changes in reproductive biomass and shoot production, normalized by plant size, for females and males of a shrub willow, Salix reinii, on Mt. Hakkoda, northeast Japan. Females maintained higher reproductive biomass than males at all altitudes; however, reproductive allocation for both sexes tended to decrease at a similar rate with an increase in altitude. Moreover, females vegetatively produced at the same rate as males at all altitudes. These findings suggest that females have a mechanism to compensate for the extra investment in reproduction irrespective of a changing environment. Shoot production did not change with altitude, suggesting that S. reinii gave priority to vegetative investment at the cost of reproductive output at higher altitudes. Inconsistent with general predictions, females did not respond more sensitively than males to severe environmental conditions in either reproductive allocation or shoot production, despite much higher resource investment in reproduction.     

Male Age Affects Female Mate Preference and Reproductive Performance in the Cabbage Beetle, <Emphasis Type="Italic">Colaphellus bowringi</Emphasis>

The influence of male age on female mate preference and reproductive performance in the cabbage beetle, Colaphellus bowringi was examined, using male and female adults of varying ages (young, middle-aged and old) after a single mating. In a simultaneous choice test, females of all age class preferentially mated with middle-aged males. Mating duration was positively related to male age. Longevity of females was not significantly affected by male age. Young females paired to middle-aged males had significantly higher egg production than those paired to old males, and the eggs of females paired to middle-aged males exhibited significantly higher hatching success than the eggs of females mated to young or old males. These results suggest that middle-aged males are more advantageous for female fitness than young and old males.     

Differences in response to defoliation between males and females of Silene dioica

Summary The response by male and female plants to herbivory was studied by experimental defoliation of the dioecious perennial herb Silene dioica in a green-house. Male and female plants were defoliated prior to and during the early flowering phase at two intensities (50% and 100% of leaf-area removed) in two consecutive years. Defoliation resulted in a decrease in the number of flowers initiated in both sexes, while a larger delay of peak flowering and a higher mortality was observed in males compared to females. In female plants, severe defoliation resulted in a reduction in seed number per capsule and in seed size compared to control. Females showed a negative correlation between the production of flowers in the first and second season in all treatments, while flowering in males the first season was not correlated with flowering in the second season. Females also showed a lower frequency of flowering than males during the two seasons studied. However, during the flowering period, males allocated significantly more biomass to flowers than did females. This outcome supports the idea that females may have a higher total reproductive expenditure than males, but males have a higher reproductive effort during flowering. Male rosette leaves were significantly preferred by the generalist herbivore Arianta arbustorum in experiments. This preference was most pronounced in trials with leaves from fertilized plants compared to nonfertilized plants. A greater storage of resources in aboveground leaves during winter by males compared to females may explain the higher preference for male leaves and the higher male mortality following early defoliation. Furthermore, males are smaller than females and may have a lower ability than females to replace lost resources needed for reproduction when defoliated early in the season.     

Sex-specific SOD levels and DNA damage in painted dragon lizards (Ctenophorus pictus)

When groups of individuals differ in activities that may influence the production of reactive molecules, such as superoxide, we expect selection to result in congruent upregulation of antioxidant production in the group(s) most at risk of suffering concomitant erosion of essential tissue and biomolecules, such as DNA. We investigate this in a (near) annual lizard species, the Australian painted dragon (Ctenophorus pictus), in which males and females have fundamentally different lifestyles, with males being overtly conspicuous and aggressive, whereas females are placid and camouflaged. When kept in identical conditions to females in captivity, males had higher levels of superoxide dismutase (SOD) through the activity season, which is consistent with selection for a higher capacity of superoxide antioxidation and a lower level of DNA damage than females. Males, however, lacked the clear negative, linear relationship between SOD and DNA erosion observed in females, suggesting that female upregulation of SOD results in a more predictable antioxidation and a more immediate target for selection. Lastly, we analysed aspects of female reproduction from a DNA erosion perspective. Females closer to ovulation, hence with less remaining, circulating vitellogenin, had higher superoxide levels. Furthermore, a multiple regression analysis showed that females that produced more clutches over time suffered more DNA erosion, whereas females with higher SOD levels suffered less DNA erosion.     

Preference of the vector thrips Frankliniella occidentalis for plants infected with thrips‐non‐transmissible Tomato spotted wilt virus

The effect of a thrips‐non‐transmissible Tomato spotted wilt virus (TSWV) on insect–host interactions between thrips and Arabidopsis thaliana was analysed. A wild‐type TSWV virulent isolate and a TSWV isolate that induces mild symptoms on inoculated plants (TSWV‐Mo) were used in this study, and TSWV‐Mo isolate was obtained by single local lesion isolation using Petunia x hybrid after several passages on Nicotiana rustica plants. In transmission test, although wild‐type TSWV (TSWV‐wt) was transmitted by two thrips species (transmission ratio; Frankliniella occidentalis, 25%; Thrips tabaci, 10%; and T. palmi, 0%), none of the thrips transmitted TSWV‐Mo. Feeding damage by F. occidentalis in A. thaliana plants was more extensive on TSWV‐wt‐infected plants than on TSWV‐Mo‐infected plants, despite comparable preference. Among the markers of plant defences, salicylic acid‐regulated genes were upregulated threefold to sixfold by TSWV‐wt or TSWV‐Mo infection. In contrast, jasmonate‐regulated genes and jasmonate/ethylene‐regulated genes were not affected by the infections. Pull assays showed that adjacent TSWV‐Mo‐infected plants were preferred over uninfected plants. In conclusion, our results showed that the transmissibility by thrips of TSWV is not related to preference of vector thrips and suggested that TSWV‐Mo‐infected plants may be used as attractants for behaviour control of thrips.     

Increased male mating rate in Drosophila is associated with Wolbachia infection

The maternally inherited bacterium Wolbachia pipientis infects 25-75% of arthropods and manipulates host reproduction to improve its transmission. One way Wolbachia achieves this is by inducing cytoplasmic incompatibility (CI), where crosses between infected males and uninfected females are inviable. Infected males suffer reduced fertility through CI and reduced sperm production. However, Wolbachia induce lower levels of CI in nonvirgin males. We examined the impact of Wolbachia on mating behaviour in male Drosophila melanogaster and D. simulans, which display varying levels of CI, and show that infected males mate at a higher rate than uninfected males in both species. This may serve to increase the spread of Wolbachia, or alternatively, may be a behavioural adaptation employed by males to reduce the level of CI. Mating at high rate restores reproductive compatibility with uninfected females resulting in higher male reproductive success thus promoting male promiscuity. Increased male mating rates also have implications for the transmission of Wolbachia.     

CTG instability in myotonic dystrophy: molecular genetic analysis of families from south-eastern France with characteristics of intergenerational variation in CGT repeat numbers.

We report clinical, genetical and genealogical findings in 149 French families from the Rh?ne-Alpes area studied over a 5-year period. There was a significant excess of DM females compared to DM males with (CTG) repeat sizes between 1-2 kb. The mean maternal (CTG) repeat size was higher than paternal repeat size. Anticipation phenomenom was significantly higher after maternal than after paternal transmission. A significant correlation between parental (CTG) repeat size and intergenerational variation both in paternal and maternal transmissions was observed. The anticipation phenomenom was more important for sons than daughters particularly after maternal transmission. The mean (CTG) repeat size in mothers of CDM cases was about twice that of mothers of NCDM children. The risk of giving birth to a CDM child increased considerably when the number of maternal (CTG) repeats was over 300 (CTG). A significant excess of DM females was observed. They had on average 24% fewer children than male patients. Paternal transmission (63.6%) of DM occurred more frequently than maternal transmission (52.7%).     

Sexually dimorphic growth in the dioecious tropical shrub, Siparuna grandiflora

1. To demonstrate evolved sex-based differences in vegetative traits of dioecious plant species, one must consider both pre-reproductive and reproductive individuals, as dimorphic patterns commonly arise secondarily from different effects of reproduction on resource balance.
2. Siparuna grandiflora , a neotropical dioecious shrub in which females allocate significantly more biomass to reproduction than males, was studied for 2 years (three reproductive events) to determine whether sex-based differences in stem growth, leaf production and allocation pattern could be detected in pre-reproductive individuals grown from cuttings in field plots or in mature naturally occurring individuals.
3. Among pre-reproductive individuals, females accumulated more stem and leaves than males, but among mature individuals, no sex-based growth differences were apparent. In mature individuals, both growth and leaf longevity were positively correlated with reproductive frequency. With regards allocation, pre-reproductive males had larger leaves than females, and mature females allocated less biomass per unit stem length than males.
4. The capacity of pre-reproductive females to grow faster than males demonstrates innate differences between the sexes. That mature females achieved equivalent growth to males, despite higher reproductive allocation, indicates that the greater growth capacity of young females is sustained in older females and enables them to compensate for greater reproductive allocation.     

Adult diet and male-female contact effects on female reproductive potential in Mexican fruit fly (Anastrepha ludens Loew) (Diptera tephritidae)

Wild strains of fruit flies (Diptera: Tephritidae) placed into laboratory rearing conditions are subjected to selection pressures caused by the diet, cages, density of flies, and other factors. Selection that changes mating behavior of the strain may result in less effective males released in sterile insect programs. Tests were performed to examine the effects of protein in diet and adult interactions on egg production and mating during sexual maturation of the Mexican fruit fly (Anastrepha ludens Loew) in laboratory cages. Flies were offspring of wild flies collected from Chiapas or Nuevo Leon, Mexico, and reared on Valencia oranges. Experiments demonstrated effects of yeast hydrolysate protein in adult diet and pairing with males on production of mature and immature eggs, numbers of females producing eggs, and mating with females aged 15 d. Addition of protein to 4% fructose in the adult diet approximately tripled mature egg production in females maintained for the total maturation period with an equal number of males. Females that matured without males produced approximately 33% more-mature eggs when fed protein than those fed no protein. Total egg production of females matured without males and fed sugar only or sugar with protein was more than twice that of females matured with males. Tests to examine the effects of male and female diet separately on female egg production showed slightly higher egg production in females fed protein, or females paired with males fed protein, but these differences were not significant. The most definitive effects were that combining wild strain females and males in cages during maturation reduced egg production. This effect was greatest when flies were not fed protein.     

Patterns of resource allocation in male and female plants of Rumex acetosa and R. acetosella

Summary Patterns of resource allocation in the dioecious Rumex acetosa and R. acetosella were investigated. Males were found to allocate more to reproduction during flower production than females, whereas females invested considerably more in reproduction during seed production. Altogether, females allocated both a higher total amount and a higher proportion of energy to reproduction than did males. By regression analysis, the influence of plant size on reproductive effort was examined separately for males and females. The results indicated that while reproductive effort is sometimes lower for tall plants than for small plants, size-independent effects have a greater influence on reproductive effort than size distribution. An analysis of variance was conducted to investigate the effects of population, season, sex and their interactions on plant size, and an analysis of covariance was used to study differences in resource allocation patterns. Different interaction effects were found to be most important in the two species of Rumex.     

Male-biased sex-ratio distortion caused by Octosporea bayeri, a vertically and horizontally-transmitted parasite of Daphnia magna

Female-biased sex-ratio distortion is often observed in hosts infected with vertically-transmitted microsporidian parasites. This bias is assumed to benefit the spread of the parasite, because male offspring usually do not transmit the parasite further. The present study reports on sex-ratio distortion in a host-parasite system with both horizontal and vertical parasite transmission: the microsporidium Octosporea bayeri and its host, the planktonic cladoceran Daphnia magna. In laboratory and field experiments, we found an overall higher proportion of male offspring in infected than in uninfected hosts. In young males, there was no parasite effect on sperm production, but, later in life, infected males produced significantly less sperm than uninfected controls. This shows that infected males are fertile. As males are unlikely to transmit the parasite vertically, an increase in male production could be advantageous to the host during phases of sexual reproduction, because infected mothers may obtain uninfected grandchildren through their sons. Life-table experiments showed that, overall, sons harboured more parasite spores than their sisters, although they reached a smaller body size and died earlier. Male production may thus be beneficial for the parasite when horizontal transmission has a large pay-off as males may contribute more effectively to parasite spread than females.     

The impact of a parasitic nematode, Thripinema fuscum, on the feeding behavior and vector competence of Frankliniella fusca

Frankliniella fusca (Hinds) (Thysanoptera: Thripidae) is the predominant thrips species found inhabiting and reproducing in peanut, Arachis hypogaea L. (Fabaceae), and is one of at least seven thrips species reported to transmit Tomato spotted wilt virus (TSWV). The entomogenous nematode Thripinema fuscum Tipping & Nguyen (Tylenchida: Allantonematidae), a natural enemy of F. fusca , parasitizes larval and adult populations under field conditions. All known Thripinema species render the host female thrips sterile and have the potential to suppress pest populations to near extinction. As a result, secondary spread of TSWV in peanut is reduced. Reduction of the virus under field conditions may also be due to lower transmission rates caused by parasite-induced alterations in host feeding behavior. Therefore, the feeding rates of healthy and parasitized F. fusca male and female cohorts on leaf discs were recorded daily for 10 days and digital images were subjected to image analysis and viral transmission rates were compared daily using double antibody sandwich enzyme-linked immunosorbent assay. Thripinema fuscum reduced the feeding of female F. fusca by nearly 65%, and the ability of females to transmit TSWV by 50%. Potential mechanisms underlying the parasite-induced alterations in feeding behavior and transmission are discussed. Parasitism by T. fuscum significantly reduced male longevity, but female longevity was not affected. These results provide further evidence that T. fuscum aids in regulating viruliferous F. fusca pest populations and suggests its potential as a biological control agent for inoculative release in peanut.     

Determinants of breeding dispersal in the sand lizard, Lacerta agilis, (Reptilia, Squamata)

We studied determinants of breeding dispersal (the distance that an animal shifts its mean home range co-ordinate between reproductive events) in an individually marked population of sand lizards (Lacerta agilis) in south west Sweden during 1987–1991. Female breeding dispersal was not determined by age, size, body condition, or number of partners, and females and males that dispersed further did not experience a higher mortality. However, females with a low reproductive output dispersed further than females that reproduced more successfully, and males that lost in bodily condition dispersed further than males that better maintained body condition. We also looked for relationships between age-differences and band-sharing similarity (DNA fingerprinting = DFP) in three categories of lizards — all females, all males, and males and females — to establish whether males would be likely to mate with close kin. Age-difference was strongly correlated with band-sharing in only one category, males and females. When males were older than females this relationship was not significant. However, when females were older than males, age-difference was strongly correlated with band-sharing. Furthermore, females that were old enough to be the mothers of the courting males shared significantly more bands with these males than did the younger females, including the females of the same age as the males. Although parental-specific DFP bands necessary for establishing paternity among our adult lizards were inaccessible to us, we suggest that our circumstantial evidence strongly suggests that some males mate with their mothers. Males that were more closely related with their neighbours also moved further when we controlled statistically for age and mating success. We suggest that by mating with many partners males not only increase their mating success, but also increase the probability of mating with females with ‘good genes’: mean heterozygosity of parents (as revealed by micro-satellites) were strongly correlated with offspring survival.     

Role des spermatheques dans l'utilisation du sperme et la stimulation de l'ovogenese chez Drosophila melanogaster

Fecundity and fertility of Drosophila females were studied after extirpation of one or two spermathecae. When extirpated females were mated only once, egg deposition was slightly accelerated but total egg production, during twelve days, was the same as in virgin operated females. Egg hatchability was high during two days but decreased more rapidly than in normal or sham-operated femalesWhen males were permanently kept with females, egg production and egg hatchability of operated females remained at a high level during the experiment. In that case, females could copulate frequently and the increase in egg production is interpreted as a consequence of paragonial fluid stimulationThe most significant rôle of spermathecae seems therefore to assume the transmission of the sperm stimulus to the brain; they probably also improve the survival capacities of sperm.     

The effect of male mating history on paternal investment, fecundity and female remating in the seed beetle Callosobruchus maculatus

1. In many organisms, males provide nutrients to females via ejaculates that can influence female fecundity, longevity and mating behaviour. The effect of male mating history on male ejaculate size, female fecundity, female longevity and female remating behaviour in the seed beetle Callosobruchus maculatus was determined.
2. The quantity of ejaculate passed to females declined dramatically with successive matings. Despite the decline, a male's ability to fertilize a female fully did not appear to decline substantially until his fourth mating.
3. When females multiply mated with males of a particular mated status, the pattern of egg production was cyclic, with egg production increasing after mating. Females multiply mated to virgins had higher fecundity than females mated to non-virgins, and females mated to twice-mated males had disproportionately increased egg production late in their life.
4. Females that mated to multiple virgins, and consequently laid more eggs, experienced greater mortality than females mated only once or mated to non-virgins, suggesting that egg production is costly, and rather than ameliorating these costs, male ejaculates may increase them by allowing or stimulating females to lay more eggs.
5. Females mating with non-virgin males remated more readily than did females mated to virgins. Females given food supplements were less likely to remate than females that were nutritionally stressed, suggesting that females remate in part to obtain additional nutrients.     

Predation by sparrowhawks Accipiter nisus on male and female pied flycatchers Ficedula hypoleuca in relation to their breeding behaviour and foraging

Bright plumage, song display, and aggressive resource defence in males may cause higher predation on males than on females during the breeding season. However, in birds, higher predation on females is sometimes observed. Parental investment may be high in females (egg-laying, incubation and feeding of offspring), which might lead to a high risk of predation. We studied predation by sparrowhawks Accipiter nisus in relation to behaviour in pied flycatchers Ficedula hypoleuca where breeding males are more conspicuous than females in plumage and behaviour. Male pied flycatchers generally occupied more exposed perches than females. Females were more mobile and foraged more than males, especially prior to and during incubation. During the incubation and nestling stages, when predation on the sexes could be directly compared, sparrowhawks took about the same number of male and female pied flycatchers. During incubation, however, females spent about 77% of the day in the nest and were 4.7 times more vulnerable than males per unit of time available (i.e. outside the nest). A comparison with the chaffinch Fringilla coelebs , where hawks took more females than males, indicates that timing of breeding, foraging behaviour and parental roles of males and females affect predation risk.