Solitary foundation or colony fission in ants: an intraspecific study shows that worker presence and number increase colony foundation success.

Dispersal and establishment strategies are highly variable. Each strategy is associated with specific costs and benefits, and understanding which factors favour or disfavour a strategy is a key issue in ecology and evolution. Ants exhibit several strategies of establishment, i.e. of colony foundation. Some species rely on winged queens that found new colonies alone when others found with accompanying workers (colony fission). The benefits conferred by these workers have been little studied and quantified, because comparing the costs and benefits of solitary foundation vs. colony fission is difficult when comparing different species. We investigated this using the ant Myrmecina graminicola, one of the few species that use both strategies. Young mated queens were allowed to found new colonies in the laboratory, with either zero (solitarily), two or four workers (colony fission). The presence of workers increased both survival and growth of the foundations over the first year, with more workers yielding higher growth. Few workers (as little as two workers) were sufficient to provide benefits, suggesting that in M. graminicola the strategy of colony fission may not dramatically decrease the number of new colonies produced compared to solitary foundation. Because queens performing solitary foundation or colony fission differ in dispersal (by flight vs. on foot), our results support the hypothesis that these two strategies of foundation coexist along a competition-colonization trade-off, where solitary foundation offers a colonization advantage, while colony fission has a competitive advantage.

Climate warming: a loss of variation in populations can accompany reproductive shifts.

The most documented response of organisms to climate warming is a change in the average timing of seasonal activities (phenology). Although we know that these average changes can differ among species and populations, we do not know whether climate warming impacts within-population variation in phenology. Using data from five study sites collected during a 13-year survey, we found that the increase in spring temperatures is associated with a reproductive advance of 10 days in natural populations of common lizards (Zootoca vivipara). Interestingly, we show a correlated loss of variation in reproductive dates within populations. As illustrated by a model, this shortening of the reproductive period can have significant negative effects on population dynamics. Consequently, we encourage tests in other species to assess the generality of decreased variation in phenological responses to climate change.

Sex-associated differences in trace metals concentrations in and on the plumage of a common urban bird species.

Urban areas encompass both favorable and stressful conditions linked with human activities and pollution. Pollutants remain of major ecological importance for synanthropic organisms living in the city. Plumage of urban birds harbour trace metals, which can result from external deposition or from internal accumulation. External and internal plumage concentrations likely differ between specific trace metals, and may further differ between males and females because of potential sex-linked differential urban use, physiology or behaviour. Here, we measured the concentrations in four trace metals (cadmium, copper, lead and zinc) in both unwashed and washed feathers of 49 male and 38 female feral pigeons (Columba livia) from Parisian agglomeration. We found that these concentrations indeed differed between unwashed and washed feathers, between males and females, and for some metals depended on the interaction between these factors. We discuss these results in the light of physiological and behavioural differences between males and females and of spatial repartition of the four trace metals in the city.

The influence of intraspecific competition on resource allocation during dependent colony foundation in a social insect.

Organisms face a trade-off between investment in fewer, larger offspring, or more, smaller offspring. Most organisms can adjust investment through variation in the size and number of offspring in response to factors such as resource availability and competition. In some social animals, established colonies divide into groups of individuals that become autonomous, a process known as colony fission (also dependent colony foundation in social insects). Resource allocation under fission can be fine-tuned by adjusting the number of new groups (offspring number) and the number of individuals in each new group (offspring size). We assessed the influence of competition on resource allocation during fission in the ant Cataglyphis cursor, by allowing colonies to fission in experimental enclosures of high or low conspecific colony density. The pattern of colony fission was similar to that observed in the field: each fissioning colony produced a few new nests comprising a highly variable number of workers and a single queen, the old queen was often replaced, and new queens were produced in excess. The number of new nests produced depended on the available workforce in the parent colony but was not affected by differences in colony density. Comparison with data from fission under natural field conditions, however, indicates that colonies in enclosures produced fewer, larger new nests, suggesting that resource investment patterns during fission are indeed subject to extrinsic factors. The density of conspecific colonies in the immediate surroundings may be an unreliable estimate of competition intensity and other factors should be considered.

Female choice for heterozygous mates changes along successive matings in a lizard.

Female mate choice and female multiple mating are major focuses of studies on sexual selection. In a multiple mating context, the benefits of mate choice can change along successive matings, and female choice would be expected to change accordingly. We investigated sequential female mate choice in the moderately polyandrous common lizard (Zootoca vivipara, synonym Lacerta vivipara). Along successive mating opportunities, we found that females were relatively unselective for the first mate, but accepted males of higher heterozygosity for subsequent mating, consistent with the trade-up choice hypothesis. We discuss the evidence of trade-up mate choice in squamates and generally trade-up for mate heterozygosity in order to motivate new studies to fill gaps on these questions.

Unequal resource allocation among colonies produced by fission in the ant Cataglyphis cursor.

How organisms allocate limited resources to reproduction is critical to their fitness. The size and number of offspring produced have been the focus of many studies. Offspring size affects survival and growth and determines offspring number in the many species where there is a trade-off between size and number. Many social insects reproduce by colony fission, whereby young queens and accompanying workers split off from a colony to form new colonies. The size of a new colony (number of workers) is set at the time of the split, and this may allow fine tuning size to local conditions. Despite the prevalence of colony fission and the ecological importance of social insects, little is known of colony fission except in honey bees. We studied colony fission in the ant Cataglyphis cursor. For clarity, "colony" and "nest" refer to colonies before and after colony fission, respectively (i.e., each colony fissions into several nests). The reproductive effort of colonies was highly variable: Colonies that fissioned varied markedly in size, and many colonies that did not fission were as large as some of the fissioning colonies. The mother queen was replaced in half of the fissioning colonies, which produced 4.0 +/- 1.3 (mean +/- SD) nests of markedly varied size. Larger fissioning colonies produced larger nests but did not produce more nests, and resource allocation among nests was highly biased. When a colony produced several nests and the mother queen was not replaced, the nest containing the mother queen was larger than nests with a young queen. These results show that the pattern of resource allocation differs between C. cursor and honey bees. They also suggest that C. cursor may follow a bet-hedging strategy with regard to both the colony size at which fission occurs and the partitioning of resources among nests. In addition, colony fission may be influenced by the age and/or condition of the mother queen, and the fact that workers allocating resources among nests have incomplete knowledge of the size and number of nests produced. These results show that the process of colony fission is more diverse than currently acknowledged and that studies of additional species are needed.

Variation in patriline reproductive success during queen production in orphaned colonies of the thelytokous ant Cataglyphis cursor.

In genetically diverse insect societies (polygynous or polyandrous queens), the production of new queens can set the ground for competition among lineages. This competition can be very intense when workers can reproduce using thelytoky as worker lineages that manage to produce new queens gain a huge benefit. Selection at the individual level might then lead to the evolution of cheating genotypes, i.e. genotypes that reproduce more than their fair share. We studied the variation in reproductive success among worker patrilines in the thelytokous and highly polyandrous ant Cataglyphis cursor. Workers produce new queens by thelytoky in orphaned colonies. The reproductive success of each patriline was assessed in 13 orphaned colonies using genetic analysis of 433 workers and 326 worker-produced queens. Our results show that patrilines contributed unequally to queen production in half of the colonies, and the success of patrilines was function of their frequencies in workers. However, over all colonies, we observed a significant difference in the distribution of patrilines between workers and worker-produced queens, and this difference was significant in three of 13 colonies. In addition, six colonies contained a low percentage of foreign workers (drifters), and in one colony, they produced a disproportionably high number of queens. Hence, we found some evidence for the occurrence of rare cheating genotypes. Nevertheless, cheating appears to be less pronounced than in the Cape Honey bee, a species with a similar reproductive system. We argue that worker reproduction by parthenogenesis might not be common in natural populations of C. cursor.

Relationship between female mating strategy, litter success and offspring dispersal.

The relationship between mating systems and dispersal has generally been studied at the population and species levels. It has hardly ever been investigated at the individual level, by studying the variations of mating and dispersal strategies between individuals. We investigated this relationship in a natural population of the common lizard (Lacerta vivipara). Assuming that dispersal has a genetic basis, juvenile dispersal would be expected to be more family-dependent in monoandrous litters than in polyandrous litters. The opposite pattern was observed. Thus, maternal effects and/or litter effects play a greater role than genetic determinism in shaping the dispersal phenotype of juveniles. Moreover, the relationship between female mating strategy and offspring dispersal depended on litter success, in a way consistent with an influence of mother-offspring competition. Such a link between mating and dispersal strategies of individuals may have major repercussions for the way we consider the roles of these processes in population functioning.

No actual conflict over colony inheritance despite high potential conflict in the social wasp Polistes dominulus.

Social insect societies are outstanding examples of cooperation and conflict. Individuals work together, yet seek to increase their inclusive fitness at each others' expense. One such conflict is over colony inheritance, when a queen inherits the colony following the death of the previous queen. Colony inheritance is common in the social wasp Polistes dominulus, and it can have dramatic fitness consequences. The subordinate inheriting the colony is often unrelated to the initial foundress (alpha) and the workers, who therefore get zero inclusive fitness. Workers are capable of mating and reproducing, so that inheritance by a subordinate rather than by a related worker is surprising. Using patterns of egg-laying and egg destruction, we show in 32 laboratory colonies that, upon the removal of alpha, workers fully accepted a subordinate as the new breeder. This new alpha monopolized reproduction to the same extent as alpha, and there was no increase in reproduction by workers and other subordinates. Why workers accept a potentially unrelated subordinate as breeder rather than a full-sister worker is unclear. They may be constrained to do so, and they may seek fitness benefits by producing males later in the season or by absconding the nest.

Antagonistic effects of a Mhc class I allele on malaria-infected house sparrows.

Genes of the Major Histocompatibility Complex (Mhc) play a fundamental role during the immune response because MHC molecules expressed on cell surface allow the recognition and presentation of antigenic peptides to T-lymphocytes. Although Mhc alleles have been found to correlate with pathogen resistance in several host-parasite systems, several studies have also reported associations between Mhc alleles and an accrued infection risk or an accelerated disease progression. The existence of these susceptibility alleles is puzzling, as the cost generated by the infection should rapidly eliminate them from the population. Here, we show that susceptibility alleles may be maintained in a population of house sparrows (Passer domesticus) if they have antagonistic effects on different malaria parasites. We found that one Mhc class I allele was associated with a 2.5-fold increase in the risk to be infected with a Plasmodium strain, but with a 6.4-fold reduction in the risk to harbour a Haemoproteus strain. We suggest that this antagonistic effect might arise because Mhc genes can alter the competitive interactions between malaria parasites within the host.

Aggressions and size-related fecundity of queenless workers in the ant Cataglyphis cursor.

In social hymenoptera, the reproductive division of labor is often linked to differences in individual body size with the reproductive caste (the queen) being larger than the workers. Likewise, the reproductive potential may vary with size within the worker caste and could affect the evolution of worker size in social insects. Here, we tested the relationship between worker size and reproductive potential in the facultative parthenogenetic ant Cataglyphis cursor. Colonies are headed by a multiply mated queen, but workers can produce gynes (virgin queens) and workers by thelytokous parthenogenesis after the queen's death. We observed the behaviour of workers (n = 357) until the production of gynes (212 h over 3 months) in an orphaned colony (mated queen not present). The size of workers was measured, and their paternal lineage determined using six microsatellite markers, to control for an effect of patriline. Larger workers were more likely to reproduce and lay more eggs indicating that individual level selection could take place. However, paternal lineage had no effect on the reproductive potential and worker size. From the behavioural and genetic data, we also show for the first time in this species, evidence of aggressive interactions among workers and a potential for nepotism to occur in orphaned colonies, as the five gynes produced belonged to a single paternal lineage.

Condition dependence of reproductive strategy and the benefits of polyandry in a viviparous lizard.

Species in which males do not contribute to reproduction beyond the provision of sperm offer good opportunities to study the potential genetic benefits that females can obtain from polyandry. Here, we report the results of a study examining the relationships between polyandry and components of female fitness in the common lizard (Lacerta vivipara). We found that polyandrous females produce larger clutches than monandrous females. Polyandrous females also lose fewer offspring during the later stages of gestation and at birth, but we did not find any relationship between polyandry and physical characteristics of viable neonates. Our results were consistent with the predictions of the intrinsic male quality hypothesis, while inbreeding avoidance and genetic incompatibility avoidance might also explain some part of the variation observed in clutch size. Moreover, the benefits of polyandry appeared to depend on female characteristics, as revealed by an interaction between reproductive strategy and female length on reproductive success. Thus, all females did not benefit equally from mating with multiple males, which could explain why polyandry and monandry coexist.

Major histocompatibility alleles associated with local resistance to malaria in a passerine.

Malaria parasites are a major cause of human mortality in tropical countries and a potential threat for wildlife, as witnessed by the malaria-induced extinction of naive Hawaiian avifauna. Identifying resistance mechanisms is therefore crucial both for human health and wildlife conservation. Patterns of malaria resistance are known to be highly polygenic in both humans and mice, with marked contributions attributed to major histocompatibility (Mhc) genes. Here we show that specific Mhc variants are linked to both increased resistance and susceptibility to malaria infection in a wild passerine species, the house sparrow (Passer domesticus). In addition, links between host immunogenetics and resistance to malaria involved population-specific alleles, suggesting local adaptation in this host-parasite interaction. This is the first evidence for a population-specific genetic control of resistance to malaria in a wild species.

Complex Mhc-based mate choice in a wild passerine.

The extreme polymorphism of the vertebrate major histocompatibility complex (Mhc) is famous for protecting hosts against constantly evolving pathogens. Mate choice is often evoked as a means of maintaining Mhc variability through avoidance of partners with similar Mhc alleles or preference for heterozygotes. Evidence for these two hypotheses mostly comes from studies on humans and laboratory mice. Here, we tested these hypotheses in a wild outbred population of house sparrows (Passer domesticus). Females were not more or less closely related to the males they paired with when considering neutral genetic variation. However, males failed to form breeding pairs when they had too few Mhc alleles and when they were too dissimilar from females at Mhc loci (i.e. had no common alleles). Furthermore, pairs did not form at random as Mhc diversity positively correlated in mating pairs. These results suggest that mate choice evolves in response to (i) benefits in terms of parasite resistance acquired from allelic diversity, and (ii) costs associated with the disruption of co-adapted genes.

Conflict over multiple-partner mating between males and females of the polygynandrous common lizards.

The optimal number of mate partners for females rarely coincides with that for males, leading to a potential sexual conflict over multiple-partner mating. This suggests that the population sex ratio may affect multiple-partner mating and thus multiple paternity. We investigate the relationship between multiple paternity and the population sex ratio in the polygynandrous common lizard (Lacerta vivipara). In six populations the adult sex ratio was biased toward males, and in another six populations the adult sex ratio was biased toward females, the latter corresponding to the average adult sex ratio encountered in natural populations. In males the frequency and the degree of polygyny were lower in male-biased populations, as expected if competition among males determines polygyny. In females the frequency of polyandry was not different between treatments, and polyandrous females produced larger clutches, suggesting that polyandry might be adaptive. However, in male-biased populations females suffered from reduced reproductive success compared to female-biased populations, and the number of mate partners increased with female body size in polyandrous females. Polyandrous females of male-biased populations showed disproportionately more mating scars, indicating that polyandrous females of male-biased populations had more interactions with males and suggesting that the degree of multiple paternity is controlled by male sexual harassment. Our results thus imply that polyandry may be hierarchically controlled, with females controlling when to mate with multiple partners and male sexual harassment being a proximate determinant of the degree of multiple paternity. The results are also consistent with a sexual conflict in which male behaviors are harmful to females.

Recovery of Na-glucose cotransport activity after renal ischemia is impaired in mice lacking vimentin.

Vimentin, an intermediate filament protein mainly expressed in mesenchyma-derived cells, is reexpressed in renal tubular epithelial cells under many pathological conditions, characterized by intense cell proliferation. Whether vimentin reexpression is only a marker of cell dedifferentiation or is instrumental in the maintenance of cell structure and/or function is still unknown. Here, we used vimentin knockout mice (Vim(-/-)) and an experimental model of acute renal injury (30-min bilateral renal ischemia) to explore the role of vimentin. Bilateral renal ischemia induced an initial phase of acute tubular necrosis that did not require vimentin and was similar, in terms of morphological and functional changes, in Vim(+/+) and Vim(-/-) mice. However, vimentin was essential to favor Na-glucose cotransporter 1 localization to brush-border membranes and to restore Na-glucose cotransport activity in regenerating tubular cells. We show that the effect of vimentin inactivation is specific and results in persistent glucosuria. We propose that vimentin is part of a structural network that favors carrier localization to plasma membranes to restore transport activity in injured kidneys.

JunD protects against chronic kidney disease by regulating paracrine mitogens.

The AP-1 transcription factor, composed of Jun and Fos proteins, plays a crucial role in the fine tuning of cell proliferation. We showed previously that AP-1 complexes are activated during the proliferative response that parallels the development of renal lesions after nephron reduction, but little is known about the specific role of individual Jun/Fos components in the deterioration process. Here we used JunD knockout (JunD-/-) mice and an experimental model of chronic renal injury (75% nephron reduction) to explore the role of JunD. Nephron reduction resulted in an initial compensatory growth phase that did not require JunD. JunD, however, was essential to inhibit a second wave of cell proliferation and to halt the development of severe glomerular sclerosis, tubular dilation, and interstitial fibrosis. We show that the effects of junD inactivation are not cell autonomous and involve upregulation of the paracrine mitogen, TGF-alpha. Expression of a transgene (REM) encoding a dominant negative isoform of the EGFR, the receptor for TGF-alpha, prevented the second wave of cell proliferation and the development of renal lesions in bitransgenic JunD-/-/REM mice. We propose that JunD is part of a regulatory network that controls proliferation to prevent pathological progression in chronic renal diseases.

Vimentin affects localization and activity of sodium-glucose cotransporter SGLT1 in membrane rafts.

It has been reported that vimentin, a cytoskeleton filament that is expressed only in mesenchymal cells after birth, is re-expressed in epithelial cells in vivo under pathological conditions and in vitro in primary culture. Whether vimentin re-expression is only a marker of cellular dedifferentiation or is instrumental in the maintenance of cell structure and/or function is a matter of debate. To address this issue, we used renal proximal tubular cells in primary culture from vimentin-null mice (Vim(-/-)) and from wild-type littermates (Vim(+/+)). The absence of vimentin did not affect cell morphology, proliferation and activity of hydrolases, but dramatically decreased Na-glucose cotransport activity. This phenotype was associated with a specific reduction of SGLT1 protein in the detergent-resistant membrane microdomains (DRM). In Vim(+/+) cells, disruption of these microdomains by methyl-beta-cyclodextrin decreased SGLT1 protein abundance in DRM, a change that was paralleled by a decrease of Na-glucose transport activity. Importantly, we showed that vimentin is located to DRM, but it disappeared after methyl-beta-cyclodextrin treatment. In Vim(-/-) cells, supplementation of cholesterol with cholesterol-methyl-beta-cyclodextrin complexes completely restored Na-glucose transport activity. Interestingly, neither cholesterol content nor cholesterol metabolism changed in Vim(-/-) cells. Our results are consistent with the view that re-expression of vimentin in epithelial cells could be instrumental to maintain the physical state of rafts and, thus, the function of DRM-associated proteins.