Evolution of sociality in spiders leads to depleted genomic diversity at both population and species levels.

Across several animal taxa, the evolution of sociality involves a suite of characteristics, a "social syndrome," that includes cooperative breeding, reproductive skew, primary female-biased sex ratio, and the transition from outcrossing to inbreeding mating system, factors that are expected to reduce effective population size (Ne). This social syndrome may be favoured by short-term benefits but come with long-term costs, because the reduction in Ne amplifies loss of genetic diversity by genetic drift, ultimately restricting the potential of populations to respond to environmental change. To investigate the consequences of this social life form on genetic diversity, we used a comparative RAD-sequencing approach to estimate genomewide diversity in spider species that differ in level of sociality, reproductive skew and mating system. We analysed multiple populations of three independent sister-species pairs of social inbreeding and subsocial outcrossing Stegodyphus spiders, and a subsocial outgroup. Heterozygosity and within-population diversity were sixfold to 10-fold lower in social compared to subsocial species, and demographic modelling revealed a tenfold reduction in Ne of social populations. Species-wide genetic diversity depends on population divergence and the viability of genetic lineages. Population genomic patterns were consistent with high lineage turnover, which homogenizes the genetic structure that builds up between inbreeding populations, ultimately depleting genetic diversity at the species level. Indeed, species-wide genetic diversity of social species was 5-8 times lower than that of subsocial species. The repeated evolution of species with this social syndrome is associated with severe loss of genomewide diversity, likely to limit their evolutionary potential.

Effects of post-hatch brooding temperature on broiler behavior, welfare, and growth.

An elevated brooding temperature during the first wk post hatch of broilers may potentially increase activity levels and reduce welfare problems in terms of non- and slow-starters, lameness, and contact dermatitis. The effects of an elevated brooding temperature the first 7 d post hatch on behavior, welfare, and growth of Ross 308 broilers were investigated. Groups of 28 broilers (14 males and 14 females) were distributed in a balanced way according to their hatching weight (below or above mean), the age of parent breeders (28 or 50 wk of age), and initial brooding temperature (normal 33°C; warm: 37°C) resulting in 8 different treatment groups. Behavioral data were collected on d zero to 6 of age, data on body weight on d zero, 7, 21, and 34 of age, and data on gait and contact dermatitis on d 21 and 34 of age. An elevated brooding temperature resulted in increased body temperature of broilers 5 h after placement (39.9 ± 0.04°C vs. 39.1 ± 0.04°C; P < 0.0001) whereas no difference was found 24 h after placement (P = 0.35). Broilers reared with elevated brooding temperature initiated feeding and drinking earlier, apart from broilers with low hatching weight from old parent breeders (P < 0.0001). They also showed higher activity levels from d one to 6 of age (P < 0.0001) and a higher inter-individual distance at d zero and one of age (P < 0.0001). Broilers with a high hatching weight reared at normal brooding temperature had a higher prevalence of hock burns at d 34 of age (P = 0.001). Broilers reared at elevated brooding temperature had lower body weight at d 7 of age (P < 0.0001); however, no difference appeared from d 21 of age (P = 0.58). No effect of brooding temperature was found on body weight uniformity (P = 0.81). In conclusion, the welfare of broilers may be improved from an elevated brooding temperature the first 7 d post hatch without affecting body weight uniformity and final body weight.

Comparative genomic study of arachnid immune systems indicates loss of beta-1,3-glucanase-related proteins and the immune deficiency pathway.

Analyses of arthropod genomes have shown that the genes in the different innate humoral immune responses are conserved. These genes encode proteins that are involved in immune signalling pathways that recognize pathogens and activate immune responses. These immune responses include phagocytosis, encapsulation of the pathogen and production of effector molecules for pathogen elimination. So far, most studies have focused on insects leaving other major arthropod groups largely unexplored. Here, we annotate the immune-related genes of six arachnid genomes and present evidence for a conserved pattern of some immune genes, but also evolutionary changes in the arachnid immune system. Specifically, our results suggest that the family of recognition molecules of beta-1,3-glucanase-related proteins (ßGRPs) and the genes from the immune deficiency (IMD) signalling pathway have been lost in a common ancestor of arachnids. These findings are consistent with previous work suggesting that the humoral immune effector proteins are constitutively produced in arachnids in contrast to insects, where these have to be induced. Further functional studies are needed to verify this. We further show that the full haemolymph clotting cascade found in the horseshoe crab is retrieved in most arachnid genomes. Tetranychus lacks at least one major component, although it is possible that this cascade could still function through recruitment of a different protein. The gel-forming protein in horseshoe crabs, coagulogen, was not recovered in any of the arachnid genomes; however, it is possible that the arachnid clot consists of a related protein, spätzle, that is present in all of the genomes.

Low genetic diversity and strong but shallow population differentiation suggests genetic homogenization by metapopulation dynamics in a social spider.

Mating systems and population dynamics influence genetic diversity and structure. Species that experience inbreeding and limited gene flow are expected to evolve isolated, divergent genetic lineages. Metapopulation dynamics with frequent extinctions and colonizations may, on the other hand, deplete and homogenize genetic variation, if extinction rate is sufficiently high compared to the effect of drift in local demes. We investigated these theoretical predictions empirically in social spiders that are highly inbred. Social spiders show intranest mating, female-biased sex ratio, and frequent extinction and colonization events, factors that deplete genetic diversity within nests and populations and limit gene flow. We characterized population genetic structure in Stegodyphus sarasinorum, a social spider distributed across the Indian subcontinent. Species-wide genetic diversity was estimated over approximately 2800 km from Sri Lanka to Himalayas, by sequencing 16 protein-coding nuclear loci. We found 13 SNPs in 6592 bp (π = 0.00045) indicating low species-wide nucleotide diversity. Three genetic lineages were strongly differentiated; however, only one fixed difference among them suggests recent divergence. This is consistent with a scenario of metapopulation dynamics that homogenizes genetic diversity across the species' range. Ultimately, low standing genetic variation may hamper a species' ability to track environmental change and render social inbreeding spiders 'evolutionary dead-ends'.

Flow cytometric sexing of spider sperm reveals an equal sperm production ratio in a female-biased species.

Producing equal amounts of male and female offspring has long been considered an evolutionarily stable strategy. Nevertheless, exceptions to this general rule (i.e. male and female biases) are documented in many taxa, making sex allocation an important domain in current evolutionary biology research. Pinpointing the underlying mechanism of sex ratio bias is challenging owing to the multitude of potential sex ratio-biasing factors. In the dwarf spider, Oedothorax gibbosus, infection with the bacterial endosymbiont Wolbachia results in a female bias. However, pedigree analysis reveals that other factors influence sex ratio variation. In this paper, we investigate whether this additional variation can be explained by the unequal production of male- and female-determining sperm cells during sperm production. Using flow cytometry, we show that males produce equal amounts of male- and female-determining sperm cells; thus bias in sperm production does not contribute to the sex ratio bias observed in this species. This demonstrates that other factors such as parental genes suppressing endosymbiont effects and cryptic female choice might play a role in sex allocation in this species.

Polyandrous females acquire indirect benefits in a nuptial feeding species.

The relative force of direct and indirect selection underlying the evolution of polyandry is contentious. When females acquire direct benefits during mating, indirect benefits are often considered negligible. Although direct benefits are likely to play a prominent role in the evolution of polyandry, post-mating selection for indirect benefits may subsequently evolve. We examined whether polyandrous females acquire indirect benefits and quantified direct and indirect effects of multiple mating on female fitness in a nuptial gift-giving spider (Pisaura mirabilis). In this system, the food item donated by males during mating predicts direct benefits of polyandry. We compared fecundity, fertility and survival of singly mated females to that of females mated three times with the same (monogamy) or different (polyandry) males in a two-factorial design where females were kept under high and low feeding conditions. Greater access to nutrients and sperm had surprisingly little positive effect on fitness, apart from shortening the time until oviposition. In contrast, polyandry increased female reproductive success by increasing the probability of oviposition, and egg hatching success indicating that indirect benefits arise from mating with several different mating partners rather than resources transferred by males. The evolution of polyandry in a male-resource-based mating system may result from exploitation of the female foraging motivation and that indirect genetic benefits are subsequently derived resulting from co-evolutionary post-mating processes to gain a reproductive advantage or to counter costs of mating. Importantly, indirect benefits may represent an additional explanation for the maintenance of polyandry.

Effects of within-colony competition on body size asymmetries and reproductive skew in a social spider.

Reproductive partitioning is a key component of social organization in groups of cooperative organisms. In colonies of permanently social spiders of the genus Stegodyphus less than half of the females reproduce, while all females, including nonreproducers, perform suicidal allo-maternal care. Some theoretical models suggest that reproductive skew is a result of contest competition within colonies, leading to size hierarchies where only the largest females become reproducers. We investigated the effect of competition on within-group body size variation over six months in S. dumicola, by manipulating food level and colony size. We found no evidence that competition leads to increased size asymmetry within colonies, suggesting that contest competition may not be the proximate explanation for reproductive skew. Within-colony body size variation was high already in the juvenile stage, and did not increase over the course of the experiment, suggesting that body size variation is shaped at an early stage. This might facilitate task specialization within colonies and ensure colony-level reproductive output by early allocation of reproductive roles. We suggest that reproductive skew in social spiders may be an adaptation to sociality selected through inclusive fitness benefits of allo-maternal care as well as colony-level benefits maximizing colony survival and production.

Task specialization in two social spiders, Stegodyphus sarasinorum (Eresidae) and Anelosimus eximius (Theridiidae).

Understanding the social organization of group-living organisms is crucial for the comprehension of the underlying selective mechanisms involved in the evolution of cooperation. Division of labour and caste formation is restricted to eusocial organisms, but behavioural asymmetries and reproductive skew is common in other group-living animals. Permanently, social spiders form highly related groups with reproductive skew and communal brood care. We investigated task differentiation in nonreproductive tasks in two permanently and independently derived social spider species asking the following questions: Do individual spiders vary consistently in their propensity to engage in prey attack? Are individual spiders' propensities to engage in web maintenance behaviour influenced by their previous engagement in prey attack? Interestingly, we found that both species showed some degree of task specialization, but in distinctly different ways: Stegodyphus sarasinorum showed behavioural asymmetries at the individual level, that is, individual spiders that had attacked prey once were more likely to attack prey again, independent of their body size or hunger level. In contrast, Anelosimus eximius showed no individual specialization, but showed differentiation according to instar, where adult and subadult females were more likely to engage in prey attack than were juveniles. We found no evidence for division of labour between prey attack and web maintenance. Different solutions to achieve task differentiation in prey attack for the two species studied here suggest an adaptive value of task specialization in foraging for social spiders.

Characterization of microsatellite loci in the subsocial spider Stegodyphus lineatus (Araneae: Eresidae).

Stegodyphus lineatus spiders live in groups consisting of closely related individuals. There appears to be no discrimination against related individuals as mates but females mate multiply, despite the fact that matings are shown to carry a cost. We have developed eight polymorphic dinucleotide microsatellite markers that allow us to assess levels of heterozygosity and relatedness among individuals of this species. These molecular markers are likely to prove highly effective tools for estimating levels of inbreeding and thus allow us to test hypotheses about the relationships between social structure, mating strategies and inbreeding avoidance.

The genetic architecture of fitness in a seed beetle: assessing the potential for indirect genetic benefits of female choice.

BACKGROUND: Quantifying the amount of standing genetic variation in fitness represents an empirical challenge. Unfortunately, the shortage of detailed studies of the genetic architecture of fitness has hampered progress in several domains of evolutionary biology. One such area is the study of sexual selection. In particular, the evolution of adaptive female choice by indirect genetic benefits relies on the presence of genetic variation for fitness. Female choice by genetic benefits fall broadly into good genes (additive) models and compatibility (non-additive) models where the strength of selection is dictated by the genetic architecture of fitness. To characterize the genetic architecture of fitness, we employed a quantitative genetic design (the diallel cross) in a population of the seed beetle Callosobruchus maculatus, which is known to exhibit post-copulatory female choice. From reciprocal crosses of inbred lines, we assayed egg production, egg-to-adult survival, and lifetime offspring production of the outbred F1 daughters (F1 productivity). RESULTS: We used the bio model to estimate six components of genetic and environmental variance in fitness. We found sizeable additive and non-additive genetic variance in F1 productivity, but lower genetic variance in egg-to-adult survival, which was strongly influenced by maternal and paternal effects. CONCLUSION: Our results show that, in order to gain a relevant understanding of the genetic architecture of fitness, measures of offspring fitness should be inclusive and should include quantifications of offspring reproductive success. We note that our estimate of additive genetic variance in F1 productivity (CVA=14%) is sufficient to generate indirect selection on female choice. However, our results also show that the major determinant of offspring fitness is the genetic interaction between parental genomes, as indicated by large amounts of non-additive genetic variance (dominance and/or epistasis) for F1 productivity. We discuss the processes that may maintain additive and non-additive genetic variance for fitness and how these relate to indirect selection for female choice.

Benefits of cooperation with genetic kin in a subsocial spider.

Interaction within groups exploiting a common resource may be prone to cheating by selfish actions that result in disadvantages for all members of the group, including the selfish individuals. Kin selection is one mechanism by which such dilemmas can be resolved. This is because selfish acts toward relatives include the cost of lowering indirect fitness benefits that could otherwise be achieved through the propagation of shared genes. Kin selection theory has been proved to be of general importance for the origin of cooperative behaviors, but other driving forces, such as direct fitness benefits, can also promote helping behavior in many cooperatively breeding taxa. Investigating transitional systems is therefore particularly suitable for understanding the influence of kin selection on the initial spread of cooperative behaviors. Here we investigated the role of kinship in cooperative feeding. We used a cross-fostering design to control for genetic relatedness and group membership. Our study animal was the periodic social spider Stegodyphus lineatus, a transitional species that belongs to a genus containing both permanent social and periodic social species. In S. lineatus, the young cooperate in prey capture and feed communally. We provide clear experimental evidence for net benefits of cooperating with kin. Genetic relatedness within groups and not association with familiar individuals directly improved feeding efficiency and growth rates, demonstrating a positive effect of kin cooperation. Hence, in communally feeding spiders, nepotism favors group retention and reduces the conflict between selfish interests and the interests of the group.

Survival benefits select for group living in a social spider despite reproductive costs.

The evolution of cooperation requires benefits of group living to exceed costs. Hence, some components of fitness are expected to increase with increasing group size, whereas others may decrease because of competition among group members. The social spiders provide an excellent system to investigate the costs and benefits of group living: they occur in groups of various sizes and individuals are relatively short-lived, therefore life history traits and Lifetime Reproductive Success (LRS) can be estimated as a function of group size. Sociality in spiders has originated repeatedly in phylogenetically distant families and appears to be accompanied by a transition to a system of continuous intra-colony mating and extreme inbreeding. The benefits of group living in such systems should therefore be substantial. We investigated the effect of group size on fitness components of reproduction and survival in the social spider Stegodyphus dumicola in two populations in Namibia. In both populations, the major benefit of group living was improved survival of colonies and late-instar juveniles with increasing colony size. By contrast, female fecundity, female body size and early juvenile survival decreased with increasing group size. Mean individual fitness, estimated as LRS and calculated from five components of reproduction and survival, was maximized for intermediate- to large-sized colonies. Group living in these spiders thus entails a net reproductive cost, presumably because of an increase in intra-colony competition with group size. This cost is traded off against survival benefits at the colony level, which appear to be the major factor favouring group living. In the field, many colonies occur at smaller size than expected from the fitness curve, suggesting ecological or life history constraints on colony persistence which results in a transient population of relatively small colonies.

Inbreeding avoidance in spiders: evidence for rescue effect in fecundity of female spiders with outbreeding opportunity.

Selection by inbreeding depression should favour mating biases that reduce the risk of fertilization by related mates. However, equivocal evidence for inbreeding avoidance questions the strength of inbreeding depression as a selective force in the evolution of mating biases. Lack of inbreeding avoidance can be because of low risk of inbreeding, variation in tolerance to inbreeding or high costs of outbreeding. We examined the relationship between inbreeding depression and inbreeding avoidance adaptations under two levels of inbreeding in the spider Oedothorax apicatus, asking whether preference for unrelated sperm via pre- and/or post-copulatory mechanisms could restore female fitness when inbreeding depression increases. Using inbred isofemale lines we provided female spiders with one or two male spiders of different relatedness in five combinations: one male sib; one male nonsib; two male sibs; two male nonsibs; one male sib and one male nonsib. We assessed the effect of mating treatment on fecundity and hatching success of eggs after one and three generations of inbreeding. Inbreeding depression in F1 was not sufficient to detect inbreeding avoidance. In F3, inbreeding depression caused a major decline in fecundity and hatching rates of eggs. This effect was mitigated by complete recovery in fecundity in the sib-nonsib treatment, whereas no rescue effect was detected in the hatching success of eggs. The rescue effect is best explained by post-mating discrimination against kin via differential allocation of resources. The natural history of O. apicatus suggests that the costs of outbreeding may be low which combined with high costs of inbreeding should select for avoidance mechanisms. Direct benefits of post-mating inbreeding avoidance and possibly low costs of female multiple mating can favour polyandry as an inbreeding avoidance mechanism.

Tissue culture supernatants from human islets of Langerhans activate the oxidative burst response of human monocytes in vitro.

Macrophages play a major role in the pathogenesis of insulin-dependent diabetes mellitus in animals. These cells are the first to invade the pancreas and macrophage-eradicating treatments reduce the incidence of the disease. In humans, however, their role is less clear. In this study we investigated the hypothesis that the pancreatic environment per se could activate macrophages. Tissue culture supernatants from human islets of Langerhans were tested for chemotactic activity and oxidative burst response in monocytes isolated from healthy adults. Preincubation with the supernatants enhanced the oxidative burst response evoked by fMLP (up to 379%) and opsonized zymosan (up to 173%). The activity decreased by dilution and was no longer detectable at 1:16. No increased activity was seen in supernatants from a number of other human endocrine and non-endocrine primary cells, suggesting a factor specific for islet tissue. The increased oxidative burst response could partially be eliminated by heat- and proteinase K treatment, suggesting that the activity could be of polypeptide nature. The factor could not be absorbed by polyvalent rabbit antibodies directed towards a variety of cytokines not by a mixture of high-titer anti-cytokine antibodies. It is possible that islet factors could also promote such monocyte activation in vivo in monocytes attracted to the islets of Langerhans by other means. This could contribute to the development of insulin-dependent diabetes in humans.

Retroperitoneal fibrosis treated with methylprednisolon pulse and disease-modifying antirheumatic drugs.

The conventional treatment of patients with ureteric obstruction due to retroperitoneal fibrosis (RF) is surgery in combination with long-term corticosteroids. This report describes 11 cases of RF with ureteric obstruction treated with methyl-prednisolon pulse therapy (MPPT) in combination with azathioprine or penicillamine following initial insertion of ureteral stents. The medial treatment suggested was successful in 7 patients, but only moderately effective in the last 4 patients. This principle of non-operative management of RF has not been previously reported.

Trends in a Danish urologic department during the years 1979 and 1984.

During recent years several international studies have shown increasing rates of prostatectomy over the past decade. In Denmark the number of prostatectomies increased by 43% in the period 1977-85. This development is still unexplained, but one of the answers is very intriguing, proposing a shift in indications for prostatectomy. Therefore we retrospectively examined the records of 207 patients in 1979 and 243 patients in 1984, admitted with a diagnosis of benign hyperplasia of the prostate (BPH). In these patients respectively 164 and 149 prostatectomies were performed in the same years, at the Department of Urology, Glostrup County hospital. We found no evidence of a change in indications towards a more liberal attitude as suspected, but in contrast, we discovered a more conservative attitude. Because the national rates since 1984 have been stable, we conclude, that the development in a specialized urologic Department can serve as a predictor for the development on a larger national scale.

A new patient weighted symptom score system (DAN-PSS-1). Clinical assessment of indications and outcomes of transurethral prostatectomy for uncomplicated benign prostatic hyperplasia.

A patient weighted symptom score system, the Danish Prostatic Symptom Score (DAN-PSS-1), including a disease specific self administered quality of life questionnaire, is presented. The model was evaluated pre- and postoperatively in 29 patients apparently suffering from uncomplicated benign prostatic hyperplasia. The score system is based on the severity of 12 symptoms related to bladder storage and voiding function, and three questions related to sexual function (symptom score). For each of these parameters the patient must also evaluate its influence on his daily life (bother score). In the 29 patients with uncomplicated benign prostatic hyperplasia (BPH) bother scores exceeded symptom scores for the irritative symptoms but not for the obstructive symptoms, and surprisingly the symptom score was less improved than the bother score 6 months after transurethral resection of the prostate (TUR-P). Furthermore the postvoiding dribble was worsened after the operation. We find that this model, DAN-PSS-1, assists in creating a solid base for the indication for and the evaluation of treatment of uncomplicated BPH.

A patient weighted symptom score system in the evaluation of uncomplicated benign prostatic hyperplasia.

A new scoring system is presented for use in patients with uncomplicated benign prostatic hyperplasia. The system is based on the patients' information on the magnitude of twelve symptoms related to bladder and voiding function (symptom score), and three questions related to sexual function. Furthermore the patients are asked to grade the impact of these symptoms on their daily lives (bother score). Multiplication of the two scores related to bladder and voiding function gives the total score. The score related to sexual function is used only for comparing the situation before and after treatment. The new patient weighted total score may assist in creating a solid base for the indication for and evaluation of surgical and non-surgical treatment of uncomplicated benign prostatic hyperplasia.

Prostatectomy in Denmark. Regional variation and the diffusion of medical technology 1977-1985.

In many countries prostatectomy is one of the most common surgical operations in elderly men. We used administrative data for the entire male population of Denmark to study temporal and regional variations in the use of prostatectomy from 1977 to 1985. The total annual number of prostatectomies increased by 43% during the period, when the transurethral procedure (TURP) gradually replaced traditional open surgery. TURP accounted for 56% of all operations in 1977 but increased its share to 92% in 1985. Substantial regional variations occurred with index values for prostatectomy in 72 recruitment areas ranging from 0.56 to 1.62 (SCV x 100 = 5.3). The amount of variation decreased during the process of technology diffusion, but remained at a relatively high level (as in other countries) even after the process had been completed (SCV x 100 = 5.1).