Social evolution in the sweat bee Halictus scabiosae

The evolution of eusociality is one of the major evolutionary transitions of life on earth. For investigating the conditions and processes that are central to the origin of such integrated social organization, it is best to study organisms in which individuals have retained some flexibility in their reproductive strategies. Halictid bees are especially well suited as model organisms, because they show huge variation in social systems, both within and between species. In this thesis, I investigated female reproductive strategies in the primitively eusocial bee Halictus scabiosae, with a focus on the role of helpers, in order to get insight into the mechanisms governing the evolution and maintenance of eusociality. This species produces two broods per year. The females from the first brood can stay in the natal nest to help raise a second brood of males and gynes that become the next-generation foundresses in spring. We first compared the morphology of females from the two broods, as well as the nutrition they receive as larvae. Then we conducted a helper- removal experiment in the field to quantify the effects of the presence of helpers on colony survival and productivity. Finally, we reconstructed pedigree relationships of individuals using microsatellite markers in order to detect who reproduces in the nest and how much individuals drift between nests. We found that first brood females had a uniformly small size and low fat reserves, which may be caused by the restricted pollen and nectar provisions on which they develop. Colony survival and productivity was increased by the presence of a single helper, but the effect was small and mostly limited to small colonies. By inferring parentage within and across colonies, we could determine that females from the first brood rarely reproduce in their natal nests. However, foundresses are frequently replaced, and foundresses and females from the first brood occasionally move to and reproduce in foreign colonies. As a result, colonies often contain offspring from unrelated individuals, and the relatedness of females to the brood they rear is low. Overall, this thesis shows that the reproductive system of H. scabiosae is highly flexible. The production of helpers in the first brood is important for colony success and productivity, but there is a high colony failure rate and part of the first brood females drift and reproduce in foreign nests. Both foundresses and helpers appear to be constrained by harsh environmental conditions or social factors limiting reproduction and independent colony founding. - L'origine des insectes sociaux est un domaine fascinant pour la recherche. Pour comprendre les mécanismes et les conditions qui sont nécessaires pour l'évolution et le maintien de la vie en société, il est judicieux d'étudier des sociétés primitives d'insectes, où toutes les femelles ont conservé la capacité de se reproduire, même si leur rôle comportemental dans la colonie est d'aider sans se reproduire. Une des familles d'abeilles, les halictes, est idéale pour cette sorte de recherche, en raison de la grande variabilité dans leur comportement social. Dans cette thèse, j'ai étudié les stratégies reproductives des femelles de Halictus scabiosae pour mieux comprendre les mécanismes qui influencent l'évolution de la vie en société. Cette espèce produit deux cohortes de couvain par année. Les femelles du premier couvain restent souvent dans leur nid natal pour aider à élever le deuxième couvain, tandis que les femelles du deuxième couvain s'accouplent et hibernent pour devenir les nouvelles fondatrices au printemps suivant. Nous avons d'abord comparé la morphologie des femelles issues des deux couvains ainsi que leur nutrition au stade de larve. Puis, dans une expérience sur le terrain, nous avons quantifié l'apport d'une ouvrière pour la survie et la productivité de la colonie. Finalement, nous avons reconstruit des pedigrees en utilisant des marqueurs génétiques, pour savoir qui se reproduit dans la colonie et combien d'individus migrent entre colonies. Les résultats montrent que les femelles du premier couvain sont uniformément plus petites et plus maigres, ce qui indique que les fondatrices réduisent les provisions de nourriture pour leur premier couvain afin de les inciter à aider dans le nid au lieu de se reproduire indépendamment. Dans l'expérience sur le terrain, la survie et la productivité de la colonie augmentaient avec la présence d'une ouvrière additionnelle, mais l'effet était petit et limité aux petites colonies. Par la reconstruction de pedigrees, nous pouvions constater que les femelles du premier couvain pondent rarement dans leurs nids natals. Les fondatrices cependant sont souvent remplacées en cours de saison, et migrent fréquemment entre nids, tandis que les femelles du premier couvain pondent parfois des oeufs dans des nids étrangers. De ce fait, les colonies contiennent souvent des descendants d'individus étrangers, et la parenté génétique entre les femelles et le deuxième couvain est basse. Cette thèse démontre que le système reproductif de H. scabiosae est très flexible. La production d'ouvrières est importante pour la survie de la colonie et sa productivité, mais le taux d'échec est élevé et une partie des femelles du premier couvain migrent et pondent dans une colonie étrangère. Autant les fondatrices que les ouvrières semblent être contraintes par des conditions environnementales ou sociales qui limitent la reproduction et les nouvelles fondations de colonie. - Die Entstehung von sozialen Lebensformen ist eines der wichtigsten Entwicklungen in der Geschichte des Lebens. Um die Bedingungen oder Prozesse zu verstehen, welche bei der Entstehung und dem Erhalt von sozialen Merkmalen wichtig sind, sollte man Lebewesen untersuchen, welche je nach Umwelteinflüßen ihr soziales Verhalten flexibel ändern können. Furchenbienen (Halictidae) gehören dazu. Diese weisen nämlich ein breites Spektrum verschiedener sozialer Organisationsformen auf, oftmals sogar innerhalb der einzelnen Arten. In meiner Doktorarbeit befasste ich mich mit den Fortpflanzungsstrategien der Weibchen der Skabiosen-Furchenbiene Halictus scabiosae. Diese Art produziert zwei Brüten pro Jahr. Die Weibchen der ersten Brut bleiben dabei meist als Arbeiterinnen in ihrem Geburtsnest, wohingegen die Weibchen der zweiten Brut nach der Paarung überwintern, um im nächsten Frühling neue Kolonien zu gründen. In einem ersten Schritt verglichen wir die beiden Brüten bezüglich der Grösse und der Fettreserven der Weibchen sowie der Pollen-Nektar-Vorräte für die Larven. Dann bestimmten wir in einem Feldexperiment, wieviel eine zusätzliche Arbeiterin zum Überleben und zur Produktiviät der Kolonie beiträgt. Schliesslich ermittelten wir durch genetische Tests die Verwandtschaftsbeziehungen zwischen den Bienen, um herauszufinden, wer in den Kolonien tatsächlich die Eier legt und ob und wieviel die Bienen zwischen verschiedenen Nestern wandern. Wir stellten fest, dass die Weibchen von der ersten Brut einheitlich kleiner sind und weniger Fettreserven besitzen. Das weist daraufhin, dass die Nestgründerin die erste Brut unterernährt, um die Wahrscheinlichkeit zu erhöhen, dass diese Weibchen als Arbeiterinnen im Nest bleiben anstatt sich unabhängig fortzupflanzen. Schon eine einzelne zusätzliche Arbeiterin verbesserte die Überlebenschancen und Produktivität der Kolonie, der Effekt war allerdings klein und auf kleine Kolonien beschränkt. Die Verwandtschaftsanalysen zeigten, dass die Arbeiterinnen nur sehr selten ein Ei in ihr Geburtsnest legen. Erstaunlicherweise wanderten die Nestgründerinnen oft zwischen verschiedenen Nestern. Einige Weibchen der ersten Brut wanderten auch in ein fremdes Nest und produzierten dort Nachkommen. Diese Doktorarbeit zeigt, dass die Fortpflanzungsstrategien der Skabiosen-Furchenbiene tatsächlich sehr flexibel sind. Die Anwesenheit von Arbeiterinnen ist wichtig für das Überleben und die Produktivität der Kolonie. Die Misserfolgsraten bleiben jedoch hoch, und ein Teil der Weibchen der ersten Brut pflanzt sich in fremden Nestern fort. Sowohl die Nestgründerinnen als auch die Weibchen der ersten Brut scheinen durch Umweltsbedingungen oder durch soziale Faktoren in der Wahl ihrer Fortpflanzungs¬strategie eingeschränkt zu sein.

Structured RNAs in the ENCODE selected regions of the human genome.

Functional RNA structures play an important role both in the context of noncoding RNA transcripts as well as regulatory elements in mRNAs. Here we present a computational study to detect functional RNA structures within the ENCODE regions of the human genome. Since structural RNAs in general lack characteristic signals in primary sequence, comparative approaches evaluating evolutionary conservation of structures are most promising. We have used three recently introduced programs based on either phylogenetic-stochastic context-free grammar (EvoFold) or energy directed folding (RNAz and AlifoldZ), yielding several thousand candidate structures (corresponding to approximately 2.7% of the ENCODE regions). EvoFold has its highest sensitivity in highly conserved and relatively AU-rich regions, while RNAz favors slightly GC-rich regions, resulting in a relatively small overlap between methods. Comparison with the GENCODE annotation points to functional RNAs in all genomic contexts, with a slightly increased density in 3'-UTRs. While we estimate a significant false discovery rate of approximately 50%-70% many of the predictions can be further substantiated by additional criteria: 248 loci are predicted by both RNAz and EvoFold, and an additional 239 RNAz or EvoFold predictions are supported by the (more stringent) AlifoldZ algorithm. Five hundred seventy RNAz structure predictions fall into regions that show signs of selection pressure also on the sequence level (i.e., conserved elements). More than 700 predictions overlap with noncoding transcripts detected by oligonucleotide tiling arrays. One hundred seventy-five selected candidates were tested by RT-PCR in six tissues, and expression could be verified in 43 cases (24.6%).

Evolutionary genomics and HIV restriction factors.

PURPOSE OF REVIEW: To provide updated insights into innate antiviral immunity and highlight prototypical evolutionary features of well characterized HIV restriction factors. RECENT FINDINGS: Recently, a new HIV restriction factor, Myxovirus resistance 2, has been discovered and the region/residue responsible for its activity identified using an evolutionary approach. Furthermore, IFI16, an innate immunity protein known to sense several viruses, has been shown to contribute to the defense to HIV-1 by causing cell death upon sensing HIV-1 DNA. SUMMARY: Restriction factors against HIV show characteristic signatures of positive selection. Different patterns of accelerated sequence evolution can distinguish antiviral strategies--offense or defence--as well as the level of specificity of the antiviral properties. Sequence analysis of primate orthologs of restriction factors serves to localize functional domains and sites responsible for antiviral action. We use recent discoveries to illustrate how evolutionary genomic analyses help identify new antiviral genes and their mechanisms of action.

New insights into the phylogenetics and biogeography of Arum (Araceae): unravelling its evolutionary history

The heat- and odour-producing genus Arum (Araceae) has interested scientists for centuries. This long-term interest has allowed a deep knowledge of some complex processes, such as the physiology and dynamics of its characteristic lure-and-trap pollination system, to be built up. However, mainly because of its large distributional range and high degree of morphological variation, species' limits and relationships are still under discussion. Today, the genus comprises 28 species subdivided into two subgenera, two sections and six subsections. In this study, the phylogeny of the genus is inferred on the basis of four plastid regions, and the evolution of several morphological characters is investigated. Our phylogenetic hypothesis is not in agreement with the current infrageneric classification of the genus and challenges the monophyly of several species. This demonstrates the need for a new infrageneric classification based on characters reflecting the evolution of this enigmatic genus. To investigate the biogeography of Arum deeply, further spatiotemporal analyses were performed, addressing the importance of the Mediterranean basin in the diversification of Arum. Our results suggest that its centre of origin was the European-Aegean region, and that major diversification happened during the last 10 Myr.

Coloration signals the ability to cope with elevated stress hormones: effects of corticosterone on growth of barn owls are associated with melanism.

Stressful situations during development can shape the phenotype for life by provoking a trade-off between development and survival. Stress hormones, mainly glucocorticoids, play an important orchestrating role in this trade-off. Hence, how stress sensitive an animal is critically determines the phenotype and ultimately fitness. In several species, darker eumelanic individuals are less sensitive to stressful conditions than less eumelanic conspecifics, which may be due to the pleiotropic effects of genes affecting both coloration and physiological traits. We experimentally tested whether the degree of melanin-based coloration is associated with the sensitivity to an endocrine response to stressful situations in the barn owl. We artificially administered the mediator of a hormonal stress response, corticosterone, to nestlings to examine the prediction that corticosterone-induced reduction in growth rate is more pronounced in light eumelanic nestlings than in darker nest mates. To examine whether such an effect may be genetically determined, we swapped hatchlings between randomly chosen pairs of nests. We first showed that corticosterone affects growth and, thus, shapes the phenotype. Second, we found that under corticosterone administration, nestlings with large black spots grew better than nestlings with small black spots. As in the barn owl the expression of eumelanin-based coloration is heritable and not sensitive to environmental conditions, it is therefore a reliable, genetically based sign of the ability to cope with an increase in blood corticosterone level.

The evolution of inbred social systems in spiders and other organisms: from short-term gains to long-term evolutionary dead-ends?

The question of why some social systems have evolved close inbreeding is particularly intriguing given expected short- and long-term negative effects of this breeding system. Using social spiders as a case study, we quantitatively show that the potential costs of avoiding inbreeding through dispersal and solitary living could have outweighed the costs of inbreeding depression in the origin of inbred spider sociality. We further review the evidence that despite being favored in the short term, inbred spider sociality may constitute in the long run an evolutionary dead end. We also review other cases, such as the naked mole rats and some bark and ambrosia beetles, mites, psocids, thrips, parasitic ants, and termites, in which inbreeding and sociality are associated and the evidence for and against this breeding system being, in general, an evolutionary dead end.

Reduction of small infrarenal abdominal aortic aneurysm expansion rate by statins.

BACKGROUND: To evaluate the effect of statins on the annual expansion rate (ER) of small infrarenal abdominal aortic aneurysms (AAA). PATIENTS AND METHODS: All patients under regular surveillance for small AAA between January 2000 and September 2007, in the Department of Angiology, Lausanne University Hospital, were included. Inclusion criteria were baseline abdominal aortic diameter between 25 and 55 mm, at least two measurements of AAA diameter and a minimum follow up of 6 months. Patients with Marfan disease, infectious or inflammatory AAA, and patients with prior AAA repair were excluded. The influence of statin use and other factors on ER were examined by bivariate and multivariate analysis. RESULTS: Among 589 patients who underwent an abdominal aorta evaluation, 94 patients (89 % men, mean age 69.1 years) were finally included in the analysis. Baseline AAA size was 39.9 ± 7.7 mm (mean±SE) and 48.7 ± 8.4 mm at end of follow-up. Patients had a regular aneurysm size assessment during 38.5 ± 27.7 months. Mean ER was 3.59 mm/y (± 2.81). The 50 patients who were treated with statin during the study period had a lower ER compared to the 44 controls (2.91 vs 4.37 mm/year, p = 0.01). CONCLUSIONS: This study confirms the considerable individual variations in the AAA expansion rate, and emphasizes the need for regular aortic diameter assessments. In this study, patients treated with statin demonstrate a significant decrease in the ER compared to controls. This finding need to be evaluated in prospective interventional studies powered to demonstrate the potential benefit of statin treatment.

Gains from switching and evolutionary stability in multi-player matrix games.

In this paper we unify, simplify, and extend previous work on the evolutionary dynamics of symmetric N-player matrix games with two pure strategies. In such games, gains from switching strategies depend, in general, on how many other individuals in the group play a given strategy. As a consequence, the gain function determining the gradient of selection can be a polynomial of degree N-1. In order to deal with the intricacy of the resulting evolutionary dynamics, we make use of the theory of polynomials in Bernstein form. This theory implies a tight link between the sign pattern of the gains from switching on the one hand and the number and stability of the rest points of the replicator dynamics on the other hand. While this relationship is a general one, it is most informative if gains from switching have at most two sign changes, as is the case for most multi-player matrix games considered in the literature. We demonstrate that previous results for public goods games are easily recovered and extended using this observation. Further examples illustrate how focusing on the sign pattern of the gains from switching obviates the need for a more involved analysis.

Stochastic demography and the neutral substitution rate in class-structured populations.

The neutral rate of allelic substitution is analyzed for a class-structured population subject to a stationary stochastic demographic process. The substitution rate is shown to be generally equal to the effective mutation rate, and under overlapping generations it can be expressed as the effective mutation rate in newborns when measured in units of average generation time. With uniform mutation rate across classes the substitution rate reduces to the mutation rate.

Epistasis and maternal effects in experimental adaptation to chronic nutritional stress in Drosophila.

Based on ecological and metabolic arguments, some authors predict that adaptation to novel, harsh environments should involve alleles showing negative (diminishing return) epistasis and/or that it should be mediated in part by evolution of maternal effects. Although the first prediction has been supported in microbes, there has been little experimental support for either prediction in multicellular eukaryotes. Here we use a line-cross design to study the genetic architecture of adaptation to chronic larval malnutrition in a population of Drosophila melanogaster that evolved on an extremely nutrient-poor larval food for 84 generations. We assayed three fitness-related traits (developmental rate, adult female weight and egg-to-adult viability) under the malnutrition conditions in 14 crosses between this selected population and a nonadapted control population originally derived from the same base population. All traits showed a pattern of negative epistasis between alleles improving performance under malnutrition. Furthermore, evolutionary changes in maternal traits accounted for half of the 68% increase in viability and for the whole of 8% reduction in adult female body weight in the selected population (relative to unselected controls). These results thus support both of the above predictions and point to the importance of nonadditive effects in adaptive microevolution.

Pregnancy-related changes in activity energy expenditure and resting metabolic rate in Switzerland.

BACKGROUND/OBJECTIVES: To measure resting metabolic rate (RMR), activity energy expenditure (AEE), total energy expenditure (TEE) and physical activity pattern, that is, duration and intensity (in metabolic equivalents, METs) of activities performed in late pregnancy compared with postpartum in healthy, well-nourished women living in Switzerland. SUBJECTS/METHODS: Weight, height, RMR, AEE, TEE and physical activity patterns were measured longitudinally in 27 healthy women aged 23-40 years at 38.2+/-1.5 weeks of gestation and 40.0+/-7.2 weeks postpartum. RESULTS: The RMR during late pregnancy was 7480 kJ per day, that is, 1320+/-760 kJ per day (21.4%) higher than the postpartum RMR (P<0.001). Absolute changes in RMR were positively correlated with the corresponding changes in body weight (r=0.61, P<0.001). RMR per kg body weight was similar in late pregnancy vs postpartum (P=0.28). AEE per kg during pregnancy and postpartum was 40+/-13 and 50+/-20 kJ/kg, respectively (P=0.001). There were significant differences in daily time spent at METs<1.5 (1067 vs 998 min, P=0.045), at 2.5< or =METs <3.0 (58 vs 82 min, P=0.002) and METs> or =6 (1 vs 6 min, P=0.014) during pregnancy and postpartum, respectively. CONCLUSIONS: Energy expenditure in healthy women living in Switzerland increases in pregnancy compared with the postpartum state. Additional energy expenditure is primarily attributed to an increase in RMR, which is partly compensated by a decrease in AEE. The decrease in physical activity-related energy costs is achieved by selecting less demanding activities and should be taken into account when defining extra energy requirements for late pregnancy in Switzerland.

Parental heart rate variability during pediatric consultation

OBJECTIVE: Research regarding communication between pediatricians and parents in pediatric consultation has mainly focused on parental satisfaction, on its influence on compliance and on communication techniques used by pediatricians. However, there is paucity in research regarding parental stress levels during pediatric consultation. Therefore, the aim of our study was to measure parental heart rate variability related as a measure of stress levels during pediatric consultation. METHODS: Video recordings with simultaneous monitoring and recording of parental heart rate were obtained from 38 pediatric consultations in the ambulatory or hospital setting of the department of pediatrics (HFR, Fribourg, Switzerland). Pulse variation was measured every 5 seconds and heart rate variability (increase or decrease were analyzed) in relation to various sections of the consultation. RESULTS: Heart rate significantly decreased at the end of the consultation compared to the beginning of the consultation (p= 0.0567). In addition, heart rate significantly decreased at the beginning of clinical examination (p= 0.0476) compared to psychosocial history taking. During the discussion of laboratory findings and diagnosis, heart rate was significantly elevated compared to the discussion of the prognosis (p=0.0505). CONCLUSION: We conclude that pediatric consultation has a significant impact on parental stress levels shown by parental heart variability. In general, it can be shown that stress levels significantly decrease at the end of the consultation compared to the beginning of the consultation. In addition, stress levels decrease at the beginning of clinical examination and increase during psychosocial history taking and discussion of laboratory findings and diagnosis. Therefore, our findings highlight the importance of a thorough consultation which include a comprehensive clinical examination with special care taken regarding psychosocial issues and information given regarding the diagnosis.