Mechanisms involved in the maintenance of inbreeding depression in gynodioecious "Silene vulgaris" (Caryophyllaceae): an experimental investigation

Summary Gynodioecy, the joint occurrence of females and hermaphrodites within natural populations, is a widely studied mating system ever since Darwin (1877). It is an exceptional mating system because continuous selection is necessary to maintain it. Since females only reproduce through ovules whereas hermaphrodites transmit genes through ovules and pollen, larger female fitness, in terms of seed output, is required to allow their maintenance. Two non-exclusive mechanisms can account for the maintenance of females. First, as females do not produce pollen they can reallocate their resources towards a higher ovule production. Second, hermaphrodites can self- and cross-fertilize whereas females are obligate outcrossers. Thus hermaphrodites should partly suffer from inbreeding depression (i.e.: the fitness decline of inbred relative to outbred individuals) and thereby produce less fit progeny than females. This thesis investigated the effects of self- and cross-fertilization of heimaphrodites over two consecutive generations. Inbreeding depression increased across the successive stages of the life- cycle (i.e.: from "seed traits" to "reproductive traits") displaying large inbreeding depression estimates (up to 0.76). This investigation not only detected large inbreeding depression estimates but also detected mechanisms involved in the maintenance of inbreeding depression. For instance cryptic self-incompatibility which is here a larger in vivo pollen performance of distant pollen compared to self-pollen; the expression of inbreeding depression especially in late life-cycle stages, and the appearance of females in the progeny of selfed hermaphrodites. The female biased sex ratio in the progeny of selfed hermaphrodites was a surprising result and could either come from the sex determining mechanisms (complex nucleo-cytoplasmic interaction(s)) and/or from inbreeding depression. Indeed, we not only got females and hermaphrodites but also partial male-sterile (PMS) individuals (i.e.: individuals with differing number of viable stamens). We detected that inbred pollen bearing plants (excluding females) have less viable stamens per flower than outbred plants. A positive correlation was detected between inbreeding depression for the number of viable stamens per flower and the difference in sex ratio between inbred and outbred individuals. A positive relationship was also detected between inbreeding depression for pollen viability and inbreeding depression for number of viable stamens per flower. Each correlation can either account for pleiotropic effects (a major gene acting on the two considered traits) or linkage disequilibrium between genes controlling each of the two related traits. If we hypothesize that these correlations are due to a major gene with pleiotropic effects, the positive relationship between inbreeding depression for number of viable stamens per flower and inbreeding depression for pollen viability showed that deleterious alleles present on a major gene coding for pollen production and viability depressed male fitness within inbred plants. The positive relationship between sex ratio difference between inbred and outbred individuals and inbreeding depression for number of viable stamens per flower indicates that (1) either number of viable stamens per flower is, in addition to inbreeding, also affected by the loci coding for sex determinism or, (2) the presence of females within the progeny of selfed hermaphrodites is a consequence of large inbreeding depression inhibiting pollen production, or (3) sex is here determined by a combination of loci coding for sex expression and inbreeding depression for male reproductive traits. In conclusion, Silene vulgaris has been shown to be a good model for understanding the evolution of mating systems that promote outbreeding. Résumé La gynodïoécie est définie comme étant la présence simultanée d'hermaphrodites et de femelles au sein de populations naturelles d'une même espèce. Ce système de reproduction a toujours fasciné le monde scientifique depuis Darwin, comme en témoigne ses écrits (1876, 1877) sur les systèmes de reproduction chez les plantes. Les femelles ne transmettent leurs gènes qu'à travers leurs ovules alors que les hermaphrodites transmettent leurs gènes à la fois par la voie mâle (le pollen) et la voie femelle (les ovules). La condition pour que la gynodïoécie se maintienne nécessite donc une fitness de la fonction femelle plus élevée chez les femelles que chez les hermaphrodites. Deux mécanismes mutuellement non exclusifs peuvent expliquer le maintien des femelles au sein de ces populations gynodioïques. D'une part, les femelles peuvent réallouer les ressources non utilisées pour la production de pollen et peuvent par conséquent produire plus d'ovules. D'autre part, la reproduction des femelles ne peut se faire que par allo-fécondation alors que les hermaphrodites, peuvent se reproduire à la fois par auto- et allo-fécondation. L'autofécondation s'accompagne en général d'une diminution de fitness de la descendance relativement à la progéniture issue d'allo-fécondation ; ce phénomène est connu sous le nom de dépression de consanguinité. Cette thèse avait pour but de mettre en évidence une éventuelle dépression de consanguinité chez Silene vulgaris, une espèce gynodioïque. Des hermaphrodites, issus de trois vallées alpines, ont été auto- et allo¬fécondés sur deux générations successives. La dépression de consanguinité pouvant s'exprimer à tous les stades de vie d'un individu, plusieurs traits de fitness, allant du nombre de graines par fruit à la production de gamètes ont été mesurés sur différents stades de vie successifs. L'estimation de la dépression de consanguinité totale atteignait des valeurs allant de 0.52 à 0.76 selon la vallée considérée, ce qui indiquerait que les hermaphrodites ont tout intérêt à limiter l'autofécondation et que les femelles ne devraient pas avoir de peine à subsister dans les vallées étudiées. Par la même occasion des mécanismes diminuant la purge potentielle du fardeau génétique, et permettant ainsi le maintien du « niveau » de dépression de consanguinité et par conséquence le maintien de la gynodïoécie ont été mis en évidence. En effet, nos résultats montrent que la dépression de consanguinité s'exprimait tard dans le cycle de vie permettant ainsi à un certain nombre individus consanguins de transmettre leurs allèles délétères à la génération suivante. D'autre part, la croissance in vivo des tubes polliniques d'auto-pollen était plus lente que celle de l'allo-pollen et donc en situation de compétition directe, les ovules devraient plutôt être issus d'allo-fécondation, diminuant ainsi les chances de purges d'allèles délétères. Enfin, l'apparition de femelles dans la progéniture d'hermaphrodites autofécondés diminue aussi les chances de purge d'allèles délétères. Il nous a été impossible de déterminer si l'apparition de femelles dans la descendance d'hermaphrodites autofécondés était due au déterminisme génétique du sexe ou si la différence de sexe ratio entre la descendance auto- et allo-fécondée était due à une éventuelle dépression de consanguinité inhibant la production de pollen. Nous avons observé que S. vulgaris ne présentaient pas uniquement des hermaphrodites et des femelles mais aussi toute sorte d'individus intermédiaires avec un nombre variable d'étamines viables. Nous avons pu mettre' en évidence des corrélations positives entre (1) la différence de sexe ratio (la proportion d'individus produisant du pollen) entre individus consanguins et non consanguins et une estimation de la dépression de consanguinité pour le nombre d'étamines viables d'individus produisant du pollen, ainsi qu'entre (2) la dépression de consanguinité pour le nombre d'étamines viables et celle estimée pour la viabilité du pollen. Chaque corrélation indique soit l'effet d'un (ou plusieurs) gène(s) pléiotropique(s), soit un déséquilibre de liaison entre les gènes. En considérant que ces corrélations sont le résultat d'effet pléiotropiques, la relation entre le nombre d'étamines viables par fleur et la viabilité du pollen, indiquerait un effet négatif de la consanguinité sur la production et la viabilité du pollen due partiellement à un gène majeur. La seconde corrélation indiquerait soit que les gènes responsables de la détermination du sexe agissent aussi sur l'expression de la fonction mâle soit que l'expression du sexe est sujette à la dépression de consanguinité, ou encore un mélange des deux. Aux regards de ces résultats, Silene vulgaris s'est avéré être un bon modèle de compréhension de l'évolution des systèmes de reproduction vers la séparation des sexes.

Evaluation of Variable Advisory Speed Limits in Work Zones, TPF-5(081), 2013

Variable advisory speed limit (VASL) systems could be effective at both urban and rural work zones, at both uncongested and congested sites. At uncongested urban work zones, the average speeds with VASL were lower than without VASL. But the standard deviation of speeds with VASL was higher. The increase in standard deviation may be due to the advisory nature of VASL. The speed limit compliance with VASL was about eight times greater than without VASL. At the congested sites, the VASL were effective in making drivers slow down gradually as they approached the work zone, reducing any sudden changes in speeds. Mobility-wise the use of VASL resulted in a decrease in average queue length, throughput, number of stops, and an increase in travel time. Several surrogate safety measures also demonstrated the benefits of VASL in congested work zones. VASL deployments in rural work zones resulted in reductions in mean speed, speed variance, and 85th percentile speeds downstream of the VASL sign. The study makes the following recommendations based on the case studies investigated: 1. The use of VASL is recommended for uncongested work zones to achieve better speed compliance and lower speeds. Greater enforcement of regulatory speed limits could help to decrease the standard deviation in speeds; 2. The use of VASL to complement the static speed limits in rural work zones is beneficial even if the VASL is only used to display the static speed limits. It leads to safer traffic conditions by encouraging traffic to slow down gradually and by reminding traffic of the reduced speed limit. A well-designed VASL algorithm, like the P5 algorithm developed in this study, can significantly improve the mobility and safety conditions in congested work zones. The use of simulation is recommended for optimizing the VASL algorithms before field deployment.

"Suddenly I get into the zone": Examining discontinuities and nonlinear changes in flow experiences at work

Work-related flow is defined as a sudden and enjoyable merging of action and awareness that represents a peak experience in the daily lives of workers. Employees" perceptions of challenge and skill and their subjective experiences in terms of enjoyment, interest and absorption were measured using the experience sampling method, yielding a total of 6981 observations from a sample of 60 employees. Linear and nonlinear approaches were applied in order to model both continuous and sudden changes. According to the R2, AICc and BIC indexes, the nonlinear dynamical systems model (i.e. cusp catastrophe model) fit the data better than the linear and logistic regression models. Likewise, the cusp catastrophe model appears to be especially powerful for modelling those cases of high levels of flow. Overall, flow represents a nonequilibrium condition that combines continuous and abrupt changes across time. Research and intervention efforts concerned with this process should focus on the variable of challenge, which, according to our study, appears to play a key role in the abrupt changes observed in work-related flow.

Design Manual for Strengthening of Continuous-Span Composite Bridges, HR-333, 1993

The need for upgrading a large number of understrength bridges in the United States has been well documented in the literature. This manual presents two methods for strengthening continuous-span composite bridges: post-tensioning of the positive moment regions of the bridge stringers and the addition of superimposed trusses at the piers. The use of these two systems is an efficient method of reducing flexural overstresses in undercapacity bridges. Before strengthening a given bridge however, other deficiencies (inadequate shear connection, fatigue problems, extensive corrosion) should be addressed. Since continuous-span composite bridges are indeterminant structures, there is longitudinal and transverse distribution of the strengthening axial forces and moments. This manual basically provides the engineer with a procedure for determining the distribution of strengthening forces and moments throughout the bridge. As a result of the longitudinal and transverse force distribution, the design methodology presented in this manual for continuous-span composite bridges is extremely complex. To simplify the procedure, a spreadsheet has been developed for use by practicing engineers. This design aid greatly simplifies the design of a strengthening system for a given bridge in that it eliminates numerous tedious hand calculations, computes the required force and moment fractions, and performs the necessary iterations for determining the required strengthening forces. The force and moment distribution fraction formulas developed in this manual are primarily for the Iowa DOT V12 and V14 three-span four-stringer bridges. These formulas may be used on other bridges if they are within the limits stated in this manual. Use of the distribution fraction formulas for bridges not within the stated limits is not recommended.

A Feasibility Study on Embedded Micro-Electromechanical Sensors and Systems (MEMS) for Monitoring Highway Structures, TR-575, 2011

Micro-electromechanical systems (MEMS) provide vast improvements over existing sensing methods in the context of structural health monitoring (SHM) of highway infrastructure systems, including improved system reliability, improved longevity and enhanced system performance, improved safety against natural hazards and vibrations, and a reduction in life cycle cost in both operating and maintaining the infrastructure. Advancements in MEMS technology and wireless sensor networks provide opportunities for long-term continuous, real-time structural health monitoring of pavements and bridges at low cost within the context of sustainable infrastructure systems. The primary objective of this research was to investigate the use of MEMS in highway structures for health monitoring purposes. This study focused on investigating the use of MEMS and their potential applications in concrete through a comprehensive literature review, a vendor survey, and a laboratory study, as well as a small-scale field study. Based on the comprehensive literature review and vendor survey, the latest information available on off-the-shelf MEMS devices, as well as research prototypes, for bridge, pavement, and traffic applications were synthesized. A commercially-available wireless concrete monitoring system based on radio-frequency identification (RFID) technology and off-the-shelf temperature and humidity sensors were tested under controlled laboratory and field conditions. The test results validated the ability of the RFID wireless concrete monitoring system in accurately measuring the temperature both inside the laboratory and in the field under severe weather conditions. In consultation with the project technical advisory committee (TAC), the most relevant MEMS-based transportation infrastructure research applications to explore in the future were also highlighted and summarized.

Effects of soil management systems on soil microbial activity, bulk density and chemical properties

The objective of this experiment was to study the effects of soil management systems on the bulk density, chemical soil properties, and on the soil microbial activity on a Latossolo Vermelho distrófico (Oxisol). Soil samples were collected from plots under the following management conditions: a) natural dense "cerrado" vegetation (savanna); b) degraded Brachiaria decumbens pasture, 20 years old; c) no-tillage treatment with annual crop sequence (bean, corn, soybean and dark-oat in continuous rotation), 8 years old; d) conventional tillage treatment with crop residues added to the soil, and annual crop sequence, 10 years old. The continuous use of no-tillage system resulted in an increase in microbial biomass and decrease in soil basal respiration, therefore displaying evident long-term effects on the increase of soil C content. The no-tillage system also provided an improvement in bulk density and chemical properties of the soil. Hence, the no-tillage management system could be an alternative for the conservation and maintenance of physical and chemical conditions and the productive potential of "cerrado" soils.

Coping with continuous human disturbance in the wild: insights from penguin heart rate response to various stressors.

ABSTRACT: BACKGROUND: A central question for ecologists is the extent to which anthropogenic disturbances (e.g. tourism) might impact wildlife and affect the systems under study. From a research perspective, identifying the effects of human disturbance caused by research-related activities is crucial in order to understand and account for potential biases and derive appropriate conclusions from the data. RESULTS: Here, we document a case of biological adjustment to chronic human disturbance in a colonial seabird, the king penguin (Aptenodytes patagonicus), breeding on remote and protected islands of the Southern ocean. Using heart rate (HR) as a measure of the stress response, we show that, in a colony with areas exposed to the continuous presence of humans (including scientists) for over 50 years, penguins have adjusted to human disturbance and habituated to certain, but not all, types of stressors. When compared to birds breeding in relatively undisturbed areas, birds in areas of high chronic human disturbance were found to exhibit attenuated HR responses to acute anthropogenic stressors of low-intensity (i.e. sounds or human approaches) to which they had been subjected intensely over the years. However, such attenuation was not apparent for high-intensity stressors (i.e. captures for scientific research) which only a few individuals experience each year. CONCLUSIONS: Habituation to anthropogenic sounds/approaches could be an adaptation to deal with chronic innocuous stressors, and beneficial from a research perspective. Alternately, whether penguins have actually habituated to anthropogenic disturbances over time or whether human presence has driven the directional selection of human-tolerant phenotypes, remains an open question with profound ecological and conservation implications, and emphasizes the need for more knowledge on the effects of human disturbance on long-term studied populations.

Modeling and control of electromechanical systems

The material presented in the these notes covers the sessions Modelling of electromechanical systems, Passive control theory I and Passive control theory II of the II EURON/GEOPLEX Summer School on Modelling and Control of Complex Dynamical Systems.We start with a general description of what an electromechanical system is from a network modelling point of view. Next, a general formulation in terms of PHDS is introduced, and some of the previous electromechanical systems are rewritten in this formalism. Power converters, which are variable structure systems (VSS), can also be given a PHDS form.We conclude the modelling part of these lectures with a rather complex example, showing the interconnection of subsystems from several domains, namely an arrangement to temporally store the surplus energy in a section of a metropolitan transportation system based on dc motor vehicles, using either arrays of supercapacitors or an electric poweredflywheel. The second part of the lectures addresses control of PHD systems. We first present the idea of control as power connection of a plant and a controller. Next we discuss how to circumvent this obstacle and present the basic ideas of Interconnection and Damping Assignment (IDA) passivity-based control of PHD systems.

Soil macrofauna under integrated crop-livestock systems in a Brazilian Cerrado Ferralsol

The objective of this work was to assess the effects of integrated crop-livestock systems, associated with two tillage and two fertilization regimes, on the abundance and diversity of the soil macrofauna. Four different management systems were studied: continuous pasture (mixed grass); continuous crop; two crop-livestock rotations (crop/pasture and pasture/crop); and native Cerrado as a control. Macrofauna was sampled using a modified Tropical Soil Biology and Fertility method, and all individuals were counted and identified at the morphospecies level for each plot. A total of 194 morphospecies were found, distributed among 30 groups, and the most representative in decreasing order of density were: Isoptera, Coleoptera larvae, Formicidae, Oligochaeta, Coleoptera adult, Diplopoda, Hemiptera, Diptera larvae, Arachnida, Chilopoda, Lepidoptera, Gasteropoda, Blattodea and Orthoptera. Soil management systems and tillage regimes affected the structure of soil macrofauna, and integrated crop-livestock systems, associated with no-tillage, especially with grass/legume species associations, had more favorable conditions for the development of "soil engineers" compared with continuous pasture or arable crops. Soil macrofauna density and diversity, assessed at morphospecies level, are effective data to measure the impact of land use in Cerrado soils.

Intercepted solar radiation by maize crops subjected to different tillage systems and water availability levels

The objective of this work was to evaluate changes in the photosynthetic photon flux density (PPFD) interception efficiency and PPFD extinction coefficient for maize crop subjected to different soil tillage systems and water availability levels. Crops were subjected to no-tillage and conventional tillage systems combined with full irrigation and non-irrigation treatments. Continuous measurements of transmitted PPFD on the soil surface and incoming PPFD over the canopy were taken throughout the crop cycle. Leaf area index and soil water potential were also measured during the whole period. Considering a mean value over the maize cycle, intercepted PPFD was higher in the conventional tillage than in the no-tillage system. During the initial stages of plants, intercepted PPFD in the conventional tillage was double the PPFD interception in the no-tillage treatment. However, those differences were reduced up to the maximum leaf area index, close to tasseling stage. The lowest interception of PPFD occurred in the conventional tillage during the reproductive period, as leaf senescence progressed. Over the entire crop cycle, the interception of PPFD by the non-irrigated plants was about 20% lower than by the irrigated plants. The no-tillage system reduced the extinction coefficient for PPFD, which may have allowed a higher penetration of solar radiation into the canopy

Variable queen number in ant colonies: no impact on queen turnover, inbreeding, and population genetic differentiation in the ant Formica selysi.

Variation in queen number alters the genetic structure of social insect colonies, which in turn affects patterns of kin-selected conflict and cooperation. Theory suggests that shifts from single- to multiple-queen colonies are often associated with other changes in the breeding system, such as higher queen turnover, more local mating, and restricted dispersal. These changes may restrict gene flow between the two types of colonies and it has been suggested that this might ultimately lead to sympatric speciation. We performed a detailed microsatellite analysis of a large population of the ant Formica selysi, which revealed extensive variation in social structure, with 71 colonies headed by a single queen and 41 by multiple queens. This polymorphism in social structure appeared stable over time, since little change in the number of queens per colony was detected over a five-year period. Apart from queen number, single- and multiple-queen colonies had very similar breeding systems. Queen turnover was absent or very low in both types of colonies. Single- and multiple-queen colonies exhibited very small but significant levels of inbreeding, which indicates a slight deviation from random mating at a local scale and suggests that a small proportion of queens mate with related males. For both types of colonies, there was very little genetic structuring above the level of the nest, with no sign of isolation by distance. These similarities in the breeding systems were associated with a complete lack of genetic differentiation between single- and multiple-queen colonies, which provides no support for the hypothesis that change in queen number leads to restricted gene flow between social forms. Overall, this study suggests that the higher rates of queen turnover, local mating, and population structuring that are often associated with multiple-queen colonies do not appear when single- and multiple-queen colonies still coexist within the same population, but build up over time in populations consisting mostly of multiple-queen colonies.

A qualitative continuous model of cellular auxin and brassinosteroid signaling and their crosstalk.

Motivation: Hormone pathway interactions are crucial in shaping plant development, such as synergism between the auxin and brassinosteroid pathways in cell elongation. Both hormone pathways have been characterized in detail, revealing several feedback loops. The complexity of this network, combined with a shortage of kinetic data, renders its quantitative analysis virtually impossible at present.Results: As a first step towards overcoming these obstacles, we analyzed the network using a Boolean logic approach to build models of auxin and brassinosteroid signaling, and their interaction. To compare these discrete dynamic models across conditions, we transformed them into qualitative continuous systems, which predict network component states more accurately and can accommodate kinetic data as they become available. To this end, we developed an extension for the SQUAD software, allowing semi-quantitative analysis of network states. Contrasting the developmental output depending on cell type-specific modulators enabled us to identify a most parsimonious model, which explains initially paradoxical mutant phenotypes and revealed a novel physiological feature.

Observation of a continuous interior crisis in the Hindmarsh-Rose neuron model.

Interior crises are understood as discontinuous changes of the size of a chaotic attractor that occur when an unstable periodic orbit collides with the chaotic attractor. We present here numerical evidence and theoretical reasoning which prove the existence of a chaos-chaos transition in which the change of the attractor size is sudden but continuous. This occurs in the Hindmarsh¿Rose model of a neuron, at the transition point between the bursting and spiking dynamics, which are two different dynamic behaviors that this system is able to present. Moreover, besides the change in attractor size, other significant properties of the system undergoing the transitions do change in a relevant qualitative way. The mechanism for such transition is understood in terms of a simple one-dimensional map whose dynamics undergoes a crossover between two different universal behaviors

Long-term continuous-flow left ventricular assist devices (LVAD) as bridge to heart transplantation.

Heart transplantation (HTx) is the treatment of choice for end-stage heart failure but the limited availability of heart's donors still represents a major issue. So long-term mechanical circulatory support (MCS) has been proposed as an alternative treatment option to assist patients scheduled on HTx waiting list bridging them for a variable time period to cardiac transplantation-the so-called bridge-to-transplantation (BTT) strategy. Nowadays approximately 90% of patients being considered for MCS receive a left ventricular assist device (LVAD). In fact, LVAD experienced several improvements in the last decade and the predominance of continuous-flow over pulsatile-flow technology has been evident since 2008. The aim of the present report is to give an overview of continuous-flow LVAD utilization in the specific setting of the BTT strategy taking into consideration the most representative articles of the scientific literature and focusing the attention on the evolution, clinical outcomes, relevant implications on the HTx strategy and future perspectives of the continuous-flow LVAD technology.

Local thermodynamics and the generalized Gibbs-Duhem equation in systems with long-range interactions

The local thermodynamics of a system with long-range interactions in d dimensions is studied using the mean-field approximation. Long-range interactions are introduced through pair interaction potentials that decay as a power law in the interparticle distance. We compute the local entropy, Helmholtz free energy, and grand potential per particle in the microcanonical, canonical, and grand canonical ensembles, respectively. From the local entropy per particle we obtain the local equation of state of the system by using the condition of local thermodynamic equilibrium. This local equation of state has the form of the ideal gas equation of state, but with the density depending on the potential characterizing long-range interactions. By volume integration of the relation between the different thermodynamic potentials at the local level, we find the corresponding equation satisfied by the potentials at the global level. It is shown that the potential energy enters as a thermodynamic variable that modifies the global thermodynamic potentials. As a result, we find a generalized Gibbs-Duhem equation that relates the potential energy to the temperature, pressure, and chemical potential. For the marginal case where the power of the decaying interaction potential is equal to the dimension of the space, the usual Gibbs-Duhem equation is recovered. As examples of the application of this equation, we consider spatially uniform interaction potentials and the self-gravitating gas. We also point out a close relationship with the thermodynamics of small systems.