Genome-wide association mapping in a wild avian population identifies a link between genetic and phenotypic variation in a life-history trait

Understanding the genetic basis of traits involved in adaptation is a major challenge in evolutionary biology but remains poorly understood. Here, we use genome-wide association mapping using a custom 50 k single nucleotide polymorphism (SNP) array in a natural population of collared flycatchers to examine the genetic basis of clutch size, an important life-history trait in many animal species. We found evidence for an association on chromosome 18 where one SNP significant at the genome-wide level explained 3.9% of the phenotypic variance. We also detected two suggestive quantitative trait loci (QTLs) on chromosomes 9 and 26. Fitness differences among genotypes were generally weak and not significant, although there was some indication of a sex-by-genotype interaction for lifetime reproductive success at the suggestive QTL on chromosome 26. This implies that sexual antagonism may play a role in maintaining genetic variation at this QTL. Our findings provide candidate regions for a classic avian life-history trait that will be useful for future studies examining the molecular and cellular function of, as well as evolutionary mechanisms operating at, these loci.

Recent developments in statistical methods for detecting genetic loci affecting phenotypic variability

A number of recent works have introduced statistical methods for detecting genetic loci that affect phenotypic variability, which we refer to as variability-controlling quantitative trait loci (vQTL). These are genetic variants whose allelic state predicts how much phenotype values will vary about their expected means. Such loci are of great potential interest in both human and non-human genetic studies, one reason being that a detected vQTL could represent a previously undetected interaction with other genes or environmental factors. The simultaneous publication of these new methods in different journals has in many cases precluded opportunity for comparison. We survey some of these methods, the respective trade-offs they imply, and the connections between them. The methods fall into three main groups: classical non-parametric, fully parametric, and semi-parametric two-stage approximations. Choosing between alternatives involves balancing the need for robustness, flexibility, and speed. For each method, we identify important assumptions and limitations, including those of practical importance, such as their scope for including covariates and random effects. We show in simulations that both parametric methods and their semi-parametric approximations can give elevated false positive rates when they ignore mean-variance relationships intrinsic to the data generation process. We conclude that choice of method depends on the trait distribution, the need to include non-genetic covariates, and the population size and structure, coupled with a critical evaluation of how these fit with the assumptions of the statistical model.

Inheritance beyond plain heritability : variance-controlling genes in Arabidopsis thaliana

The phenotypic effect of a gene is normally described by the mean-difference between alternative genotypes. A gene may, however, also influence the phenotype by causing a difference in variance between genotypes. Here, we reanalyze a publicly available Arabidopsis thaliana dataset [1] and show that genetic variance heterogeneity appears to be as common as normal additive effects on a genomewide scale. The study also develops theory to estimate the contributions of variance differences between genotypes to the phenotypic variance, and this is used to show that individual loci can explain more than 20% of the phenotypic variance. Two well-studied systems, cellular control of molybdenum level by the ion-transporter MOT1 and flowering-time regulation by the FRI-FLC expression network, and a novel association for Leaf serration are used to illustrate the contribution of major individual loci, expression pathways, and gene-by-environment interactions to the genetic variance heterogeneity.

High female mortality resulting in herd collapse in free-ranging domesticated reindeer (Rangifer tarandus tarandus) in Sweden

Reindeer herding in Sweden is a form of pastoralism practised by the indigenous Sami population. The economy is mainly based on meat production. Herd size is generally regulated by harvest in order not to overuse grazing ranges and keep a productive herd. Nonetheless, herd growth and room for harvest is currently small in many areas. Negative herd growth and low harvest rate were observed in one of two herds in a reindeer herding community in Central Sweden. The herds (A and B) used the same ranges from April until the autumn gathering in October-December, but were separated on different ranges over winter. Analyses of capture-recapture for 723 adult female reindeer over five years (2007-2012) revealed high annual losses (7.1% and 18.4%, for herd A and B respectively). A continuing decline in the total reindeer number in herd B demonstrated an inability to maintain the herd size in spite of a very small harvest. An estimated breakpoint for when herd size cannot be kept stable confirmed that the observed female mortality rate in herd B represented a state of herd collapse. Lower calving success in herd B compared to A indicated differences in winter foraging conditions. However, we found only minor differences in animal body condition between the herds in autumn. We found no evidence that a lower autumn body mass generally increased the risk for a female of dying from one autumn to the next. We conclude that the prime driver of the on-going collapse of herd B is not high animal density or poor body condition. Accidents or disease seem unlikely as major causes of mortality. Predation, primarily by lynx and wolverine, appears to be the most plausible reason for the high female mortality and state of collapse in the studied reindeer herding community.

On processing GPS tracking data of spatiotemporal car movements : a case study

The advancement of GPS technology has made it possible to use GPS devices as orientation and navigation tools, but also as tools to track spatiotemporal information. GPS tracking data can be broadly applied in location-based services, such as spatial distribution of the economy, transportation routing and planning, traffic management and environmental control. Therefore, knowledge of how to process the data from a standard GPS device is crucial for further use. Previous studies have considered various issues of the data processing at the time. This paper, however, aims to outline a general procedure for processing GPS tracking data. The procedure is illustrated step-by-step by the processing of real-world GPS data of car movements in Borlänge in the centre of Sweden.

GRASP and statistical bounds for heuristic solutions to combinatorial problems

The quality of a heuristic solution to a NP-hard combinatorial problem is hard to assess. A few studies have advocated and tested statistical bounds as a method for assessment. These studies indicate that statistical bounds are superior to the more widely known and used deterministic bounds. However, the previous studies have been limited to a few metaheuristics and combinatorial problems and, hence, the general performance of statistical bounds in combinatorial optimization remains an open question. This work complements the existing literature on statistical bounds by testing them on the metaheuristic Greedy Randomized Adaptive Search Procedures (GRASP) and four combinatorial problems. Our findings confirm previous results that statistical bounds are reliable for the p-median problem, while we note that they also seem reliable for the set covering problem. For the quadratic assignment problem, the statistical bounds has previously been found reliable when obtained from the Genetic algorithm whereas in this work they found less reliable. Finally, we provide statistical bounds to four 2-path network design problem instances for which the optimum is currently unknown.

Increasing the power of genome wide association studies in natural populations using repeated measures : evaluation and implementation

1. Genomewide association studies (GWAS) enable detailed dissections of the genetic basis for organisms' ability to adapt to a changing environment. In long-term studies of natural populations, individuals are often marked at one point in their life and then repeatedly recaptured. It is therefore essential that a method for GWAS includes the process of repeated sampling. In a GWAS, the effects of thousands of single-nucleotide polymorphisms (SNPs) need to be fitted and any model development is constrained by the computational requirements. A method is therefore required that can fit a highly hierarchical model and at the same time is computationally fast enough to be useful. 2. Our method fits fixed SNP effects in a linear mixed model that can include both random polygenic effects and permanent environmental effects. In this way, the model can correct for population structure and model repeated measures. The covariance structure of the linear mixed model is first estimated and subsequently used in a generalized least squares setting to fit the SNP effects. The method was evaluated in a simulation study based on observed genotypes from a long-term study of collared flycatchers in Sweden. 3. The method we present here was successful in estimating permanent environmental effects from simulated repeated measures data. Additionally, we found that especially for variable phenotypes having large variation between years, the repeated measurements model has a substantial increase in power compared to a model using average phenotypes as a response. 4. The method is available in the R package RepeatABEL. It increases the power in GWAS having repeated measures, especially for long-term studies of natural populations, and the R implementation is expected to facilitate modelling of longitudinal data for studies of both animal and human populations.

Residential planning, driver mobility and CO2 emission

The literature on residences and citizens’ transports has focused on either reforming traffic managing in response to residential relocation or post-evaluation of urban planning policies or the evolution of the urban spatial form. In a city there are hotspots that attract the citizens and most of the transportation in the city arises as the citizens’ movement between their residence and the hotspots. Little scholarly attention has been devoted to the possibility to minimize citizens’ transportation in the city by the urban planning of residential areas. In this paper we propose a method to evaluate the environmental impact (in terms of CO2-emissions) of urban plans of residential areas. The method is illustrated in a Swedish case of a midsize city which is presently preoccupied with urban planning of new residential areas in response to substantial population growth due to immigration. The residential plans aims to increase the compactness and residential density in the current center and sub centers leads to less CO2 emissions compare to urban expansion to the edge of the city. The plans of concentrated apartment buildings are more effective in meeting residential needs and mitigating CO2 emissions than dispersed single-family houses.

Spatial modeling of data with excessive zeros applied to reindeer pellet-group counts

We analyze a real data set pertaining to reindeer fecal pellet-group counts obtained from a survey conducted in a forest area in northern Sweden. In the data set, over 70% of counts are zeros, and there is high spatial correlation. We use conditionally autoregressive random effects for modeling of spatial correlation in a Poisson generalized linear mixed model (GLMM), quasi-Poisson hierarchical generalized linear model (HGLM), zero-inflated Poisson (ZIP), and hurdle models. The quasi-Poisson HGLM allows for both under- and overdispersion with excessive zeros, while the ZIP and hurdle models allow only for overdispersion. In analyzing the real data set, we see that the quasi-Poisson HGLMs can perform better than the other commonly used models, for example, ordinary Poisson HGLMs, spatial ZIP, and spatial hurdle models, and that the underdispersed Poisson HGLMs with spatial correlation fit the reindeer data best. We develop R codes for fitting these models using a unified algorithm for the HGLMs. Spatial count response with an extremely high proportion of zeros, and underdispersion can be successfully modeled using the quasi-Poisson HGLM with spatial random effects.

Renar och vindkraft II : Vindkraft i drift och effekter på renar och renskötsel

Utbyggnaden av vindkraft inom renskötselområdet har ökat markant det senaste decenniet, trots att kunskapen om påverkan av vindkraftsetableringar ännu inte är fullt utredd och dokumenterad. I den här rapporten beskriver vi framförallt hur vindkraftparker i driftsfas påverkar renarna och renskötseln i tre olika områden. I Malå sameby har vi studerat kalvningsområdet kring Storliden och Jokkmokkslidens vindkraftparker. I Vilhelmina Norra sameby har vi studerat vinterbetesområdet kring Stor-Rotlidens vindkraftpark, samt Lögdeålandets betesområde med Gabrielsbergets vindkraftpark som används av Byrkije reinbetesdistrikt från Norge. För att få en helhetsbild av hur renarna använder sitt betesområde är det viktigt att studera renarnas betes- och förflyttningsmönster långsiktigt och över hela deras betesområde och inte bara inom det lokala området nära parken. Det är också viktigt att ta hänsyn till att renarnas betesutnyttjande skiftar från år till år och mellan olika årstider beroende på väderlek och andra yttre förutsättningar. Vi vill också understryka vikten av att kombinera den traditionella kunskapen från renskötarna med vedertagna vetenskapliga analysmetoder för att besvara de frågor som är viktiga för att kunna bedriva en hållbar renskötsel. Vi har undersökt renarnas användning av områdena genom att utföra spillningsinventeringar under åren 2009-2015 (endast i Malå sameby), och genom att följa renar utrustade med GPS-halsband under åren 2005-2015. Datat är insamlat före och under byggfas och under driftsfas (för Gabrielsberget finns GPS-data endast för driftsfasen). Vi har analyserat data genom att utveckla statistiska modeller för val av betesområde för varje område där vi har beräknat hur renarna förhåller sig till vindkraftparksområdet före, under och efter byggnation, och på Gabrielsberget när parken varit avstängd under 40 dagar och under drift vid olika renskötselsituationer. Genom intervjuer, möten och samtal, samt information från Gabrielsbergets vindkraftparks kontrollprogram, har vi tagit del av renskötarnas erfarenheter av hur renarnas beteende, och därmed även renskötseln, påverkats av vindkraftsutbyggnaden i respektive område. Våra resultat visar att renarna både på kalvnings- och på vinterbetesområden påverkas negativt av vindkraftsetableringarna (Tabell a). Renarna undviker att beta i områden där de kan se och/eller höra vindkraftsverken och föredrar att vistas i områden där vindkraftverken är skymda. I kalvningsområdet i Malå ökade användningen av skymda områden med 60 % under driftsfas. I vinterbetesområdet på Gabrielsberget, när renarna utfodrades i parken och kantbevakades intensivt för att stanna i parkområdet under driftsfas, ökade användningen av skymda områden med 13 % jämfört med när de inte var utfodrade och fick ströva mer fritt. Resultaten visar också att renarna minskar sin användning av området nära vindkraftparkerna. I kalvningslandet i Malå minskar renarna sin användning av områden inom 5 km från parkerna med 16-20 %. Vintertid vid Gabrielsbergets vindkraftpark undvek renarna parken med 3 km. Våra resultat visar även att renarnas betesro minskar inom en radie på 4 km från vindkraftparkerna under kalvningsperioden och tiden därefter i jämförelse med perioden före byggfas. Exakta avstånd som renarna påverkas beror på förutsättningarna i respektive område, exempelvis hur topografin ser ut eller om området är begränsat av stängsel eller annan infrastruktur. Förändringarna i habitatutnyttjande i våra studieområden blev tydligare när parkerna var centralt belägna i renarnas betesområde, som i kalvningsområdet i Malå eller i vinterbeteslandet på Gabrielsberget, medan det inte var lika tydliga effekter kring Stor-Rotlidens park, som ligger i utkanten av ett huvudbetesområde. Oftast är snöförhållandena bättre ur betessynpunkt högre upp i terrängen än nere i dalgångarna, på grund av stabilare temperatur, vind som blåser bort snötäcket och mer variation i topografin. Därför kan etablering av vindkraftparker i höglänta områden försämra möjligheten att använda sådana viktiga reservbetesområden under vintrar med i övrigt dåliga snöförhållanden, vilka blir allt vanligare i och med klimatförändringarna. Våra resultat tyder inte direkt på att renarna påverkats negativt under dåliga betesvintrar men fler år av studier behövs för att ytterligare klargöra hur vindkraft påverkar renarna under dessa vintrar. Våra studier har visat att etablering av vindkraft har konsekvenser för renskötseln under både barmarkssäsongen och under vintern, men effekterna förmodas få störst inverkan inom vinterbetesområdet där det är svårt att hitta alternativa betesområden för renarna. Under sommaren är betestillgången oftast mindre begränsad och renarna kan lättare hitta alternativa områden. En direkt konsekvens av Gabrielsbergets vindkraftpark som är placerad mitt i ett vinterbetesområde har blivit att renarna behöver tillskottsutfodras och bevakas intensivare för att de inte ska gå ut ur området. När den naturliga vandringen mellan olika betesområden störs för att renarna undviker att vistas i ett område kan det leda till att den totala tillgången till naturligt bete minskar och att man permanent måste tillskottsutfodra, alternativt minska antalet renar. Annan infrastruktur som vägar och kraftledningar påverkar också renarna. Vid Storliden och Jokkmokksliden och vid Stor-Rotliden där data samlats in innan vindkraftparken uppfördes visar våra resultat att renarna undviker de omkringliggande landsvägarna redan innan parkerna etablerades. Vid Stor-Rotliden ökar dock renarna användningen av områden nära vägarna efter att parken är byggd. På Gabrielsberget, där vi endast har data under drifttiden, är renarna närmare vägarna (även stora vägar som E4) när renskötarna minskar på kantbevakningen för att inte hålla renarna nära parken. Detta ökar naturligtvis risken för trafikolyckor och innebär att renskötarna måste bevaka dessa områden intensivare. Sist i rapporten presenterar vi förslag till åtgärder som kan användas för att underlätta arbetet för renskötseln om det är så att en vindkraftpark redan är byggd. Några exempel på åtgärder som är direkt kopplat till parken är att stänga av vägarna in i vindkraftparken för att förhindra nöjeskörning med skoter och bil under den tiden renarna vistas i området samt tät dialog mellan vindkraftsbolag och sameby angående vinterväghållningen av vägarna till och inom vindkraftparken. Andra mer regionala åtgärder för att förbättra förutsättningarna för renskötselarbetet på andra platser för samebyn, kan vara att sätta stängsel längst större vägar och järnvägar (t.ex. E4:an eller stambanan) i kombination med strategiskt utplacerade ekodukter. Detta för att underlätta och återställa möjligheterna till renarnas fria strövning och renskötarnas flytt av renar mellan olika betesområden.   

A Microdata Analysis Approach to Transport Infrastructure Maintenance

Maintenance of transport infrastructure assets is widely advocated as the key in minimizing current and future costs of the transportation network. While effective maintenance decisions are often a result of engineering skills and practical knowledge, efficient decisions must also account for the net result over an asset's life-cycle. One essential aspect in the long term perspective of transport infrastructure maintenance is to proactively estimate maintenance needs. In dealing with immediate maintenance actions, support tools that can prioritize potential maintenance candidates are important to obtain an efficient maintenance strategy. This dissertation consists of five individual research papers presenting a microdata analysis approach to transport infrastructure maintenance. Microdata analysis is a multidisciplinary field in which large quantities of data is collected, analyzed, and interpreted to improve decision-making. Increased access to transport infrastructure data enables a deeper understanding of causal effects and a possibility to make predictions of future outcomes. The microdata analysis approach covers the complete process from data collection to actual decisions and is therefore well suited for the task of improving efficiency in transport infrastructure maintenance. Statistical modeling was the selected analysis method in this dissertation and provided solutions to the different problems presented in each of the five papers. In Paper I, a time-to-event model was used to estimate remaining road pavement lifetimes in Sweden. In Paper II, an extension of the model in Paper I assessed the impact of latent variables on road lifetimes; displaying the sections in a road network that are weaker due to e.g. subsoil conditions or undetected heavy traffic. The study in Paper III incorporated a probabilistic parametric distribution as a representation of road lifetimes into an equation for the marginal cost of road wear. Differentiated road wear marginal costs for heavy and light vehicles are an important information basis for decisions regarding vehicle miles traveled (VMT) taxation policies. In Paper IV, a distribution based clustering method was used to distinguish between road segments that are deteriorating and road segments that have a stationary road condition. Within railway networks, temporary speed restrictions are often imposed because of maintenance and must be addressed in order to keep punctuality. The study in Paper V evaluated the empirical effect on running time of speed restrictions on a Norwegian railway line using a generalized linear mixed model.

Reindeer habitat selection under the risk of brown bear predation during calving season

The depredation of semi-domesticated reindeer by large carnivores reflects an important human-wildlife conflict in Fennoscandia. Recent studies have revealed that brown bears (Ursus arctos) may kill substantial numbers of reindeer calves (Rangifer tarandus tarandus) in forest areas in Sweden. Several authors have suggested that predation risk is an important driver of habitat selection in wild Rangifer populations where predation is a limiting factor, but little is known about these mechanisms in semi-domesticated populations. We examined the habitat selection of female reindeer in relation to spatial and temporal variations in brown bear predation risk on the reindeer calving grounds and evaluated the simultaneous responses of brown bears and reindeer to landscape characteristics. We used GPS data from 110 reindeer years (97 individuals) and 29 brown bear years (19 individuals), from two reindeer herding districts in the forest area of northern Sweden. Our results did not indicate that reindeer alter their behavior in response to spatiotemporal variation in brown bear predation risk, on the scale of the calving range. Instead, we suggest that spatiotemporal behavioral adjustments by brown bears were the main driver of prey-predator interactions in our study system. Contrasting responses by brown bears and reindeer to clear-cuts and young forest indicate that forestry can influence species interactions and possibly yield negative consequences for the reindeer herd. Even if clear-cuts may be beneficial in terms of calf survival, logging activity will eventually cause greater abundance of young regenerating forest, reducing available reindeer habitats and increasing habitat preferred by brown bears. Domestication may have made semi-domesticated reindeer in Fennoscandia less adapted to cope with predators. Areal restrictions, limiting the opportunity for dispersion and escape, possibly make the calves more susceptible to predation. Also, a generally higher population density in semi-domesticated herds compared to wild populations can make dispersion a less efficient strategy and the reindeer calves easier prey. Overall, the lack of ability of the reindeer females to reduce brown bear encounter risk on the scale of the calving range is probably an important reason for the high brown bear predation rates on reindeer calves documented in our study areas. 

The Power of the Synthetic Control Method

The synthetic control method (SCM) is a new, popular method developed for the purpose of estimating the effect of an intervention when only one single unit has been exposed. Other similar, unexposed units are combined into a synthetic control unit intended to mimic the evolution in the exposed unit, had it not been subject to exposure. As the inference relies on only a single observational unit, the statistical inferential issue is a challenge. In this paper, we examine the statistical properties of the estimator, study a number of features potentially yielding uncertainty in the estimator, discuss the rationale for statistical inference in relation to SCM, and provide a Web-app for researchers to aid in their decision of whether SCM is powerful for a specific case study. We conclude that SCM is powerful with a limited number of controls in the donor pool and a fairly short pre-intervention time period. This holds as long as the parameter of interest is a parametric specification of the intervention effect, and the duration of post-intervention period is reasonably long, and the fit of the synthetic control unit to the exposed unit in the pre-intervention period is good.