TY - JOUR A1 - Romero-Mujalli, Daniel A1 - Rochow, Markus A1 - Kahl, Sandra M. A1 - Paraskevopoulou, Sofia A1 - Folkertsma, Remco A1 - Jeltsch, Florian A1 - Tiedemann, Ralph T1 - Adaptive and nonadaptive plasticity in changing environments: Implications for sexual species with different life history strategies JF - Ecology and Evolution N2 - Populations adapt to novel environmental conditions by genetic changes or phenotypic plasticity. Plastic responses are generally faster and can buffer fitness losses under variable conditions. Plasticity is typically modeled as random noise and linear reaction norms that assume simple one-to- one genotype–phenotype maps and no limits to the phenotypic response. Most studies on plasticity have focused on its effect on population viability. However, it is not clear, whether the advantage of plasticity depends solely on environmental fluctuations or also on the genetic and demographic properties (life histories) of populations. Here we present an individual-based model and study the relative importance of adaptive and nonadaptive plasticity for populations of sexual species with different life histories experiencing directional stochastic climate change. Environmental fluctuations were simulated using differentially autocorrelated climatic stochasticity or noise color, and scenarios of directiona climate change. Nonadaptive plasticity was simulated as a random environmental effect on trait development, while adaptive plasticity as a linear, saturating, or sinusoidal reaction norm. The last two imposed limits to the plastic response and emphasized flexible interactions of the genotype with the environment. Interestingly, this assumption led to (a) smaller phenotypic than genotypic variance in the population (many-to- one genotype–phenotype map) and the coexistence of polymorphisms, and (b) the maintenance of higher genetic variation—compared to linear reaction norms and genetic determinism—even when the population was exposed to a constant environment for several generations. Limits to plasticity led to genetic accommodation, when costs were negligible, and to the appearance of cryptic variation when limits were exceeded. We found that adaptive plasticity promoted population persistence under red environmental noise and was particularly important for life histories with low fecundity. Populations produing more offspring could cope with environmental fluctuations solely by genetic changes or random plasticity, unless environmental change was too fast. KW - developmental canalization KW - environmental change KW - genetic accommodation KW - Individual-based models KW - limits KW - many-to-one genotype–phenotype map KW - noise color KW - phenotypic plasticity KW - reaction norms KW - stochastic fluctuations Y1 - 2020 SN - 2045-7758 VL - 11 IS - 11 PB - John Wiley & Sons, Inc. CY - New Jersey ER - TY - GEN A1 - Romero-Mujalli, Daniel A1 - Rochow, Markus A1 - Kahl, Sandra M. A1 - Paraskevopoulou, Sofia A1 - Folkertsma, Remco A1 - Jeltsch, Florian A1 - Tiedemann, Ralph T1 - Adaptive and nonadaptive plasticity in changing environments: Implications for sexual species with different life history strategies T2 - Postprints der Universität Potsdam : Mathematisch-Naturwissenschaftliche Reihe N2 - Populations adapt to novel environmental conditions by genetic changes or phenotypic plasticity. Plastic responses are generally faster and can buffer fitness losses under variable conditions. Plasticity is typically modeled as random noise and linear reaction norms that assume simple one-to- one genotype–phenotype maps and no limits to the phenotypic response. Most studies on plasticity have focused on its effect on population viability. However, it is not clear, whether the advantage of plasticity depends solely on environmental fluctuations or also on the genetic and demographic properties (life histories) of populations. Here we present an individual-based model and study the relative importance of adaptive and nonadaptive plasticity for populations of sexual species with different life histories experiencing directional stochastic climate change. Environmental fluctuations were simulated using differentially autocorrelated climatic stochasticity or noise color, and scenarios of directiona climate change. Nonadaptive plasticity was simulated as a random environmental effect on trait development, while adaptive plasticity as a linear, saturating, or sinusoidal reaction norm. The last two imposed limits to the plastic response and emphasized flexible interactions of the genotype with the environment. Interestingly, this assumption led to (a) smaller phenotypic than genotypic variance in the population (many-to- one genotype–phenotype map) and the coexistence of polymorphisms, and (b) the maintenance of higher genetic variation—compared to linear reaction norms and genetic determinism—even when the population was exposed to a constant environment for several generations. Limits to plasticity led to genetic accommodation, when costs were negligible, and to the appearance of cryptic variation when limits were exceeded. We found that adaptive plasticity promoted population persistence under red environmental noise and was particularly important for life histories with low fecundity. Populations produing more offspring could cope with environmental fluctuations solely by genetic changes or random plasticity, unless environmental change was too fast. T3 - Zweitveröffentlichungen der Universität Potsdam : Mathematisch-Naturwissenschaftliche Reihe - 1170 KW - developmental canalization KW - environmental change KW - genetic accommodation KW - Individual-based models KW - limits KW - many-to-one genotype–phenotype map KW - noise color KW - phenotypic plasticity KW - reaction norms KW - stochastic fluctuations Y1 - 2020 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:kobv:517-opus4-523201 SN - 1866-8372 IS - 1170 ER - TY - GEN A1 - Folkertsma, Remco A1 - Westbury, Michael V. A1 - Eccard, Jana A1 - Hofreiter, Michael T1 - The complete mitochondrial genome of the common vole, Microtus arvalis (Rodentia: Arvicolinae) T2 - Postprints der Universität Potsdam : Mathematisch-Naturwissenschaftliche Reihe N2 - The common vole, Microtus arvalis belongs to the genus Microtus in the subfamily Arvicolinae. In this study, the complete mitochondrial genome of M. arvalis was recovered using shotgun sequencing and an iterative mapping approach using three related species. Phylogenetic analyses using the sequence of 21 arvicoline species place the common vole as a sister species to the East European vole (Microtus levis), but as opposed to previous results we find no support for the recognition of the genus Neodon within the subfamily Arvicolinae, as this is, as well as the genus Lasiopodomys, found within the Microtus genus. T3 - Zweitveröffentlichungen der Universität Potsdam : Mathematisch-Naturwissenschaftliche Reihe - 481 KW - Microtus arvalis KW - Arvicolinae KW - mitochondrial genome KW - common vole KW - phylogeny Y1 - 2018 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:kobv:517-opus4-412994 SN - 1866-8372 IS - 481 ER - TY - GEN A1 - Breedveld, Merel Cathelijne A1 - Folkertsma, Remco A1 - Eccard, Jana T1 - Rodent mothers increase vigilance behaviour when facing infanticide risk T2 - Postprints der Universität Potsdam Mathematisch- Naturwissenschaftliche Reihe N2 - Infanticide, the killing of unrelated young, is widespread and frequently driven by sexual conflict. especially in mammals with exclusive maternal care, infanticide by males is common and females suffer fitness costs. Recognizing infanticide risk and adjusting offspring protection accordingly should therefore be adaptive in female mammals. Using a small mammal (Myodes glareolus) in outdoor enclosures, we investigated whether lactating mothers adjust offspring protection, and potential mate search behaviour, in response to different infanticide risk levels. We presented the scent of the litter’s sire or of a stranger male near the female’s nest, and observed female nest presence and movement by radiotracking. While both scents simulated a mating opportunity, they represented lower (sire) and higher (stranger) infanticide risk. compared to the sire treatment, females in the stranger treatment left their nest more often, showed increased activity and stayed closer to the nest, suggesting offspring protection from outside the nest through elevated alertness and vigilance. females with larger litters spent more time investigating scents and used more space in the sire but not in the stranger treatment. Thus, current investment size affected odour inspection and resource acquisition under higher risk. Adjusting nest protection and resource acquisition to infanticide risk could allow mothers to elicit appropriate (fitness-saving) counterstrategies, and thus, may be widespread. T3 - Zweitveröffentlichungen der Universität Potsdam : Mathematisch-Naturwissenschaftliche Reihe - 766 KW - vole clethrionomys-glareolus KW - male bank voles KW - maternal aggression KW - reproductive strategies KW - offspring-defense KW - myodes-glareolus KW - predation risk KW - prairie vole KW - recognition KW - costs Y1 - 2019 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:kobv:517-opus4-438074 SN - 1866-8372 IS - 766 ER - TY - JOUR A1 - Breedveld, Merel Cathelijne A1 - Folkertsma, Remco A1 - Eccard, Jana T1 - Rodent mothers increase vigilance behaviour when facing infanticide risk JF - Scientific Reports N2 - Infanticide, the killing of unrelated young, is widespread and frequently driven by sexual conflict. especially in mammals with exclusive maternal care, infanticide by males is common and females suffer fitness costs. Recognizing infanticide risk and adjusting offspring protection accordingly should therefore be adaptive in female mammals. Using a small mammal (Myodes glareolus) in outdoor enclosures, we investigated whether lactating mothers adjust offspring protection, and potential mate search behaviour, in response to different infanticide risk levels. We presented the scent of the litter’s sire or of a stranger male near the female’s nest, and observed female nest presence and movement by radiotracking. While both scents simulated a mating opportunity, they represented lower (sire) and higher (stranger) infanticide risk. compared to the sire treatment, females in the stranger treatment left their nest more often, showed increased activity and stayed closer to the nest, suggesting offspring protection from outside the nest through elevated alertness and vigilance. females with larger litters spent more time investigating scents and used more space in the sire but not in the stranger treatment. Thus, current investment size affected odour inspection and resource acquisition under higher risk. Adjusting nest protection and resource acquisition to infanticide risk could allow mothers to elicit appropriate (fitness-saving) counterstrategies, and thus, may be widespread. KW - vole clethrionomys-glareolus KW - male bank voles KW - maternal aggression KW - reproductive strategies KW - offspring-defense KW - myodes-glareolus KW - predation risk KW - prairie vole KW - recognition KW - costs Y1 - 2019 U6 - https://doi.org/10.1038/s41598-019-48459-9 SN - 2045-2322 VL - 9 PB - Macmillan Publishers Limited, part of Springer Nature CY - London ER - TY - JOUR A1 - Eccard, Jana A1 - Reil, Daniela A1 - Folkertsma, Remco A1 - Schirmer, Annika T1 - The scent of infanticide risk? BT - Behavioural allocation to current and future reproduction in response to mating opportunity and familiarity with intruder JF - Behavioral ecology and sociobiology N2 - The killing of young by unrelated males is widespread in the animal kingdom. In short-lived small rodents, females can mate immediately after delivery (post-partum oestrus) and invest in future reproduction, but infanticide may put the nestlings, their current reproductive investment, at risk. Here, we investigated the behavioural trade-offs between mating interest and nest protection in an arena experiment with bank voles (Myodes glareolus). Non-gravid females (n=33) were housed at one end of a large structured arena with their nestlings. Different scents (cage bedding) were presented to each female in a replicated design. Three combinations of mating opportunities and male-female familiarity were simulated using different scent donors: mating opportunity with the sire of the nestlings with whom the female was familiar; mating opportunity with a male unrelated to the offspring and unfamiliar to the female, thus posing a higher risk to the offspring; and neither risk nor mating opportunity (clean control). Most females investigated male scents, regardless of familiarity, leaving their litter unprotected. During control treatment, females with larger litters spent less time at the scent area, indicating increasing nursing demands or better protection. Females with older litters visited scents more often, suggesting an increased interest in reproduction while they are non-gravid alongside the decreased risk of infanticide for older young. In the presence of unfamiliar scents, females spent more time protecting their nests, supporting the perceived association of unfamiliarity with infanticide risk. Thus, rodent females flexibly allocate time spent between searching for a mate and protecting their nest, which is modulated by their familiarity with a potential intruder.Significance statementInfanticide by conspecific males is an extreme form of sexual conflict and has large costs on females, abolishing their investment into current offspring. In an experimental approach, we exposed lactating female bank voles to different combinations of mating opportunity and familiarity to a (simulated) intruder: (1) the sire of the nestlings with whom the female was familiar and, therefore, potentially less risky in terms of infanticide; (2) a male which was unrelated and unfamiliar to the female and thus posed a higher risk to the offspring; or (3) as a control, cage bedding, which posed neither risk of infanticide nor a mating opportunity. We show that females flexibly allocated pup protection and mating interest based on their familiarity with the male, indicating that the unfamiliar males pose a threat to offspring, which is perceived by the females. Females further adjusted their behaviour to the size and/or age of their current litter, investing more time in male scents when offspring were older, thus balancing future and current investments into reproduction. KW - Reproductive strategy KW - Infanticide risk KW - Odour recognition KW - Familiarity KW - Counterstrategy KW - Sexual conflict Y1 - 2018 U6 - https://doi.org/10.1007/s00265-018-2585-4 SN - 0340-5443 SN - 1432-0762 VL - 72 IS - 175 PB - Springer Nature Switzerland AG CY - New York ER - TY - JOUR A1 - Folkertsma, Remco A1 - Westbury, Michael V. A1 - Eccard, Jana A1 - Hofreiter, Michael T1 - The complete mitochondrial genome of the common vole, Microtus arvalis (Rodentia: Arvicolinae) JF - Mitochondrial DNA Part B N2 - The common vole, Microtus arvalis belongs to the genus Microtus in the subfamily Arvicolinae. In this study, the complete mitochondrial genome of M. arvalis was recovered using shotgun sequencing and an iterative mapping approach using three related species. Phylogenetic analyses using the sequence of 21 arvicoline species place the common vole as a sister species to the East European vole (Microtus levis), but as opposed to previous results we find no support for the recognition of the genus Neodon within the subfamily Arvicolinae, as this is, as well as the genus Lasiopodomys, found within the Microtus genus. KW - Microtus arvalis KW - Arvicolinae KW - mitochondrial genome KW - common vole KW - phylogeny Y1 - 2018 U6 - https://doi.org/10.1080/23802359.2018.1457994 SN - 2380-2359 VL - 3 IS - 1 SP - 446 EP - 447 ER - TY - THES A1 - Folkertsma, Remco T1 - Evolutionary adaptation to climate in microtine mammals N2 - Understanding how organisms adapt to their local environment is a major focus of evolutionary biology. Local adaptation occurs when the forces of divergent natural selection are strong enough compared to the action of other evolutionary forces. An improved understanding of the genetic basis of local adaptation can inform about the evolutionary processes in populations and is of major importance because of its relevance to altered selection pressures due to climate change. So far, most insights have been gained by studying model organisms, but our understanding about the genetic basis of local adaptation in wild populations of species with little genomic resources is still limited. With the work presented in this thesis I therefore set out to provide insights into the genetic basis of local adaptation in populations of two voles species: the common vole (Microtus arvalis) and the bank vole (Myodes glareolus). Both voles species are small mammals, they have a high evolutionary potential compared to their dispersal capabilities and are thus likely to show genetic responses to local conditions, moreover, they have a wide distribution in which they experience a broad range of different environmental conditions, this makes them an ideal species to study local adaptation. The first study focused on producing a novel mitochondrial genome to facilitate further research in M. arvalis. To this end, I generated the first mitochondrial genome of M. arvalis using shotgun sequencing and an iterative mapping approach. This was subsequently used in a phylogenetic analysis that produced novel insights into the phylogenetic relationships of the Arvicolinae. The following two studies then focused on the genetic basis of local adaptation using ddRAD-sequencing data and genome scan methods. The first of these involved sequencing the genomic DNA of individuals from three low-altitude and three high-altitude M. arvalis study sites in the Swiss Alps. High-altitude environments with their low temperatures and low levels of oxygen (hypoxia) pose considerable challenges for small mammals. With their small body size and proportional large body surface they have to sustain high rates of aerobic metabolism to support thermogenesis and locomotion, which can be restricted with only limited levels of oxygen available. To generate insights into high-altitude adaptation I identified a large number of single nucleotide polymorphisms (SNPs). These data were first used to identify high levels of differentiation between study sites and a clear pattern of population structure, in line with a signal of isolation by distance. Using genome scan methods, I then identified signals of selection associated with differences in altitude in genes with functions related to oxygen transport into tissue and genes related to aerobic metabolic pathways. This indicates that hypoxia is an important selection pressure driving local adaptation at high altitude in M. arvalis. A number of these genes were linked with high-altitude adaptation in other species before, which lead to the suggestion that high-altitude populations of several species have evolved in a similar manner as a response to the unique conditions at high altitude The next study also involved the genetic basis of local adaptation, here I provided insights into climate-related adaptation in M. glareolus across its European distribution. Climate is an important environmental factor affecting the physiology of all organisms. In this study I identified a large number of SNPs in individuals from twelve M. glareolus populations distributed across Europe. I used these, to first establish that populations are highly differentiated and found a strong pattern of population structure with signal of isolation by distance. I then employed genome scan methods to identify candidate loci showing signals of selection associated with climate, with a particular emphasis on polygenic loci. A multivariate analysis was used to determine that temperature was the most important climate variable responsible for adaptive genetic variation among all variables tested. By using novel methods and genome annotation of related species I identified the function of genes of candidate loci. This showed that genes under selection have functions related to energy homeostasis and immune processes. Suggesting that M. glareolus populations have evolved in response to local temperature and specific local pathogenic selection pressures. The studies presented in this thesis provide evidence for the genetic basis of local adaptation in two vole species across different environmental gradients, suggesting that the identified genes are involved in local adaptation. This demonstrates that with the help of novel methods the study of wild populations, which often have little genomic resources available, can provide unique insights into evolutionary processes. N2 - Ein Schwerpunkt der Evolutionsbiologie besteht darin, zu verstehen, wie sich Organismen an ihre lokale Umgebung anpassen. Lokale Anpassung tritt ein, wenn die Kräfte der divergierenden natürlichen Selektion im Vergleich zu anderen evolutionären Kräften stark genug sind. Ein verbessertes Verständnis der genetischen Grundlagen der lokalen Anpassung kann Informationen über die Evolutionsprozesse in Populationen liefern und ist durch seine Relevanz für durch den Klimawandel bedingte veränderte Selektionsdrücke von großer Bedeutung. Bisher wurden die meisten Erkenntnisse durch Untersuchungen an Modellorganismen gewonnen. Jedoch ist das Verständnis der genetischen Grundlagen der lokalen Anpassung in Wildpopulationen von Arten mit geringen genomischen Ressourcen noch immer begrenzt. Mit den in dieser Doktorarbeit vorgestellten Untersuchungen war es daher mein Ziel, Einblicke in die genetischen Grundlagen der lokalen Anpassung in Populationen von zwei Wühlmausarten zu geben: der Feldmaus (Microtus arvalis) und der Rötelmaus (Myodes glareolus). Bei beiden handelt es sich um kleine Säugetiere mit einem, im Vergleich zu ihrer Ausbreitungsfähigkeit, hohen Evolutionspotential. Daher ist anzunehmen, dass sie genetische Reaktionen auf lokale Bedingungen zeigen. Hinzu kommt, dass sie aufgrund ihrer großen Verbreitung ein großes Spektrum an verschiedenen Umweltbedingungen erfahren, was sie zu einer idealen Spezies, für die Untersuchung lokaler Anpassung macht. Die erste Studie dieser Arbeit konzentrierte sich auf die Erstellung eines bisher nicht verfügbaren mitochondriellen Genoms, um die weitere Forschung an M. arvalis zu erleichtern. Dies wurde mittels Shotgun-Sequenzierung und eines iterativen Kartierungsansatzes erreicht. Anschließend wurde es in einer phylogenetischen Analyse verwendet, die neue Erkenntnisse über die phylogenetischen Beziehungen der Arvicolinae lieferte. Die folgenden zwei Studien konzentrierten sich auf die genetische Basis der lokalen Anpassung unter Verwendung von ddRAD-Sequenzierungsdaten und Genom-Scan-Methoden. Die erste umfasste die Sequenzierung der genomischen DNA von Individuen aus drei M. arvalis-Untersuchungsgebieten in geringer Höhe und drei in großer Höhe in den Schweizer Alpen. Umgebungen in großer Höhe mit niedrigen Temperaturen und niedrigem Sauerstoffgehalt (Hypoxie) stellen kleine Säugetiere vor erhebliche Herausforderungen. Aufgrund ihrer geringen Körpergröße und proportional großen Körperoberfläche müssen sie hohe aerobe Stoffwechselraten aufrechterhalten, um die Thermogenese und Fortbewegung zu unterstützen, die mit begrenzter Sauerstoffverfügbarkeit eingeschränkt sein können. Um Einblicke in die Höhenanpassung zu erhalten, habe ich eine große Anzahl von Einzelnukleotidpolymorphismen (SNPs) identifiziert. Mit Hilfe dieser Daten wurden ein hohes Maß an Differenzierung zwischen den Untersuchungsorten und ein klares Muster der Populationsstruktur zusammen mit einem isolation-by-distance Signal identifiziert. Unter Verwendung von Genom-Scan-Methoden identifizierte ich Selektionssignale in Genen, die mit Höhenunterschieden verbunden werden. Diese besitzen Funktionen, die mit dem Sauerstofftransport in das Gewebe sowie mit aeroben Stoffwechselwegen zusammenhängen. Dies weist darauf hin, dass Hypoxie ein wichtiger Selektionsdruck für die lokale Anpassung in großer Höhe für M. arvalis ist. Einige dieser Gene sind bereits früher mit der Höhenanpassung bei anderen Arten in Verbindung gebracht worden. Dies führte zu der Annahme, dass sich Populationen in großer Höhe lebender verschiedener Arten in Anpassung an die einzigartigen Bedingungen in großer Höhe auf ähnliche Weise entwickelt haben. Die nächste Studie befasste sich ebenfalls mit den genetischen Grundlagen der lokalen Anpassung. Hier stellte ich Erkenntnisse über die klimabedingte Anpassung von M. glareolus in ihrem europäischen Verbreitungsgebiet vor. Das Klima ist ein wichtiger Umweltfaktor, der die Physiologie aller Organismen beeinflusst. In dieser Studie identifizierte ich zehntausende SNPs bei Individuen aus zwölf in ganz Europa verteilten M. glareolus-Populationen. Diese ergaben eine starke Differenzierung der Populationen mit deutlicher Populationsstruktur und einem Signal für isolation-by-distance. Anschließend verwendete ich Genom-Scan-Methoden, um mögliche Loci zu identifizieren, die mit dem Klima verbundene Selektionssignale aufweisen, wobei der Schwerpunkt dabei auf polygenen Loci lag. Eine Multivariaten Analysemethode ermittelte, dass die Temperatur die wichtigste Klimavariable unter allen getesteten Variablen ist, die für die adaptive genetische Variation verantwortlich ist. Mit Hilfe neuartiger Methoden und der Annotation von Genomen verwandter Spezies identifizierte ich die Funktion von Genen an Kandidatenloci. Diese zeigten, dass die unter Selektion stehenden Gene Funktionen im Zusammenhang mit der Energiehomöostase und den Immunprozessen ausüben. Dies wiederum deutet darauf hin, dass sich die Populationen von M. glareolus in Reaktion auf die lokale Temperatur und den spezifischen lokalen Selektionsdruck für Krankheitserreger entwickelt haben. Die in dieser Arbeit vorgestellten Studien liefern Belege für die genetische Basis der lokalen Anpassung auf verschiedene Umweltgradienten in zwei Wühlmausarten. Dies deutet darauf hin, dass die identifizierten Gene an der lokalen Anpassung beteiligt sind. Darüber hinaus zeigt dies, dass Untersuchungen wildlebender Populationen mit geringen genomischen Ressourcen durch den Einsatz neuartiger Methoden einzigartige Einblicke in evolutionäre Prozesse ermöglichen können. T2 - Evolutionäre Klimaanpassungen bei Wühlmausarten KW - Genomics KW - Local adaptation KW - Altitude KW - Climate KW - Microtus arvalis KW - Myodus glareolus KW - Höhe KW - Klima KW - Genomik KW - lokale Anpassung KW - Feldmaus KW - Rötelmaus Y1 - 2020 U6 - http://nbn-resolving.de/urn/resolver.pl?urn:nbn:de:kobv:517-opus4-476807 ER -