A quantitative genetic signature of senescence in a short-lived perennial plant.

The evolution of senescence (the physiological decline of organisms with age) poses an apparent paradox because it represents a failure of natural selection to increase the survival and reproductive performance of organisms. The paradox can be resolved if natural selection becomes less effective with age, because the death of postreproductive individuals should have diminished effects on Darwinian fitness [1, 2]. A substantial body of empirical work is consistent with this prediction for animals, which transmit their genes to progeny via an immortal germline. However, such evidence is still lacking in plants, which lack a germline and whose reproduction is diffuse and modular across the soma. Here, we provide experimental evidence for a genetic basis of senescence in the short-lived perennial plant Silene latifolia. Our pedigree-based analysis revealed a marked increase with age in the additive genetic variance of traits closely associated with fitness. This result thus extends to plants the quantitative genetic support for the evolutionary theory of senescence.

Predation drives interpopulation differences in parental care expression.

Expressing parental care after oviposition or parturition is usually an obligate (evolved) trait within a species, despite evolutionary theory predicting that widespread species should vary in whether or not they express parental care according to local selection pressures. The lizard Eutropis longicaudata expresses maternal care only in a single population throughout its large geographical range, but why this pattern occurs is unknown. We used reciprocal translocation and predator exclusion experiments to test whether this intraspecific variation is a fixed trait within populations and whether predator abundance explains this perplexing pattern. Wild-caught female lizards that were reciprocally translocated consistently guarded or abandoned eggs in line with their population of origin. By contrast, most lizards raised in a common garden environment and subsequently released as adults adopted the maternal care strategy of the recipient population, even when the parents originated from a population lacking maternal care. Egg predation represents a significant fitness cost in the populations where females display egg-guarding behaviour, but guarding eggs outweighs this potential cost by increasing hatching success. These results imply that predators can be a driving force in the expression of parental care in instances where it is normally absent and that local selection pressure is sufficient to cause behavioural divergence in whether or not parental care is expressed.

Relationships of Scincid Lizards (Mabuya spp; Reptilia: Scincidae) from the Cape Verde Islands Based on Mitochondrial and Nuclear DNA Sequences

Partial DNA sequences from two mitochondrial (mt) and one nuclear gene (cytochrome b, 12S rRNA, and C-mos) were used to estimate the phylogenetic relationships among the six extant species of skinks endemic to the Cape Verde Archipelago. The species form a monophyletic unit, indicating a single colonization of the islands, probably from West Africa. Mabuya vaillanti and M. delalandii are sister taxa, as indicated by morphological characters. Mabuya fogoensis and M. stangeri are closely related, but the former is probably paraphyletic. Mabuya spinalis and M. salensis are also probably paraphyletic. Within species, samples from separate islands always form monophyletic groups. Some colonization events can be hypothesized, which are in line with the age of the islands. C-mos variation is concordant with the topology derived from mtDNA.

Proto-South-East Asia as a trigger of early angiosperm diversification

Darwin described as an 'abominable mystery' the abrupt origin of angiosperms in the mid-Cretaceous and the high diversification rates in their early history. The father of evolutionary theory could not fathom this rapid diversification and rather invoked that 'there was during long ages a small isolated continent in the S. hemisphere, which served as the birthplace of the higher plants'. In this essay, we comment on the spatial origin of angiosperms, but focus primarily on understanding the abiotic factors that promoted the early diversification of angiosperms by reviewing palaeobotanical, palaeogeographical, phylogenetics and biogeographical evidence. We argue that islands located in the region today occupied by South-East Asia played a major role in angiosperm diversification during the Late Jurassic and Early Cretaceous.

Gibbard's expressivism: an interdisciplinary critical analysis

This paper examines key aspects of Allan Gibbard's psychological account of moral activity. Inspired by evolutionary theory, Gibbard paints a naturalistic picture of morality mainly based on two specific types of emotion: guilt and anger. His sentimentalist and expressivist analysis is also based on a particular conception of rationality. I begin by introducing Gibbard's theory before testing some key assumptions underlying his system against recent empirical data and theories. The results cast doubt on some crucial aspects of Gibbard's philosophical theory, namely his reduction of morality to anger and guilt, and his theory of 'normative governance'. Gibbard's particular version of expressivism may be undermined by these doubts.

On the evolution of harming and recognition in finite panmictic and infinite structured populations.

Natural selection may favor two very different types of social behaviors that have costs in vital rates (fecundity and/or survival) to the actor: helping behaviors, which increase the vital rates of recipients, and harming behaviors, which reduce the vital rates of recipients. Although social evolutionary theory has mainly dealt with helping behaviors, competition for limited resources creates ecological conditions in which an actor may benefit from expressing behaviors that reduce the vital rates of neighbors. This may occur if the reduction in vital rates decreases the intensity of competition experienced by the actor or that experienced by its offspring. Here, we explore the joint evolution of neutral recognition markers and marker-based costly conditional harming whereby actors express harming, conditional on actor and recipient bearing different conspicuous markers. We do so for two complementary demographic scenarios: finite panmictic and infinite structured populations. We find that marker-based conditional harming can evolve under a large range of recombination rates and group sizes under both finite panmictic and infinite structured populations. A direct comparison with results for the evolution of marker-based conditional helping reveals that, if everything else is equal, marker-based conditional harming is often more likely to evolve than marker-based conditional helping.

Inbreeding and selection on sex ratio in the bark beetle Xylosandrus germanus.

ABSTRACT: BACKGROUND: Local Mate Competition (LMC) theory predicts a female should produce a more female-biased sex ratio if her sons compete with each other for mates. Because it provides quantitative predictions that can be experimentally tested, LMC is a textbook example of the predictive power of evolutionary theory. A limitation of many earlier studies in the field is that the population structure and mating system of the studied species are often estimated only indirectly. Here we use microsatellites to characterize the levels of inbreeding of the bark beetle Xylosandrus germanus, a species where the level of LMC is expected to be high. RESULTS: For three populations studied, genetic variation for our genetic markers was very low, indicative of an extremely high level of inbreeding (FIS = 0.88). There was also strong linkage disequilibrium between microsatellite loci and a very strong genetic differentiation between populations. The data suggest that matings among non-siblings are very rare (3%), although sex ratios from X. germanus in both the field and the laboratory have suggested more matings between non-sibs, and so less intense LMC. CONCLUSIONS: Our results confirm that caution is needed when inferring mating systems from sex ratio data, especially when a lack of biological detail means the use of overly simple forms of the model of interest.

Studying the molecular causes of ageing in ants

RÉSUMÉ DE THÈSE Au cours de ma thèse, je me suis intéressée aux causes physiologiques du vieillissement en utilisant les fourmis comme modèle. Les trois castes de fourmis - les mâles, les ouvrières et les reines - présentent des longévités très différentes, tout en étant génétiquement identiques. Ceci implique que les différences de longévité sont dues à des variations entre castes dans le pattern d'expression de gènes. Mon travail chez la fourmi a consisté d'une part à mettre en place les outils pour identifier de tels gènes à grande échelle, de l'autre à étudier le rôle de gènes et de mécanismes qui affectent la longévité chez d'autres espèces. Pour identifier de nouveaux gènes potentiellement impliqués dans le vieillissement, nous avons développé des puces à ADN. Cette technique permet la comparaison du niveau d'expression de milliers de gènes entre deux échantillons. L'application de cette méthode aux reines et ouvrières adultes nous a jusqu'à présent permis d'identifier neuf gènes surexprimés chez les reines. Trois d'entre eux sont potentiellement impliqués dans le maintien et la réparation du soma, deux processus qui sont supposés avoir un impact crucial sur la longévité. Parmi les mécanismes impliqués dans le vieillissement chez d'autres espèces, nous nous sommes principalement intéressés aux télomères, qui sont les extrémités des chromosomes. Chez les vertébrés, les télomères se raccourcissent à chaque division cellulaire, entre autres parce que l'ADN polymérase ne peut répliquer cette partie des chromosomes en entier. Or des télomères courts entravent la prolifération des cellules et peuvent même induire l'apoptose, ce qui pourrait se répercuter sur la capacité des organismes à régénérer des tissus. J'ai pu montrer que chez les fourmis mâles (la caste qui vit le moins longtemps) les télomères se raccourcissent beaucoup plus vite que chez les reines et les ouvrières. L'explication la plus plausible pour cette différence est que les mâles, étant adapté à une vie très éphémère, n'investissent qu'un minimum d'énergie dans la machinerie de maintenance qui assure le bon fonctionnement des cellules. Ces résultats sont intéressants car ils permettent pour la première fois de faire le lien entre les théories évolutives du vieillissement et la biologie des télomères. THESIS ABSTRACT During my thesis I used ants as a model to study the proximate (i.e., molecular) causes of ageing and lifespan determination. Ant queens, workers and males differ tremendously in lifespan, although all three castes are genetically identical. Importantly, this implies that genes and molecular pathways responsible for modulating lifespan are regulated in a caste-specific manner. To find new genes potentially involved in ageing, we first constructed 371-gene-cDNA microarrays for the ant L. niger. This molecular tool can be used to survey the relative gene expression levels of two samples for thousands of genes simultaneously. By applying this method to adult queens and workers we identified nine genes that are overexpressed in queens. Three of them are putatively involved in somatic maintenance and repair, two processes that have been previously suggested as important for ageing and lifespan determination. We expect to identify many more candidate genes in the near future by using the 9000-gene fire ant microarrays we have recently developed. We also investigated whether factors linked to ageing in other organisms could affect lifespan determination in ants. One project was on telomeres, the ends of linear chromosomes. For various reasons telomeres shorten with every cell division. Since short telomeres can lead to cellular defects such as impaired cell division, telomeres have been hypothesized as playing a role in ageing. We tested whether telomere length in ant somatic tissues correlates with caste-specific lifespan in young adults. The short-lived L. niger mates did indeed have significantly shorter telomeres than the longer-lived queens and workers, probably because telomere attrition is faster in males than in queens and workers. Queens did not, however, have longer telomeres than the shorter-lived workers. These findings are consistent with the idea that telomere length may play a role in ageing under some circumstances, but they also clearly demonstrate that other factors must be involved. We argue that sex-specific telomere length patterns in ants ultimately reflect adaptive differences in the level of somatic maintenance between males and females, and thus create a link between telomere biology and the evolutionary theory of ageing.

Evidence of recombination in putative ancient asexuals.

Ancient asexuals have been considered to be a contradiction of the basic tenets of evolutionary theory. Barred from rearranging genetic variation by recombination, their reduced number of gene arrangements is thought to hamper their response to changing environments. For the same reason, it should be difficult for them to avoid the build-up of deleterious mutations. Several groups of taxonomically diverse organisms are thought to be ancient asexuals, although clear evidence for or against the existence of recombination events is scarce. Several methods have recently been developed for predicting recombination events by analyzing aligned sequences of a given region of DNA that all originate from one species. The methods are based on phylogenetic, substitution, and compatibility analyses. Here we present the results of analyses of sequence data from different loci studied in several groups of evolutionarily distant species that are considered to be ancient asexuals, using seven different types of analysis. The groups of organisms were the arbuscular mycorrhizal fungi (Glomales), Darwinula stevensoni (Darwinuloidea crustacean ostracods) and the bdelloid rotifers (Bdelloidea), which are thought to have been asexual for the last 400, 25-100, and 35-40 Myr, respectively. The seven different analytical methods evaluated the evolutionary relationships among haplotypes, and these methods had previously been shown to be reliable for predicting the occurrence of recombination events. Despite the different degree of genetic variation among the different groups of organisms, at least some evidence for recombination was found in all species groups. In particular, predictions of recombination events in the arbuscular mycorrhizal fungi were frequent. Predictions of recombination were also found for sequence data that have previously been used to infer the absence of recombination in bdelloid rotifers. Although our results have to be taken with some caution because they could signal very ancient recombination events or possibly other genetic variation of nonrecombinant origin, they suggest that some cryptic recombination events may exist in these organisms.

Long live the queen: studying aging in social insects

Aging is a fascinating, albeit controversial, chapter in biology. Few other subjects have elicited more than a century of ever-increasing scientific interest. In this review, we discuss studies on aging in social insects, a group of species that includes ants and termites, as well as certain bee and wasp species. One striking feature of social insects is the lifespan of queens (reproductive females), which can reach nearly 30 years in some ant species. This is over 100 times the average lifespan of solitary insects. Moreover, there is a tremendous variation in lifespan among castes, with queens living up to 500 times longer than males and 10 times longer than workers (non-reproductive individuals). This lifespan polymorphism has allowed researchers to test the evolutionary theory of aging and Y more recently Y to investigate the proximate causes of aging. The originality of these studies lies in their use of naturally evolved systems to address questions related to aging and lifespan determination that cannot be answered using the conventional model organisms.

Risk assessment tool for radical technologies – Case Technology A and Technology B

Työssä testataan radikaalien teknologioiden liiketoimintariskien evaluointityökalua. Työn teoreettinen tausta koostuu teknologia- ja innovaatioteorioista hyödyntäen myös resurssipohjaista yritysteoriaa täydennettynä evolutionäärisellä teorialla. Teoreettisessa osuudessa rakennetaan viitekehys, jolla liiketoimintariskejä voidaan arvioida ja muodostaa riskiprofiili. Liiketoimintariskien muuttujina ovat markkina-, teknologia- ja organisaatioriskit. Primäärisenä tietolähteenä käytettiin teema- ja strukturoituja haastatteluita. Ensimmäinen haastattelu käsitti evaluointityökalun käytettävyyttä ja riskienhallintaa yleensä. Loput haastattelut liittyivät teknologian A ja teknologian B liiketoimintariskien arvioimiseen. Tulokset osoittavat molemmat teknologiat sisältävän radikaaleille teknologioille ominaisia epävarmuustekijöitä. Riskiprofiilin hyödyllisyys liittyy liiketoimintariskien samanaikaiseen havaitsemiseen auttaen näin päätöksenteossa. Tärkeää evaluoinnissa on kiinnittää huomiota näkökulmaan, josta riskejä tarkastellaan riskiprofiilin validiteetin parantamiseksi.

On Innovative Search: the use of internal and external sources of innovation among Finnish innovators

This dissertation explores the use of internal and external sources of knowledge in modern innovation processes. It builds on a framework that combines theories such as a behavioural theory of the firm, the evolutionary theory of economic change, and modern approaches to strategic management. It follows the recent increase in innovation research focusing on the firm-level examination of innovative activities instead of traditional industry-level determinants. The innovation process is seen as a problem- and slack- driven search process, which can take several directions in terms of organizational boundaries in the pursuit of new knowledge and other resources. It thus draws on recent models of technological change, according to which firms nowadays should build their innovative activities on both internal and external sources of innovation rather than relying solely on internal resources. Four different research questions are addressed, all of which are empirically investigated via a rich dataset covering Finnish innovators collected by Statistics Finland. Firstly, the study examines how the nature of problems shapes the direction of any search for new knowledge. In general it demonstrates that the nature of the problem does affect the direction of the search, although under resource constraints firms tend to use external rather than internal sources of knowledge. At the same time, it shows that those firms that are constrained in terms of finance seem to search both internally and externally. Secondly, the dissertation investigates the relationships between different kinds of internal and external sources of knowledge in an attempt to find out where firms should direct their search in order to exploit the potential of a distributed innovation process. The concept of complementarities is applied in this context. The third research question concerns how the use of external knowledge sources – openness to external knowledge – influences the financial performance of firms. Given the many advantages of openness presented in the current literature, the focus is on how it shapes profitability. The results reveal a curvilinear relationship between profitability and openness (taking an inverted U-shape), the implication being that it pays to be open up to a certain point, but being too open to external sources may be detrimental to financial performance. Finally, the dissertation addresses some challenges in CISbased innovation research that have received relatively little attention in prior studies. The general aim is to underline the fact that comprehensive understanding of the complex process of technological change requires the constant development of methodological approaches (in terms of data and measures, for example). All the empirical analyses included in the dissertation are based on the Finnish CIS (Finnish Innovation Survey 1998-2000).

Enabling Change in Universities: Enhancing Education for Sustainable Development with Tools for Quality Assurance

Enabling Change in Universities: Enhancing Education for Sustainable Development with Tools for Quality Assurance This thesis deals with enabling change in universities, more explicitly enhancing education for sustainable development with tools for quality assurance. Change management is a discipline within management that was developed in the 1980s because business changed from being predictable to unpredictable. The PEST mnemonic is a method to categorize factors enabling change; such as political, economic, socio-cultural and technological factors, which all affect higher education. A classification of a change, in either hard or soft, can help understanding the type of change that an organization is facing. Hard changes are more applied to problems that have clear objectives and indicators, with a known cause of the problem. Soft changes are applied to larger problems that affect the entire organization or beyond it. The basic definition for sustainable development is: the future generations should have similar opportunities as the previous. The UN has set as a global goal an integration of education for sustainable development (ESD) at all levels of education during 2005- 2014. The goal is set also in universities, the graduates of which are future leaders for all labor markets. The objective for ESD in higher education is that graduates obtain the competence to take economic, social and environmental costs and benefits into account when making decisions. Knowledge outcomes should aim for systematic and holistic thinking, which requires cross disciplinary education. So far, the development of ESD has not achieved its goals. The UN has identified a need for more transdisclipnary research in ESD. A joint global requirement for universities is quality assurance, the aim of which is to secure and improve teaching and learning. Quality, environmental and integrated management systems are used by some universities for filling the quality assurance requirements. The goal of this thesis is to open up new ways for enhancing ESD in universities, beyond the forerunners; by exploring how management systems could be used as tools for promoting ESD. The thesis is based on five studies. In the first study, I focus on if and how tools for quality assurance could be benefitted for promoting ESD. It is written from a new perspective, the memetic, for reaching a diversity of faculty. A meme is an idea that diffuses from brain to brain. It can be applied for cultural evolution. It is a theory that is based on the evolutionary theory by Darwin, applied for social sciences. In the second Paper, I present the results from the development of the pilot process model for enhancing ESD with management systems. The development of the model is based on a study that includes earlier studies, a survey in academia and an analysis of the practice in 11 universities in the Nordic countries. In the third study, I explore if the change depends on national culture or if it is global. It is a comparative study on both policy and implementation level, between the Nordic countries and China. The fourth study is a single case study based on change management. In this study, I identify what to consider in order to enable the change: enhancing ESD with tools for quality assurance in universities. In the fifth Paper, I present the results of the process model for enhancing ESD with management systems. The model was compared with identified drivers and barriers for enhancing ESD and for implementing management systems. Finally, the process model was piloted and applied for identifying sustainability aspects in curricula. Action research was chosen as methodology because there are not already implemented approaches using quality management for promoting ESD, why the only way to study this is to make it happen. Another reason for choosing action research is since it is essential to involve students and faculty for enhancing ESD. Action based research consists of the following phases: a) diagnosing, b) planning action, c) taking action and d) evaluating action. This research was made possible by a project called Education for Sustainable Development in Academia in the Nordic countries, ESDAN, in which activities were divided into these four phases. Each phase ended with an open seminar, where the results of the study were presented. The objective for the research project was to develop a process for including knowledge in sustainable development in curricula, which could be used in the quality assurance work. Eleven universities from the Nordic countries cooperated in the project. The aim was, by applying the process, to identify and publish examples of relevant sustainability aspects in different degree programs in universities in the Nordic countries. The project was partly financed by the Nordic Council of Ministers and partly by the participating pilot universities. Based on the results of my studies, I consider that quality, environmental and integrated management systems can be used for promoting ESD in universities. Relevant sustainability aspects have been identified in different fields of studies by applying the final process model. The final process model was compared with drivers and barriers for enhancing ESD and for implementing management systems in universities and with succeeding with management systems in industry. It corresponds with these, meaning that drivers are taken into account and barriers tackled. Both ESD and management systems in universities could be considered successful memes, which can reflect an effective way of communication among individuals. I have identified that management systems could be used as tools for hard changes and to support the soft change of enhancing ESD in universities with management system. Based on the change management study I have summarized recommendations on what to consider in order to enable the studied change. The main practical implications of the results are that the process model could be applied for assessment, benchmarking and communication of ESD, connected to quality assurance, when applied. This is possible because the information can be assembled in one picture, which facilitates comparison. The memetic approach can be applied for structuring. It is viable to make comparative studies between cultures, for getting insight in special characteristics of the own culture. Action based research is suitable for involving faculty. Change management can be applied for planning a change, which both enhancing ESD and developing management systems are identified to be.

Synecological comparisons sustained by ecophysiological fingerprinting of intrinsic photosynthetic capacity of plants as assessed by measurements of light response curves

In some literature variations in photosynthetic rates are considered to be of little relevance for individual fitness. This depends among other things on how one defines fitness, i.e. if one takes strictly Darwinian fitness as seed production or if one needs to evaluate particular traits and consider plant establishment. It also matters if one takes the Darwinian "organism individual" as the central entity in evolution ("individual fitness") or the "species individual" in a modified "Structure of Evolutionary Theory" sensu Stephen Jay Gould. A phenotypically expressed trait like photosynthetic rate, even if intra- and interspecific differences may be small, can matter in habitat performance and niche acquisition. Light dependence curves (LCs) of photosynthetic rates are now readily measured under field conditions using miniaturized equipment of pulse amplitude modulated fluorometers. In contrast to actual momentary measurements of quantum yield of photosynthesis under actually prevailing ambient conditions, LC measurements reflect the expressed intrinsic capacity of photosynthesis. In this review we explore the power of LC measurements yielding cardinal points such as maximum apparent electron transport rate of photosystem II (ETRmax) and saturating photosynthetically active radiation (PARsat) in making intra- and interspecific comparisons of plant performance and synecological fingerprinting in ecophysiological studies across species, sites, habitats and ecosystems.

Inclusive fitness maximization: an axiomatic approach

Kin selection theorists argue that evolution in social contexts will lead organisms to behave as if maximizing their inclusive, as opposed to personal, fitness. The inclusive fitness concept allows biologists to treat organisms as akin to rational agents seeking to maximize a utility function. Here we develop this idea and place it on a firm footing by employing a standard decision-theoretic methodology. We show how the principle of inclusive fitness maximization and a related principle of quasi-inclusive fitness maximization can be derived from axioms on an individual’s ‘as if preferences’ (binary choices). Our results help integrate evolutionary theory and rational choice theory, help draw out the behavioural implications of inclusive fitness maximization, and point to a possible way in which evolution could lead organisms to implement it.