AMP-activated protein kinase plays an important evolutionary conserved role in the regulation of glucose metabolism in fish skeletal muscle cells

AMPK, a master metabolic switch, mediates the observed increase of glucose uptake in locomotory muscle of mammals during exercise. AMPK is activated by changes in the intracellular AMP:ATP ratio when ATP consumption is stimulated by contractile activity but also by AICAR and metformin, compounds that increase glucose transport in mammalian muscle cells. However, the possible role of AMPK in the regulation of glucose metabolism in skeletal muscle has not been investigated in other vertebrates, including fish. In this study, we investigated the effects of AMPK activators on glucose uptake, AMPK activity, cell surface levels of trout GLUT4 and expression of GLUT1 and GLUT4 as well as the expression of enzymes regulating glucose disposal and PGC1α in trout myotubes derived from a primary muscle cell culture. We show that AICAR and metformin significantly stimulated glucose uptake (1.6 and 1.3 fold, respectively) and that Compound C completely abrogated the stimulatory effects of the AMPK activators on glucose uptake. The combination of insulin and AMPK activators did not result in additive nor synergistic effects on glucose uptake. Moreover, exposure of trout myotubes to AICAR and metformin resulted in an increase in AMPK activity (3.8 and 3 fold, respectively). We also provide evidence suggesting that stimulation of glucose uptake by AMPK activators in trout myotubes may take place, at least in part, by increasing the cell surface and mRNA levels of trout GLUT4. Finally, AICAR increased the mRNA levels of genes involved in glucose disposal (hexokinase, 6-phosphofructokinase, pyruvate kinase and citrate synthase) and mitochondrial biogenesis (PGC-1α) and did not affect glycogen content or glycogen synthase mRNA levels in trout myotubes. Therefore, we provide evidence, for the first time in non-mammalian vertebrates, suggesting a potentially important role of AMPK in stimulating glucose uptake and utilization in the skeletal muscle of fish.

Combining conservative and variable markers to inferthe evolutionary history of Prunus subgen. Amygdalus s.l.under domestication

The genus Prunus L. is large and economically important. However, phylogenetic relationships within Prunus at low taxonomic level, particularly in the subgenus Amygdalus L. s.l., remain poorly investigated. This paper attempts to document the evolutionary history of Amygdalus s.l. and establishes a temporal framework, by assembling molecular data from conservative and variable molecular markers. The nuclear s6pdh gene in combination with the plastid trnSG spacer are analyzed with bayesian and maximum likelihood methods. Since previous phylogenetic analysis with these markers lacked resolution, we additionally analyzed 13 nuclear SSR loci with the δµ2 distance, followed by an unweighted pair group method using arithmetic averages algorithm. Our phylogenetic analysis with both sequence and SSR loci confirms the split between sections Amygdalus and Persica, comprising almonds and peaches, respectively. This result is in agreement with biogeographic data showing that each of the two sections is naturally distributed on each side of the Central Asian Massif chain. Using coalescent based estimations, divergence times between the two sections strongly varied when considering sequence data only or combined with SSR. The sequence-only based estimate (5 million years ago) was congruent with the Central Asian Massif orogeny and subsequent climate change. Given the low level of differentiation within the two sections using both marker types, the utility of combining microsatellites and data sequences to address phylogenetic relationships at low taxonomic level within Amygdalus is discussed. The recent evolutionary histories of almond and peach are discussed in view of the domestication processes that arose in these two phenotypically-diverging gene pools: almonds and peaches were domesticated from the Amygdalus s.s. and Persica sections, respectively. Such economically important crops may serve as good model to study divergent domestication process in close genetic pool.

Functional and evolutionary analysis of DXL1, a non-essential gene encoding a 1-deoxy-D-xylulose 5-phosphate synthase like protein in Arabidopsis thaliana

The synthesis of 1-deoxy-D-xylulose 5-phosphate (DXP), catalyzed by the enzyme DXP synthase (DXS), represents a key regulatory step of the 2-C-methyl-D-erythritol 4-phosphate (MEP) pathway for isoprenoid biosynthesis. In plants DXS is encoded by small multigene families that can be classified into, at least, three specialized subfamilies. Arabidopsis thaliana contains three genes encoding proteins with similarity to DXS, including the well-known DXS1/CLA1 gene, which clusters within subfamily I. The remaining proteins, initially named DXS2 and DXS3, have not yet been characterized. Here we report the expression and functional analysis of A. thaliana DXS2. Unexpectedly, the expression of DXS2 failed to rescue Escherichia coli and A. thaliana mutants defective in DXS activity. Coherently, we found that DXS activity was negligible in vitro, being renamed as DXL1 following recent nomenclature recommendation. DXL1 is targeted to plastids as DXS1, but shows a distinct expression pattern. The phenotypic analysis of a DXL1 defective mutant revealed that the function of the encoded protein is not essential for growth and development. Evolutionary analyses indicated that DXL1 emerged from DXS1 through a recent duplication apparently specific of the Brassicaceae lineage. Divergent selective constraints would have affected a significant fraction of sites after diversification of the paralogues. Furthermore, amino acids subjected to divergent selection and likely critical for functional divergence through the acquisition of a novel, although not yet known, biochemical function, were identified. Our results provide with the first evidences of functional specialization at both the regulatory and biochemical level within the plant DXS family.

Complex patterns of global spread in invasive insects: eco-evolutionary and management consequences

The advent of simple and affordable tools for molecular identification of novel insect invaders and assessment of population diversity has changed the face of invasion biology in recent years. The widespread application of these tools has brought with it an emerging understanding that patterns in biogeography, introduction history and subsequent movement and spread of many invasive alien insects are far more complex than previously thought. We reviewed the literature and found that for a number of invasive insects, there is strong and growing evidence that multiple introductions, complex global movement, and population admixture in the invaded range are commonplace. Additionally, historical paradigms related to species and strain identities and origins of common invaders are in many cases being challenged. This has major consequences for our understanding of basic biology and ecology of invasive insects and impacts quarantine, management and biocontrol programs. In addition, we found that founder effects rarely limit fitness in invasive insects and may benefit populations (by purging harmful alleles or increasing additive genetic variance). Also, while phenotypic plasticity appears important post-establishment, genetic diversity in invasive insects is often higher than expected and increases over time via multiple introductions. Further, connectivity among disjunct regions of global invasive ranges is generally far higher than expected and is often asymmetric, with some populations contributing disproportionately to global spread. We argue that the role of connectivity in driving the ecology and evolution of introduced species with multiple invasive ranges has been historically underestimated and that such species are often best understood in a global context.

The use of future-oriented knowledge in regional innovation processes: Research on knowledge generation, transfer and conversion

The importance of the regional level in research has risen in the last few decades and a vast literature in the fields of, for instance, evolutionary and institutional economics, network theories, innovations and learning systems, as well as sociology, has focused on regional level questions. Recently the policy makers and regional actors have also began to pay increasing attention to the knowledge economy and its needs, in general, and the connectivity and support structures of regional clusters in particular. Nowadays knowledge is generally considered as the most important source of competitive advantage, but even the most specialised forms of knowledge are becoming a short-lived resource for example due to the accelerating pace of technological change. This emphasizes the need of foresight activities in national, regional and organizational levels and the integration of foresight and innovation activities. In regional setting this development sets great challenges especially in those regions having no university and thus usually very limited resources for research activities. Also the research problem of this dissertation is related to the need to better incorporate the information produced by foresight process to facilitate and to be used in regional practice-based innovation processes. This dissertation is a constructive case study the case being Lahti region and a network facilitating innovation policy adopted in that region. Dissertation consists of a summary and five articles and during the research process a construct or a conceptual model for solving this real life problem has been developed. It is also being implemented as part of the network facilitating innovation policy in the Lahti region.

Evolutionary analysis of mitogenomes from parasitic and free-living flatworms

Mitochondrial genomes (mitogenomes) are useful and relatively accessible sources of molecular data to explore and understand the evolutionary history and relationships of eukaryotic organisms across diverse taxonomic levels. The availability of complete mitogenomes from Platyhelminthes is limited; of the 40 or so published most are from parasitic flatworms (Neodermata). Here, we present the mitogenomes of two free-living flatworms (Tricladida): the complete genome of the freshwater species Crenobia alpina (Planariidae) and a nearly complete genome of the land planarian Obama sp. (Geoplanidae). Moreover, we have reanotated the published mitogenome of the species Dugesia japonica (Dugesiidae). This contribution almost doubles the total number of mtDNAs published for Tricladida, a species-rich group including model organisms and economically important invasive species. We took the opportunity to conduct comparative mitogenomic analyses between available free-living and selected parasitic flatworms in order to gain insights into the putative effect of life cycle on nucleotide composition through mutation and natural selection. Unexpectedly, we did not find any molecular hallmark of a selective relaxation in mitogenomes of parasitic flatworms; on the contrary, three out of the four studied free-living triclad mitogenomes exhibit higher A+T content and selective relaxation levels. Additionally, we provide new and valuable molecular data to develop markers for future phylogenetic studies on planariids and geoplanids.

Brain size and evolutionary diversification in birds

The role of behavior in evolution remains controversial, despite that some ideas are over 100 years old. Changes in behavior are generally believed to enhance evolution by exposing individuals to new selective pressures and by facilitating range expansions. However, this hypothesis lacks firm empirical evidence. Moreover, behavioral changes can also inhibit evolution by hiding heritable variation from natural selection. Taking advantage of the complete phylogeny of extant birds, a new species-level measure of past diversification rate and the best existing measures of brain size (n = 1326 species), I show here that relative brain size is associated (albeit weakly) with diversification rates. Assuming that brain relative size reflects behavioral flexibility, an assumption well-supported by evidence, this finding supports the idea that behavior can enhance evolutionary diversification. This view is further supported by the discovery that the most important factor influencing diversification rates is ecological generalism, which is believed to require behavioral flexibility. Thus, behavioral changes that expose animals to a variety of environments can have played an important role in the evolution of birds.

Analysis of metabolites from plants of the Swartzia genus using chemical indexes: evolutionary tendencies

The chemical indexes, suggested by Gottlieb et al., have not been used before regarding evolutionary tendency of species in the Swartzia genus. However, the importance of this work encouraged for an analysis of the Swartzia genus using the metabolites isolated from nine species. The analysis, based on calculated chemical indexes, provided an evolutionary tendency for these plants, which correlates with the classification based on morphological analysis.

The molecular mechanisms and evolutionary significance of plumage colour variation in pied flycatchers (Ficedula hypoleuca)

A common feature of natural populations is that individuals differ in morphology, physiologyand behavior (i.e .phenotype). A thorough understanding of the molecular mechanisms and evolutionary forces behind this phenotypic variation is a prerequisite for understanding evolution.This thesis examines the molecular mechanism and the roles of the different evolutionary forces in plumage colour variation in pied flycatchers (Ficedulahypoleuca). Malepied flycatchers exhibit marked variation in both pigmentary and structural plumage colourand the trait has repeatedly been suggested to be of adaptive significance. An examination of plumage colour variation on reproductive output trevealed that structural colouration, and more specifically the degree of ultraviolet (UV) reflectance had an effect on number of young sired. Paternity analyses of breeding males revealed that males that had been cuckolded by their social mate tended to be less UV reflectant than males that had not been cuckolded.Neither pigment-based norstructural colouration was found to affect the probability of siring young in other nests. Phenotypic differentiation was found to be markedly greater than differentiation at neutralgenetic markers across the pied flycatcher breeding range. Furthermore patterns of differentiationin phenotypes and selectively neutral genes were not uniform. Outlier tests searching for genomic footprints of selection revealed elevated levels of genetic divergence in a gene associated with feather development (and thus potentially structural colouration) and ultraviolet vision. Th eobserved differentiation in allelic frequencies was particularly pronounced in the Spanish piedflycatcher populations. Examining gene expression during feather development indicated that the TYRP1 gene (known to be involved in the production of black pigment) may be relevant in generating phenotypic variation in pied flycatcher plumage. Also, energy homeostasis related genesfeatured prominently among the genes found to be expressed in one extreme phenotype but not the other. This is of particular interest in light of what is known about the pleiotropy ofthe melanocortin system which underlies brown-black pigment production. The melanocortinsystem is also associated with energy homeostasis (among a number of other physiological functions) and thus the results could be pointing to the signalling function of brown-blackplumage. Plumage colour variation in pied flycatchers, both structural and pigmentary, can thus beconcluded to be exhibiting signals of non-neutral evolution. Structural colouration was found to play a role in sexual selection and putative signals of selection were further detected in acandidate gene for this trait. Evidence for non-neutral evolution of pigmentary colouration was also detected. These findings, together with the fact that preliminary evidence for an energy balance associated signalling function for plumage was found, present good starting points for further investigations into the meaning and mechanisms of plumage colour variation in piedflycatchers.

Convergence in Industry Evolution

The objective of the thesis is to enhance understanding of the evolution of convergence. Previous research has shown that the technological interfaces between distinct industries are one of the major sources of new radical cross-industry innovations. Despite the fact that convergence in industry evolution has attracted a substantial managerial interest, the conceptual confusion within the field of convergence exists. Firstly, this study clarifies the convergence phenomenon and its impact to industry evolution. Secondly, the study creates novel patent analysis methods to analyze technological convergence and provide tools for anticipating the early stages of convergence. Overall the study combines the industry evolution perspective and the convergence view of industrial evolution. The theoretical background for the study consists of the industry life cycle theories, technology evolution, and technological trajectories. The study links several important concepts in analyzing industry evolution, technological discontinuities, path-dependency, technological interfaces as a source of industry transformation, and the evolutionary stagesof convergence. Based on reviewing the literature a generic understanding of industry transformation and industrial dynamics was generated. In the convergence studies, the theoretical basis is in the discussion of different convergence types and their impacts on industry evolution, and in anticipating and monitoring the stages of convergence. The study is divided in two parts. The first part gives a general overview, and the second part comprises eight research publications. Our case study is based historically on two very distinct industries of the paper and electronics companies as a test environment to evaluate the importance of emerging business sectors and technological convergence as a source of industry transformation. Both qualitative and quantitative research methodology are utilized. The results of this study reveal that technological convergence and complementary innovations from different fields have significant effect to the emerging new business sector formation. The patent-based indicators in the analysis of technological convergence can be utilized on analyzing technology competition, capability and competence development, knowledge accumulation, knowledge spill-overs, and technology-based industry transformation. The patent-based indicators can provide insights to the future competitive environment. Results and conclusions from empirical part seem not be in conflict with real observations in the industry.

Suomen Ilmavoimien johtamisjärjestelmän evoluutio ilmasotateorian, kansallisten instituutioiden ja johtamisjärjestelmän ulkomaisen kehityksen näkökulmasta

Ilmavoimien johtamisjärjestelmä on osa Ilmavoimien järjestelmäkokonaisuutta, jonka kaksi muuta osaa ovat taistelujärjestelmä ja tukeutumisjärjestelmä. Ilmavoimien materiaalista suorituskykyä rakennetaan tämän järjestelmäajattelun pohjalta. tässä tutkimuksessa Ilmavoimien johtamisjärjestelmää tutkitaan kolmen kokonaisuuden, ilmavalvontajärjestelmän, ilmatilannekuvan muodostamisjärjestelmän ja tulenkäytön johtamisjärjestelmän, näkökulmasta. Ilmavoimien johtamisjärjestelmän laajuuden vuoksi tutkimusaluetta on jouduttu rajaamaan. Tutkimus perustuu evoluutioparadigmaan, jonka mukaisesti kaikki olevainen on evolutionaarista. Mikään tässä ajassa oleva ilmiö ei ole historiaton. Jokaisella ilmiöllä on nykyisyytensä lisäksi historia ja tulevaisuus. Evoluutioparadigman avulla laajennetaan Ilmavoimien johtamisjärjestelmän nykyisyyden ymmärtämistä kuvaamalla ja analysoimalla sen evoluutiota. Tutkimusaineistoa analysoidaan käyttäen hyväksi polkuriippuvuutta evolutionaarisena mallina. tätä mallia on käytetty uusinstitutionaalisessa ja evolutionaarisessa taloustieteessä ja taloushistoriassa tutkittaessa yritysten, toimialojen tai tuotteiden pysyvyyttä markkinoilla sekä erilaisten innovaatioiden vaikuttavuutta menestymiseen eri markkinatilanteissa. Tutkimusasetelman lähtökohtana on Ilmavoimien johtamisjärjestelmäevoluution kuvaaminen kolmen tekijän tasapainoasetelman suhteen, joita ovat instituutiot, ilmasotateoria ja kansainvälinen ilmavoimien johtamisjärjestelmän kehitys. tutkimuksen tavoitteena on löytää institutionaalinen logiikka Ilmavoimien johtamisjärjestelmän evoluutiolle sekä sen eri kehitysprosesseihin liittyvä mahdollinen polkuriippuvuuden logiikka. Tutkittavina instituutioina ovat kansallinen poliittinen päätöksenteko, joka ilmentyy erilaisina komiteamietintöjä, raportteina ja selontekoina. Sotilaallista instituutiota edustavat eri operatiiviset ohjeet, ohjesäännöt ja doktriinit, jotka ovat ohjanneet johtamisjärjestelmäkehitystä. Ilmasotateorian vaikuttavuuden analyysiä varten tutkimuskohteiksi on valittu seitsemän merkittävää ilmasotateoreetikkoa. Kenraalimajuri Giulio douhet, ilmamarsalkka Hugh Trenchard ja kenraalimajuri William Mitchell edustavat ilmasotateorian varhaista kautta. Kansallista ilmasotateorian kehitystä edustavat eversti Richard Lorentz ja kenraalimajuri Gustaf Erik Magnusson. Yhdysvaltalaiset everstit John Boyd ja John Warden III ovat uuden ajan ilmasotateoreetikkoja. Näiden henkilöiden tuottamien teorioiden avulla voidaan piirtää kuva muutoksesta, jota ilmasodankäynnin teoreetti- sessa ajattelussa on tapahtunut. Ilmavoimien johtamisjärjestelmän evoluutiolle haetaan vertailua kehityksestä, jota on tapahtunut Yhdysvalloissa, Isossa-Britanniassa ja Saksassa. Ilmavoimat on saanut vaikutteita muistakin maista, mutta näiden maiden kehityksen avulla voidaan selittää Suomessa tapahtunutta kehitystä. Tutkimuksessa osoitetaan, että kansainvälisellä johtamisjärjestelmäevoluutiolla on ollut merkittävä vaikutus suomalaiseen kehitykseen. Tämä tutkimus laajentaa prosessuaalista tutkimusteoriaa ja polkuriippuvuusmallin käyttöä sotatieteelliseen tutkimuskenttään. tutkimus yhdistää toisiinsa aivan uudella tavalla sotilasorganisaation institutionaalisia tekijöitä pitkässä evoluutioketjussa. Tutkimus luo pohjaa prosessuaaliseen, havaintoihin perustuvaan evoluutioajatteluun, jossa eri tekijöiden selitysmalleja ja kausaalisuutta eri periodien aikana voidaan kuvata. Tutkimuksen tuloksena ilmavoimien johtamisjärjestelmäevoluutiossa paljastui merkittäviä piirteitä. Teknologia on ollut voimakas katalysaattori ilmapuolustuksen evoluutiossa. Uusien teknologisten innovaatioiden ilmestyminen taistelukentälle on muuttanut oleellisesti taistelun kuvaa. Sodankäynnin revoluutiosta huolimatta sodankäynnin tai operaatiotaidon ja taktiikan perusperiaatteissa ei ole tapahtunut perustavanlaatuista muutosta. Ilmavoimien johtamisjärjestelmän kehitys on voimakkaasti linkittynyt ulkomaiseen johtamisjärjestelmäkehitykseen, jossa teknologiaimplementaatiot perustuvat usean eri ilmiön paljastumiseen ja hyväksikäyttöön. Sotilas- ja siviili-instituutiot ovat merkittävästi vaikuttaneet Ilmavoimien johtamisjärjestelmän kansalliseen kehitykseen. Ne ovat antaneet poliittisen ohjauksen, taloudellisten resurssien ja strategis-operatiivisten käskyjen ja suunnitelmien avulla perusteet, joiden pohjalta johtamisjärjestelmää on kehitetty. Tutkimus osoittaa, että Suomen taloudellisten resurssien rajallisuus on ollut merkittävin institutionaalinen rajoite Ilmavoimien johtamisjärjestelmää kehitettäessä. Useat poliittiset ohjausasiakirjat ovat korostaneet, ettei Suomella pienenä kansakuntana ole taloudellisia resursseja seurata kansainvälistä sotilasteknologiakehitystä. Lisäksi ulko- ja turvallisuuspoliittinen liikkumavapaus on vaikuttanut kehittämismahdollisuuksiin. Ilmasotateorian evoluutio on luonut johtamisjärjestelmän kehitykselle välttämättömän konseptuaalisen viitekehyksen, jotta ilmasota on voitu viedä käytännön tasolle. Teoria, doktriini ja instituutiot toimivat vuorovaikutuksessa, jossa ne interaktiivisesti vaikuttavat toinen toisiinsa. Tutkimus paljasti kuusi merkittävää sokkia, jotka saivat aikaan radikaaleja muutoksia johtamisjärjestelmän evoluutiopolulla. tutkimuksen perusteella vaikuttavimmat muutoksia aiheuttavat sokit olivat radikaalit turvallisuuspoliittiset muutokset kuten sota ja voimakkaat kansantalouden muutokset kuten lama. Sokkeja aiheuttaneet kuusi ajankohtaa olivat: 1. Puolustusvoimien rakentamisen aloittaminen vapaussodan jälkeen 1918 2. Maailmanlaajuinen lama 1929–1933 ja eurooppalainen rauhanaate 1928–1933 3. Talvi- ja jatkosota 1939–1944 4. Uusi alku Pariisin rauhansopimuksen 1947 ja YYA-sopimuksen 1948 varjossa 5. Kylmän sodan päättyminen ja Suomen lama 1990–1993 6. Maailmanlaajuinen lama 2008- Tutkimuksen perusteella voidaan todeta, että Suomen ilmavoimien johtamisjärjestelmän kehittäminen on perustunut rationaalisiin päätöksiin, jotka ovat saaneet vaikutteita ulkomaisesta ilmasotateorian ja -doktriinien kehityksestä sekä kansainvälisestä johtamisjärjestelmäkehityksestä. Johtamisjärjestelmän evoluutioon on vaikuttanut globaali konvergenssi, johon on tehty kansallisen tason ratkaisuja järjestelmien adaptaation ja implementaation yhteydessä.

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).