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.

Avian Evolutionary Genomics: Studies of Ficedula Flycatchers

In this thesis, different genetic tools are used to investigate both natural variation and speciation in the Ficedula flycatcher system: pied (Ficedula hypoleuca) and collared (F. albicollis) flycatchers. The molecular evolution of a gene involved in postnatal body growth, GH, has shown high degree of conservation at the mature protein between birds and mammals, whereas the variation observed in its signal peptide seems to be adaptive in pied flycatcher (I & II). Speciation is the process by which reproductive barriers to gene flow evolve between populations, and understanding the mechanisms involved in pre- and post-zygotic isolation have been investigated in Ficedula flycatchers. The Z chromosome have been suggested to be the hotspot for genes involved in speciation, thus sequencing of 13 Z-linked coding genes from the two species in allopatry and sympatry have been conducted (III). Surprisingly, the majority of Z-linked genes seemed to be highly conserved, suggesting instead a potential involvement of regulatory regions. Previous studies have shown that genes involved in hybrid fitness, female preferences and male plumage colouration are sex-linked. Hence, three pigmentation genes have been investigated: MC1R, AGRP, and TYRP1. Of these three genes, TYRP1 was identified as a strong candidate to be associated with black-brown plumage variation in sympatric populations, and hence is a strong candidate for a gene contributing to pre-zygotic isolation (IV). In sympatric areas, where pied and collared flycatchers have overlapping breeding areas, hybridization sometimes occurs leading to the production of unfit hybrids. By using a proteomic approach a novel expression pattern in hybrids was revealed compared to the parental species (V) and differentially expressed proteins subsequently identified by sequence similarity (VI). In conclusion, the Z chromosome appears to play an important role in flycatcher speciation, but probably not at the coding level. In addition the novel expression patterns might give new insights into the maladaptive hybrids.

Yksityisrahoitteinen aluerakentaminen

Diplomityön tavoitteena oli tutkia yksityisrahoitusmallin soveltuvuutta julkisen sektorin palvelutuotantoon sekä kuvata yksityisrahoitteisen aluerakentamishankkeen sisältö ja liiketoimintaprosessin eteneminen hankkeen ideasta konsessiosopimuksen päättymiseen asti.Huoltosuhteen muuttumisen lisäksi muuttoliike kasvukeskuksiin lisää julkisen sektorin paineita tuottaa skandinaavisen hyvinvointiyhteiskuntamme vaatimusten mukaisia palveluita. Julkinen sektori onkin ajautunut tilanteeseen, jossa vaihtoehtoina ovat sen tuottamien palveluiden supistaminen ja maksullistaminen tai tuotantomuotojen kehittäminen.Työssä esitetty yksityisrahoitteinen aluerakentamismalli mahdollistaa laadukkaiden julkisten palveluiden tuottamisen oikea-aikaisesti ja kustannustehokkaasti. Tässä uudessa aluerakentamisen mallissa yksityinen sektori rahoittaisi, rakentaisi sekä operoisi ja ylläpitäisi määräajan perinteisesti kunnalle kuuluneita infrastruktuuri- ja palveluhankkeita.Tutkimuksen keskeinen tulos on yksityisrahoitteisen aluerakentamisen liiketoimintaprosessikaavio. Liiketoimintaprosessin kuvaus edesauttaa monimutkaisen prosessin ymmärtämistä ja mahdollistaa yksityisrahoitteisen aluerakentamismallin edelleen kehittämisen sekä sen markkinoinnin julkiselle ja yksityiselle sektorille. Liiketoimintaprosessikaaviossa kuvataan prosessin osapuolet ja prosessin eteneminen julkisen ja yksityisen sektorin tarpeesta ja YIT:n liikeideasta aina alueen rakentamiseen, operointiin, ylläpitoon ja palvelutuotannon siirtoon julkiselle ja yksityiselle sektorille

Ett rätt barna-klenodium

Motto [I]: Psal. 19: v. 11. Herrans bod äro klara, ... Motto [II]: Rom. I: v. 16. Evangelium är Guds kraft ... Motto [III]: Matth. 18: v. 3. Vthan j omwänden eder och warden såsom barn, .. Nimiön kehyslauselma: Psal. 34. v. 12. Kommer hijt, barn, hörer mig: .. Priv.: Henrik Christofer Merckell. Arkit: A-D12 E6.

Bland åsnor och elefanter : en utvärdering av USA:s presidenter

Arbetet går ut på en systematisering rörande bedömningen av USA:s presidenter. Detta sker genom att utarbeta fyra värderingskriterier. Dessa kriterier benämns \"problem\" (= vilken typ av utmaningar presidenten har ställts inför samt på vilket sätt presidenten har löst dessa utmaningar), \"ämbete\" (= på vilket sätt presidentämbetets tyngd har påverkats av ämbetsinnehavaren), \"popularitet\" (= på vilket sätt presidenten har fått gehör för sina åtgärder hos allmänhet och de politiska eliterna) samt \"USA\" (= på vilket sätt presidenten har påverkat USA:s styrka som nation rent generellt). Tidigare utvärderingar av USA:s presidenter har tenderat att vara något jippo-betonade, även i de fall där akademiker ligger bakom. Vanligen har man haft en expertpanel bestående av allt från några dussin bedömare till närmare 1.000. Antingen har man inte haft några värderingskriterier alls att gå efter eller så har de varit något oklart formulerade - för att inte tala om de fall där de har varit så pass många att de har överlappat varandra. Vidare finns en tradition där en del forskare förkastar hela tanken på att kunna bedöma presidenter från olika tidsperioder. Strävan har varit att råda bot på tidigare missförhållanden i den diskussion som har förts på området. Eftersom samtliga presidenter dessutom har verkat under samma konstitution bör en jämförelse/utvärdering vara fullt möjlig att genomföra. De som klarar sig bäst i min utvärdering är Washington, Lincoln och F. Roosevelt. Vidare klarar sig t.ex. McKinley och Reagan bättre än de vanligen klarar sig i motsvarande bedömningar. Till de som klarar sig sämre hör t.ex. L. Johnson och J. Madison. Allra sämst klarar sig de presidenter som verkade strax före Lincoln (Pierce och Buchanan) alternativt strax efter (A. Johnson). Den sittande presidenten (Obama) samt de två mest kortvariga presidenterna (Garfield och W.H. Harrison) har lämnats bort från arbetet. Själva tyngdpunkten i arbetet ligger dock i själva systematiseringen av värderingskriterierna även om arbetet rent volymmässigt domineras av då kriterierna appliceras på de enskilda presidenterna.