Muotia, mukavuutta ja mielipiteitä. Kaakeliuuni yhteiskunnallisten muutosten ilmentäjänä keskiajan ja uuden ajan alun Suomessa

Fashion, Comf ort and Conviction is a study investigating the adoption and significance of the ceramic tile stove in medieval and early modern Finland. It presents stove tiles found in archaeological excavations, places the tiles in their cultural contexts and examines them as indicators of various social changes. The early history of the ceramic tile stove goes back to the 12th century in the German-speaking Alpine regions, from where it spread to the Baltic Sea region during the Middle Ages. On the basis of the archaeological finds presented in this study, the introduction of the tile stove in Finland dates to the early 15th century. Its adoption was connected to the increase in foreign trade links and social emulation, the demand for comfort, and innovations in building technology and interior design. Stove tiles also reflect the political and religious convictions of their time and open a fascinating window on everyday life in medieval and early modern Finland.

The Integrin Tail: A Tale of Cell Motility and Division

Integrin transmembrane receptor functions are regulated by adaptor molecules binding to their alpha and beta subunit intracellular domains, or tails, thus affecting integrin traffic and adhesion during e.g. cell motility. Interestingly, many cellular proteins function in both cell motility and cell division, thus raising the possibility that integrins might be involved in regulating the cell cycle. A thorough understanding of cell division is essential in cell biology and in human malignancies. It is well established that failures to complete cell cycle can give rise to genetically unstable cells with tumorigenic properties. Transformed cells promote the disruption of intercellular adhesions such as tight junctions, and this correlates with the onset of cell motility, invasion and unfavorable prognosis in cancer. In this study, we analyzed integrin regulation, mediated by adaptor binding to the  subunit tail, during cell motility and cell division. We revealed a novel molecular mechanism by which Rab21, through association with the integrin alpha subunits, drives integrin endosomal traffic during mitotic phases. In addition, we found indications for this finding in vivo, as RAB21 gene deletions were mapped in ovarian and prostate cancer samples. Importantly, the multinucleated phenotype of cultured ovarian cancer cells could be reverted by Rab21 overexpression. In this thesis work, we also show how the tight junction protein ZO-1 unexpectedly interacts with the 5 integrin cytoplasmic domain in the lamellipodia to promote cell motility and at the cleavage furrow to support separation of the daughter cells. The alpha5-ZO-1 complex formation was dependent on PKC which regulates ZO-1 phosphorylation and its subcellular localization. In addition, by an in situ detection method, we showed that a subset of metastatic human lung cancers expressed the alpha5beta-ZO-1 complex. Taken together, we were able to identify new molecular pathways that regulate integrin functions in an alpha tail-mediated fashion. These findings firmly suggest that genetic alterations in integrin traffic may lead to progression of tumorigenesis as a result of failed cell division. Also, the interplay of integrins and ZO-1 in forming spatially regulated adhesive structures broadens our view of crosstalk between pathways and distinct adhesive structures that can be involved in cancer cell biology.

Taktiset toimenpiteet ja niiden käyttö muotikaupan tuoteryhmän kannattavuuden kehittämisessä - case: Ketjuohjattu naisten muodin myymälä

Muotikauppaketjun yksittäisellä myymälällä on tyypillisesti varsin rajoitetusti mahdollisuuksia päättää myymälässä myytävistä tuotteista, niiden hankinnasta sekä hinnoittelusta. Tämän tutkielman tavoitteena oli selvittää, millä toimenpiteillä muotialalla toimivan yrityksen ketjuohjattu myymälä voi vaikuttaa oman yksikkönsä myyntiin ja tavoitellun kannattavuuden saavuttamiseen. Lähempään tarkasteluun rajattiin visuaaliseen markkinointiin ja hinnanalennuksiin liittyvät toimenpiteet. Empiirinen osuus toteutettiin kvalitatiivisena tutkimuksena yhdessä ketjuohjatussa muotikaupan myymälässä. Aineiston hankinnassa käytettiin kolmea eri aineistonkeruumuotoa: havainnointia, teemahaastattelua sekä myyntitapahtuma-aineiston keruuta. Tutkimustulokset osoittivat, että myyntikauden aikana tehtävillä tuotteiden esillepanojen uudistuksilla ja tuotteen paikan vaihtamisella myymälässä on merkitystä myynnin edistämisessä ja hinnanalennusten välttämisessä. Myymälöitä tulisi rohkaista aloittamaan hinnanalentaminen huonosti myyville tuotteille aikaisemmin myyntikauden aikana ja tuotteiden kauden jälkeistä varastointia tulisi välttää. Tutkimustulosten perusteella yksittäisellä ketjuohjatulla myymälällä on varsin paljon mahdollisuuksia vaikuttaa eri toimenpiteillään oman yksikkönsä kannattavuuteen, mutta se vaatii onnistuakseen tiivistä yhteistyötä ketjuohjauksen kanssa.

JUSTIFYING LIBERAL RETRIBUTIVE JUSTICE: PUNISHMENT, CRIMINALIZATION, AND HOLISTIC RETRIBUTIVISM

ABSTRACT In this article I explore whether liberal retributive justice should be conceived of either individualistically or holistically. I critically examine the individualistic account of retributive justice and suggest that the question of retribution – i.e., whether and when punishment of an individual is compatible with just treatment of that individual – must be answered holistically. By resorting to the ideal of sensitive reasons, a model of legitimacy at the basis of our best normative models of democracy, the article argues that in modern liberal democracies, punishment of an offender A for f is compatible with just treatment of A only if punishment of an individual for f can be legitimate in A's and A's fellow citizens' eyes. Only once retributive justice is understood in this holistic fashion the imposition of punishment can be made compatible with just treatment of individuals.

Development of research networks : the case of social capital

The aim of this study is to explore how a new concept appears inscientific discussion and research, how it diffuses to other fields and out of the scientific communities, and how the networks are formed around the concept. Text and terminology take the interest of a reader in the digital environment. Texts create networks where the terminology used is dependent on the ideas, viewsand paradigms of the field. This study is based mainly on bibliographic data. Materials for bibliometric studies have been collected from different databases. The databases are also evaluated and their quality and coverage are discussed. The thesauri of those databases that have been selected for a more in depth study have also been evaluated. The material selected has been used to study how long and in which ways an innovative publication, which can be seen as a milestone in a specific field, influences the research. The concept that has been chosen as a topic for this research is Social Capital, because it has been a popular concept in different scientific fields as well as in everyday speech and the media. It seemed to be a `fashion concept´ that appeared in different situations at the Millennium. The growth and diffusion of social capital publications has been studied. The terms connected with social capital in different fields and different stages of the development have also been analyzed. The methods that have been used in this study are growth and diffusion analysis, content analysis, citation analysis, coword analysis and cocitation analysis. One method that can be used tounderstand and to interpret results of these bibliometric studies is to interview some key persons, who are known to have a gatekeeper position in the diffusion of the concept. Thematic interviews with some Finnish researchers and specialists that have influenced the diffusion of social capital into Finnish scientificand social discussions provide background information. iv The Milestone Publications on social capital have been chosen and studied. They give answers to the question "What is Social Capital?" By comparing citations to Milestone Publications with the growth of all social capital publications in a database, we can drawconclusions about the point at which social capital became generally approved `tacit knowledge´. The contribution of the present study lies foremost in understanding the development of network structures around a new concept that has diffused in scientific communities and also outside them. The network means both networks of researchers, networks of publications and networks of concepts that describe the research field. The emphasis has been on the digital environment and onthe socalled information society that we are now living in, but in this transitional stage, the printed publications are still important and widely used in social sciences and humanities. The network formation is affected by social relations and informal contacts that push new ideas. This study also gives new information about using different research methods, like bibliometric methods supported by interviews and content analyses. It is evident that interpretation of bibliometric maps presupposes qualitative information and understanding of the phenomena under study.

Palveluprosessin suorituskyvyn mittaaminen

Uudelle toimialalle siirtyminen tuo mukanaan yrityksille monenlaisia haasteita. Tämän työn tavoitteena on rakentaa SOL Palvelut Oy:n uusien turvatarkastuspalveluiden johdolle helppokäyttöinen suorituskykymittaristo nykytilanteen kartoittamiseen sekä päätöksenteon tukemiseen. Mittariston rakentamiseen käytetään Lappeenrannan teknillisen yliopiston kehittämää Excel-pohjaista Sake-ohjelmistoa. Lisäksi työn tarkoituksena on tarjota mittaustulosten, teorioiden sekä oman kokemuksen pohjalta muutamia kehitysehdotuksia palvelun tuottamisen parantamiseksi. Oma kokemuspohja syntyy turvatarkastajan peruskurssin suorittamisen sekä itse turvatarkastajan työn tekemisen kautta. Teoriaosuudet käsittelevät palveluiden laatujohtamista, viestintää, motivointia sekä suorituskyvyn mittaamista. Aihealueet ovat määräytyneet mittariston rakentamisen tarpeen, mittaustulosten sekä havaittujen kehityskohteiden kautta. Teorioiden lisäksi työssä tutkitaan turvatarkastuspalveluiden tuottamisen taustalla vaikuttavia tekijöitä kuten lainsäädäntöä, ilmailumääräyksiä sekä kilpailua. Työn empiriaosuus koostuu puolestaan mittariston rakentamisen vaiheista sekä varsinaisten mittaustulosten käsittelemisestä ja hyödyntämisestä. Työlle asetetut tavoitteet täyttyivät hyvin ja yrityksen johdolle kyettiin rakentamaan hyvällä pohjalla oleva suorituskyvyn mittausjärjestelmä. Nykyarvojen perusteella pystyttiin kartoittamaan yrityksen nykytilaa ja sitä kautta voitiin luoda kehitysehdotuksia toimintojen tehostamiseksi. Erityisiksi kehityskohteiksi nousivat talous, koulutus sekä henkilöstö. Näiden tulosten sekä oman työkokemuksen pohjalta työssä on käsitelty kehitysehdotuksia, jotka koskevat mm. viestintää, palkitsemista, palveluohjaajia, koulutuksen kannattavuutta sekä työvuoroja.

Vapautta muodista!

Kirjallisuusarvostelu

Mitochondrial disease in Southwestern Finland. Population-based molecular genetic and clinical studies

Mitochondria are present in all eukaryotic cells. They enable these cells utilize oxygen in the production of adenosine triphosphate in the oxidative phosphorylation system, the mitochondrial respiratory chain. The concept ‘mitochondrial disease’ conventionally refers to disorders of the respiratory chain that lead to oxidative phosphorylation defect. Mitochondrial disease in humans can present at any age, and practically in any organ system. Mitochondrial disease can be inherited in maternal, autosomal dominant, autosomal recessive, or X-chromosomal fashion. One of the most common molecular etiologies of mitochondrial disease in population is the m.3243A>G mutation in the MT-TL1 gene, encoding mitochondrial tRNALeu(UUR). Clinical evaluation of patients with m.3243A>G has revealed various typical clinical features, such as stroke-like episodes, diabetes mellitus and sensorineural hearing loss. The prevalence and clinical characteristics of mitochondrial disease in population are not well known. This thesis consists of a series of studies, in which the prevalence and characteristics of mitochondrial disease in the adult population of Southwestern Finland were assessed. Mitochondrial haplogroup Uk was associated with increased risk of occipital ischemic stroke among young women. Large-scale mitochondrial DNA deletions and mutations of the POLG1 gene were the most common molecular etiologies of progressive external ophthalmoplegia. Around 1% of diabetes mellitus emerging between the ages 18 – 45 years was associated with the m.3243A>G mutation. Moreover, among these young diabetic patients, mitochondrial haplogroup U was associated with maternal family history of diabetes. These studies demonstrate the usefulness of carefully planned molecular epidemiological investigations in the study of mitochondrial disorders.

Kasvatus on kasvamaan saattamista. Kasvatusfilosofinen tutkimus J. A. Hollon sivistyskasvatusajattelusta

Tutkimukseni käsittelee J. A. Hollon (1885–1967) sivistyskasvatusajattelua. Hollo oli monitoiminen kulttuurivaikuttaja, joka toimi kriitikkona, kirjailijana, suomentajana ja kasvatustieteilijänä. Häntä voidaan pitää J. V. Snellmanin rinnalla yhtenä merkittävimpänä suomalaisena kasvatusajattelijana. Hänen kasvatusajattelustaan ei ole kuitenkaan aiemmin tehty väitöskirjatason tutkimusta. Tutkimuskysymykseni ovat seuraavat: 1. Millainen on Hollon näkemys kasvatuksesta, kasvatuksen maailmasta ja kasvatuksen teoriasta? 2. Mikä on Hollon käsitys kasvattajan ja kasvatettavan merkityksestä kasvatustapahtumassa? 3. Mitä asioita sisältyy sivistyskasvatuksen eli kasvamaan saattamisen elementteihin? Tutkimukseni on kasvatusfilosofinen. Tutkimusmenetelmäni on systemaattinen analyysi ja lähestymistapani on hermeneuttinen. Tutkimukseni pääaineistona ovat Hollon kasvatusta koskevat kirjoitukset, joista tärkeimmät ovat Mielikuvitus ja sen kasvattaminen I-II (1918, 1919), Kasvatuksen maailma (1927), Kasvatuksen teoria (1927) ja Itsekasvatus ja elämisen taito (1931). Hollon mukaan kasvatuksen maailma on suhteellisen itsenäinen elämänmuoto (Lebensform), jolla on oma ontologinen erityislaatunsa, so. sui generis. Kasvatusoppia ei pidä redusoida psykologiaan tai filosofiaan, koska sillä tavoin se menettää tieteellisen itsenäisyytensä. Hollon mielestä kasvatuksen teoria on teoria käytäntöä varten. Kasvatuksen teorian luomisessa tulee ottaa huomioon kasvatuksen maailman erityispiirteenä oleva kokonaisvaltainen näkökulma ja elämän palvelemisen päämäärä. Kasvattaminen on aina myös eettistä toimintaa. Kasvatuksen tavoitteena on hyvä elämä. Hollon mukaan kasvattajan tehtävä on luoda kasvatettavalleen eheä sivistyksellinen perusta. Tämä voi tapahtua vain laaja-alaisen sivistyskasvatuksen avulla, jonka runkona on antiikin humanistinen sivistysperinne. Sivistyskasvatukseen kuuluvat älyllinen, eettinen, uskonnollinen, esteettinen ja toiminnallinen kasvatus. Mielikuvituksen avulla kasvattaja voi yhdistää kasvatuksen osa-alueet eheäksi kokonaisuudeksi. Ilman mielikuvitusta erilaiset ilmiöt olisivat pirstaleisina, toisistaan erillisinä osina ihmisen mielessä. Opettajan persoona on merkittävä tekijä kasvatuksessa. Se tulee ottaa huomioon opettajankoulutuksen eli kasvattajan kasvattamisen valinnoissa. Opettaja-kasvattajan on tärkeää opiskella laajasti humanistisia opintoja, koska kasvatuksessa on kysymys ihmisestä. Ennen kaikkea kasvattajan eettistä ja esteettistä kykyä tulee harjoituttaa. Näin hän oppii käyttämään mielikuvitustaan kasvatustapahtumassa siten, että hän tulee kasvatuksellisesti näkeväksi kasvamaan saattajaksi, joka ymmärtää sen, mikä kussakin tilanteessa vaatii erityistä huomiota. Tutkimukseni osoittaa, että Hollon henkitieteellinen ja fenomenologis-hermeneuttinen kasvatusnäkemys ei ole vain vastaparadigma empiiriselle kasvatustieteelle, vaan myös nykyajan teknis-taloudelliselle eetokselle, joka yhtäältä uhkaa välineellistää kasvatuksen ja toisaalta väärällä tavoin tieteellistää kasvatuksen tutkimuksen. Tämän takia kasvatusoppi kysymyksineen uhkaa siirtyä kasvatuskeskustelussa syrjemmälle, jopa hävitä kokonaan. Kasvatuksen ja kasvatuksen tutkimuksen vaarana on niiden liiallinen sitouttaminen tuotantoelämän jatkeeksi, minkä seurauksena on ihmisyyden toteuttamisen vaikeutuminen. Tutkimuksen lopuksi esitän ideaalikoulunäkemykseni, joka perustuu osittain Hollon kasvatusnäkemykseen. Hollon näkemys on yhä ajankohtainen ja merkittävä kontribuutio kasvatusta, sen teoriaa ja käytäntöä koskevaan keskusteluun.

Jeg strikker, derfor er jeg! : læring og identitet i uformelle læringsrum

The dissertation ´I knit, therefore I am!´ Learning and identity in informal space has two main purposes. The first purpose being an investigation of how new value attributions and thinking can generate novel and usable knowledge to the field of craftsmanship, and the second purpose being a display of a different and overlooked philosophical and cultural potential in a reflexive mode of expression, which is able to reflect the normative comprehension of craftsmanship. The dissertation focuses on learning and identity in informal spaces of learning and how it is possible to relate such a learning perspective to crafts training in educational establishments. The empirical foundation of this dissertation is ‘craftivism’. In the dissertation activists from the Nordic countries have been interviewed about what they do when they put up their textile graffiti on lamp posts and house walls. Three research problems are presented: 1) What stories do people who work as crafts activists, tell about ways of relating and methods of action when they make crafts? 2) What do these stories tell about learning and identity? 3) How may the research results influence training and education in craftsmanship? These questions are being asked in order to acquire new knowledge in two aspects; first aspect being knowledge about crafts in relation to techniques, tradition and the objects in crafts, and the second aspect being knowledge about learning and identity in informal spaces of learning. The dissertations theoretical foundation is post structural and sociocultural combined with hermeneutical-inspired qualitative interviews. The author’s position and pre-understanding is subject to discussion in relation to the informant; the performing activist, as the background for both of them is craftsmanship. Starting from cultural studies, it is possible to see the activist subject’s conditions of possibilities in the culture, as the activism of the sub-cultural phenomenon’s craft lights up through a performing approach to the individual’s actions. First the research material has been analysed for events of textile graffiti and possible themes in the events, after which the results have been summarised. Next the research material has been analysed for events about learning and identity due to the author’s wish of comprehending the background of and motivational force in activism. The analysis is divided in main perspectives with different dimensions. The results of the analysis show the activist subject’s construction of an individual who actively takes part in a community by e.g. creating joy, changing the world’s perception of sustainability or by feminizing the public space. By taking crafts over the borders (and away from the class room) crafts become contextualized in a novel fashion thus obtaining an independent status. In this fashion the dissertation writes itself into a new method of comprehending and performing traditional craftsmanship techniques.

Skill Development in Music Reading. The Eye-Movement Approach

This dissertation examined skill development in music reading by focusing on the visual processing of music notation in different music-reading tasks. Each of the three experiments of this dissertation addressed one of the three types of music reading: (i) sight-reading, i.e. reading and performing completely unknown music, (ii) rehearsed reading, during which the performer is already familiar with the music being played, and (iii) silent reading with no performance requirements. The use of the eye-tracking methodology allowed the recording of the readers’ eye movements from the time of music reading with extreme precision. Due to the lack of coherence in the smallish amount of prior studies on eye movements in music reading, the dissertation also had a heavy methodological emphasis. The present dissertation thus aimed to promote two major issues: (1) it investigated the eye-movement indicators of skill and skill development in sight-reading, rehearsed reading and silent reading, and (2) developed and tested suitable methods that can be used by future studies on the topic. Experiment I focused on the eye-movement behaviour of adults during their first steps of learning to read music notation. The longitudinal experiment spanned a nine-month long music-training period, during which 49 participants (university students taking part in a compulsory music course) sight-read and performed a series of simple melodies in three measurement sessions. Participants with no musical background were entitled as “novices”, whereas “amateurs” had had musical training prior to the experiment. The main issue of interest was the changes in the novices’ eye movements and performances across the measurements while the amateurs offered a point of reference for the assessment of the novices’ development. The experiment showed that the novices tended to sight-read in a more stepwise fashion than the amateurs, the latter group manifesting more back-and-forth eye movements. The novices’ skill development was reflected by the faster identification of note symbols involved in larger melodic intervals. Across the measurements, the novices also began to show sensitivity to the melodies’ metrical structure, which the amateurs demonstrated from the very beginning. The stimulus melodies consisted of quarter notes, making the effects of meter and larger melodic intervals distinguishable from effects caused by, say, different rhythmic patterns. Experiment II explored the eye movements of 40 experienced musicians (music education students and music performance students) during temporally controlled rehearsed reading. This cross-sectional experiment focused on the eye-movement effects of one-bar-long melodic alterations placed within a familiar melody. The synchronizing of the performance and eye-movement recordings enabled the investigation of the eye-hand span, i.e., the temporal gap between a performed note and the point of gaze. The eye-hand span was typically found to remain around one second. Music performance students demonstrated increased professing efficiency by their shorter average fixation durations as well as in the two examined eye-hand span measures: these participants used larger eye-hand spans more frequently and inspected more of the musical score during the performance of one metrical beat than students of music education. Although all participants produced performances almost indistinguishable in terms of their auditory characteristics, the altered bars indeed affected the reading of the score: the general effects of expertise in terms of the two eye- hand span measures, demonstrated by the music performance students, disappeared in the face of the melodic alterations. Experiment III was a longitudinal experiment designed to examine the differences between adult novice and amateur musicians’ silent reading of music notation, as well as the changes the 49 participants manifested during a nine-month long music course. From a methodological perspective, an opening to research on eye movements in music reading was the inclusion of a verbal protocol in the research design: after viewing the musical image, the readers were asked to describe what they had seen. A two-way categorization for verbal descriptions was developed in order to assess the quality of extracted musical information. More extensive musical background was related to shorter average fixation duration, more linear scanning of the musical image, and more sophisticated verbal descriptions of the music in question. No apparent effects of skill development were observed for the novice music readers alone, but all participants improved their verbal descriptions towards the last measurement. Apart from the background-related differences between groups of participants, combining verbal and eye-movement data in a cluster analysis identified three styles of silent reading. The finding demonstrated individual differences in how the freely defined silent-reading task was approached. This dissertation is among the first presentations of a series of experiments systematically addressing the visual processing of music notation in various types of music-reading tasks and focusing especially on the eye-movement indicators of developing music-reading skill. Overall, the experiments demonstrate that the music-reading processes are affected not only by “top-down” factors, such as musical background, but also by the “bottom-up” effects of specific features of music notation, such as pitch heights, metrical division, rhythmic patterns and unexpected melodic events. From a methodological perspective, the experiments emphasize the importance of systematic stimulus design, temporal control during performance tasks, and the development of complementary methods, for easing the interpretation of the eye-movement data. To conclude, this dissertation suggests that advances in comprehending the cognitive aspects of music reading, the nature of expertise in this musical task, and the development of educational tools can be attained through the systematic application of the eye-tracking methodology also in this specific domain.

Evaluation of follicular lymphoid depletion in the Bursa of Fabricius: an alternative methodology using digital image analysis and artificial neural networks

Fifty Bursa of Fabricius (BF) were examined by conventional optical microscopy and digital images were acquired and processed using Matlab® 6.5 software. The Artificial Neuronal Network (ANN) was generated using Neuroshell® Classifier software and the optical and digital data were compared. The ANN was able to make a comparable classification of digital and optical scores. The use of ANN was able to classify correctly the majority of the follicles, reaching sensibility and specificity of 89% and 96%, respectively. When the follicles were scored and grouped in a binary fashion the sensibility increased to 90% and obtained the maximum value for the specificity of 92%. These results demonstrate that the use of digital image analysis and ANN is a useful tool for the pathological classification of the BF lymphoid depletion. In addition it provides objective results that allow measuring the dimension of the error in the diagnosis and classification therefore making comparison between databases feasible.

Product placement on personal blogs: a multiple case study from companies’ perspective

As traditional advertising is losing its value in corporations’ marketing, new and growing social media marketing channels are increasing their importance. The growing interest in the media and the lack of research on the subject create a clear need for this study. The objectives are to increase understanding about product placement in personal blogs and examine the benefits as well as the downsides created by this marketing channel and present the requirements for success. The study also takes a stand on what kind of things a successful product placement strategy in personal blogs includes and what points should be considered when creating a strategy. The study’s empirical part consists of seven thematic interviews with case companies’ representatives and one agent. The study’s personal blogs were delimited to consist only lifestyle and fashion blogs. The results show that product placement on personal lifestyle and fashion blogs is a strongly growing marketing channel and it is best suited to reach young women. It is also a very good channel to change and improve brand image. Via personal blogs it is possible to reach a large number of consumers with a very cost-efficient manner. Thus utilizing the channel is suitable for all companies regardless of the size of the marketing budget. Of course, companies have to consider the suitability of their product to the themes of blogs. Executing a successful campaign in this media requires good relationship management skills and understanding of this particular media as the final content is not in advertiser’s hands. As a marketing channel, personal blogs differ highly from traditional marketing channels because of their constantly changing nature and multidirectional quality where several parties are in interaction.

Stop bugging me! : diversity in appearance of warning coloration

In nature, many animals use body coloration to communicate with each other. For example, colorations can be used as signals between individuals of the same species, but also to recognise individuals of other species, and if they may comprise a threat or not. Many animals use protective coloration to avoid predation. The two most common strategies of protective coloration are camouflage and aposematism. Camouflaged animals have coloration that minimises detection, usually by matching colours or structures in the background. Aposematic animals, on the other hand, signal to predators that they are defended. The defence can be physical structures, such as spikes and hairs, or chemical compounds that make the animal distasteful or even deadly toxic. In order for the warning signal to be effective, the predator has to recognise it as such. Studies have shown that birds for example, that are important visual predators on insects, learn to recognise and avoid unpalatable prey faster if they contrast the background or have large internal contrasts. Typical examples of aposematic species have conspicuous colours like yellow, orange or red, often in combination with black. My thesis focuses on the appearance and function of aposematic colour patterns. Even though researchers have studied aposematism for over a century, there is still a lot we do not know about the phenomenon. For example, as it is crucial that the predators recognise a warning signal, aposematic colorations should assumingly evolve homogeneously and be selected for maximal conspicuousness. Instead, there is an extensive variation of colours and patterns among warning colorations, and it is not uncommon to find typical cryptic colours, such as green and brown in aposematic colour patterns. One hypothesis to this variation is that an aposematic coloration does not have to be maximally signalling in order to be effective, instead it is sufficient to have distinct features that can be easily distinguished from edible prey. To be maximally conspicuous is one way to achieve this, but not the only way. Another hypothesis is that aposematic prey that do not exhibit maximal conspicuousness can exploit both camouflage and aposematism in a distance-dependent fashion, by being signalling when seen close up but camouflaged at a distance. Many prey animals also make use of both strategies by shifting colour at different ecological conditions such as seasonal variations, fluctuations in food resources or between life stages. Yet another explanation for the variation may be that prey animals are usually exposed to several predator species that vary in visual perception and tolerance towards various toxins. The aim with this thesis is, by studying their functions, to understand why aposematic warning signals vary in appearance, specifically in the level of conspicuousness, and if warning coloration can be combined with camouflage. In paper I, I investigated if the colour pattern of the aposematic larva of the Apollo butterfly (Parnassius apollo) can switch function with viewing distance, and be signalling at close range but camouflaged at a distance, by comparing detection time between different colour variants and distances. The results show that the natural coloration has a dual distance-dependent function. Moreover, the study shows that an aposematic coloration does not have to be selected for maximal conspicuousness. A prey animal can optimise its coloration primarily by avoiding detection, but also by investing in a secondary defence, which presence can be signalled if detected. In paper II, I studied how easily detected the coloration of the firebug (Pyrrhocoris apterus), a typical aposematic species, is at different distances against different natural backgrounds, by comparing detection time between different colour variants. Here, I found no distance-dependent switch in function. Instead, the results show that the coloration of the firebug is selected for maximal conspicuousness. One explanation for this is that the firebug is more mobile than the butterfly larva in study I, and movement is often incompatible with efficient camouflage. In paper III, I investigated if a seasonal related colour change in the chemically defended striated shieldbug (Graphosoma lineatum) is an adaptation to optimise a protective coloration by shifting from camouflage to aposematism between two seasons. The results confirm the hypothesis that the coloration expressed in the late summer has a camouflage function, blending in with the background. Further, I investigated if the internal pattern as such increased the effectiveness of the camouflage. Again, the results are in accordance with the hypothesis, as the patterned coloration was more difficult to detect than colorations lacking an internal pattern. This study shows how an aposematic species can optimise its defence by shifting from camouflage to aposematism, but in a different fashion than studied in paper I. The aim with study IV was to study the selection on aposematic signals by identifying characteristics that are common for colorations of aposematic species, and that distinguish them from colorations of other species. I compared contrast, pattern element size and colour proportion between a group of defended species and a group of undefended species. In contrast to my prediction, the results show no significant differences between the two groups in any of the analyses. One explanation for the non-significant results could be that there are no universal characteristics common for aposematic species. Instead, the selection pressures acting on defended species vary, and therefore affect their appearance differently. Another explanation is that all defended species may not have been selected for a conspicuous aposematic warning coloration. Taken together, my thesis shows that having a conspicuous warning coloration is not the only way to be aposematic. Also, aposematism and camouflage is not two mutually exclusive opposites, as there are prey species that exploit both strategies. It is also important to understand that prey animals are exposed to various selection pressures and trade-offs that affect their appearance, and determines what an optimal coloration is for each species or environment. In conclusion, I hold that the variation among warning colorations is larger and coloration properties that have been considered as archetypically aposematic may not be as widespread and representative as previously assumed.