Anti-predator adaptations in aquatic environments

Predation is an important selective force that has led to the evolution of a variety of fascinating anti-predator adaptations, such as many types of protective coloration and prey behaviours. Because the evolution of life has begun in the aquatic environment and many anti-predator adaptations are found already in relative primitive taxa, it is likely that many of these adaptations evolved initially in the aquatic environment. Yet, there has been surprisingly little research on the mechanisms and function of antipredator adaptations in aquatic systems. To understand the function of anti-predator adaptations and natural selection imposed on prey appearance and behaviour, I have investigated how protective coloration can be used, either as such or together with behavioural adaptations, to manipulate predator behaviour and decrease predation risk. To this end I conducted a series of behaviour ecological laboratory experiments in which I manipulated the visual appearance of artificial backgrounds and prey items. In paper I of this thesis, I investigated background choice as an anti-predator strategy, by observing the habitat choice of the least killifish (Heterandria formosa) between pairs of artificial backgrounds, both in the presence and absence of predation threat. It has been suggested that prey could decrease their risk of being detected by predators either by preferring backgrounds into which they blend or by preferring visually complex backgrounds. The least killifish preferred a background that matched their patterning to a background that mismatched it, showing that they are able to respond to cues of visual similarity between their colour pattern and the surrounding environment. Interestingly however, in female least killifish visual complexity of the background was a more important cue for habitat safety and may override or act together with background matching when searching for a safe habitat. It is possible that in females, preference for visually complex backgrounds is associated with lower opportunity costs than preference for matching backgrounds would be. Generally, the least killifish showed stronger preference while under predation threat, indicating that their background choice behaviour is an antipredator adaptation. Many aquatic prey species have eyespots, which are colour patterns that consist of roughly concentric rings and have received their name because they for humans often resemble the vertebrate eye. I investigated the anti-predator function of eyespots against predation by fish in papers II, III and IV. Some eyespots have been suggested to benefit prey by diverting the strikes of predators away from vital parts of the prey body or towards a direction that facilitates prey escape. Although proposed over a century ago, the divertive effect of eyespots has proven to be difficult to show experimentally. In papers II and III, I tested for divertive effect of eyespots towards attacking fish by presenting artificial prey with eyespots to laboratory reared three-spined sticklebacks (Gasterosteus aculeatus). I found that eyespots strongly influenced the behaviour of attacking sticklebacks and effectively drew their strikes towards the eyespots. To further investigate this divertive effect and whether the specific shape of eyespots is important for it, I tested in paper III the response of fish also to other markings than eyespots. I found that eyespots were generally more effective in diverting the first strikes of attacking fish compared to other prey markings. My findings suggest that the common occurrence of eyespots in aquatic prey species can at least partly be explained by the divertive effect of the eyespot shape, possibly together with the relative simple developmental mechanisms underlying circular colour patterns. An eyebar is a stripe that runs through the eye, and this pattern has been suggested to obscure the real eyes of the prey by visually blending parts of the eyes and head of the prey and by creating false edges. In paper III, I show that an eyebar effectively disrupts an eyelike shape. This suggests that eyebars provide an effective way to conceal the eyes and consequently obstruct detection and recognition of prey. This experiment also demonstrates that through concealment of the eyes, eyebars could be used to enhance the divertive effect of eyespots, which can explain the common occurrence of eyebars in many species of fish that have eyespots. Larger eyespots have been shown to intimidate some terrestrial predators, such as passerine birds, either because they resemble the eyes of the predator’s own enemy or because highly salient features may have an intimidating effect. In papers II and IV, I investigated whether the occurrence of eyespots in some aquatic prey could be explained by their intimidating effect predatory fish. In paper IV, I also investigated the reason for the intimidating effect of eyelike prey marks. In paper II, I found no clear intimidating effect of eyespots, whereas in paper IV, using a different approach, I found that sticklebacks hesitated to attack towards eyelike but not towards non-eyelike marks. Importantly, paper IV therefore presents the first rigorous evidence for the idea that eye mimicry, and not merely conspicuousness, underlies the intimidating effect. It also showed that the hesitation shown by fish towards eyelike marks is partly an innate response that is reinforced by encounters with predators. Collectively, this thesis shows that prey colour pattern and the visual appearance of the habitat influence the behaviour of fish. The results demonstrate that protective coloration provides numerous distinctive ways for aquatic prey to escape predation. Thus, visual perception and behaviour of fish are important factors shaping the appearance and behaviours of aquatic prey.

Trophic cascades in boreal landscapes: top predator protection on tree seedlings and forest grouse

Changes in the abundance of top predators have brought about notable, cascading effects in ecosystems around the world. In this thesis, I examined several potential trophic cascades in boreal ecosystems, and their separate interspecific interactions. The main aim of the thesis was to investigate whether predators in the boreal forests have direct or indirect cascading effects on the lower trophic levels. First, I compared the browsing effects of different mammalian herbivores by excluding varying combinations of voles, hares and cervids from accessing the seedlings of silver birch (Betula pendula), Scots pine (Pinus sylvestris) and Norway spruce (Picea abies). Additionally, I studied the effect of simulated predation risk on vole browsing by using auditory cues of owls. Moving upwards on the trophic levels, I examined the intraguild interactions between the golden eagle (Aquila chrysaetos), and its mesopredator prey, the red fox (Vulpes vulpes) and the pine marten (Martes martes). To look at an entire potential trophic cascade, I further studied the combined impacts of eagles and mesopredators on the black grouse (Tetrao tetrix) and the hazel grouse (Tetrastes bonasia), predicting that the shared forest grouse prey would benefit from eagle presence. From the tree species studied, birch appears to be the most palatable one for the mammalian herbivores. I observed growth reductions in the presences of cervids and low survival associated with hares and voles, which suggests that they all weaken regeneration in birch stands. Furthermore, the simulated owl predation risk appeared to reduce vole browsing on birches in late summer, although the preferred grass forage is then old and less palatable. Browsing by voles and hares had a negative effect on the condition and survival of Scots pine, but in contrast, the impact of mammalian herbivores on spruce was found to be small, at least when more preferred food is available. I observed that the presence of golden eagles had a negative effect on the abundance of adult black grouse but a positive, protective effect on the proportion of juveniles in both black grouse and hazel grouse. Yet, this positive effect was not dependent on the abundance foxes or martens, nor did eagles seem to effectively decrease the abundance of these mesopredators. Conversely, the protection effect on grouse could arise from fear effects and also be mediated by other mesopredators. The results of this thesis provide important new information about trophic interactions in the boreal food webs. They highlight how different groups of mammalian herbivores vary in their effects on the growth and condition of different tree seedlings. Lowered cervid abundances could improve birch regeneration, which indirectly supports the idea that the key predators of cervids could cause cascading effects also in Fennoscandian forests. Owls seem to reduce vole browsing through an intimidation effect, which is a novel result of the cascading effects of owl vocalisation and could even have applications for protecting birch seedlings. In the third cascade examined in this thesis, I found the golden eagle to have a protective effect on the reproducing forest grouse, but it remains unclear through which smaller predators this effect is mediated. Overall, the results of this thesis further support the idea that there are cascading effects in the forests of Northern Europe, and that they are triggered by both direct and non‐lethal effects of predation.

The effect of brand trust on satisfaction, trust, and loyalty on a consumer magazine website

Internetin käyttö on yleistynyt valtavasti viimeisen kymmenen vuoden aikana. Internetissä kuluttajalta puuttuu kasvokkainen kontakti myyjään, mikä voi vähentää kuluttajan luottamusta myyjää kohtaan. Tällaisessa tilanteessa kuluttaja voi käyttää brändejä viitteenä myyjän luotettavuudesta. Tämän tutkielman tarkoituksena on tutkia brändiluottamuksen vaikutusta kuluttajan tyytyväisyyteen, luottamukseen ja uskollisuuteen Internet-sivuja kohtaan. Tutkielman kysely (n=875) toteutettiin Vauva-lehden Internet-sivuilla. Tutkielman tulokset paljastavat, että brändiluottamuksella on merkittävä myönteinen vaikutus kuluttajan tyytyväisyyteen, luottamukseen ja uskollisuuteen Internet-sivuja kohtaan. Internet-sivujen aikaisempikäyttö sekä rekisteröityminen sivuille vaikuttavat myös luottamukseen Internet-sivuja kohtaan. Lisäksi brändiluottamuksen vaikutus brändiuskollisuuden osoittautui erittäin merkittäväksi.

The role of trust and its evolution in distributed teams in a multinational organisation

Tutkimuksen päätavoitteena on ollut selvittää miten kansallinen ja organisaatiokulttuuri, niihin liittyvät normit ja arvot edesauttavat tai vaikeuttavat luottamuksen kehittymistä monikulttuurisissa tiimeissä maailmanlaajuisessa organisaatiossa. Tutkimuksen avulla haluttiin myös selvittää miten luottamus kehittyy hajautetuissa monikansallisissa tiimeissä WorldCom Internationalissa. Empiirinen tutkimusmenetelmä perustuu kvalitatiivisiin teemahaastatteluihin, jotka tehtiin WorldComin työntekijöille. Tutkimuksessa havaittiin, ettei yhteisten sosiaalisten normien merkitys luottamuksen syntymiselle ole niin merkittävä, koska WorldComin yhtenäiset toimintatavat sekä hallitseva amerikkalaisen emoyhtiön "kotikulttuuri" muodostavat yhtenäiset toimintalinjat tiimeissä. Tietokonevälitteisen kommunikoinnin jatkuva käyttö on edesauttanut työntekijöiden ns. sosiaalisen älyn kehittymistä, sillä henkilökohtaisen tapaamisen puuttuminen kehittää vastaavasti taitoja aistia ja tulkita sähköpostien tai puhelinneuvotteluiden aikana välittyviä vastapuolen "näkymättömiä" vihjeitä.

Thylakoid Protein Phosphorylation in Regulation of Photosynthesis

Once the seed has germinated, the plant is forced to face all the environmental changes in its habitat. In order to survive, plants have evolved a number of different acclimation systems. The primary reaction behind plant growth and development is photosynthesis. Photosynthesis captures solar energy and converts it into chemical form. Photosynthesis in turn functions under the control of environmental cues, but is also affected by the growth, development, and metabolic state of a plant. The availability of solar energy fluctuates continuously, requiring non-stop adjustment of photosynthetic efficiency in order to maintain the balance between photosynthesis and the requirements and restrictions of plant metabolism. Tight regulation is required, not only to provide sufficient energy supply but also to prevent the damage caused by excess energy. The very first reaction of photosynthesis is splitting of water into the form of oxygen, hydrogen, and electrons. This most fundamental reaction of life is run by photosystem II (PSII), and the energy required for the reaction is collected by the light harvesting complex II (LHCII). Several proteins of the PSII-LHCII complex are reversibly phosphorylated according to the energy balance between photosynthesis and metabolism. Thylakoid protein phosphorylation has been under extensive investigation for over 30 years, yet the physiological role of phosphorylation remains elusive. Recently, the kinases behind the phosphorylation of PSII-LHCII proteins (STN7 and STN8) were identified and the knockout mutants of these kinases became available, providing powerful tools to elucidate the physiological role of PSII-LHCII phosphorylation. In my work I have used the stn7 and stn8 mutants in order to clarify the role of PSII-LHCII phosphorylation in regulation and protection of the photosynthetic machinery according to environmental cues. I show that STN7- dependent PSII-LHCII protein phosphorylation is required to balance the excitation energy distribution between PSII and PSI especially under low light intensities when the excitation energy transfer from LHC to PSII and PSI is efficient. This mechanism differs from traditional light quality-induced “state 1” – “state 2” transition and ensures fluent electron transfer from PSII to PSI under low light, yet having highest physiological relevance under fluctuating light intensity. STN8-dependent phosphorylation of PSII proteins, in turn, is required for fluent turn-over of photodamaged PSII complexes and has the highest importance upon prolonged exposure of the photosynthetic apparatus to excess light.

The Importance of Semantics: Visual World Studies on Drawing Inferences and Resolving Anaphors

The present thesis investigated the importance of semantics in generating inferences during discourse processing. Three aspects of semantics, gender stereotypes, implicit causality information and proto-role properties, were used to investigate whether semantics is activated elaboratively during discourse comprehension and what its relative importance is in backward inferencing compared to discourse/structural cues. Visual world eye-tracking studies revealed that semantics plays an important role in both backward and forward inferencing: Gender stereotypes and implicit causality information is activated elaboratively during online discourse comprehension. Moreover, gender stereotypes, implicit causality and proto-role properties of verbs are all used in backward inferencing. Importantly, the studies demonstrated that semantic cues are weighed against discourse/structural cues. When the structural cues consist of a combination of cues that have been independently shown to be important in backward inferencing, semantic effects may be masked, whereas when the structural cues consist of a combination of fewer prominent cues, semantics can have an earlier effect than structural factors in pronoun resolution. In addition, the type of inference matters, too: During anaphoric inferencing semantics has a prominent role, while discourse/structural salience attains more prominence during non-anaphoric inferencing. Finally, semantics exhibits a strong role in inviting new inferences to revise earlier made inferences even in the case the additional inference is not needed to establish coherence in discourse. The findings are generally in line with the Mental Model approaches. Two extended model versions are presented that incorporate the current findings into the earlier literature. These models allow both forward and backward inferencing to occur at any given moment during the course of processing; they also allow semantic and discourse/structural cues to contribute to both of these processes. However, while the Mental Model 1 does not assume interactions between semantic and discourse/structural factors in forward inferencing, the Mental Model 2 does assume such a link.

Fiktiivisen kirjallisuuden maailmasta monikulttuuriseen Suomeen. Ennakointikertomus kirjallisuudenopetuksen ja monikulttuurisuuskasvatuksen välineenä

From the world of fiction literature into multi-cultural Finland. Anticipatory story as an educational tool in teaching of literature and multiculturalism The research clarifies the relationship between reading fiction literature and multicultural value education in basic education. The research focuses on the subject didactics of mother tongue and literature and on the literature teaching in particular. The objective is to develop a method that is easily transferable into a teaching context so as to intensify the educational discussion based on fiction literature. In essence, understanding fiction literature and the ethical and moral thinking resemble one another, because both of them aim at empathizing with the thinking of a person or with a situation foreign to oneself. For this reason fiction literature is ideally suited for the discussion on ethical and moral values within a subject entity in the basic education. The empirical unit of the research consists of two parts. The first part explains how youth novels published in the years 1993 – 2007 describe multiculturalism in Finnish society. Books on multiculturalism are still few in number within youth literature, and people with a foreign background are mainly minor or background characters in such literary works. Nevertheless, youth novels serve well as a starting point for an educational discussion about multicultural issues. More often than not characters in youth novels are stereotypes and even opposite to each other. The juxtaposing makes a young reader question the stereotypes associated with immigrants. Besides the stereotype, reference to a prototype or a model is possible. The second part tests the usefulness of an anticipatory story based on a fictive text for an educational discussion about multicultural issues. The empirical material was collected from the eighth-grade teaching groups in basic education as follows: one teaching group was an immigrant group, the second one a group of Finns with experience of immigrants while the third group of students had hardly any experience of immigrants. The anticipatory stories were written on the basis of extracts from youth novels with multicultural themes. The material collected for a total of 120 anticipatory stories was analysed by using meaning cue analysis. Using the meaning cue analysis, the anticipatory stories were divided into three groups: stories with predominantly positive meaning cues of interculturalism, ambivalent stories with both positive and negative meaning cues of interculturalism and the stories with predominantly negative meaning cues of interculturalism. The meaning cues produced by girls and boys differ from one another, in particular, by the negative meaning cues of interculturalism. For girls, the predominant meaning cue is fear whereas for boys, it is that of violence. It would also seem that the students, in particular, boys with little experience of immigrants produce more negative meaning cues of interculturalism than do immigrants or Finnish students with experience of immigrants. Further still, it seems that active reading of fiction literature affects the meaning cues of interculturalism in an ambivalent direction. In the way of youth novels this is understandable, because youth novels in general are made up of opposite characters and meaning cues. The less the student takes an interest in reading, the more he used meaning cues from outside the parent text for his anticipatory story. No doubt it would be possible to use fiction literature in the literature education to a much higher extent than it is being used today whereby the literature could be used in basic education for reviewing subject entities or study contents of other study subjects. By way of an anticipatory story and the meaning cue analysis, it is possible to intensify the educational discussions based on fiction literature. However, using fiction literature in the literature education requires consideration of the specific genre of fiction literature.

Miehittämättömien ilma-alusten käyttömahdollisuudet ilmasta maahan -aseiden tarvitseman maalitiedon hankinnassa ilmatorjunnan suojaamassa ympäristössä

Vuoden 2004 iskukykytutkimus esittää lentotiedustelujärjestelmän hankkimista tukemaan HN-kalustoon perustuvaa ilmasta maahan -järjestelmää. Tässä tutkimustyössä on tutkittu kolmen erilaisen miehittämättömän ilma-aluksen käyttömahdollisuuksia ilmasta maahan -aseiden tarvitseman maalitiedon hankkimisessa. Tutkimuksessa mukana olleet ilmasta maahan -aseet olivat JDAM (Joint Direct Attack Munition), AGM-154 JSOW (Joint Standoff Weapon) ja toistaiseksi epävarman hankintansa vuoksi vähäisemmässä määrin AGM-158 JASSM (Joint Air to Surface Standoff Weapon). Miehittämättömiksi ilmaaluksiksi valittiin Predator-koneen kolme eri kehitysversiota, MQ-1 Predator A, MQ-9 Reaper (Predator B) ja Avenger (Predator C). Tutkimuksen pääasiallinen tutkimusmenetelmä on ollut laadullinen kirjallisuusselvitys julkisia lähteitä käyttäen. Suoritusarvoista johdettuihin päätelmiin on käytetty soveltuvin osin määrällisiä tutkimusmenetelmiä. Lähestymistapa tutkimukseen on ollut hermeneuttinen. Alakysymysten osalta kyseessä on ollut deskriptiivinen tutkimus ja tavoitteena on ollut löytää avainasiat miehittämättömien ilma-alusten käytön kannalta. Tutkimusraportin ensimmäinen luku on johdanto. Toisessa luvussa on selvitetty toimintaympäristön yleiset piirteet ja kolmannessa luvussa ilmasta maahan -aseiden ominaisuudet. Neljäs luku käsittelee tutkimuksessa olleiden miehittämättömien ilma-alusten ominaisuuksia maalitiedon hankkimismahdollisuuksien kannalta. Tutkimustyön johtopäätökset on kirjattu viidenteen lukuun. Tulosten perusteella perinteisten, ilman häiveominaisuuksia rakennettujen miehittämättömien ilma-alusten ei ole mahdollista hankkia kaikkea ilmasta maahan -aseiden tarvitsemaa maalitietoa. Lähialueen ilmatorjunnan vaikutuksen väistämiseksi häiveominaisuudet ja jonkin asteinen tunkeutumiskyky ovat välttämättömät erityisesti pitkän kantaman ilmasta maahan -aseiden tarvitseman maalitiedon hankkimisessa.

Miehittämättömän ilma-aluksen hyödyt ja haitat ilmatiedustelussa operaatio Enduring Freedomissa

Tämän tutkielman tarkoituksena on kirjallisuuskatsauksen keinoin luoda käsitys miehittämättömän ilma-aluksen ilmatiedustelukäytön hyödyistä ja haitoista operaatio Enduring Freedomissa. Tutkielmassa pohditaan lisäksi UAV:iden tulevaisuuden näkymiä ja kehityssuuntia ilmatiedustelun kannalta. Tämä tutkielma sisältää kvalitatiivisen tutkimuksen piirteitä ja tutkimusmetodina käytetään narratiivista kirjallisuuskatsausta. Tutkielman pääluvuissa tutustutaan UAV-konseptiin sekä tutkitaan UAV:n hyötyjä ja haittoja ilmatiedustelussa operaatio Enduring Freedomissa. Tutkielmassa on pyritty tutustumaan aihealueeseen kokonaisvaltaisesti ja sen pohjalta luomaan teoreettinen kokonaisuus tutkimuskysymyksiin vastaten. Operaatio Enduring Freedom alkoi 7.10.2001, kun Yhdysvaltain johtama liittouma hyökkäsi Afganistaniin vastauksena 11.9.2001 tehtyihin terrori-iskuihin. Operaation tavoitteena oli tuhota terroristien koulutusleirit ja infrastruktuuri Afganistanin alueella, pidättää al-Qaidan johto sekä lopettaa terrorismitoiminta Afganistanissa. Yhdysvaltain ilmavoimat on käyttänyt operaation alusta alkaen ilmatiedustelussa pääasiassa kahta UAVtyyppiä, jotka ovat RQ/MQ-1 Predator ja RQ-4 Global Hawk. Tutkielmassa perehdytään näiden kahden UAV:n hyötyihin ja haittoihin. Vertailupohjana käytetään miehitettyjä ilma-aluksia. Tutkielmassa käytettyjen lähteiden perusteella voidaan todeta, että miehittämättömillä ilma-aluksilla on etuja, joita ei voi saavuttaa miehitettyjä ilma-aluksia käyttäen. Hyödyt on jaettu tässä tutkielmassa rakenteellisiin, kustannuksellisiin ja tehtäväkohtaisiin hyötyihin. UAVkäytön merkittävin haitta lienee maa-aseman ja ilma-aluksen välisen datalinkin aiheuttamat haasteet ja rajoitteet. Lisäksi miehittämättömän ilma-aluksen haavoittuvuus ilmatorjuntatulten ja ilmasta ilmaan -ohjusten vaikutuspiirissä on vielä tällä hetkellä haaste. Tutkielmassa käsitellyt hyödyt ja haitat on todettu operaatio Enduring Freedomin kaltaisessa epäsymmetrisessä toimintaympäristössä, joten niiden soveltuvuutta symmetriseen toimintaympäristöön ei ole juurikaan arvioitu. Miehittämättömien ilma-alusten kehitystä todennäköisesti jatketaan tulevaisuudessa, ja kehityksen painopiste tullee olemaan monikäyttöisten UAV:iden kehityksessä. UAV:t tullee tulevaisuudessa korvaamaan miehitetyt ilma-alukset ilmatiedustelun ja ilmasta maahan -toiminnan osalta.

Tehokalastuksen vaikutukset Lappajärvessä : Kalastus elinkeinoksi Lappajärvellä -hankkeen vuosien 2001-2006 seurannan loppuraportti

Lappajärven, Etelä-Pohjanmaan suurimman järven, ekologinen tila on heikentynyt ja se luokitellaan nykyisin tyydyttäväksi. Rehevöityminen, 1990-luvun muikkukato ja yhteiskunnan rakennemuutos ovat vähentäneet kalastajien ja kalansaaliin määrää. Ammattikalastuksen edellytysten parantamiseksi toteutettiin vuosina 2001–2004 ”Kalastus elinkeinoksi Lappajärvellä” – hanke. Hankkeen vaikutuksia kalastoon ja järven ekosysteemiin seurattiin vuosina 2001–2006. Kokonaissaalis vuosina 2001–2004 oli 1655 tonnia (n. 29 kg/ha/vuosi). Vaikka saalistavoitetta (2100 tn) ei täysin saavutettu, oli hankkeella runsaasti myönteisiä, joskin osin vähäisiä tai epävarmoja vaikutuksia järven tilaan ja kalastoon. Kalastajien yksikkösaaliit etenkin kuhan ja isokokoisen ahvenen osalta nousivat hankkeen alussa. Loppuvuosina yksikkösaaliiden kehitys oli osittain ristiriitaista. Ulapan taloudellisesti vähempiarvoinen kalasto, varsinkin kuorekanta harveni selvästi. Muikku puolestaan lisääntyi ja pysyi runsaana. Kuhan ja isokokoisen ahvenen lisääntymistä selittivät pääosin lämpimät kesät ja kuhan onnistuneet istutukset. Petokalat, etenkin kuha, voivat osaltaan säädellä Lappajärven ulapan planktonsyöjäkalojen kantoja ja toisaalta estää särjen siirtymistä ulapalle. Tehokalastuksen loputtua alkoi näkyä merkkejä kalakantojen palautumisesta takaisin kohti lähtötilannetta. Rantavyöhykkeellä hankkeen vaikutukset jäivät selvästi vähäisemmiksi kuin ulapalla. Eläinplankton lisääntyi hieman ylemmissä vesikerroksissa, mutta tilanne alkoi palautua ennalleen hankkeen jälkeen. Sinileväkukinnot vaikuttivat vähentyneen hankkeen aikana, tosin kukintojen voimakas vaihtelu vaikeutti arviointia. Tehokalastus ja vesiensuojelun tehostuminen vähensivät fosforipitoisuuksia, tosin myös vähäsateiset vuodet vaikuttivat vähenemiseen. Pitoisuuksien lasku jatkui hankkeen jälkeen. Hanketta edeltänyt klorofyllipitoisuuksien kasvusuuntaus pysähtyi ja pitoisuustaso vakiintui. Tehokalastus pienensi klorofylli/fosfori-suhdetta ja vähensi sisäistä kuormitusta, mutta vaikutus alkoi heiketä hankkeen loputtua. Typpipitoisuuksiin, veden väriin tai näkösyvyyteen kalastus ei vaikuttanut. Tehokalastuksella voidaan parantaa Lappajärven tilaa, mutta kalastuksen on oltava jatkuvaa. Saalistavoitteeksi voisi riittää n. 20-30 kg/ha vuodessa, mutta samalla on jatkettava ulkoisen kuormituksen vähentämistä. Kalastuksen tulisi kohdistua arvokalojen lisäksi myös vähempiarvoisiin kalalajeihin ja samalla on tuettava petokalakantojen vahvistumista.

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.

Miehittämättömien ilma-alusten tekninen kehitys 1990-luvulta nykypäivään

Tutkielman päätutkimuskysymys on, miten miehittämättömät ilma-alukset ovat teknisesti kehittyneet 1990-luvulta nykypäivään. Työssä keskitytään Yhdysvaltain asevoimien käyttämiin miehittämättömiin ilma-aluksiin, koska Yhdysvallat ovat edelläkävijä niiden kehittämisessä ja valmistamisessa. Tutkittavat miehittämättömät ilma-alukset ovat Pioneer, Predator, Hunter, Global Hawk ja Shadow. Tutkielman alatutkimuskysymykset ovat, mitä teknisiä muutoksia eri miehittämättömien ilma-alusten versioiden välillä on tapahtunut ja miten miehittämättömien ilma-alusten tekninen kehitys on vaikuttanut niiden suorituskykyyn. Tutkimusmenetelmänä työssä käytetään laadullista aineistoanalyysia eli kirjallisuustutkimusta. Lähdeaineistona hyödynnetään kirjallisuutta, vanhoja tutkielmia, lehtiä, julkaisuja sekä internetlähteitä, mitkä liittyvät miehittämättömiin ilma-aluksiin. Tutkimukseen käytetty materiaali on suurimmaksi osaksi sähköisessä muodossa, koska aihe on niin tuore ja kehittyy edelleen. Tutkielman tuloksena voidaan havaita, että miehittämättömien ilma-alusten tekninen kehitys on ollut nopeaa viime vuosien aikana ja jatkuu edelleen. Niihin on tehty paljon rakenteellisia muutoksia sekä koneiden runkoihin on tuotu uudempia ja tehokkaampia sensorijärjestelmiä. Myös aseistusta on kehitetty osassa koneista. Teknisellä kehityksellä on ollut vaikutusta miehittämättömien ilma-alusten suorituskykyyn etenkin taisteluun tarkoitetuilla koneilla. Melkein kaikissa miehittämättömissä ilma-aluksissa on kyetty kasvattamaan koneen maksiminopeutta ja toimintakorkeutta. Muutoksia on myös tapahtunut joidenkin koneiden toiminta-ajassa. Johtopäätöksenä voidaan todeta, että suurimmat teknisen kehityksen kohteet miehittämättömissä ilma-aluksissa ovat olleet rakenteellisten muutosten ansiosta uusien sensorijärjestelmien tuonti koneisiin sekä niiden aseistaminen. Tekninen kehitys jatkuu edelleen miehittämättömissä ilma-aluksissa ja niillä pyritään yhä enemmän korvaamaan miehitettyjä ilmaaluksia ja niiden tehtäviä. Tällä voidaan välttää inhimillisiä virheitä sekä käyttää ilmaaluksia yhä vaarallisemmissa tehtävissä.

Tuning cell motility : roles of nestin and vimentin in cancer cell invasion and motility

The cytoskeleton is a key feature of both prokaryotic and eukaryotic cells. Itis comprised of three protein families, one of which is the intermediate filaments (IFs). Of these, the IFs are the largest and most diverse. The IFs are expressed throughout life, and are involved in the regulation of cell differentiation, homeostasis, ageing and pathogenesis. The IFs not only provide structural integrity to the cell, they are also involved in a range of cellular functions from organelle trafficking and cell migration to signalling transduction. The IFs are highly dynamic proteins, able to respond and adapt their network rapidly in response to intra- and extra- cellular cues. Consequently they interact with a whole host of cellular signalling proteins, regulating function, and activity, and cellular localisation. While the function of some of the better-known IFs such as the keratins is well studied, the understanding of the function of two IFs, nestin and vimentin, is poor. Nestin is well known as a marker of differentiation and is expressed in some cancers. In cancer, nestin is primarily described as is a promoter of cell motility, however, how it fulfils this role remains undefined. Vimentin too is expressed in cancer, and is known to promote cell motility and is used as a marker for epithelial to mesenchymal transition (EMT). It is only in the last decade that studies have addressed the role that vimentin plays in cell motility and EMT. This work provides novel insight into how the IFs, nestin and vimentin regulate cell motility and invasion. In particular we show that nestin regulates the cellular localisation and organisation of two key facilitators of cell migration, focal adhesion kinase and integrins. We identify nestin as a regulator of extracellular matrix degradation and integrin-mediated cell invasion. Two further studies address the specific regulation of vimentin by phosphorylation. A detailed characterisation study identified key phosphorylation sites on vimentin, which are critical for proper organisation of the vimentin network. Furthermore, we show that the bioactive sphingolipids are vimentin network regulators. Specifically, the sphingolipids induced RhoA kinasedependent (ROCK) phosphorylation at vimentin S71, which lead to filament reorganisation and inhibition of cell migration. Together these studies shed new light into the regulation of nestin and vimentin during cell motility.

Physical Attractiveness as a Signal of Biological Quality

It has been commonly thought that standards of beauty are arbitrary cultural conventions that vary between cultures and time. In my thesis I found that it is not so. Instead, I show that attractiveness and preferred traits serve as cues to phenotypic qualities that provide selective benefits for those who choose their mates based on these criteria. In the first study I show that attractive men have a stronger antibody response to the hepatitis b vaccine and higher levels of testosterone than their less attractive peers. Men with low levels of testosterone also tend to have high levels of the stress hormone cortisol, suggesting that their immune responses may have been inhibited by stress hormones. Thus, facial attractiveness may serve as an honest cue of the strength of immune defence in men. In the second study, I show that the attractiveness of the male body is also a cue of better immunity. In addition, I show that adiposity, both in men’s faces and bodies, is a better cue of the strength of immunity and attractiveness than of masculinity. In the third study, I test the preferences of women from 13 countries for facial cues of testosterone and cortisol. I show that there is cross-cultural variation in women’s preference for cues of testosterone and cortisol in male faces. I found a relationship between the health of a nation and women’s preferences for cues of testosterone in the male face and the interaction between preferences for cues of testosterone and cortisol. I show also a relationship between preferences for cues of testosterone and a societal-level measure of parasite stress. Thus, it seems that societal-level ecological factors influence the relative value of traits as revealed by combinations of testosterone and stress hormones. In the fourth study, I show that women’s immune responsiveness (amount of antibodies produced) does not predict facial attractiveness. Instead, plasma cortisol level is negatively associated with attractiveness, indicating that stressed women look less attractive. Fat percentage is curvilinearly associated with facial attractiveness, indicating that being too thin or too fat reduces attractiveness. This study suggests that in contrast to men, facial attractiveness in women does not indicate the strength of immune defence, but is associated with other aspects of long-term health and fertility: circulating levels of the stress hormone cortisol and the percentage of body fat. In the last study I show that the attractiveness of men’s body odor is positively correlated with stress hormone levels, suggesting also that the attractiveness of body odors may signal the phenotypic quality of males to females. However, the attractiveness of men’s body odor was not associated with testosterone levels. My thesis suggests that the standard of beauty is not in the eye of the beholder. Instead, our standard of beauty is hardwired in our brains by genes that are selected by natural selection and also influenced by current environmental conditions.