Models of Dispersal and Diversification

Ecology and evolutionary biology is the study of life on this planet. One of the many methods applied to answering the great diversity of questions regarding the lives and characteristics of individual organisms, is the utilization of mathematical models. Such models are used in a wide variety of ways. Some help us to reason, functioning as aids to, or substitutes for, our own fallible logic, thus making argumentation and thinking clearer. Models which help our reasoning can lead to conceptual clarification; by expressing ideas in algebraic terms, the relationship between different concepts become clearer. Other mathematical models are used to better understand yet more complicated models, or to develop mathematical tools for their analysis. Though helping us to reason and being used as tools in the craftmanship of science, many models do not tell us much about the real biological phenomena we are, at least initially, interested in. The main reason for this is that any mathematical model is a simplification of the real world, reducing the complexity and variety of interactions and idiosynchracies of individual organisms. What such models can tell us, however, both is and has been very valuable throughout the history of ecology and evolution. Minimally, a model simplifying the complex world can tell us that in principle, the patterns produced in a model could also be produced in the real world. We can never know how different a simplified mathematical representation is from the real world, but the similarity models do strive for, gives us confidence that their results could apply. This thesis deals with a variety of different models, used for different purposes. One model deals with how one can measure and analyse invasions; the expanding phase of invasive species. Earlier analyses claims to have shown that such invasions can be a regulated phenomena, that higher invasion speeds at a given point in time will lead to a reduction in speed. Two simple mathematical models show that analysis on this particular measure of invasion speed need not be evidence of regulation. In the context of dispersal evolution, two models acting as proof-of-principle are presented. Parent-offspring conflict emerges when there are different evolutionary optima for adaptive behavior for parents and offspring. We show that the evolution of dispersal distances can entail such a conflict, and that under parental control of dispersal (as, for example, in higher plants) wider dispersal kernels are optimal. We also show that dispersal homeostasis can be optimal; in a setting where dispersal decisions (to leave or stay in a natal patch) are made, strategies that divide their seeds or eggs into fractions that disperse or not, as opposed to randomized for each seed, can prevail. We also present a model of the evolution of bet-hedging strategies; evolutionary adaptations that occur despite their fitness, on average, being lower than a competing strategy. Such strategies can win in the long run because they have a reduced variance in fitness coupled with a reduction in mean fitness, and fitness is of a multiplicative nature across generations, and therefore sensitive to variability. This model is used for conceptual clarification; by developing a population genetical model with uncertain fitness and expressing genotypic variance in fitness as a product between individual level variance and correlations between individuals of a genotype. We arrive at expressions that intuitively reflect two of the main categorizations of bet-hedging strategies; conservative vs diversifying and within- vs between-generation bet hedging. In addition, this model shows that these divisions in fact are false dichotomies.

Investigating online reading with eye tracking and EEG : The influence of text format, reading task and parafoveal stimuli on reading processes

Research on reading has been successful in revealing how attention guides eye movements when people read single sentences or text paragraphs in simplified and strictly controlled experimental conditions. However, less is known about reading processes in more naturalistic and applied settings, such as reading Web pages. This thesis investigates online reading processes by recording participants eye movements. The thesis consists of four experimental studies that examine how location of stimuli presented outside the currently fixated region (Study I and III), text format (Study II), animation and abrupt onset of online advertisements (Study III), and phase of an online information search task (Study IV) affect written language processing. Furthermore, the studies investigate how the goal of the reading task affects attention allocation during reading by comparing reading for comprehension with free browsing, and by varying the difficulty of an information search task. The results show that text format affects the reading process, that is, vertical text (word/line) is read at a slower rate than a standard horizontal text, and the mean fixation durations are longer for vertical text than for horizontal text. Furthermore, animated online ads and abrupt ad onsets capture online readers attention and direct their gaze toward the ads, and distract the reading process. Compared to a reading-for-comprehension task, online ads are attended to more in a free browsing task. Moreover, in both tasks abrupt ad onsets result in rather immediate fixations toward the ads. This effect is enhanced when the ad is presented in the proximity of the text being read. In addition, the reading processes vary when Web users proceed in online information search tasks, for example when they are searching for a specific keyword, looking for an answer to a question, or trying to find a subjectively most interesting topic. A scanning type of behavior is typical at the beginning of the tasks, after which participants tend to switch to a more careful reading state before finishing the tasks in the states referred to as decision states. Furthermore, the results also provided evidence that left-to-right readers extract more parafoveal information to the right of the fixated word than to the left, suggesting that learning biases attentional orienting towards the reading direction.

Maidontuottajatila ja ravinnepolitiikka: Tilan toimintamekanismit, politiikan tarve ja ohjauskeinojen vaikuttavuus

Tutkielman tarkoituksena on rakentaa malli, jolla voidaan arvioida maitotilan toimintaa monipuolisesti ravinnehuuhtouman kannalta. Tavoitteena on ennen muuta analyyttinen tarkastelu siten, että kuitenkin huomioiden maidontuotannon ominaispiirteet, keskeisinä esimerkkeinä ravinnon, maidontuotannon ja lannan ominaisuuksien väliset riippuvuudet. Analyysissa tarkastellaan tilanomistajan yksityisen hyödynmaksimoinnin ja yhteiskunnan tavoitteiden eroavaisuutta. Lisäksi johdetaan optimaaliset ohjauskeinot ja arvioidaan eräiden yksinkertaisten ohjauskeinojen vaikuttavuutta. Mallin analysoinnin perusteella yksityinen ja yhteiskunnallinen optimiratkaisu eroavat toisistaan kaikkien päätösmuuttujien osalta. Ei voida kuitenkaan yleispätevästi sanoa muutosten merkittävyyttä tai suuntaa ottamatta kantaa mallin funktiomuotoihin ja parametriarvoihin. Optimaaliset ohjauskeinot tulisi asettaa keinolannoitteen ja lannan levitykselle, väkirehuruokinnalle ja säilörehun viljelylle, mutta ei eläinten määrälle. Toisaalta optimaaliset ohjauskeinot ovat hyvin monimutkaisia. Tarkasteltujen yksinkertaisempien ohjauskeinojen vaikuttavuutta ei voida tarkasti arvioida analyyttisella tasolla. Numeeristen tulosten perusteella yhteiskunnallisessa optimissa eläinmäärä olisi hieman yksityistä optimia pienempi, väkirehun käyttö vähäisempää, enemmän peltoa allokoitaisiin säilörehun viljelyyn ja lannoitustasot olisivat kautta linjan hieman pienemmät. Lannanlevitykset eroavat etäisyyden suhteen: molemmissa tapauksissa lannanlevityksen intensiteetti kasvaa kohti tilan keskusta kuljettaessa, mutta yksityisessä optimissa lähimmälle pellolle dumpataan kaikki ylimääräinen lanta, kun taas yhteiskunnallisessa optimissa lanta levitetään tasaisemmin eri pelloille. Ero kokonaishyvinvoinnissa jää pieneksi, mutta eläinmäärän kasvu kärjistäisi lannan dumppauksen aiheuttamia huuhtoumahaittoja. Yksinkertaisista ohjauskeinoista lannoitusrajoitus sekä ravinne- ja keinolannoitevero osoittautuivat kohtalaisen käyttökelpoisiksi numeeristen tulosten perusteella. Sen sijaan eläinmäärän rajoittaminen ja lannan kuljetuskustannuksia kompensoivat tuet vaikuttavat tulosten perusteella huonoilta ratkaisuilta.