<p>Recent changes in the timing of annual events are a sign that climate change is already impacting ecosystems. Carbon sequestration by forests increases with longer growing seasons. Biodiversity can be affected by mis-timing of events through shading interactions and frost damage. Projecting forests' ability to provide these ecosystem services in the future requires an understanding of trees' phenological responses to a new climate. I begin by proposing a first order definition of an `optimal' phenological response to warming: that the mean temperature following budburst should remain essentially constant. Analogously, the temperature preceding senescence can serve the same role. </p><p>To understand which environmental cues will drive future changes in phenology, I assimilate clues from observational and experimental literature. For budburst in woody plants, spring warmth, over-winter chilling and light drive nearly all behavior, but species' responses vary widely. Species using chilling or light as safety mechanisms against budburst during mid-winter thaws are thought to be less able to phenologically track a warming climate. However, I show that even species cued solely by spring warmth are likely to under-track temperature changes. Fall cues are more idiosyncratic, and a plant's driver of senescence is likely to vary from year to year. </p><p>Models are a tempting method to untangle species budburst cues and forecast phenology under warmer climate scenarios. I tested two models' ability to recover parameters used to simulate budburst data. The simpler model was cued only by spring warmth while the complex one modulated warmth requirements with chilling exposure. For the simple model, parameters could be recovered consistently from some, but not all, regions of parameter space. The complex model's parameters were largely unrecoverable. To understand the consequences of parameter uncertainty, I applied both models to an 18 year phenological record of 13 deciduous tree species. While a few species fell into identifiable regions of the simple model's parameter space, most did not, and projected budburst dates had wide parameter-derived uncertainty intervals. These bands were wider still under a 5°C warming scenario. Even greater uncertainty resulted from the complex model.</p><p>To better understand plants' potential for growing season extension I subjected seedlings to warmer climates in a series of open-topped chambers in sites at each end of the eastern deciduous biome. Soil and air were heated to 3 or 5°C above ambient, or left unheated. For nearly all species, warming hastened budburst and germination and delayed senescence. However, these events failed to track temperature changes, happening at warmer temperatures in hotter chambers. Individual species showed a remarkable variability of all events' dates within treatments, and even within chambers. Because phenological traits are heritable, this offers a potential for evolutionary response to climate change.</p><p>This research has shown that while individual trees extend their growing seasons under warmer temperatures, they typically under-respond to the magnitude of warming, suggesting forests' capacity for increased carbon sequestration may reach a limit. However, within populations, trees vary substantially in their phenological responses, forming a possibility for evolutionarily adaptation to changing cues.</p> / Dissertation

Climate changes have been observed in the Northern Hemisphere during the last century, causing a longer growing season and upslope expansion of forest, tree, and shrub limits. Here, a long-term historical perspective was used as a tool for investigating how climate change has impacted woody plants at the forest, tree, and shrub limits across the study region. For this, historical data about plant communities were used, including the position of the forest, tree, and shrub limits, gathered by the Reindeer Commission in Troms County (1914-1915). The historical data were compared to contemporary data from a re-visit study in 2022. In addition, variations in microclimatic factors were investigated by looking at the growing season length, distance to coast, slope, and aspect. Importantly, the growing season length was found to have increased across the study region with the largest increase along the coast. The increase in the growing season length led to an upslope shift in the vegetation limits, although not significantly so for the forest limit. The field layer had shifted from meadow to heath around the historical shrub limit, and non-significantly for the forest and tree limit zones, indicating that the alpine tundra is the most sensitive to the observed warming. Distance to coast and slope did not impact the shift in vegetation limits, while aspect impacted the forest limit shift. The results from this thesis indicate that future studies should expand their research beyond climate variables and explore factors such as herbivory and land use change

No description available.

Abstract Adaptation to local environment is important for all organisms to guarantee survival and to maximize reproduction. Populations of the same species may live in environments that differ markedly. Due to differential selection pressures this can lead to population differentiation, which can be studied both at the phenotypic and at the gene level. The growing season cued by long days is typically short in the north, whereas southern populations have long growing seasons and are adapted to short days. Seasonal fluctuations in temperature also differ between northern and southern environments. Daylength and temperature regulate the timing of flowering in plants. Environmental regulation of flowering and its genetic basis has been extensively studied in the annual model species Arabidopsis thaliana. The perennial growth and flowering habit has been studied especially in trees, but studies on herbaceous plants species have been lacking. In this thesis, I have studied adaptation to growing season length in a perennial herbaceous model species Arabidopsis lyrata. Individuals from populations adapted to northern and southern environments in Europe were grown in same conditions in the growth chambers and in the field. Differentiation between the populations was studied by observing their flowering phenotypes and by studying the expression of genes that are candidates for governing the phenotypic differentiation. The main result in the thesis was that adaptation to short growing season in north can be seen as long daylength requirement for flowering and as fast developmental rate. Critical daylength for flowering likely regulates especially the timing of flowering cessation in the end of the growing season. Flowering time of individuals from northern populations also responded more strongly to cold treatment (representing winter) than that of the southern population. The cold requirement for flowering guarantees that the plants only flower after the winter in the spring with suitable conditions. Expression studies indicated that population differentiation in flowering could be at least partly governed by the expression variation in a few candidate genes. The results in this thesis are valuable for instance for understanding perennial species in general, including tree and crop species, and for predicting how plants response to changing climate. / Tiivistelmä Ympäröiviin oloihin sopeutuminen on tärkeää kaikille organismeille selviytymisen ja jälkeläistuoton kannalta. Saman lajin eri populaatiot saattavat elää ympäristöissä, joiden olosuhteet poikkeavat toisistaan huomattavasti. Tällöin populaatioihin kohdistuvat erilaiset valintapaineet ja populaatiot erilaistuvat. Erilaistuminen havaitaan tarkastelemalla yksilöiden ilmiasuja ja geenejä. Kasvukauden pituus määrittää eteläisten ja pohjoisten kasvuympäristöjen valintapaineita. Pohjoisessa kasvukausi on lyhyt ja sen alkamisesta ja loppumisesta kertoo pitkä päivänpituus. Etelässä on pitkä kasvukausi ja siellä elävät populaatiot ovat sopeutuneet lyhyeen päivänpituuteen. Myös vuodenaikaiset lämpötilavaihtelut eroavat pohjoisten ja eteläisten alueiden välillä. Muutokset päivänpituudessa ja lämpötilassa säätelevät kasveilla kukkimisen ajankohtaa. Kukkimiseen vaikuttavia ympäristötekijöitä ja kukkimista sääteleviä geenejä on tutkittu paljon yksivuotisella mallilajilla lituruoholla (Arabidopsis thaliana). Monivuotisten kasvien kasvun ja kukkimisen säätelyä on tutkittu etenkin puilla, mutta hyvin vähän ruohovartisilla kasveilla. Tässä väitöskirjatyössä tutkin kasvukauden pituuteen sopeutumista monivuotisella ruohovartisella lajilla, idänpitkäpalolla (Arabidopsis lyrata). Pohjoisiin ja eteläisiin ympäristöoloihin sopeutuneiden eurooppalaisten populaatioiden yksilöitä kasvatettiin samanlaisissa olosuhteissa kontrolloiduissa kasvatushuoneissa ja kenttäolosuhteissa. Populaatioiden erilaistumista tarkkailtiin kukkimiseen liittyvissä ominaisuuksissa sekä eroja selittävien kandidaattigeenien ekspressiossa. Tutkimuksen päätulos oli, että pohjoisen populaation lyhyeen kasvukauteen sopeutuminen voidaan havaita pitkän päivän vaatimuksena kukkimiselle ja nopeana kehityksenä. Kenttäolosuhteissa päivänpituus sääteli etenkin kukkimisen lopetusta kasvukauden lopussa. Pitkä kylmäkäsittely nopeutti kukkimista etenkin pohjoisessa populaatiossa. Kukkimista edeltävä kylmävaatimus takaa, etteivät kasvit kuki syksyllä epäsuotuisissa olosuhteissa, vaan vasta keväällä talven jälkeen. Populaatioiden erilaistuminen kukkimisen päivänpituus- ja kylmävaatimuksessa selittyy todennäköisesti ainakin osittain kandidaattigeenien ekspressioeroilla. Tämän väitöskirjatyön tuloksia voidaan soveltaa monivuotisten viljelykasvien jalostuksessa tai ennustettaessa kasvipopulaatioiden sopeutumista ilmastonmuutokseen.