Species distribution models predict a wholesale redistribution of trees in the next century, yet migratory responses necessary to spatially track climates far exceed maximum post-glacial rates. The extent to which populations will adapt will depend upon phenotypic variation, strength of selection, fecundity, interspecific competition, and biotic interactions. Populations of temperate and boreal trees show moderate to strong clines in phenology and growth along temperature gradients, indicating substantial local adaptation. Traits involved in local adaptation appear to be the product of small effects of many genes, and the resulting genotypic redundancy combined with high fecundity may facilitate rapid local adaptation despite high gene flow. Gene flow with preadapted alleles from warmer climates may promote adaptation and migration at the leading edge, while populations at the rear will likely face extirpation. Widespread species with large populations and high fecundity are likely to persist and adapt, but will likely suffer adaptational lag for a few generations. As all tree species will be suffering lags, interspecific competition may weaken, facilitating persistence under suboptimal conditions. Species with small populations, fragmented ranges, low fecundity, or suffering declines due to introduced insects or diseases should be candidates for facilitated migration.

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Adaptation, migration or extirpation: climate change outcomes for tree populations

Although trees have responded to global warming in the past - to temperatures higher than they are now - the rate of change predicted in the 21st century is likely to be unprecedented. Greenhouse gas emissions could cause a 3-6°C increase in mean land surface temperature at high and temperate latitudes. Despite this, few experiments have isolated the effects of temperature for this scenario on trees and forests. This review focuses on tree and forest responses at boreal and temperate latitudes, ranging from the cellular to the ecosystem level. Adaptation to varying temperatures revolves around the trade-off between utilizing the full growing season and minimizing frost damage through proper timing of hardening in autumn and dehardening in spring. But the evolutionary change in these traits must be sufficiently rapid to compensate for the temperature changes. Many species have a positive response to increased temperature - but how close are we to the optima? Management is critical for a positive response of forest growth to a warmer climate, and selection of the best species for the new conditions will be of vital importance. Contents Summary 369 I. Introduction 370 II. Photosynthesis and respiration 370 III. Soil organic matter decomposition and mineralization 373 IV. Phenology and frost hardiness 376 V. Whole tree experimental responses to warming 380 VI. Changes in species distribution at warmer temperatures 381 VII. Adaptation and evolution 383 VIII. Ecosystem level responses to warming 387 Acknowledgements 390 References 390 Appendix I. Temperature response functions 399.

https://nph.onlinelibrary.wiley.com/doi/pdf/10.1046/j.1469-8137.2001.00057.x

Tree and forest functioning in response to global warming

In tree species native to temperate and boreal regions, the activity-dormancy cycle is an important adaptive trait both for survival and growth. We discuss recent research on mechanisms controlling the overlapping developmental processes that define the activity-dormancy cycle, including cessation of apical growth, bud development, induction, maintenance and release of dormancy, and bud burst. The cycle involves an extensive reconfiguration of metabolism. Environmental control of the activity-dormancy cycle is based on perception of photoperiodic and temperature signals, reflecting adaptation to prevailing climatic conditions. Several molecular actors for control of growth cessation have been identified, with the CO/FT regulatory network and circadian clock having important coordinating roles in control of growth and dormancy. Other candidate regulators of bud set, dormancy and bud burst have been identified, such as dormancy-associated MADS-box factors, but their exact roles remain to be discovered. Epigenetic mechanisms also appear to factor in control of the activity-dormancy cycle. Despite evidence for gibberellins as negative regulators in growth cessation, and ABA and ethylene in bud formation, understanding of the roles that plant growth regulators play in controlling the activity-dormancy cycle is still very fragmentary. Finally, some of the challenges for further research in bud dormancy are discussed.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1365-3040.2012.02552.x

The dynamic nature of bud dormancy in trees: environmental control and molecular mechanisms.

http://www.uws.or.ug/wp-content/themes/UWS/PDFs/FINAL/Priscilla%20-Nakasongola%20presentation%20october%202011.pdf

Global warming and sexual plant reproduction

Plants grow in dynamic environments where they can be exposed to a multitude of stressful factors, all of which affect their development, yield, and, ultimately, reproductive success. Plants are adept at rapidly acclimating to stressful conditions and are able to further fortify their defenses by retaining memories of stress to enable stronger or more rapid responses should an environmental perturbation recur. Indeed, one mechanism that is often evoked regarding environmental memories is epigenetics. Yet, there are relatively few examples of such memories; neither is there a clear understanding of their duration, considering the plethora of stresses in nature. We propose that this field would benefit from investigations into the processes and mechanisms enabling recovery from stress. An understanding of stress recovery could provide fresh insights into when, how, and why environmental memories are created and regulated. Stress memories may be maladaptive, hindering recovery and affecting development and potential yield. In some circumstances, it may be advantageous for plants to learn to forget. Accordingly, the recovery process entails a balancing act between resetting and memory formation. During recovery, RNA metabolism, posttranscriptional gene silencing, and RNA-directed DNA methylation have the potential to play key roles in resetting the epigenome and transcriptome and in altering memory. Exploration of this emerging area of research is becoming ever more tractable with advances in genomics, phenomics, and high-throughput sequencing methodology that will enable unprecedented profiling of high-resolution stress recovery time series experiments and sampling of large natural populations.

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Reconsidering plant memory: Intersections between stress recovery, RNA turnover, and epigenetics

The temperature during maternal reproduction affects adaptive traits in progenies of Norway spruce (Picea abies (L) Karst.). Seed production in a cold environment advances bud set and cold acclimation in the autumn and dehardening and flushing in spring, whereas a warm reproductive environment delays timing of these traits. We repeated crosses between the same parents and produced seeds under contrasting temperatures. Elevated temperatures were applied at different time points from female meiosis to embryogenesis, followed by full-sib progeny tests in common environments. We measured timing of terminal bud formation, cold acclimation in the autumn and transcription levels of conifer phytochromes PhyO, PhyN, PhyP, and the class IV chitinase PaChi4 in these tests. No progeny differences were found that could be related to temperature differences during prezygotic stages and fertilization. In contrast, progeny performance was strongly associated with the degree-days from proembryo to mature seeds. Progenies with a warm embryonic history formed terminal buds later, were less hardy and expressed lower transcription levels of the Phy and PaChi4 genes. We hypothesize that temperature during zygotic embryogenesis and seed maturation regulates an 'epigenetic memory' in the progeny, involving differential expression of genes that may regulate bud phenology, cold acclimation and embryogenesis in Norway spruce.

https://onlinelibrary.wiley.com/doi/pdf/10.1111/j.1365-3040.2005.01356.x

Climatic adaptation in Picea abies progenies is affected by the temperature during zygotic embryogenesis and seed maturation

Adaptive traits in Picea abies (Norway spruce) progenies are influenced by the maternal temperatures during seed production. Here, we have extended these studies by testing the effects of maternal photoperiod and temperature on phenology and frost hardiness on progenies. Using eight phytotron rooms, seeds from three unrelated crosses were made in an environmental 2 x 2 factorial combination of long and short days and high and low temperatures. The progenies were then forced to cease growth rapidly at the end of the first growing season. An interactive memory effect was expressed the second growth season. Progenies from high temperature and short days, and from low temperatures and long days, started growth later in spring, ceased shoot growth later in summer, grew taller and were less frost hardy in the autumn than their full siblings from low temperatures and short days, and from high temperatures and long days. Norway spruce has developed a memory mechanism, regulating adaptive plasticity by photoperiod and temperature, which could counteract harmful effects of a rapidly changing climate.

Daylength and temperature during seed production interactively affect adaptive performance of Picea abies progenies.

Two experiments were designed to test possible effects of photoperiod and temperature during microsporogensis to anthesis on early autumn frost-hardiness of Picea abies progenies. Pollen lots were produced in phytotron rooms and used in crosses in a seed orchard. No biologically important differences in progeny performance were evident either between high and low temperature or between long and short-day treatments, and no significant interaction between photoperiod and temperature was found. In a third experiment, however, an effect of the environment during female flowering was obtained. Crosses performed in early spring (March) inside a heated greenhouse (short day, high temperature) produced progenies which were less hardy than their full-sibs reproduced from crosses indoors (long day, high temperature) and outdoors (long day, low temperature) in May. The most hardy siblings originated from the late-spring outdoor crosses. These results indicate that some stages in reproduction during female flowering, such as female meiosis, pollen tube growth, syngamy, early embryogenesis and embryo competition, may be sensitive to temperature and/or photoperiodic signals which can be transmitted to the progeny. We suspect that the altered performance of the progenies could be due to an activation of a regulatory mechanism affecting the expression of genes controlling adaptive traits. Both the present and earlier results have implications for the genetic interpretation of provenance differences in Norway spruce.

Ola Gram Dæhlen

Papers

Climatic adaptation in Picea abies progenies is affected by the temperature during zygotic embryogenesis and seed maturation

Daylength and temperature during seed production interactively affect adaptive performance of Picea abies progenies.

Influence of the female flowering environment on autumn frost-hardiness of Picea abies progenies

An epigenetic memory from time of embryo development affects climatic adaptation in Norway spruce.