Heikki Hänninen

Potential for evolutionary responses to climate change – evidence from tree populations

Evolutionary responses are required for tree populations to be able to track climate change. Results of 250 years of common garden experiments show that most forest trees have evolved local adaptation, as evidenced by the adaptive differentiation of populations in quantitative traits, reflecting environmental conditions of population origins. On the basis of the patterns of quantitative variation for 19 adaptation-related traits studied in 59 tree species (mostly temperate and boreal species from the Northern hemisphere), we found that genetic differentiation between populations and clinal variation along environmental gradients were very common (respectively, 90% and 78% of cases). Thus, responding to climate change will likely require that the quantitative traits of populations again match their environments. We examine what kind of information is needed for evaluating the potential to respond, and what information is already available. We review the genetic models related to selection responses, and what is known currently about the genetic basis of the traits. We address special problems to be found at the range margins, and highlight the need for more modeling to understand specific issues at southern and northern margins. We need new common garden experiments for less known species. For extensively studied species, new experiments are needed outside the current ranges. Improving genomic information will allow better prediction of responses. Competitive and other interactions within species and interactions between species deserve more consideration. Despite the long generation times, the strong background in quantitative genetics and growing genomic resources make forest trees useful species for climate change research. The greatest adaptive response is expected when populations are large, have high genetic variability, selection is strong, and there is ecological opportunity for establishment of better adapted genotypes.

Plant Development Models

In this chapter we provide a brief overview of plant phenology modeling, focusing on mechanistic phenological models. After a brief history of plant phenology modeling, we present the different models which have been described in the literature so far and highlight the main differences between them, i.e. their degree of complexity and the different types of response function to temperature they use. We also discuss the different approaches used to build and parameterize such models. Finally, we provide a few examples of applications mechanistic plant phenological models have been successfully used for, such as frost hardiness modeling, tree growth modeling, tree species distribution modeling and temperature reconstruction of the last millennium.

Relationship between temperature and the seasonal course of photosynthesis in Scots pine at northern timberline and in southern boreal zone

In earlier studies the seasonal dynamics of photosynthetic capacity in northern conifers has been explained as a slow response to the ambient temperature. We tested this concept with Scots pine (Pinus sylvestris L.). We analysed the seasonal dynamics of photosynthetic efficiency in Scots pine at the timberline in Finnish Lapland, and in a southern boreal forest in Southern Finland. The relationship between the daily photosynthetic efficiency and leaf temperature history was determined from continuous measurements of shoot CO2 exchange. The shoot CO2 exchange and photosynthetic efficiency showed similar seasonal patterns in the northern and in the southern locations, following daily mean temperature with a delay. The relationship between the temperature history and photosynthetic efficiency appeared to be near sigmoidal both in the northern and in the southern trees. The relationship was also consistent from year-to-year, thus the seasonal course of photosynthetic efficiency can be predicted accurately from the ambient temperature using a sigmoidal relationship. A rapid decrease of photosynthetic efficiency was observed when daytime temperature dropped below zero or frost had occurred in the previous night. The difference in the rate of acclimation of photosynthetic efficiency between the north and the south was small.

Computations on Frost Damage to Scots Pine Under Climatic Warming in Boreal Conditions

To investigate the risk of frost damage to Scots pine (Pinus sylvestris L.) in northern regions under climatic warming, a submodel for such damage to trees was included in a forest ecosystem model of the gap type. An annual growth multiplier describing the effects of frost was calculated with the help of simulated daily frost hardiness and daily minimum temperature. The annual growth multiplier was used in the main ecosystem model when simulating the development of a tree stand using a time step of one year. Simulations of the growth and development of Scots pine stands in southern Finland (61{degrees} N) under an elevating temperature indicated that climatic warming could increase the risk of frost damage due to premature onset of growth during warm spells in the late winter and early spring. Risk of frost damage implies uncertainty in yield expectations from boreal forest ecosystems in the event of climatic warming. 38 refs., 9 figs., 4 tabs.

Different growth sensitivity to enhanced UV-B radiation between male and female Populus cathayana

We investigated sex-related morphological and physiological responses to enhanced UV-B radiation in the dioecious species Populus cathayana Rehd. Cuttings were subjected to two UV-B radiation regimes: ambient (4.5 kJ m⁻² day⁻¹) and enhanced (12.5 kJ m⁻² day⁻¹) biologically effective UV-B radiation for one growing season. Enhanced UV-B radiation was found to significantly decrease the shoot height and basal diameter and to reduce the leaf area, dry matter accumulation, net photosynthesis rate (P(n)), chlorophyll a/b ratio (Chl a/b) and anthocyanin content. Enhanced UV-B radiation also increased chlorophyll pigment, leaf nitrogen, malondialdehyde and abscisic acid (ABA) content, superoxide dismutase and peroxidase activities and UV-B-absorbing compounds. No significant effects of enhanced UV-B radiation were found on biomass allocation, gas exchange (except for P(n)), photochemical efficiency of photosystem II or water use efficiency. Moreover, different sensitivity to enhanced UV-B radiation between males and females was detected. Under enhanced UV-B radiation, males exhibited significantly higher basal diameter and leaf nitrogen, and lower Chl a/b, ABA content, UV-B-absorbing compounds, as well as less decrement of leaf area and dry matter accumulation than did females. However, no significant sexual differences in these traits were found under ambient UV-B radiation. Our results suggest that males may possess a greater UV-B resistance than do females, with males having a more efficient antioxidant system and higher anthocyanin content to alleviate UV-B penetration stress than females.

Increased heat requirement for leaf flushing in temperate woody species over 1980–2012: effects of chilling, precipitation and insolation

Recent studies have revealed large unexplained variation in heat requirement-based phenology models, resulting in large uncertainty when predicting ecosystem carbon and water balance responses to climate variability. Improving our understanding of the heat requirement for spring phenology is thus urgently needed. In this study, we estimated the species-specific heat requirement for leaf flushing of 13 temperate woody species using long-term phenological observations from Europe and North America. The species were defined as early and late flushing species according to the mean date of leaf flushing across all sites. Partial correlation analyses were applied to determine the temporal correlations between heat requirement and chilling accumulation, precipitation and insolation sum during dormancy. We found that the heat requirement for leaf flushing increased by almost 50% over the study period 1980-2012, with an average of 30 heat units per decade. This temporal increase in heat requirement was observed in all species, but was much larger for late than for early flushing species. Consistent with previous studies, we found that the heat requirement negatively correlates with chilling accumulation. Interestingly, after removing the variation induced by chilling accumulation, a predominantly positive partial correlation exists between heat requirement and precipitation sum, and a predominantly negative correlation between heat requirement and insolation sum. This suggests that besides the well-known effect of chilling, the heat requirement for leaf flushing is also influenced by precipitation and insolation sum during dormancy. However, we hypothesize that the observed precipitation and insolation effects might be artefacts attributable to the inappropriate use of air temperature in the heat requirement quantification. Rather than air temperature, meristem temperature is probably the prominent driver of the leaf flushing process, but these data are not available. Further experimental research is thus needed to verify whether insolation and precipitation sums directly affect the heat requirement for leaf flushing.

Impacts of Climate Change on Cold Hardiness of Conifers

Due to increased anthropogenic emissions into the atmosphere, concentrations of CO2 and other greenhouse gases have been increasing globally during the past decades. Despite realized and planned control measures, this increase is predicted to continue in the future. Due to the accompanying changes in the physical properties of the atmosphere, the air temperature of the globe is predicted to rise dramatically in the future. According to most meteorological scenarios, the increase of the global mean temperature will be 1 to 4.5°C by the year 2100. However, it is also predicted that the level of warming will be more pronounced in the north than in the south and greater during winter than during summer months (IPCC 1996).

Recovery of photosynthesis of boreal conifers during spring: a comparison of two models

Two models of the effects of air temperature on the recovery of photosynthetic capacity of boreal conifers during spring were compared with respect to three aspects: (i) An examination of the ecophysiological assumptions of the models revealed several differences between the models, the most important ones being related to photosynthetic dormancy during winter, reversibility of the recovery, and effects of short term frosts. (ii) Model simulations were applied to deduce the predictions of the models for the springtime recovery of photosynthetic capacity. Both models predicted an increase in the photosynthetic capacity as a result of rising air temperatures. However, the model developed and parameterised for Scots pine predicted earlier recovery than the model developed and parameterised for Norway spruce, and there was also more variation in the predicted recovery with the former than with the latter. Furthermore, in the case of the Scots pine model the photosynthetic capacity fluctuated with fluctuations of air temperature, i.e. the recovery was drastically reversed in several cases, whereas the reverse played generally only a minor role in the quite straightforwardly increasing photosynthetic capacity predicted by the Norway spruce model. A review of the empirical findings available in the literature suggests that a new model unifying some of the ecophysiological assumptions of the current models should be developed. (iii) Additional simulations were carried out to examine the implications of the ecophysiological differences between the models for the photosynthetic production of Scots pine in central Finnish conditions. For the photosynthetic production calculated for the first half of the year, the Norway spruce model as parameterised for Scots pine predicted on an average 23.3% higher values than the Scots pine model. Thus, also with respect to estimation of the carbon sequestration of forests, further unifying model development is called for.

The minimum temperature for budburst in Betula depends on the state of dormancy

Vegis has put forward the theory that the range of growth-promoting temperatures changes during the induction and the release of dormancy. We have tested the response of buds of Betula pubescens Ehrh. and B. pendula Roth. on temperature during the induction and release of dormancy. Betula seedlings were exposed to dormancy-inducing high-temperature and short-day conditions and subsequently to dormancy-releasing chilling conditions in darkness. To monitor the dormancy status of the seedlings, subsets of them were transferred to five forcing temperatures and their budburst was observed. The results show that the expression of dormancy was temperature dependent, so that the minimum temperature for 100% budburst rose during the induction and dropped during the release of dormancy. These responses may explain previous contradictions between experimental and modelling studies, but that needs to be verified with more extensive experiments, some of which are identified in this study. The results provide further evidence for the concept of gradual change in bud dormancy. They also suggest that global change studies modelling budburst phenology should address the changing expression of bud dormancy.

Bud burst in Norway spruce (Picea abies): preliminary evidence for age-specific rest patterns

This study examines the effect of chilling and photoperiod on rest completion and bud burst in Norway spruce [Picea abies (L.) Karst.] using twigs from both 15-year-old (“young trees”) and 56-year-old (“old trees”) trees. The material was transferred between September and May from outdoors to experimental forcing conditions with four different photoperiods. The bud burst percentage of the twigs from young trees generally increased in all photoperiods until the end of the year. After that it decreased until vernal equinox (March 20) and then increased steeply towards spring. This new observation of transient rest completion during autumn suggests that young trees have (I) a transient time window during late autumn when ontogenetic development is possible, and (ii) a secondary rest culminating approximately at the time of vernal equinox. In twigs from old trees the transient rest completion was much weaker as the bud burst percentages generally remained under 20 during autumn and winter. At vernal equinox there were no burst buds in twigs from the old trees in any photoperiod and after that the bud burst percentage increased basically in the same manner as in the young trees. The bud burst percentage of the twigs from young trees was generally higher as the photoperiod increased. However, no evidence for absolute long photoperiod requirement of rest completion was observed.