Erja Taulavuori

Biodiversity, distributions and adaptations of arctic species in the context of environmental change

Abstract The individual of a species is the basic unit which responds to climate and UV-B changes, and it responds over a wide range of time scales. The diversity of animal, plant and microbial species appears to be low in the Arctic, and decreases from the boreal forests to the polar deserts of the extreme North but primitive species are particularly abundant. This latitudinal decline is associated with an increase in super-dominant species that occupy a wide range of habitats. Climate warming is expected to reduce the abundance and restrict the ranges of such species and to affect species at their northern range boundaries more than in the South: some Arctic animal and plant specialists could face extinction. Species most likely to expand into tundra are boreal species that currently exist as outlier populations in the Arctic. Many plant species have characteristics that allow them to survive short snow-free growing seasons, low solar angles, permafrost and low soil temperatures, low nutrient availability and physical disturbance. Many of these characteristics are likely to limit species. responses to climate warming, but mainly because of poor competitive ability compared with potential immigrant species. Terrestrial Arctic animals possess many adaptations that enable them to persist under a wide range of temperatures in the Arctic. Many escape unfavorable weather and resource shortage by winter dormancy or by migration. The biotic environment of Arctic animal species is relatively simple with few enemies, competitors, diseases, parasites and available food resources. Terrestrial Arctic animals are likely to be most vulnerable to warmer and drier summers, climatic changes that interfere with migration routes and staging areas, altered snow conditions and freeze-thaw cycles in winter, climate-induced disruption of the seasonal timing of reproduction and development, and influx of new competitors, predators, parasites and diseases. Arctic microorganisms are also well adapted to the Arctics climate: some can metabolize at temperatures down to −39°C. Cyanobacteria and algae have a wide range of adaptive strategies that allow them to avoid, or at least minimize UV injury. Microorganisms can tolerate most environmental conditions and they have short generation times which can facilitate rapid adaptation to new environments. In contrast, Arctic plant and animal species are very likely to change their distributions rather than evolve significantly in response to warming.

Responses to projected changes in climate and UV-B at the species level

Abstract Environmental manipulation experiments showed that species respond individualistically to each environmental-change variable. The greatest responses of plants were generally to nutrient, particularly nitrogen, addition. Summer warming experiments showed that woody plant responses were dominant and that mosses and lichens became less abundant. Responses to warming were controlled by moisture availability and snow cover. Many invertebrates increased population growth in response to summer warming, as long as desiccation was not induced. CO2 and UV-B enrichment experiments showed that plant and animal responses were small. However, some microorganisms and species of fungi were sensitive to increased UV-B and some intensive mutagenic actions could, perhaps, lead to unexpected epidemic outbreaks. Tundra soil heating, CO2 enrichment and amendment with mineral nutrients generally accelerated microbial activity. Algae are likely to dominate cyanobacteria in milder climates. Expected increases in winter freeze-thaw cycles leading to ice-crust formation are likely to severely reduce winter survival rate and disrupt the population dynamics of many terrestrial animals. A deeper snow cover is likely to restrict access to winter pastures by reindeer/caribou and their ability to fiee from predators while any earlier onset of the snow-free period is likely to stimulate increased plant growth. Initial species responses to climate change might occur at the sub-species level: an Arctic plant or animal species with high genetic/racial diversity has proved an ability to adapt to different environmental conditions in the past and is likely to do so also in the future. Indigenous knowledge, air photographs, satellite images and monitoring show that changes in the distributions of some species are already occurring: Arctic vegetation is becoming more shrubby and more productive, there have been recent changes in the ranges of caribou, and “new” species of insects and birds previously associated with areas south of the treeline have been recorded. In contrast, almost all Arctic breeding bird species are declining and models predict further quite dramatic reductions of the populations of tundra birds due to warming. Species–climate response surface models predict potential future ranges of current Arctic species that are often markedly reduced and displaced northwards in response to warming. In contrast, invertebrates and microorganisms are very likely to quickly expand their ranges northwards into the Arctic.

Accelerated dehardening in bilberry (Vaccinium myrtillus L.) induced by a small elevation in air temperature

The effect of climatic warming on the dehardening potential of bilberry (Vaccinium myrtillus L.) in a northern boreal environment (65 degrees N) was studied. Natural stands of bilberry were heated artificially in winter. No reference point for the heating was set, since the purpose was to follow the fluctuations in ambient air temperatures. These were 2-3 degrees C higher in the heated plots than in the control plots from October to May. Frost resistance (LT50) and the pH of cell effusate were monitored throughout. Bud phenology was assessed in May and related to various biochemical analyses, including glucose, fructose, sucrose, starch and total and reduced glutathione. Frost resistance started to decrease earlier in the heated plants, as did the pH of the cell effusate. Bud phenology was in accordance with the LT50 and pH results, since new growth had emerged in the heated plants by the beginning of May, when the controls still displayed dormancy. Concentrations of glucose, fructose and sucrose were significantly lower in the heated bilberries while concentrations of starch were higher. The heated plants also exhibited the lowest glutathione concentrations, but the difference was only marginal. The redox state of glutathione showed no difference between the treatments. The results suggest that a small elevation in air temperature can accelerate dehardening in the bilberry. It is thus concluded that climatic warming may entail a real risk of early dehardening and further frost damage for the bilberry.

Impacts of extreme winter warming events on plant physiology in a sub-Arctic heath community.

Insulation provided by snow cover and tolerance of freezing by physiological acclimation allows Arctic plants to survive cold winter temperatures. However, both the protection mechanisms may be lost with winter climate change, especially during extreme winter warming events where loss of snow cover from snow melt results in exposure of plants to warm temperatures and then returning extreme cold in the absence of insulating snow. These events cause considerable damage to Arctic plants, but physiological responses behind such damage remain unknown. Here, we report simulations of extreme winter warming events using infrared heating lamps and soil warming cables in a sub-Arctic heathland. During these events, we measured maximum quantum yield of photosystem II (PSII), photosynthesis, respiration, bud swelling and associated bud carbohydrate changes and lipid peroxidation to identify physiological responses during and after the winter warming events in three dwarf shrub species: Empetrum hermaphroditum, Vaccinium vitis-idaea and Vaccinium myrtillus. Winter warming increased maximum quantum yield of PSII, and photosynthesis was initiated for E. hermaphroditum and V. vitis-idaea. Bud swelling, bud carbohydrate decreases and lipid peroxidation were largest for E. hermaphroditum, whereas V. myrtillus and V. vitis-idaea showed no or less strong responses. Increased physiological activity and bud swelling suggest that sub-Arctic plants can initiate spring-like development in response to a short winter warming event. Lipid peroxidation suggests that plants experience increased winter stress. The observed differences between species in physiological responses are broadly consistent with interspecific differences in damage seen in previous studies, with E. hermaphroditum and V. myrtillus tending to be most sensitive. This suggests that initiation of spring-like development may be a major driver in the damage caused by winter warming events that are predicted to become more frequent in some regions of the Arctic and that may ultimately drive plant community shifts.

In vivo Chlorophyll fluorescence is not always a good indicator of cold hardiness

Summary This paper demonstrates that Chlorophyll fluorescence in vivo is not always a true indicator of plant cold hardiness. In this experiment, frost resistance and chlorophyll fluorescence of needles of Scots pine seedlings were followed under natural and controlled phytotron conditions during the most active cold hardening period.

Growth Responses of Trees to Arctic Light Environment

Currently, climate is warming at a considerable rate. The warming climate shifts the vegetation toward the poles. This causes competition between southern and northern plant species and ecotypes. Light environment obviously has an important role in controlling the competition. The controlling factors include daylength, spectral composition (especially the ratio of red/far-red light, blue light and UV-radiation), and the amount of photosynthetically active radiation.

MODELLING FROST RESISTANCE OF SCOTS PINE SEEDLINGS USING TEMPERATURE, DAYLENGTH AND PH OF CELL EFFUSATE

The annual course of frost resistance (LT50) and the pH of the cell effusate in needles of two-year-old Scots pine seedlings were monitored in a field experiment in Oulu, Northern Finland (65° N, 25° E) during 1995. The aim of the work was to to develop model to predict the annual variation in frost resistance by pH of the cell effusate and meteorological data. The seedlings were covered with a fibre cloth shelter which transmitted sufficient light for them to experience the photoperiod, but prevented the accumulation of snow over them. The shelter above the seedlings was removed at the beginning of May and erected again at the end of September. The seedlings were watered only for the time when the shelter was removed, and received fertilizer only during the previous summer (1994).Frost resistance was only -5° C during the growing season but more than -100° C during the winter rest period. It was about -10° C at the end of August, increased to -55° C in the next three weeks, and reached -100° C at the beginning of October. The pH of the cell effusate was lowest during the growing season and highest in winter, the difference being about one and half pH unit. Needles exposed to -196° C showed pH from 4.0 in summer to 5.5 in winter, while pH of the non-frozen needles varied from 5.0 to 6.5, respectively. Seasonal variation in frost resistance was explained by a regression model well (R2 = 0.9) when day length, minimum air temperature and pH were entered as variables.

Drought tolerance of juvenile and mature leaves of a deciduous dwarf shrub Vaccinium myrtillus L. in a boreal environment.

The difference between drought tolerance of juvenile and mature leaves of the winter-deciduous dwarf shrub bilberry (Vaccinium myrtillus L.) from a northern boreal environment was investigated. It was hypothesised that mature leaves are more drought sensitive than juvenile leaves. Bilberry plants were allowed to dry out by excluding irrigation when leaves were at juvenile and mature stages. Tissue water content decreased at both phenological stages, but the response was more pronounced in the mature leaves. Anthocyanin concentrations increased as the tissue water content decreased, and again this occurred to a greater extent in the mature leaves. Chlorophyll concentrations decreased only marginally at the juvenile stage, while the decrease was significant in the mature leaves. Chlorophyll degradation was enhanced by drought stress. Soluble proteins decreased and protein oxidation increased in the mature leaves, and degradation of oxidised proteins increased in the drought-stressed plants. The results suggest that leaves of bilberry are more sensitive to drought stress at the mature stage, and that drought stress accelerates senescence at the mature stage. The significance of the results is that dry periods during the juvenility of leaves are not as detrimental as they may be later in summer. In addition, the strategy of a winter-deciduous plant is obviously to protect its perennial parts from severe drought by accelerated leaf senescence at the mature stage. Therefore, the deciduous life form may provide an excellent adaptation against drought also in northern ecosystems. The role of anthocyanins in photoprotection under drought stress is also discussed.

Improved elongation of Scots pine seedlings under blue light depletion is not dependent on resource acquisition

Removal of blue light (400-500 nm) induced shoot elongation of 2-year-old Scots pine (Pinus sylvestris L.) seedlings, which was not related to resource acquisition (carbohydrates, C/N ratio and soluble proteins) and frost hardening. The seedlings were grown in northern Finland (64°N) in plexiglass chambers, either orange in colour or transparent, during elongation and cold hardening periods in 2001. The orange chamber removed the blue wavelengths. The results suggest that the growth inhibiting effect of blue light on Scots pine elongation is probably a photomorphogenic regulation response; the removal of blue light did not affect the gas exchange and accumulation of growth resources. In addition, the removal of blue light also did not affect the physiological parameters (pigment composition, chlorophyll fluorescence and lipid peroxidation) measured during the preparation for winter.

Polyamines and Glutathione Metabolism in N Fertilized Scots pine Seedlings during Cold Hardening

Summary Scots pine (Pinus sylvestris L.) seedlings in an experimental field in northern Finland (65 °N) were subjected to two doses (0.5 or 1.0 g per seedling) of ammonium nitrate (NH4NO3 × Ca(OH)2) fertilizer on two different dates (June 6 or July 6, 1996). The effect of an increased supply of nitrogen (N) on certain compounds related to N metabolism and cold hardening were investigated. Samples for the analysis of free polyamines (PAs), glutathione, glutathione reductase (GR; EC 1.6.4.2) and γ-glutamyltranspeptidase (γ-GT; EC 2.3.2.2) were harvested on three occasions during cold hardening in autumn 1996 (on Aug. 5th, Aug. 19th and Sept. 30th). The levels of PAs generally increased as a function of dose and delay in fertilization, while responses of glutathione concentration, proportion of reduced glutathione (GSH %), GR and γ-GT activities varied. PA levels generally decreased during hardening while responses of the compounds of glutathione metabolism varied. Glutathione concentration in fertilized plants tended to decrease during hardening, while that in non-fertilized ones showed no clear response. The GR activity increased considerably and the γ-GT activity showed no significant response during hardening.