Early-stage detection of root freezing injuries of Scots pine (Pinus sylvestris L.) seedlings by impedance loss factor and hydraulic conductance.

The condition of the root system affects the quality of seedlings in forestry and horticulture. Previously, the electrical impedance loss factor (δ) and the reverse-flow hydraulic conductance (Kr ) of the roots of Scots pine seedlings were found to increase when assessed a few days after frost damage. How these variables change with time after the root damage is unknown. We arranged an experiment with 1.5-year-old Scots pine seedlings exposed to -5°C or - 30°C, with the control seedlings kept at 3°C. Then, δ and Kr of roots were monitored for 5 weeks in favorable growing conditions. The properties of the roots were observed to be in a dynamic state after the damage. A significant difference in δ was found between the test temperatures -30°C versus -5°C and 3°C (p = 0.004 and p < 0.001, respectively). The clearest effect of freezing injuries on δ of roots was observed in the first measurement 1 week after the freezing test. The temperature significantly affected Kr, too, with a significant difference between the low-temperature treated plants -30°C versus -5°C and control (p < 0.001, respectively). The difference in Kr between -30°C and the other two temperatures increased with time and was the largest in the last samples, taken after 5 weeks. We conclude that the impedance loss factor may detect root damage if the measurements occur early enough after the damage, but a longer time difference (3-5 weeks) is needed according to the reverse-flow hydraulic conductance.

Separating the effects of air and soil temperature on silver birch. Part I. Does soil temperature or resource competition determine the timing of root growth?

The aboveground parts of boreal forest trees mostly grow earlier, and the roots later, in the growing season. We aimed to experimentally test whether the extrinsic driver of soil temperature or the intrinsic driver (resource competition between plant parts) is a more important control for the root and shoot growth of silver birch (Betula pendula Roth) seedlings. Sixteen two-year-old seedlings were grown in controlled environment rooms for two simulated growing seasons (GS1, GS2). In GS1, all the seedlings were acclimatized under the same conditions, but in GS2, the soil temperature treatments were: (i) constant 10 °C (Cool); (ii) constant 18 °C (Warm); (iii) early growing season at 10 °C, switched to 18 °C later (Early Cool Late Warm, ECLW) and (iv) early growing season 18 °C, switched to 10 °C later (Early Warm Late Cool, EWLC). The treatments did not affect growth allocation between shoots and roots. Warm soil benefitted shoot elongation as it slowed down in EWLC and accelerated in ECLW after the soil temperature switch. However, whole-tree biomasses were similar to Cool and the seedlings grew largest in Warm. Phenology was not strongly affected by soil temperature, and root and shoot growth did not usually peak simultaneously. Short root mortality increased strongly in ECLW and decreased in EWLC after the soil temperature switch. Long root longevity was not significantly affected but long root growth ceased earliest in ECLW. Soil warming increased foliar nutrient contents. Growth dynamics were not solely driven by soil temperature, but resource competition also played a significant role. The study showed the importance of soil temperature for fine root dynamics not only through root growth but also via root mortality, as soil warming increased mortality even more than growth. Soil temperature has complex effects on tree and soil functioning, which further affects carbon dynamics in forest ecosystems that have a climate feedback.

Separating the effects of air and soil temperature on silver birch. Part II. The relation of physiology and leaf anatomy to growth dynamics.

The aboveground parts of boreal forest trees grow earlier in the growing season, the roots mostly later. The idea was to examine whether root growth followed soil temperature, or whether shoot growth also demanded most resources in the early growing season (soil temperature vs internal sink strengths for resources). The linkage between air and soil temperature was broken by switching the soil temperature. We aimed here to identify the direct effects of different soil temperature patterns on physiology, leaf anatomy and their interactions, and how they relate to the control of the growth dynamics of silver birch (Betula pendula Roth). Sixteen 2-year-old seedlings were grown in a controlled environment for two 14-week simulated growing seasons (GS1, GS2). An 8-week dormancy period interposed the GSs. In GS2, soil temperature treatments were applied: constant 10 °C (Cool), constant 18 °C (Warm), early growing season at 10 °C switched to 18 °C later (Early Cool Late Warm) and 18 °C followed by 10 °C (Early Warm Late Cool) were applied during GS2. The switch from cool to warm enhanced the water status, net photosynthesis, chlorophyll content index, effective yield of photosystem II (ΔF/Fm') and leaf expansion of the seedlings. Warm treatment increased the stomatal number per leaf. In contrast, soil cooling increased glandular trichomes. This investment in increasing the chemical defense potential may be associated with the decreased growth in cool soil. Non-structural carbohydrates were accumulated in leaves at a low soil temperature showing that growth was more hindered than net photosynthesis. Leaf anatomy differed between the first and second leaf flush of silver birch, which may promote tree fitness in the prevailing growing conditions. The interaction of birch structure and function changes with soil temperature, which can further reflect to ecosystem functioning.

Dynamics of above- and belowground responses of silver birch saplings and soil gases to soil freezing and waterlogging during dormancy.

Winter precipitation and soil freeze-thaw events have been predicted to increase in boreal regions with climate change. This may expose tree roots to waterlogging (WL) and soil freezing (Fr) more than in the current climate and therefore affect tree growth and survival. Using a whole-tree approach, we studied the responses of silver birch (Betula pendula Roth.) saplings, growing in mineral soil, to 6-week Fr and WL in factorial combinations during dormancy, with accompanying changes in soil gas concentrations. Physiological activation (dark-acclimated chlorophyll fluorescence and chlorophyll content index) and growth of leaves and shoot elongation and stem diameter growth started earlier in Fr than NoFr (soil not frozen). The starch content of leaves was temporarily higher in Fr than NoFr in the latter part of the growing season. Short and long root production and longevity decreased, and mortality increased by soil Fr, while there were no significant effects of WL. Increased fine root damage was followed by increased compensatory root growth. At the beginning of the growing season, stem sap flow increased fastest in Fr + WL, with some delay in both NoWL (without WL) treatments. At the end of the follow-up growing season, the hydraulic conductance and impedance loss factor of roots were higher in Fr than in NoFr, but there were no differences in above- and belowground biomasses. The concentration of soil carbon dioxide increased and methane decreased by soil Fr at the end of dormancy. At the beginning of the growing season, the concentration of nitrous oxide was higher in WL than in NoWL and higher in Fr than in NoFr. In general, soil Fr had more consistent effects on soil greenhouse gas concentrations than WL. To conclude, winter-time WL alone is not as harmful for roots as WL during the growing season.

Are arbuscular-mycorrhizal Alnus incana seedlings more resistant to drought than ectomycorrhizal and nonmycorrhizal ones?

Arbuscular mycorrhizas (AMs) prevail in warm and dry climates and ectomycorrhizas (EMs) in cold and humid climates. We suggest that the fungal symbionts benefit their host plants especially in the corresponding conditions. The hypothesis tested was that AM plants are more drought-resistant than EM or nonmycorrhizal (NM) plants. Grey alder (Alnus incana (L.) Moench) seedlings were inoculated with two species of either AM or EM fungi or none. In one controlled-environment experiment, there was a watering and a drought treatment. Another set of seedlings were not watered until permanent wilting. The AM plants were somewhat smaller than EM and NM, and at the early stage of the drought treatment, the soil-moisture content was slightly higher in the AM pots. Shoot water potential was highest in the AM treatment during severe drought, while stomatal conductance and photosynthesis did not show a mycorrhizal effect. In the lethal-drought set, the AM plants maintained their leaves longer than EM and NM plants, and the AM seedlings survived longer than NM seedlings. Foliar phosphorus and sulfur concentrations remained higher in AM plants than EM or NM, but potassium, copper and iron increased in EM during drought. The root tannin concentration was lower in AM than EM and drought doubled it. Although the difference in drought resistance was not large, the hypothesis was supported by the better performance of AM plants during a severe short-term drought. Sustained phosphorus nutrition during drought in AM plants was a possible reason for this. Moreover, the higher foliar sulfur and lower metal-nutrient concentrations in AM may reflect differences in nutrient uptake or (re)translocation during drought, which merit further research. The much larger tannin concentrations in EM root systems than AM did not appear to protect the EM plants from drought. The differential tannin accumulation in AM and EM plants needs further attention.