Tapani Repo

The relation between growth cessation and frost hardening in Scots pines of different origins

The cessation of shoot elongation, diameter growth and needle elongation were compared with the initiation of frost hardening of the stems and needles in an 8-year-old provenance trial of Scots pine (Pinus sylvestris L.) established in central Finland. The saplings were of six different origins ranging from Estonia to northern Finland, forming a latitudinal gradient of ca. 10°N. The frost hardiness of the stems of current-year shoots was assessed by electrical impedance analysis and that of current-year needles by electrolyte leakage and visual scoring of damage. Artificial freezing tests were used in the assessments. The pattern of growth cessation (shoot and needle elongation, diameter growth) tended to follow the latitude of origin, i.e. growth ceased in the northernmost provenance first and in the southernmost one last. Both stems and needles of the northern provenances hardened earlier than the southern ones, but the differences in hardiness disappeared as hardening progressed. Growth cessation and initial hardening to –15°C were clearly correlated at the provenance level, indicating that growth must cease prior to hardening, and that earlier cessation of growth predicts earlier frost hardening of stems and needles. No differences in frost hardiness of stems were found at the provenance level at the end of the growing period in August. At that time, the frost hardiness of needles of the northernmost provenance was higher than that of other origins. Within the provenance, the stems were less hardy than the needles.

Seasonal variation in the frost hardiness of Scots pine and Norway spruce in old provenance experiments in Finland

Abstract Temperature is one of the major factors regulating the acclimation of forest trees to winter conditions in the boreal zone. The projected climate warming may therefore affect the overwintering pattern of trees. In this study, old provenance trials with Scots pine ( Pinus sylvestris L.) and Norway spruce ( Picea abies (L.) Karst) were used to simulate the effects of a warmer climate on the seasonal variation in frost hardiness. In both species, there were differences between northern and southern provenances in the timing of bud and needle hardening during autumn. The northern provenances hardened earlier. During the dehardening phase in spring, no significant difference was found between the provenances. For buds of Norway spruce, fluctuations in hardiness that coincided rather well with fluctuations in the ambient temperature were observed during mid-winter. The results of this study do not support the theory that climate warming will increase the risk of frost damage in Scots pine and Norway spruce in boreal areas. Further research is recommended.

Analysis of the willow root system by electrical impedance spectroscopy

Information on plant roots is increasingly needed for understanding and managing plants under various environmental conditions, including climate change. Several methods have been developed to study fine roots but they are either destructive or cumbersome, or may not be suitable for studies of fine root functionality. Electrical impedance, resistance, and capacitance have been proposed as possible non-destructive measures for studying roots. Their use is limited by a lack of knowledge concerning the electrical circuit of the system. Electrical impedance spectroscopy (EIS) was used for hydroponically raised willows (Salix schwerinii) to estimate the root system size. The impedance spectra were investigated in three experimental set-ups and the corresponding appropriate lumped models were formulated. The fit of the proposed lumped models with the measured impedance spectra data was good. The model parameters were correlated with the contact area of the roots and/or stems raised in the hydroponic solution. The EIS method proved a useful non-destructive method for assessing root surface area. This work may be considered to be a new methodological contribution to understanding root systems and their functions in a non-destructive manner.

The effect of late summer fertilization on the frost hardening of second-year Scots pine seedlings

In this study the effect of summer fertilization on the initiation of frost hardening of containerized second-year Scots pine (Pinus sylvestris L.) seedlings is studied. During the second growing season three different fertilization programs (water soluble NPK with micronutrients) determined by electrical conductivity of peat water extract (0.2, 0.5 and 1.2 mS cm-1) were initiated. The growth and nutrient concentrations of needles were monitored during the fertilization period. The frost hardiness of seedlings was assessed on four separate occasions at two week intervals from August 7 to September 18. This assessment was based on artificial freezing tests and visual damage scoring of tissue browning on current-year needles. Clear differences in foliar N, P and K concentrations were observed between the fertilization treatments. Fertilization prolonged the growing period of needles and increased root collar diameter. In all the tests, the highest fertilization level resulted in the highest level of frost hardiness. The difference between the fertilization treatments ranged from 1 °C to 2.2 °C. Frost hardiness increased with an increase in foliar nitrogen concentration and slightly less consistently with increases in foliar phosphorus and potassium concentrations.

Estimation of standard error of impedance‐estimated frost resistance

The magnitude of measurement errors of the specific impedance difference was estimated and a formula to approximate the variance of the estimated frost resistance was derived. The measurements of specific impedance difference include the measurement errors of impedance before and after frost treatment and cross-sectional area. These errors in connection with the population variation cause variation in the estimated frost resistance. The frost resistance is estimated by first expressing the specific impedance difference values as a logistic sigmoid function of the treatment temperature, and then evaluating the inverse function at a given value of the specific impedance difference. The error variance between the estimated and the measured frost resistance was calculated using the estimated parameters, their standard deviations and correlations. In an example the impedance estimated frost resistance (LT.i0Qm) was - 10.9°C and its standard deviation 0.8°C.

Autumn colouration of first year Pinus sylvestris seedlings during frost hardening

Autumn colouration, dry matter content and frost hardiness of Pinus sylvestris seedlings were monitored during their first autumn. Seedlings of southern (62° N, 24° E) and northern (67° N, 26° E) Finland origin were grown under two fertilization regimes in Suonenjoki (63°N, 27°E). Northern seedlings turned purple earlier than southern ones, and low fertilized earlier than normal fertilized. Northern seedlings also hardened earlier than southern ones, while fertilization had no effect on hardening. Neither purple autumn colour nor dry matter content was clearly related to frost hardiness of seedlings when treatment means were compared. Comparison of seedlings within treatments suggested, however, that purple seedlings had higher dry matter content than green seedlings at the beginning of hardening period but the relationship between colouring and frost hardiness was not clear.

An appraisal of the electrical resistance method for assessing root surface area

Electrical resistances of roots and stems of hydroponically raised willows (Salix schwerinii) were studied and related to root morphology. Willow cuttings with and without roots were set in a constant electric field (effective voltage of 0.1 V, sine-AC, 128 Hz) in a hydroponic solution. The electrical resistance of different components in the measurement system was measured and analysed in relation to root surface area in contact with the cultivation solution. Axial resistivities of single root segments and of stems were measured. The results showed that the resistance decreased in relation to an increase in the contact surface area of the roots with the solution. The resistance depended strongly on the contact area of the stem with the solution, however, thus causing bias in the evaluation of root surface area. This work is a new contribution for the understanding of current pathways in the root system as exposed to an external electric field and for developing a non-destructive method to study plant roots accordingly. It may be concluded that the electrical resistance method is a useful non-destructive method to study roots and their physiological properties. Electrical analogues for roots and stem comprising resistors are discussed in relation to in situ measurements.

The effects of artificial soil frost on cambial activity and xylem formation in Norway spruce

We studied the effects of artificial soil frost on cambial activity and xylem formation on 47-year-old Norway spruce [Picea abies (L.) Karst.] trees grown on medium fertile site type (with moraine soil) in eastern Finland (62°42′N; 29°45′E). Different soil frost treatments applied were: (1) natural snow accumulation and melting (control, CTRL); (2) artificial removal of snow from soil surface during two consecutive winters (OPEN); and (3) snow clearing and insulation (FROST), which was in other ways similar to OPEN, but the ground was insulated in early spring to delay soil thawing. Each treatment was replicated in three blocks, and two sample trees in each plot were repeatedly microcored during growing seasons of 2006–2007 for the analysis of the onset, cessation and the duration of xylem formation. The phases of tracheid differentiation (tracheids in radial enlargement, secondary cell wall formation, and mature tracheids) were measured from the microcores of 2007. The intra-ring growth and wood density variables were analysed based on X-ray densitometry. In FROST in 2006, xylem formation started a week later than in the other treatments. In 2007, no difference was found between the treatments. The discrepancy in results between the two study years may be explained by between-years variation in weather, i.e., the winter was colder in 2005/2006 than in 2006/2007. No effects of soil frost treatments on tracheid differentiation and on most of the intra-ring growth and density variables were discovered. Our results suggest that the delayed thawing of moraine soil may slightly affect the onset, timing and duration of xylem formation in Norway spruce. However, the effects of delayed soil frost may depend also on the soil type and become more evident with increasing water holding capacity of the soil.

Invading and resident defoliators in a changing climate: cold tolerance and predictions concerning extreme winter cold as a range‐limiting factor

1. Winter temperatures in northern latitudes are predicted to increase markedly as a result of ongoing climate change, thus making the invasion of new insect defoliators possible. The establishment of new outbreak pest species may have major effects on northern ecosystems that are particularly sensitive to disturbances.

Freezing tolerance of ectomycorrhizal fungi in pure culture.

The ability to survive freezing and thawing is a key factor for the existence of life forms in large parts of the world. However, little is known about the freezing tolerance of mycorrhizal fungi and their role in the freezing tolerance of mycorrhizas. Threshold temperatures for the survival of these fungi have not been assessed experimentally. We grew isolates of Suillus luteus, Suillus variegatus, Laccaria laccata, and Hebeloma sp. in liquid culture at room temperature. Subsequently, we exposed samples to a series of temperatures between +5°C and −48°C. Relative electrolyte leakage (REL) and re-growth measurements were used to assess the damage. The REL test indicated that the lethal temperature for 50% of samples (LT50) was between −8.3°C and −13.5°C. However, in the re-growth experiment, all isolates resumed growth after exposure to −8°C and higher temperatures. As many as 64% of L. laccata samples but only 11% in S. variegatus survived −48°C. There was no growth of Hebeloma and S. luteus after exposure to −48°C, but part of their samples survived −30°C. The fungi tolerated lower temperatures than was expected on the basis of earlier studies on fine roots of ectomycorrhizal trees. The most likely freezing tolerance mechanism here is tolerance to apoplastic freezing and the concomitant intracellular dehydration with consequent concentrating of cryoprotectant substances in cells. Studying the properties of fungi in isolation promotes the understanding of the role of the different partners of the mycorrhizal symbiosis in the freezing tolerance.