Effect of maternal soil treatment on first year growth and autumn frost hardiness of Pinus sylvestris L. full‐sib families

After‐effects of maternal soil treatment, controlling for photoperiod and weather conditions, were tested in a nursery experiment with Pinus sylvestris L. seedlings. Three full‐sib crosses were replicated in 4 different soil treatment plots of a grafted experimental seed orchard (63"54'N). Weight of filled seeds, percentage of empty seeds, first year hypocotyl and epicotyl length, and autumn frost hardiness (from artificial freeze testing) were evaluated. Systematic effects from maternal soil treatment were small compared to family differences for all characters studied. However, epicotyl length of the progenies showed a positive relationship with N needle content of the mother grafts. The results also indicated an important interaction between maternal soil treatment and autumn frost hardiness of the families. It is suggested that the altered mother tree treatment could lead to a change in progeny performance, corresponding to a 1.5 degree difference in latitudinal origin of natural stand seedlings. The results demonstrate the advantages of providing the mother tree with uniform conditions in early progeny testing.     

Morphological and physiological differences in Scots pine seedlings of six seed origins

One-year-old seedlings of Scots pine (Pinus sylvestris L.) offour native seed origins (Loch Maree Islands, Glengarry/GlenMorriston, Glen Affric and Abernethy), a commercial Britishseedlot, and a seedlot from Hedesunda, in middle Sweden, werecompared at monthly intervals from October 1993 to April 1994.Seedling morphology, root condition, root frost hardiness andbud dry matter were determined at each date. There were clear morphological differences among seed origins.Seedlings raised from the commercial seedlot (A70) were largerbut had a poorer root:shoot ratio than the other seed origins.Of the native pines tested, the Loch Maree Islands origin allocateda larger proportion of its photosynthate to fine roots and needlesand smaller proportion to woody structures. Seedlings raised from the commercial British seedlot tendedto have poorer bud lignification than the other origins andalso, in autumn, higher electrolyte leakage rates from its fineroots. During winter, the Swedish origin had the lowest fineroot electrolyte leakage. Seedlings of all origins showed aprogressive increase in fine root hardiness towards mid-winterwith maximum hardiness (–7°C) in January. Dehardeningoccurred over subsequent months reaching –3°C in April.Differences among origins were evident. The Swedish seedlotdeveloped greater frost resistance than the other origins, hardeningbegan earlier in autumn and dehardening began later in spring.The commercial seedlot hardened later than the other originsbut reached a similar level of frost hardiness by January. Ofthe native pines, seedlings of the Loch Maree Islands originwere slowest to develop root hardiness.     

Characterizing the frost sensitivity of black spruce photosynthesis during cold acclimation

We used photosynthetic light response curves to measure and model the responses of two provenances of 3-year-old black spruce (Picea mariana (Mill.) BSP) seedlings to severe artificial frost treatments applied at 2-week intervals during cold acclimation. Black spruce seedlings responded to cold acclimation with long-term suppression of photosynthetic capacity (Amax) and apparent quantum-use efficiency (alpha'). Short-term reductions in both photosynthetic parameters following frost treatments were dependent on the extent of cold acclimation of the seedlings and the severity of the frost treatments. Large reductions in Amax in response to the frost treatments were observed in seedlings that had undergone little cold acclimation and these reductions were associated with an irreversible reduction in alpha'. Such seedlings recovered only partially during the subsequent 23 days, whereas seedlings in most other treatments showed complete recovery of Amax after 13 days. The impact of frost treatments on Amax and alpha' did not vary with seedling provenance. We propose an algorithm that predicts the combined effects of cold acclimation and severe freezing temperatures on the extent of the suppression of A(max) during autumn. The algorithm is based on (1) the maximum Amax observed during the growing season, (2) the accumulation of cold degree-days, based on a minimum nocturnal temperature < 5 degrees C, and (3) the severity of freezing temperatures during autumn. The parameters developed in the algorithm showed that cold acclimation of black spruce seedlings had a greater impact on the reduction of Amax in autumn than did the severe frost treatments. Mean Amax of seedlings subjected to artificial frosts showed a strong correlation with values predicted by the algorithm (r2 = 0.91).     

Assessment of seedling storability of Quercus robur and Pinus sylvestris

Pedunculate oak (Quercus robur L.) and Scots pine (Pinus sylvestris L.) seedlings were lifted on several occasions during autumn 1997 to determine the relationships between storability and frost hardiness. On each lifting date their physiological status was determined by assessment of shoot and root electrolyte leakage and frost hardiness, assessed as freeze-induced electrolyte leakage. Additional seedlings were simultaneously cold-stored for field planting and assessment of preplanting root growth potential in April 1998. First year field performance was determined the following winter. Storability and cold acclimation patterns differed between the two species. Both were negatively affected by early lifting, but oak was less sensitive with respect to survival, and pine attained tolerance to cold storage more rapidly and earlier with respect to growth increment. The correlations between shoot frost hardiness and performance suggest that freeze-induced shoot electrolyte leakage (SEL_diff?20) below a threshold of 5% is a good storability predictor for Scots pine in Denmark. A completely reliable criterion for pedunculate oak could not be established.     

No difference in frost hardiness between high and low altitude Pinus sylvestris (L.) offspring

Seedlings representing offspring from 46 Swedish natural stands of Pinus sylvestris (L.) from latitude 63° to 68° and altitude 75 to 675 m were artifically frost hardened and tested for autumn frost hardiness using artifical freeze testing in a programmable freezing chamber. A clinal variation in frost hardiness was observed over latitude. Altitude had no effect on the frost hardiness. The results are discussed in relation to the reproduction‐ and migration biology of P. sylvestris.     

Chlorophyll fluorescence and CO(2) assimilation of black spruce seedlings following frost in different temperature and light conditions

Effects of artificial frosts on light-saturated photosynthesis (A(max)) and ground, maximal and variable fluorescence variables (F(o), F(m), and F(v) and F(v)/F(m)) were monitored on 1-year-old foliage of black spruce seedlings (Picea mariana (Mill.) BSP) grown at high (25 degrees C), moderate (15 degrees C) and low (5 degrees C) temperatures and moderate (240 &mgr;mol m(-2) s(-1)) and low (80 &mgr;mol m(-2) s(-1)) irradiances. Photoinhibition of 1-year-old foliage was greater in seedlings grown in moderate light than in seedlings grown in low light. Photoinhibition increased with decreasing growth chamber temperature at both irradiances. Most changes in F(v)/F(m) were caused by changes in F(v). Exposure to -4 degrees C decreased both F(v)/F(m) and A(max) compared with control values. The effect of the -4 degrees C frost treatment was greater in seedlings grown in low light than in seedlings grown in moderate light, probably because seedlings grown in moderate light were already partially photoinhibited before the frost treatment. Following -4 degrees C treatment, neither F(v)/F(m) nor A(max) recovered in seedlings grown in low light. Light-saturated photosynthesis decreased with decreasing growth chamber temperature. Light-saturated photosynthesis was more sensitive to the -3 and -4 degrees C frost treatments in seedlings grown at 25 degrees C than in seedlings grown at 15 and 5 degrees C. The A(max) of seedlings grown at 15 degrees C was sensitive only to the -4 degrees C frost treatment, whereas A(max) of seedlings grown at 5 degrees C was not sensitive to any of the frost treatments. Recovery of A(max) following frost took longer in seedlings grown at high temperatures than in seedlings grown at low temperatures. For seedlings grown at the same temperature but under different irradiances, both A(max) and F(v)/F(m) reflected damage to the photosynthetic system following a moderate frost. However, for seedlings grown at the same irradiance but different temperatures, A(max) provided a more sensitive indicator of frost damage to the photosynthetic system than F(v)/F(m) ratio.     

Timing of cold temperature exposure affects root and shoot frost hardiness of Picea Mariana container seedlings

Seventeen‐week‐old black spruce seedlings were hardened under short daylengths and one of three short day length environments, which were either warm (24/16°C, day/night) throughout a 10 week hardening period (WW), cool (10/5°C) throughout hardening (CC), or warm for three weeks followed by seven weeks of cool temperatures (WC). Greatest root and shoot frost hardiness resulted from the exposure of seedlings to three weeks of warm followed by seven weeks of cool temperatures. Seedlings receiving warm temperatures throughout hardening increased in root and shoot frost hardiness, but to a lesser extent than seedlings exposed to cool temperatures. The frost hardiness of woody roots was generally greater than that of fine roots, but the extent of the difference in frost hardiness depended on the time since bud initiation and on the hardening treatment.     

Assimilation and stomatal conductance responses of red spruce to midwinter frosts and the constituent ions of acid mist

Red spruce (Picea rubens Sarg.) seedlings growing outside in open-top chambers were sprayed twice weekly with artificial mists at either pH 2.5 or 5.6, for five months during the 1988 growing season. The mists contained one of the following: water, pH 5.6 (control); (NH(4))(2)SO(4), pH 5.6; NH(4)NO(3), pH 5.6; HNO(3), pH 2.5; H(2)SO(4), pH 2.5; or (NH(4))(2)SO(4) + NH(4)NO(3), pH 2.5. During January 1989, the light responses of assimilation and stomatal conductance were assessed in the laboratory following a 4-day equilibration at 12 degrees C. The aerial portions of the intact trees were then subjected to a mild (-10 degrees C) frost for three hours during the night and the rate of recovery of light-saturated assimilation (A(max)) was determined the following day using the same branches as were used for the assimilation studies before the frost treatment. The same trees were then subjected to a second frost of -18 degrees C for three hours during the following night and the recovery of A(max) of the same branches was measured the next day. All of the acid mist treatments increased A(max) and apparent quantum yield relative to the control treatment when measured before the frost treatments. Frosts of -10 and -18 degrees C resulted in a significant decline in A(max) of seedlings in all treatments except the control. Stomatal conductance increased with increasing irradiance in seedlings in the acid mist treatments that did not contain SO(4) (2-) ion. Stomatal conductance of seedlings in acid mist treatments containing SO(4) (2-) ion was insensitive to changes in irradiance over the range 50-1500 micro mol m(-2) s(-1). It is concluded that acid precipitation increased the sensitivity of the assimilation response to midwinter frosts that follow a brief warm period. The SO(4) (2-) ion appears to be significant in causing increased sensitivity to frost and in causing stomatal insensitivity to light flux density.     

Phenotypic Differences Between Natural and Selected Populations of Picea Abies. II. Apical Mitotic Activity and Growth Related Parameters

Apical mitotic index (MI) and growth of non-juvenile Norway spruce [Picea abies (L.) Karst.] populations growing in northern Sweden (63°54' N) were monitored in 1996. Annual leader shoot lengths and shoot growth components for the period 1990-1997 were measured in 1997. In 1997 populations transferred more than approximately 3° in latitude were found to be shorter than local populations. MI levels were initially high in all populations in mid-April. In spring, populations originating north of 63°30' N showed higher MI levels, and started shoot growth earlier, than populations originating further south. In autumn, MI levels were higher in populations originating south of 63°30' N than in populations originating further north and higher in seed orchard populations than in natural populations of similar latitudinal origin. The number of stem-units (NSU) was correlated with tree height and leader shoot length. NSU and elongation of stem-units appeared to be primarily influenced by summer temperature. Prolonged MI activity in autumn did not result in high NSU but appeared to be associated with a higher risk of frost damage to buds. The results are discussed in relation to climatic factors, seed transfer, selection and inherent growth rhythms.     

Variation in spring and autumn frost tolerance among provenances of Russian larches (Larix Mill.)

Abstract

Spring and autumn frost tolerance was measured using material from a range-wide (50–67° N, 38–158° E) provenance trial of four Russian larch species (Larix sukaczewii Dyl., L. sibirica Ledeb., L. gmelinii Rupr. and L. cajanderi Mayr.) growing in northern Sweden. Shoots were collected in early May and late September and frozen at ?8, ?12, ?16 and ?20°C. Cambial damage was assessed visually after development under ideal conditions for 2 weeks. Differences in frost damage among provenances were highly significant in both spring and autumn. Autumn frost damage was significantly correlated with provenance latitude and longitude and spring frost damage was significantly correlated with provenance longitude but not latitude. Frost damage was not correlated with provenance elevation. North-western provenances showed the least damage and far-eastern provenances the greatest damage in both spring and autumn. A possible explanation for less spring frost damage to north-western provenances is adaptation to maritime conditions in proximity to the Barents Sea, which is often ice free in late winter. This would counteract early loss of frost tolerance and bud flushing if warm spells occurred in late winter. North-eastern Siberian provenances did not show similar adaptation and may exhibit increased spring frost damage if global warming eventually results in the Arctic Ocean north of Siberia becoming ice free in late winter.     

Effects of Seed Weight and Seed Type on Early Seedling Growth of Pinus sylvestris under Harsh and Optimal Conditions

The effects of seed weight and seed type on seedling growth of Pinus sylvestris (L.) were studied by seeding individually weighed orchard and stand seed in different mixtures under harsh (direct seeding in field) and optimal (seeding in nursery) conditions. In the nursery experiment an increase in the seed weight from 3 to 7 mg increased the seedling height by 10-27% and total weight by 27-113%, and decreased the height/diameter ratio by 5-6% after 2 yrs. With elimination of seed weight effects, orchard seedlings were 2% taller than stand seedlings in year 2. Without elimination of seed weight effects, orchard seedlings were 7-13% taller. In the field experiment an increase in the seed weight from 3 to 7 mg increased seedling height by 18-65%, stem volume by 81-274% and the number of top-buds by 23-34% in year 5. After elimination of seed weight effects, orchard seedlings were 7-13% taller than stand seedlings and without elimination of seed weight effects 20-21% taller after 5 yrs. Even after elimination of both seed weight and genetic effects orchard seedlings were 3-9% larger than stand seedlings in the field experiment. In conclusion, the influence of seed weight and seed type on growth traits and slenderness is highly significant and the influence seems to be greater in harsh conditions.     

黄竹细胞悬浮培养和原生质体分离*

以牡竹属黄竹（ＤｅｎｄｒｏｃａｌａｍｕｓｍｅｍｂｒａｎｃｅｕｓＭｕｎｒｏ）的竹节及无菌苗根颈为外植体，培养于含有５ｍｇ／Ｌ２，４－Ｄ、０．２ｍｇ／ＬＫＴ和２００ｍｇ／ＬＬＨ的ＭＳ培养基上诱导愈伤组织，然后移至相同成份或不同浓度２，４－Ｄ的液体培养基中，进行悬浮培养，以建立分散性良好的悬浮细胞系。利用悬浮细胞和无菌苗叶片经酶解获大量原生质体，其产量分别约为每克鲜重２．５×１０５个和５×１０５个，活力均可达８０％。     

Frost tolerance of two-year-old Picea glauca seedlings grown under different irrigation regimes in a forest nursery

Abstract

This study examined the impact of increased irrigation efficiency on the hardening and frost tolerance of 2-year-old containerized white spruce seedlings in the context of groundwater protection, irrigation management and the maintenance of seedling quality in northern climates. The seedlings were grown under three different irrigation regimes (IR =30%, 40% and 55% v/v; cm³ H₂O/cm³ substrate) and were hardened under conditions of natural photoperiod and temperature. After being subjected to artificial frost tests on four sampling dates during autumn, the seedlings were compared for bud development and frost tolerance. IR had no influence on frost tolerance as determined by measurements of physiological (electrolyte leakage, root water loss) and morphological (shoot damage, root initiation) variables. At the end of the second growing season, there was no significant difference between IRs in seedling height, root collar diameter, shoot dry mass and root dry mass. The results indicate that the amount of water applied to large-dimension 2-year-old white spruce seedlings during the growing season can be significantly decreased without prematurely impeding their growth or hindering their acquisition of frost tolerance.     

Temperature control of the development of frost hardiness in two populations of Leptospermum scoparium

Seedlings of Leptospermum scoparium J.R. et G. Forst (manuka) originating from seed from a low altitude coastal site (Auckland) and from a high altitude inland site (Desert Road) were grown for 96 days in four controlled environments to compare the relationship between growth temperature and frost hardening. Day/night temperature treatments were 12/6, 12/3, 12/0 and 12/-3 degrees C. Frost hardiness was determined at 14-day intervals by exposing whole seedlings to temperatures ranging from -2 to -8 degrees C. Frost damage differed significantly between the two populations: Desert Road seedlings were less affected than Auckland seedlings. At all growth temperatures, the time courses of frost hardiness of both populations followed curvilinear relationships reaching a maximum hardiness at about Day 50, after which the seedlings spontaneously dehardened. The rate of frost hardening increased linearly with decreasing temperature from 6 to 0 degrees C, but thereafter, no further increase occurred with decreasing temperature to -3 degrees C. The frost hardening process was more sensitive to temperature in the Desert Road seedlings than in the Auckland seedlings, and this difference may account for the intraspecific variation in frost hardening capacity of this species. Comparisons with Pinus radiata D. Don and Lolium perenne L. indicated that interspecific variation in frost hardening capacity can also be accounted for by differences in the sensitivity of the hardening process to temperature.     

Fungicidal Control of Lophodermium seditiosum on Pinus sylvestris Seedlings in Swedish Forest Nurseries

During the 1990s, there were serious outbreaks of the pathogen Lophodermium seditiosum on pine seedlings in Swedish forest nurseries, even though the seedlings had been treated with the fungicide propiconazole. The present experiment was carried out to evaluate two other fungicides, fluazinam and azoxystrobin, as possible alternatives to propiconazole. In the tests, which were all carried out in the same forest nursery, seedlings were treated with either propiconazole, fluazinam or azoxystrobin, and the proportion of needles with ascocarps of L. seditiosum and the number of ascocarps per needle were recorded over the following 2 yrs. Seedlings treated with azoxystrobin already appeared healthier than control seedlings in September of the first year, and by November all azoxystrobin-treated seedlings had fewer ascocarps per needle compared with control seedlings. In autumn of the second year, there were no ascocarps on seedlings treated with fluazinam or azoxystrobin, whereas seedlings treated with propiconazole had similar numbers of ascocarps to non-treated control seedlings.     

Early season short-day treatment did not affect the field performance of Pinus sylvestris container seedlings in summer planting

Abstract

When Scots pine (Pinus sylvestris L.) seedlings are short-day (SD) treated in May or early June, height growth ceases but seedlings reflush within a few weeks and grow secondary needles (later referred to as early SD treatment). These “2-year-old-like” seedlings have been thought to be more stress tolerant than traditional 1-year-old pine seedlings, and thus suitable for summer and autumn plantings. To test this, seedlings sown in April were early SD treated over a 3-week period from the beginning of June and the seedlings were planted at 10-day intervals from the beginning of July until the end of September, and also during the following spring. Neither survival nor height growth of early SD-treated seedlings differed from untreated (1-year-old) seedlings. Moreover, for both early SD-treated seedlings and 1-year-old seedlings, those planted in summer grew and survived better after planting but had more multiple leaders than seedlings planted in autumn or spring.     

Frost damage to planted Norway spruce seedlings — influence of site preparation and seedling type

Damage to Norway spruce (Picea abies (L.) Karst.) seedlings by summer frost was monitored on 10 clear-cuttings of various ages to which slash removal and four types of site preparation had been applied. The clear-cuttings were established on two sites (5 on each) in southern Sweden from 1989 through 1993 and planted each year from the year of cutting until 1993. In total, 7680 seedlings were analysed for frost injuries during the first three years following planting. The site preparation methods evaluated were: application of herbicide to ground vegetation; mowing of ground vegetation; scarification (mounding); and control. Two seedling types, bare-rooted and containerised seedlings, were evaluated.Periods of frost susceptibility were calculated using data on bud and shoot development during 1997 together with weather data for each of the studied years and sites. The date of flushing was well correlated to the air temperature sum (+5°C threshold value) in spring. Frost damage varied considerably between sites and study years. The lowest measured minimum air temperature and the frost-day sum during the frost-susceptible period were correlated to the frequency of frost damage. Soil scarification reduced frost damage in the first growing season. Bare-rooted seedlings had a considerably lower frequency of frost injury compared with containerised ones, possibly because flushing of the former was delayed. There was no statistically significant effect of herbicide application, mowing or slash removal on the frequency of frost damage. Seedling growth was significantly reduced by frost injury, especially in cases where injuries were sustained during several years. Survival was only slightly affected by frost damage.     

Autumn Frost Damage on Young Picea sitchensis 2. Shoot Frost Hardening, and the Probability of Frost Damage in Scotland

The natural increase in frost hardiness of detached shoots ofPicea sltchensis during August to November was measured usinga programmable freezing chamber. Oregon, Queen Charlotte Islandsand Alaskan provenances were compared, and the effects on hardeningof long days, warm temperatures and frosts were determined.A computer model was constructed to mimic the observed patternsof autumn frost hardening, as functions of air minimum temperatures,daylengths and the occurrence of frosts. The model was used(a) to describe the pattern of autumn frost hardening at differentsites in northern Britain, using past meteorological records,and hence (b) to determine when frosts occurred that might havedamaged young trees. The model accurately predicted known instancesof autumn frost damage at Kirroughtree and Carnwath. The predicted probability of autumn frost damage on young treesof P. sitchensis in upland areas of Scotland was much lowerthan that previously predicted for spring frost damage. Theestimated return time for autumn frost damage to an Oregon provenanceat Eskdalemuir was 8.3 years, and the return time for a Q.C.I.provenance was longer than 10 years. Most damaging frosts occurredin October, but frosts like those on 13–15 October 1971,which followed warm weather and caused wide spread damage inScotland, have been quite rare. Alaskan provenances would rarelybe damaged by autumn frosts, nor would trees of Q.C.I. provenancegrowing in lowland areas of Scotland, or at Masset on the QueenCharlotte Islands.     

Ice nucleation and frost resistance of Pinus canariensis seedlings bearing needles in three different developmental states

Frost resistance and ice formation in different developmental states of needles of P. canariensis seedlings were assessed. Regrowth after frost damage was used to determine the overall frost survival capacity. Two distinct freezing exotherms (E1, E2) were registered. E1 was between ?1.7 and ?2.0 °C. Initial frost damage (LT₁₀) was 1.5–2.7 °C below E1. E2 was between ?5.6 and ?6.0 °C, and either corresponded with LT₅₀ or occurred in between LT₁₀ and LT₅₀. Current year needles were less frost resistant than 1-year-old needles. The overall recuperation capacity of seedlings revealed that frost survival may be underestimated when only needle damage is assessed. Freezing of seedlings with or without roots had no effect on the frost resistance of needles but recuperation capacity was significantly affected. Seedlings survived ?10 °C during summer indicating that they withstand the lowest naturally occurring frosts in Tenerife.     

Influence of Stem Diameter on the Survival and Growth of Containerized Norway Spruce Seedlings Attacked by Pine Weevils (Hylobius spp.)

Pine weevils (Hylobius spp.) feeding on stem bark of young conifer seedlings pose a serious threat to forest regeneration-planting programmes in Nordic countries. This study was designed to determine the threshold diameter for planted, untreated containerized seedlings, above which pine weevils cause little or no damage. The effects of sublethal weevil damage on seedling growth were also assessed. In total, 5320 containerized spruce seedlings were planted on scarified and unscarified plots on three sites in southern Sweden. Seedlings in six size classes, which differed with regard to age (1.5-3.5 yrs) and cultivation density (28-446 seedlings m 2) were grown using the Combicell system. None of the seedlings was treated with insecticides, except for those in the smallest class, where both untreated and treated seedlings were used. Inspections were made periodically during the first 3 yrs and after both 5 and 7 yrs. A statistically significant relationship was found between seedling losses due to pine weevil attack and seedling stem-base diameter at the time of planting out, on both scarified and unscarified plots. For seedlings with a stem-base diameter of around 10 mm, mortality due to pine weevil attack on scarified plots was low enough to be considered negligible. This threshold diameter was several millimetres greater for seedlings planted on unscarified plots. An analysis of the relationship between the extent of weevil damage and seedling growth rate showed that among surviving seedlings, those that grew fast tended to show low levels of damage. On unscarified plots, the mortality rate amongst seedlings treated once with a permethrin insecticide was only one-third that of untreated seedlings. On scarified plots, the corresponding difference was somewhat smaller. Repeated insecticide treatment resulted in a pronounced reduction in seedling mortality on the unscarified plots, whereas the effect was weaker on scarified plots.