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.     

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.     

Use of short-day treatment in the production of Norway spruce mini-plug seedlings under plant factory conditions

Mini-plug transplant seedlings of Norway spruce have been cultivated in closed growth systems, so-called plant factories, for few years. The aim of the experiment was to define a short-day treatment (SD) that harden seedlings to sustain 3 months of cold storage, but does not have adverse effects on growth, morphology, and vitality. The seedlings were subjected to one of the following treatments: (1) 12?h photoperiod + 3 weeks duration; (2) 8?h photoperiod + 3?wk duration; (3) 12?h photoperiod + 5?wk duration; and (4) 8?h photoperiod + 5?wk SD. All the SD treatments yielded healthy seedlings that grew well after the cold storage. The frost hardiness of the seedlings improved when the photoperiod was reduced from 12 to 8?h, and when the SD duration was increased from 3 weeks to 5 weeks, but reducing the photoperiod from 12?h to 8?h caused growth reductions. The root and shoot regrowth after cold storage was highest in seedlings that had received 12?h photoperiod and 5?wk duration. However, 12?h photoperiod and 3?wk duration may be an adequate practice for nurseries that treat multiple crops in their SD facilities.     

Effects of nitrogen fertilization and temperature on frost hardiness of Aleppo pine (Pinus halepensis Mill.) seedlings assessed by chlorophyll fluorescence

In a 14-week study, 1-year-old Aleppo pine seedlings were grownin two growth chambers. Seedlings were artificially hardenedby decreasing photoperiod and temperature. In each chamber halfof the seedlings were fertilized with nitrogen (8.4 mg seedling^–1).In order to determine the relative importance of the hardeningenvironment versus fertilization, each chamber was programmedto decrease night temperatures down to a low of 8 or 4°C.Chlorophyll fluorescence and frost hardiness was measured fivetimes during the experiment. A sample of seedlings from eachtreatment was exposed to an artificial frost at –5°Cand the freezing effects were assessed by measurements of chlorophyllfluorescence and visual evaluation of needle damage. Seedlingsincreased their frost hardiness during the experiment in allthe treatments but the ratio of variable to maximal chlorophyllfluorescence (F_v/F_m) measured before freezing did not vary duringthe experiment. This indicates that Aleppo pine maintains itsphotosynthetic ability during hardening in contrast to otherconiferous species from colder climates. The effect of nitrogenfertilization on frost hardiness was small in comparison withchamber effect. Nitrogen fertilization slightly delayed theacquisition of hardening in the coldest chamber. Seedlings inthe warmest chamber did not become fully resistant to –5°C,but in the coldest chamber, where night temperature reached4°C, all the seedlings were resistant to the frost. Severedamage caused by frost could be related to a rapid rise of minimalfluorescence (F₀) but the best index of damage was the dropof F_v/F_m after freezing.     

Frost hardening of Pinus radiata seedlings: effects of temperature on relative growth rate,carbon balance and carbohydrate concentration

Pinus radiata (D. Don) seedlings were grown for 100 days at day/night temperatures of 10/1, 15/1, 20/1 and 25/1 degrees C, to determine whether temperatures above a threshold of 5 degrees C influence frost hardiness development. Relationships between hardening and relative growth rate, carbohydrate concentration and net carbon balance were also investigated. Seedlings hardened at a nearly constant rate in each treatment, although the rate of hardening was strongly temperature dependent. It increased as the temperature declined, but in a curvilinear fashion. Temperatures below 9.5 degrees C were effective in hardening the seedlings. During the daily temperature cycle, dehardening occurred at temperatures above the threshold, whereas hardening occurred at temperatures below the threshold. The net difference between the two processes determined the development of frost hardiness. The development of frost hardiness was negatively correlated with relative growth rate and positively correlated with the accumulation of starch and sugars. We conclude that frost hardening is a complex process that is causally linked to carbohydrate concentrations.     

Effects of early long-night treatment on diameter and height growth,second flush and frost tolerance in two-year-old Picea abies container seedlings

Abstract

The object of this study was to obtain Norway spruce seedlings with buds set, ready for summer planting from 1 July. An early long-night treatment prevented flushing of the newly formed terminal buds and ceased height growth, but slightly reduced hardiness in buds and needles. Nevertheless, a sufficient hardiness level in the autumn was acquired at a Norwegian nursery at 59°46′ N, with plants of the local provenance given a long-night treatment (14 h) for 13 days from 25 June. Similar treatment at a nursery at 64°30′ N did not give the same result; all treatments led to a second flush with resumed growth of the local provenance. A trial with seed lots from several provenances was therefore performed at this nursery, and a significant correlation was found between the critical night length of the seed lot and their ability to produce non-flushing buds; the longer the critical night length of the seed lot, the fewer non-flushing buds. Responses at the northern nursery are probably due to the lack of a dark period after termination of the treatment, and too short a treatment period to attain bud dormancy. An early and successful long-night treatment will also produce shorter seedlings with a larger root collar diameter.     

Aftereffects of maternal environment on autumn frost hardiness in Pinus sylvestris seedlings in relation to cultivation techniques

Full-sib families of Pinus sylvestris (L.) from genetically identical parent clones, grown at three geographic locations (64, 59.5, and 56 degrees N), were tested for autumn frost hardiness. At one of the locations (64 degrees N), maternal clones were grown in plots subjected to different site treatments. Progenies raised in a heated greenhouse with additional artificial light were induced to undergo two shoot elongation periods, separated by a period with long nights and non-hardening temperatures, followed by a hardening period. Frost hardiness was tested by exposing seedlings to -17, -19, or -21 degrees C during the hardening period. Progenies produced at the northernmost locality (64 degrees N) were the most hardy. The hardiness ranking among localities and the magnitude of differences in hardiness were in accordance with earlier results obtained from the same crosses that had undergone only one shoot elongation period. Effects of maternal site treatments were significant but small compared with family differences. The growth regimes used in the present study did not eliminate maternal effects induced by geographic location, but maternal effects related to site treatment decreased substantially in relation to family variation when compared with seedlings that had undergone only a single shoot elongation period. The results are discussed in relation to other studies of the same crosses raised under different growth conditions.     

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.     

Frost hardening spruce container stock for overwintering in Ontario

Difficulties overwintering container stock in northern Ontario led to the development of the extended greenhouse culture hardening regime for spruce seedlings. Laboratories to measure shoot frost hardiness and evaluate terminal bud development were established to monitor nursery crops being hardened using this regime. Information on frost hardiness and bud development provided by these laboratories has been used by nursery managers to determine readiness of container seedlings for overwintering. Since 1982, over 200 stock lots have been monitored by these operational laboratories. This database can be used to determine the importance of nursery cultural factors and seed source on frost hardening. The database shows large differences between nurseries in approach to hardening seedlings which were reflected in levels of freezing damage, winter desiccation and overwintering success. Rates of frost hardening (i.e., the interval between terminal bud initiation and attainment of a --15°C level of shoot frost hardiness) of crops produced in north central Ontario failed to show significant seed source effects. The rate of frost hardening was faster in crops producing fewer needle primordia in terminal buds.     

Frost Hardiness of Red Alder (Alnus rubra) Provenances in Britain

The phenology and frost hardiness of shoots of 15 provenancesof Alnus rubra growing in Scotland were measured over one autumn,winter and spring. Dates of budset (in September) and the onsetof rapid frost hardening (in October-November) occurred about2 days earlier for each degree latitude of origin northwards,except for an Idaho provenance. However, all provenances dehardenedat about the same time in March and burst their buds between8 and 14 April. Assuming that rapid frost hardening in the autumnwas triggered primarily by shortening daylengths, Alaskan provenancesof A. rubra seemed better adapted to British conditions thansouthern British Columbian provenances, which have been mostcommonly planted. However, even Alaskan provenances are proneto spring frost damage. Scottish A. glutinosa and Alaskan A.sinuata set buds and frost hardened 1–2 weeks before eventhe Alaskan A. rubra, and burst their buds 2–3 weeks laterin April-May. All three species were hardy to below –30°Cfrom December to mid-March.     

Phenotypic Differences Between Natural and Selected Populations of Picea Abies. I. Frost Hardiness

The frost hardiness of non-juvenile Norway spruce [Picea abies (L.) Karst.] populations growing in northern Sweden (63°54' N) was monitored during 1996-1997. The investigated progenies originated from 12 natural populations and six seed orchards located between 58° N and 68° N in Sweden. Frost hardiness of needles was assessed by measuring chlorophyll fluorescence and electrolyte leakage after freezing. The loss of frost hardiness in 1-yr-old needles during spring occurred slightly earlier in populations originating north of 63°30' N than in those originating further south. Dehardening was slightly delayed in selected populations compared with natural populations of similar origin. The level of frost hardiness during autumn was higher in populations originating north of 63°30' N than in those originating south of this latitude, but there were no clear differences in frost hardiness between selected and natural populations of similar origin. The results are discussed in relation to climatic factors and inherent growth rhythms.     

华山松抗寒性的地理变异

本文报道了1980－1988年间26个产地的华山松实生苗和幼树的冻害情况，看到原产云贵高原亚热带山区的华山松，引种到暖温带气候区种植极不抗寒，基本不能越冬，云贵高原区域内种源的抗寒性也有某种程度的差异，尤以西部（云南保山地区）最弱，东部和北部（云南北部、四川西南部和贵州西部）较强，因而不经试验，不宜把云贵高原华山松种子调入暖温带使用。     

Frost hardening in first-year eastern larch (Larix laricina) container seedlings

Eastern larch (Larix laricina [du Roi] K. Koch) container seedlings were tested to determine shoot frost hardiness development under short or long days and warm (15 to 25 °C) or cool (10/5 °C, day/night) temperatures, to aid in the development of greenhouse hardening strategies. Seedlings were sampled sequentially over time (25 seedlings per week) from a population of 1000 trees. Frost hardiness increased significantly after one week of fluctuated over the next 6 weeks, and increased thereafter through week 14. Seven weeks of warm, intermittent short days, followed by 6 weeks of cool, continuous short days, resulted in greater frost hardiness than 13 weeks of warm, intermittent short days. In contrast, seedlings exposed to 7 weeks of warm, intermittent short days, followed by six weeks of warm, long days were significantly less frost hardy. Stems with needles attached had lower Index of Injury than stems without needles.     

Effects of elevated CO(2) and temperature on cold hardiness and spring bud burst and growth in Douglas-fir (Pseudotsuga menziesii)

We examined effects of elevated CO(2) and temperature on cold hardiness and bud burst of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings. Two-year-old seedlings were grown for 2.5 years in semi-closed, sunlit chambers at either ambient or elevated (ambient + ~ 4 degrees C) air temperature in the presence of an ambient or elevated (ambient + ~ 200 ppm) CO(2) concentration. The elevated temperature treatment delayed needle cold hardening in the autumn and slowed dehardening in the spring. At maximum hardiness, trees in the elevated temperature treatment were less hardy by about 7 degrees C than trees in the ambient temperature treatment. In general, trees exposed to elevated CO(2) were slightly less hardy during hardening and dehardening than trees exposed to ambient CO(2). For trees in the elevated temperature treatments, date to 30% burst of branch terminal buds was advanced by about 6 and 15 days in the presence of elevated CO(2) and ambient CO(2), respectively. After bud burst started, however, the rate of increase in % bud burst was slower in the elevated temperature treatments than in the ambient temperature treatments. Time of bud burst was more synchronous and bud burst was completed within a shorter period in trees at ambient temperature (with and without elevated CO(2)) than in trees at elevated temperature. Exposure to elevated temperature reduced final % bud burst of both leader and branch terminal buds and reduced growth of the leader shoot. We conclude that climatic warming will influence the physiological processes of dormancy and cold hardiness development in Douglas-fir growing in the relatively mild temperate region of western Oregon, reducing bud burst and shoot growth.     

短日照处理在夏季造林中的应用及研究进展

北欧、北美的许多地区在夏季造林前对云杉苗木进行2 ~ 3周的短日照处理,以促进苗木提前形成顶芽和进入休眠,提高苗木的木质化程度,增强苗木抗机械损伤和抵御生物或非生物胁迫的能力;通过抑制苗木的高生长,使碳水化合物更多地向根系分配,可促进根系的生长发育,增强苗木吸收水分、养分的能力,显著改善夏季造林苗木的质量,取得了较好的造林效果。目前,应用短日照处理调控夏季造林苗木质量的相关研究受到广泛关注。文中重点分析短日照处理的起始时间、持续时间和日照长度等主要技术参数对苗木木质化及其抗旱性和抗寒性的影响,并对短日照处理技术在我国夏季造林中的应用提出展望。     

Short-day treatment during the growing period limits shoot growth and increases frost hardiness of hybrid aspen plants in the nursery

In Finland, under nursery conditions hybrid aspen may continue their shoot growth until early September. Thus, frost hardening is usually delayed. To solve this problem, we used a three-week period of short-day (SD) treatment between late July and mid-August. During autumn after frost exposure, frost hardiness (FH) was assessed three times with a stem-browning test. The re-sults showed that after SD treatment shoot growth ceased and FH increased when compared to untreated hybrid aspen. Furthermore, the height of SD-treated hybrid aspen varied much less than that of the control plants. We conclude that SD treatment in the nursery during the growing period can be used as a supplementary method for producing well-hardened and uniform hybrid aspen plants.     

Autumn frost hardiness in Norway spruce plus tree progeny and trees of the local and transferred provenances in central Sweden

Reforestation with provenances from locations remote from the planting site (transferred provenances) or the progeny of trees of local provenances selected for superior form and vigor (plus trees) offer alternative means to increase yield over that obtained by the use of seed from unselected trees of the local provenance. Under Swedish conditions, Norway spruce (Picea abies (L.) Karst.) of certain transferred provenances generally has an advantage in productivity relative to the local provenance comparable to that of progeny of plus trees. The aim of this study was to explore the extent to which productivity gains achieved by provenance transfer or the use of plus tree progeny are associated with reductions in autumn frost hardiness, relative to that of trees of the local provenance. In a field trial with 19-year-old trees in central Sweden, bud hardiness was tested on four occasions during the autumn of 2002. Trees of the local provenance were compared with trees of a south Swedish provenance originating 3 degrees of latitude to the south, a Belarusian provenance and the progeny of plus trees of local origin. The Belarusian provenance was the least hardy and the local provenance the most hardy, with plus tree progeny and the south Swedish provenance being intermediate in hardiness. Both the Belarusian provenance and the plus tree progeny were significantly taller than trees of the other populations. Within provenances, tree height was negatively correlated with autumn frost hardiness. Among the plus tree progeny, however, no such correlation between tree height and autumn frost hardiness was found. It is concluded that although the gain in productivity achieved by provenance transfer from Belarus was comparable to that achieved by using the progeny of plus trees of the local provenance, the use of trees of the Belarus provenance involved an increased risk of autumn frost damage because of later hardening.     

Dormancy release and chilling requirement of buds of latitudinal ecotypes of Betula pendula and B. pubescens

Bud burst and dormancy release of latitudinal ecotypes of Betula pendula Roth and B. pubescens Ehrh. from Denmark ( approximately 56 degrees N), mid-Norway ( approximately 64 degrees N) and northern Norway ( approximately 69 degrees N) were studied in controlled environments. Dormant seedlings were chilled at 0, 5 or 10 degrees C from October 4 onward and then, at monthly intervals from mid-November to February, batches of seedlings were held at 15 degrees C in an 8-h (SD) or 24-h (LD) photoperiod to permit flushing. A decline in days to bud burst occurred with increasing chilling time in all ecotypes. In November, after 44 chilling days, time to bud burst was least in plants chilled at 0 and 5 degrees C. The difference diminished with increasing chilling time, and in February, after 136 chilling days, bud burst was earliest in plants chilled at 10 degrees C. Long photoperiods during flushing significantly reduced thermal time after short chilling periods (44 and 74 days), but had no effect when the chilling requirement was fully met after 105 or more chilling days. No significant difference in these responses was found between the two species. In both species, chilling requirement decreased significantly with increasing latitude of origin. Bud burst was normal in seedlings overwintered at 12 degrees C, but was erratic and delayed in seedlings overwintered at 15 and especially at 21 degrees C, indicating that the critical overwintering temperature is between 12 and 15 degrees C. We conclude that there is little risk of a chilling deficit in birch under Scandinavian winter conditions even with a climatic warming of 7-8 degrees C. The likely effects of a climatic warming include earlier bud burst, a longer growing season and increased risk of spring frost injury, especially in high latitude ecotypes.     

Influence of nutrient supply on spring frost hardiness and time of bud break in Norway spruce (Picea abies (L.) Karst.) seedlings

When spring frosts occur on recently planted forest sites, severe damage may occur to the seedlings. The aim of the present study was to test how different low levels of nutrient concentrations in Norway spruce (Picea abies (L.) Karst.) seedlings affected spring frost hardiness and time of bud break. Seedlings were grown in a greenhouse for one season and supplied with fertiliser containing 22, 43 and 72 mg N l^–1, respectively. The treatments resulted in needle nitrogen concentrations ranging from 0.9 to 1.8% in autumn. After winter storage at 0 °C, bud break was recorded on seedlings growing in the greenhouse, outdoors and in growth chambers at 12 °C and at 17 °C. Freezing tests were performed on seedlings directly removed from winter storage and following one week growth in the greenhouse. Seedlings receiving fertiliser with 43 mg N l^–1 had less freezing injury than the two other fertilisation treatments in the present study. The earliest bud break occurred in seedlings receiving 72 mg N l^–1.     

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.