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.     

Seedling cold hardiness, bud set, and bud break in nine provenances of Pinus greggii Engelm.

Cold hardiness and timing of bud set and bud break are important processes that provide protection of nursery seedlings against low temperatures. Seedlings of 9 provenances of Pinus greggii from two different regions of Mexico were tested to determine cold hardiness, bud set, and bud break timing differences. Needle sections were exposed to freezing temperatures to determine an injury index of each provenance. In addition, bud set and bud break timing were recorded through the fall, winter and spring. There were significant differences in cold hardiness between seedlings from northern and southern provenances. At the maximum cold hardiness, the index of injury (LT₅₀) for northern provenances was LT₅₀ = −18 °C, compared to −12 °C for southern provenances. There was a considerable variation among the provenances in the proportion of seedlings that set terminal buds. Seedlings from northern provenances had greater proportions of seedlings that set a terminal bud than seedlings from southern provenances. There were also significant differences in the bud break timing in the following spring among the 9 provenances. Seedlings from northern provenances broke bud earlier than southern provenances. Cold hardiness, bud set, and bud break timing results may be useful to determine how far a specific seed source can be moved from its natural environment.     

杉木苗期抗寒性的遗传变异和家系选择

近20年来,我国杉木遗传改良工作进展很快,但杉木育种目标多局限于速生丰产,过于单一。目前杉木的栽培区域不断从中心产区向边缘产区扩展延伸,出现了对不良气候、土壤条件的适应性问题。例如,1987年冬浙江各地苗圃遭受早霜冻害十分严重,使上百万株杉苗     

Needle growth in Sitka spruce (Picea sitchensis): effects of nutrient deficiency and needle position within shoots

Needle development and shoot growth were studied in 14- and 20-year-old Sitka spruce (Picea sitchensis (Bong.) Carr.) planted on an oligotrophic peat and fertilized with N, P and K (control), N and P (-K), N and K (-P) or unfertilized (-NPK). Shoot extension, needle number, and the size and number of needle cells were observed throughout the season. Beginning with bud burst in early May, needle growth continued for nine weeks in the control treatment. Needle growth in the mineral deficiency treatments terminated at the same time as in the controls, although it did not begin until up to three weeks later than in the control trees. Needles developed acropetally along the shoot, with basal needles completing their development first. Cell division in needles of control trees lasted about 3 weeks, with cell expansion continuing for a further 3 weeks. Initial mean cell cycle times were shortest for proximal needles following bud burst, but lengthened as the season progressed. Cell number increased from 6600 in the primordia to 200 000 in mature needles. Final needle dimensions and cell number varied according to the position of the needle on the shoot. The largest needles with the most cells were near the middle of the shoot. Relative to the controls, all mineral deficiency treatments reduced shoot length, bud and needle dimensions, and needle cell number, particularly the -P treatment. Potassium and P deficiency treatments reduced primordial cell number by up to 42%, whereas final mean cell size was 30% greater in the -P treatment, 17% greater in the -NPK treatment, but 14% smaller in the -K treatment. Mean cell size was constant for needles at all positions in any particular treatment at any time, so that final needle size was determined by cell number alone. Needle dry weight/fresh weight ratio continued to increase until early August, with significantly higher ratios in the -K and -P treatments than in the controls.     

Mechanisms of Douglas-fir resistance to western spruce budworm defoliation: bud burst phenology, photosynthetic compensation and growth rate

We compared growth rates among mature interior Douglas-fir (Pseudotsuga menziesii var. glauca (Beissn.) Franco) trees showing resistance or susceptibility to defoliation caused by western spruce budworm (Choristoneura occidentalis Freeman), and among clones and half-sib seedling progeny of these trees in a greenhouse. We also investigated bud burst phenology and photosynthetic responses of clones to budworm defoliation in greenhouse experiments. Resistant mature trees had a higher radial growth rate than susceptible trees, especially during periods of budworm defoliation. Clones from resistant trees grew larger crowns than clones from susceptible trees, whereas stem base diameter at the ground line and height did not differ. Half-sib seedling progeny from resistant trees had larger stem diameter, height, and total biomass than progeny from susceptible trees. Mean 5-year radial growth increment of mature trees was more strongly correlated with growth of seedlings than with growth of clones. Clones from resistant trees had later bud burst than clones from susceptible trees, and budworm defoliation of clones depended on the degree of synchrony between bud burst phenology and budworm larval feeding. Clones of resistant and susceptible mature trees showed similar responses of net photosynthetic rate to 2 years of budworm defoliation. We conclude that phenotypic differences in crown condition of Douglas-fir trees following western spruce budworm defoliation are influenced by tree genotype and that high growth rate and late bud burst phenology promote tree resistance to budworm defoliation.     

Frost hardiness of 16 European provenances of sessile oak growing in Scotland

The frost hardiness of 16 European provenances of sessile oakQuercus petraea (Matt.)Liebl. originating from six Europeancountries was examined from autumn until budburst in springusing the method of relative conductivity. There were significant differences of frost hardiness betweenprovenances and a strong relationship between phenology andfrost hardiness. In spring, provenances that burst bud earlydehardened earlier than provenances that burst bud later. Inautumn, provenances that stopped growing early were more frosthardy than provenances that continued to grow. The interprovenance ranges of frost hardiness were greatestin spring and autumn and least in midwinter. Consequently, itwas not possible to demonstrate significant differences of frosthardiness between provenances in December and January. Overall, German, Polish and Danish provenances were more frosthardy than French, Austrian and British provenances. Frenchprovenances were least hardy at all times.     

Evaluation of a portable chlorophyll meter to estimate chlorophyll and nitrogen contents in sugar maple (Acer saccharum Marsh.) leaves

Portable, non-destructive chlorophyll meters could be a valuable tool for forest managers and researchers. The ability of such meters to assess both chlorophyll and nitrogen content has been well established for many agricultural species, but not well-studied for North American forest tree species. The objective of this study was to evaluate the ability of a portable chlorophyll meter to estimate chlorophyll (CHL) and nitrogen (N) content in sugar maple (Acer saccharum Marsh.) leaves by testing correlations between chlorophyll content index values (CCI) and extractable CHL and N in large heterogeneous samples. Significant correlations were observed between CCI and CHL (P ≤ 0.001, r² = 76%) and CCI and N (P ≤ 0.001, r² = 64%). These data indicate that the meter tested can be an effective tool for estimating relative chlorophyll and nitrogen content in sugar maple leaves.     

Observations on bud burst phenology in a field trial established with Poplar (Populus spp.)

Bud flushing is very important for the survival and growth of trees, a phenomenon matched each year with the annual course of temperature and the timing of bud flushing in the spring. Essentially it represents a serious ecological and evolutionary tradeoff between survival and growth. The most suitable timing of bud burst permits trees to begin growth sufficiently early to take advantage of favorable spring conditions, but late enough to decrease the risks of tissue damage from late frost. In the present study bud burst spring phenology of poplar (Populus tremula and P. tremuloides) from eight different provenances, originating from Europe and the USA, was observed during March and April, 2009. The experimental plot was located at Solling, Germany (51°44′0″ N, 9°36′0″ E). A six stage subjective scoring system of bud burst phenology was used to identify the phenological stages of the seedlings, where each plant was observed twice a week. The aim of the study was to predict phenotypic variation in poplar, originating from regions between 42° and 60° N latitude, growing in similar environments. Timing of bud flushing of poplar was recorded. It was found that seedlings of provenance 3, which originated from 42.35° N latitude, started and completed flushing significantly earlier than those of other provenances, while seedlings of provenance 5, originating from 54.29° N latitude, started flushing very late and only a few plants reached top scoring at the end of the experimental period. Analysis of variance showed statistically highly significant differences (p < 0.05) in bud flushing among the provenances. The correlation between scoring and flushing periods was very strong within provenances although the flushing pattern differed among provenances (origin of the planted seedlings). Bud flushing showed a negative correlation with the origin of the planted seedlings. Given the field experience gained with this experiment, it is recommended that seedlings from provenances 5 and 8 could be used for future plantations where late frost may be a problem for the young shoots of seedlings.     

Impact of elevated carbon dioxide concentration and temperature on bud burst and shoot growth of boreal Norway spruce

Effects of elevated temperature and atmospheric CO2 concentration ([CO2]) on spring phenology of mature field-grown Norway spruce (Picea abies (L.) Karst.) trees were followed for three years. Twelve whole-tree chambers (WTC) were installed around individual trees and used to expose the trees to a predicted future climate. The predicted climate scenario for the site, in the year 2100, was 700 micromol mol-1 [CO2], and an air temperature 3 degrees C higher in summer and 5 degrees C higher in winter, compared with current conditions. Four WTC treatments were imposed using combinations of ambient and elevated [CO2] and temperature. Control trees outside the WTCs were also studied. Bud development and shoot extension were monitored from early spring until the termination of elongation growth. Elevated air temperature hastened both bud development and the initiation and termination of shoot growth by two to three weeks in each study year. Elevated [CO2] had no significant effect on bud development patterns or the length of the shoot growth period. There was a good correlation between temperature sum (day degrees>or=0 degrees C) and shoot elongation, but a precise timing of bud burst could not be derived by using an accumulation of temperature sums.     

Temperature sum accumulation effects on within-population variation and long-term trends in date of bud burst of European white birch (Betula pendula)

Within-population variation in phenology of boreal trees indicates their adaptability to climatic variations. Although interannual variations in date of bud burst have been widely discussed, little is known about within-population variation, the key determinants for this variation and the effects of this variation on estimates of trends in bud burst date. Over a period of nine years, we monitored timing of bud burst daily in 30 mature white birch (Betula pendula Roth) trees in a naturally regenerated stand. Our results revealed not only large interannual variation but also considerable intraannual variation among individual trees in date of bud burst, the maximum within-population variation being four weeks. Bud burst can be accurately predicted by the date when a threshold value of temperature sum in spring is reached (base temperature +5 degrees C). Based on this temperature sum and past temperature records, we estimated the trend in date of bud burst. The linear trend estimate based on the years 1926-2005 is an advancement of 1.2 days per decade (95% confidence interval, +/- 0.7 days), which is much less than that predicted by time series based on coarser time intervals. We conclude that, because of large interannual differences, and large annual within-population variations in bud burst, estimates of bud burst date based on measurements made over a period of only a few decades are unreliable.     

Chlorophyll fluorescence and variations in tissue cold hardiness in response to freezing stress in Douglas-fir seedlings

Two-year-old coastal Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) seedlings from two seed lots were exposed to controlled freezing temperatures every 4 weeks from October 1993 through April 1994. Freezing effects were assessed by measuring chlorophyll fluorescence emissions 1 day after freezing and by evaluating damage to the bud, cambium, and needle tissues 7 days after freezing. Differences between the seed lots as well as changes in cold hardiness among the bud, cambium, and needle tissues were evident throughout the duration of the study. Tissue damage was higher with increased freezing stress. Severity of damage to each of the tissues varied seasonally. Chlorophyll fluorescence emissions were lower with higher freezing stress (except during November and December, when test temperatures were not low enough to significantly damage the seedlings) and showed a strong relationship with morphological assessments of freezing stress. The slope of the slow kinetics phase of the chlorophyll fluorescence curve tended to be less steep (i.e., quenching was reduced) with higher freezing stress. Nonfrozen chlorophyll fluorescence measurements showed no obvious relationship with LT₅₀ for either seed lot. However, chlorophyll fluorescence measurements are useful for determining cold hardiness and resistance to stress, because they provide a rapid assessment of seedling vigor following exposure to freezing.     

Variation in cold hardiness and carbohydrate concentration from dormancy induction to bud burst among provenances of three European oak species

Although cold hardiness is known to be a major determinant of tree species distribution, its dynamics and the factors that regulate it remain poorly understood. Variation in cold hardiness and carbohydrate concentration, from dormancy induction until bud burst, were investigated in populations of two deciduous (Quercus robur L. and Quercus pubescens Willd.) and one evergreen (Quercus ilex L.) European oak. Mean cold hardiness values in January were -56, -45 and -27 degrees C for Q. robur, Q. pubescens and Q. ilex, respectively. Soluble carbohydrate concentrations were closely related to instantaneous cold hardiness, estimated by the electrolyte leakage method, whereas total carbohydrate concentration was related to maximum cold hardiness. Both cold hardiness and carbohydrate concentration showed a close linear relationship with temperatures at the location of the sampled population. Our results show that temporal variation in both the inter- and intraspecific cold hardiness in European oaks can be related to variations in the concentrations of soluble carbohydrates and that these relationships appear to be driven by temperature.     

Root-zone temperatures affect phenology of bud break, flower cluster development, shoot extension growth and gas exchange of 'Braeburn' (Malus domestica) apple trees

We investigated the effects of root-zone temperature on bud break, flowering, shoot growth and gas exchange of potted mature apple (Malus domestica (Borkh.)) trees with undisturbed roots. Soil respiration was also determined. Potted 'Braeburn' apple trees on M.9 rootstock were grown for 70 days in a constant day/night temperature regime (25/18 degrees C) and one of three constant root-zone temperatures (7, 15 and 25 degrees C). Both the proportion and timing of bud break were significantly enhanced as root-zone temperature increased. Rate of floral cluster opening was also markedly increased with increasing root-zone temperature. Shoot length increased but shoot girth growth declined as root-zone temperatures increased. Soil respiration and leaf photosynthesis generally increased as root-zone temperatures increased. Results indicate that apple trees growing in regions where root zone temperatures are < or = 15 degrees C have delayed bud break and up to 20% fewer clusters than apple trees exposed to root zone temperatures of > or = 15 degrees C. The effect of root-zone temperature on shoot performance may be mediated through the mobilization of root reserves, although the role of phytohormones cannot be discounted. Variation in leaf photosynthesis across the temperature treatments was inadequately explained by stomatal conductance. Given that root growth increases with increasing temperature, changes in sink activity induced by the root-zone temperature treatments provide a possible explanation for the non-stomatal effect on photosynthesis. Irrespective of underlying mechanisms, root-zone temperatures influence bud break and flowering in apple trees.     

Frost hardiness, bud phenology and growth of containerized Picea mariana seedlings grown at three nitrogen levels and three temperature regimes

We studied the influence of temperature and near- and sub- optimal mineral nutrition of black spruce seedlings (Picea mariana [Mill.] B.S.P.) during their second growing period on bud set, bud development, growth, mineral content and cold tolerance. Bud break and growth after bud break were also studied. Seedlings were grown for 106 d in growth chambers under three temperature regimes in combination with three concentrations of a fertilizer. They were then cold hardened for 56 d and dehardened for 66 d.Under these near- and sub-optimal N levels, bud formation occurred during the growing season. Bud formation was accelerated with decreasing fertilization, but was not affected by temperature treatments. Needles from seedlings with 0.64% N (dry mass basis) before hardening did not harden. Those with 0.87% N showed a lesser degree of hardiness than those with 1.28% N. Stem diameter increased at the beginning of the hardening period. During this acclimation period, shoot dry mass decreased with time at a constant rate and at the same rate over time for all treatments whereas root dry mass was more variable. Total number of needle primordia was low and no difference was observed among growing conditions. Bud break was similar in all treatments. Following bud break, shoot height and stem diameter increases were small but their magnitude varied with the nutritional regimes applied during the previous growing period. During hardening, nitrogen concentration of shoot tissues first increased and then decreased; phosphorus concentration first increased and then remained stable; potassium concentration remained stable. Concentration of these three elements generally decreased in the roots during this hardening.     

Growth and phenology variation in progeny of Scots pine seed orchards and commercial seed stands

Tree improvement in Poland has been most advanced for Scots pine, but existing seed orchards have not been progeny-tested yet. We examined variation in growth traits—tree height at ages 4 and 8 years, and diameter at age 13 years—in the common garden experiment testing open-pollinated progenies of 31 seed orchards and 5 commercial seed stands (referred to as populations) at 5 locations. We also examined bud burst phenology at two to five sites at three growing seasons. At one experimental site during the 5th growing season, we measured shoot growth rhythm on all populations. Similar measurements of shoot growth were done on a subset of populations during the 6th growing season together with the analysis of needle growth and foliar chemistry. We found significant variation among populations in growth traits, but also significant population × site (G × E) interactions. We used the regression approach and ecovalence analysis to examine populations’ performance stability. Most populations had average responsiveness to environment, and a set of least-responsive poor-growing populations contributed the most to the G × E interaction. Variation in bud burst phenology was associated with geographical distribution of tested progenies. The early bud-bursting populations originated from the north-eastern to north-central Poland, and a group of late bud-bursting populations originated mainly from the south-eastern region. Correlations between bud burst and growth traits were weak to medium and varied by site, but early bud-bursting populations tended to show stronger growth on height and diameter. We found significant differences among populations in final leader length, shoot elongation time and relative growth rate (RGR). However, RGR and shoot elongation time explained less than 30% of variation in leader length and were weakly correlated with tree height. Populations varied in needle length, specific leaf area and foliar nitrogen concentration, but time trends in these traits did not vary among populations or predefined groups of populations. Therefore, the analysis of growth rhythm or needle traits did not help resolve variation in tree growth to support selection decisions. Contrary to our expectation, progeny of seed orchards did not perform significantly better than that of commercial seed stands. This finding, however, should not be extrapolated beyond our set of populations. Nonetheless, the local seed sources were not always the best. From a selection standpoint, our results help culling the worst populations rather than selecting the best ones. Therefore, testing individual family progeny and genetic roguing of existing seed orchards is highly recommended.     

Within‐population variation in autumn frost hardiness and its relationship to bud‐set and height growth in Picea abies

Artificial freezing tests were performed with two sets of 12 full‐sib families of Picea abies. Each set was selected from a complete diallel cross performed within a natural population. Significant differences in autumn frost hardiness were observed between the two populations, which originate from the same altitude and longitude approximately 60 km apart. Substantial variation in frost hardiness was observed within both populations. Significant pheno‐typic correlations at the individual level were found between freezing injury and the two traits terminal bud‐set and height growth year one. However, no significant relationships were present between freezing injury and bud‐set at the family level, indicating that bud‐set cannot be used to predict autumn frost hardiness for families.     

Seasonal variation in photochemical activity and hardiness in clones of Norway spruce (Picea abies)

We investigated changes in photochemical activity and cold hardiness of detached needles of three clones of Picea abies (L.) Karst. by measuring variable chlorophyll fluorescence (F(v)/F(m)), before and after artificial freezing, from September to June. Photochemical activity varied considerably during the study, but only minor differences in photochemical activity among the clones were observed before freezing. Photochemical activity was high during early fall and then declined from November until April. Photochemical activity was at a minimum in April and then increased quickly to high values in May. During the period from late September to October, and also during the winter, differences in F(v)/F(m) ratios after artificial freezing to below -10 degrees C were observed among clones, indicating clonal differences in cold hardiness and hardiness development. The clone having an average height of 2.3 m after 11 years showed consistently lower cold hardiness than clones that had reached average heights of 4.0 and 5.0 m. There were also differences in the temperature requirement for bud flushing among clones.     

香榧花芽分化与核酸的关系研究初报

香榧花芽分化与核酸的关系研究初报苏梦云周国璋关键词香榧花芽分化核酸香榧（ＴｏｒｅｙａｇｒａｎｄｉｓＦｏｒｔ．ｅｘＬｉｎｄｌ．）是我国著名的珍稀干果,雌雄异株,一般在４月中下旬开花授粉。由于香榧童期较长（大约需２０ａ左右才开花）,如何促进提早开花结实已...     

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.