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.     

Intrapopulation variation in flowering phenology and fecundity of silver birch, implications for adaptability to changing climate

Earlier modeling has suggested that long distance gene flow is of importance in increasing the adaptability of tree populations in a changing climate. In times of warming temperatures, early flowering phenotypes may be favored, because early flowering may be connected to early onset of growth. Long term direct measurements of flowering, pollen cloud and intrapopulation fecundity variations are needed to test this hypothesis. Having one of the furthest transported types of pollen grains, birch may have good potential for long distance gene flow. Our daily observations over eight years, of 30 silver birches (Betula pendula Roth), showed that the onset and duration of flowering phenology was determined by the accumulation of spring temperature sums, but the window for potential long distance gene flow was narrow. The interannual phenology variations were large, the order of timing of female and male catkins in various trees of this monoecious species tended to differ, and the timing of generative and vegetative spring phenologies were not correlated at tree level. Early flowering trees tended to have a higher variation in germinability than later flowering trees. No other connection between timing of phenology of flowering and seed quantity or quality was found. Although stochastic climatic conditions produce interannually variable phenological windows, probability for long distance gene flow from areas differing in timing of spring temperature accumulations may be low in silver birch. Intertree variations in fecundity were high, and the majority of seeds and pollen were produced by only a few trees. Moreover, the amount of seeds produced was positively related to seed germinability, thus large phenotypic fecundity variations may decrease interannual genetic variations in seed sets.     

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.     

Genetic variation and inheritance in a 9 × 9 diallel in silver birch (Betula pendula)

A complete diallel cross was made among nine Betula pendula trees growing in a natural population and a trial was planted on agricultural soil at one site. This exceptional trial has provided estimates of genetic parameters that can only be estimated in complete diallels. Traits measured were height and diameter during a period of 37 years, and assessments were made of bud burst, leaf abscission and rust infection at the early ages. All traits showed genetic variation and the variance components of general combining ability (GCA) effects were dominating, with heritability estimates of 0.16 and 0.23 for height and diameter at age six years. The best-growing families could be identified at that age. At age 37 years, when the trial had been thinned twice, the offspring from the highest and lowest ranked parent for growth contributed with 19% and 6% of the total volume of the stand, respectively. The GCA effects were also highly significant for the assessment traits, but with an interaction with year for bud burst. High values of estimates of genetic correlations proved that bud burst, leaf abscission and rust infection are interrelated, and also to some extent with growth traits. Families with an early bud burst were tallest, were less affected by the rust fungus and kept their leaves later in the autumn.     

Effects of dormancy and environmental factors on timing of bud burst in Betula pendula

We tested three theories predicting the timing of bud burst in mature birch (Betula pendula Roth) trees utilizing a 60-year phenological time series together with meteorological temperature observations. Predictions of the timing of bud burst based on light conditions in addition to temperature were more accurate than predictions based on dormancy development and temperature (prediction standard error of 2.4 days versus 4.3 days). The signal from light conditions, represented by fixed calendar date, determined the start of bud ontogenesis rather than dormancy release. We suggest that models developed to predict the timing of bud burst be utilized in the analysis of plant responses to climate change and of climate change itself.     

美洲黑杨无性系的苗期生长比较

从美国田纳西州、路易斯安那州引进599个无性系,通过3年苗期试验,对苗期生长表现较好的13个无性系进行生长量测定,结果表明：一根一干苗各无性系之间地径生长存在显著性差异,二根一干苗和三根一干苗各无性系之间地径生长存在着极显著性差异;苗高生长在各无性系之间均存在着极显著性差异;参试无性系的地径、苗高表型变异系数分别为12.638%和5.570%,广义遗传力分别为0.835和0.860,遗传变异系数分别为11.562%和5.150%。说明各性状遗传潜力较大,存在着广泛的遗传变异,具有获得较大遗传增益选择的潜力。     

Susceptibility of silver birch (Betula pendula) to herbivorous insects is associated with the size and phenology of birch – implications for climate warming

Susceptibility of trees to herbivorous insects is often associated with the tree growth and phenology, both of which are sensitive to a warming climate. Therefore, changes in these traits may affect the intensity of herbivory on trees in the future. We grew silver birch (Betula pendula) populations differing in their growth and phenology in a common garden experiment, and studied the effect of these plant traits on the leaf herbivory, insect herbivore density and community composition of herbivores in two consecutive years, 2011 and 2012. Differences in size and the timing of growth cessation among the birch genotypes were related to differences in herbivore community compositions and herbivory. In 2011, the most intensive herbivore damage was observed on birch genotypes of small size, late bud burst and early growth cessation, and these genotypes originated from high-latitude populations. The differences among birch genotypes in their susceptibility to herbivore damage may lead to differences in their competitive abilities. While climate warming will affect phenology and growth of plants, our study shows, that it may indirectly affect herbivory and herbivore communities as well.     

Genetic variation of Prunus avium in susceptibility to cherry leaf spot (Blumeriella jaapii) in spatially heterogeneous infected seed orchards

Cherry leaf spot (Blumeriella jaapii (Rehm) Arx.) is a very serious disease of wild cherry (Prunus avium L.), which produces premature leaf defoliation and vigor decrease. In two clonal seed orchards of P. avium naturally infected by B. jaapii, spatial heterogeneity and autocorrelation of neighbor damage caused by cherry leaf spot impeded proper analysis of the fungus incidence. The iterative spatial analysis (ISA), based on variography and kriging, was successfully used to eliminate the effect of this spatial heterogeneity in analysis of genetic variation in susceptibility to B. jaapii. Significant differences among P. avium clones were found, with moderate to high broad-sense heritability estimates. Genetic by environment interactions, although significant, were not quantitatively important. A strong relationship between leaf spot susceptibility and bud burst was found. However, other factors must be affecting the genetic variation in leaf spot susceptibility, as differences among clones remained highly significant when considering the bud burst as a covariate in the genetic model.     

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.     

Genetic and environmental determinants of silver birch growth and herbivore resistance

Plant growth and herbivore resistance is affected by both environmental and genetic factors. The aim of this study was to examine (1) whether genotype explains variation in herbivore resistance, plant growth and tolerance of feeding within natural silver birch populations, (2) whether there is significant interaction between birch genotype and environment or the study year, and (3) whether there are defence costs in field grown juvenile silver birch saplings. The 22 genotypes studied were assigned insect exposure and insect removal treatments in two different environments. Resistance to insect herbivory was measured as the amount of undamaged leaves of the saplings and tolerance of feeding as a difference in relative height increment of the saplings between insect exposure and removal treatments in relation to the amount of damage (i.e. opposite number of resistance). Our study shows that a local silver birch population has substantial genotypic variation in resistance to insect herbivores and this variation is not affected by the environment or the study year. Genotypic variation was not observed for tolerance of feeding. We also showed that, in contrast to genotypic variation in resistance, the genotypic variation in growth among silver birch genotypes strongly depended on the environment, i.e. silver birch genotypes have a high degree of phenotypic plasticity in their growth. Therefore, defence costs in terms of lower growth rate appear to be highly dependent on the environment. To conclude, our results indicate that silver birch populations have a good potential to adapt to possibly increasing herbivory in boreal forests in warmer future climates.     

香椿无性系苗期生长及早期选择研究

[目的]对香椿无性系的苗期生长进行分析,筛选优良香椿无性系,奠定遗传改良的基础。[方法]以49个香椿无性系为对象,分别于2015、2016、2017年测量其1、2、3 a生株高、胸径,计算材积,对表型性状及其增长量进行方差分析和遗传参数估算,采用独立淘汰法筛选排名前20%的无性系为优良,对中选无性系进行遗传增益等参数估算,并将试验地不同年份特征气象因子与无性系表型性状进行相关分析。[结果]株高、胸径及材积的生长连续3 a在无性系间存在显著差异,且生长性状的遗传变异系数、表型变异系数逐年减小,而变异幅度、重复力逐年增大;3 a生时,株高、胸径及材积的无性系重复力分别为0.55、0.50、0.67,重复力高。2015—2017年,49个香椿无性系之间胸径、材积增长量分别呈显著、极显著差异,3 a生材积与材积、胸径增长量呈高度正相关关系。以3 a生香椿无性系材积及其增长量为2组指标,独立筛选出优良无性系10个,中选无性系群体平均材积大于0.010 4 m~3·株~(-1),增长量大于0.009 7 m~3·株~(-1)。3 a生材积的遗传增益为17.38%,中选香椿无性系的稳定性系数b值均大于1,对年份环境敏感。气象因子-表型性状相关性分析显示,材积与年总降水量、日均降水量有显著正相关关系。[结论]参试香椿无性系间材积差异极显著,选择潜力大;中选的优良香椿无性系遗传增益超过15%,但其在不同年份差异大且稳定性差,这可能与年份降水量有关。     

杉木无性系生长表现和优良无性系选择

对营造在福建省洋口国有林场的10年生杉木无性系测定林进行调查分析得出:试验林总体生长表现较好,树高、胸径和材积生长在无性系间存在明显差异,达极显著水平;无性系的重复力均比较大,其中胸径和材积的无性系重复力达0.691和0.757,无性系的生长稳定性较好;通过综合比较,从试验林选择出优良无性系5个,10年生时平均材积遗传增益达23%以上,材积现实增益达30%以上,表现出明显的生长优势。     

High autumn temperature delays spring bud burst in boreal trees, counterbalancing the effect of climatic warming

The effect of temperature during short-day (SD) dormancy induction was examined in three boreal tree species in a controlled environment. Saplings of Betula pendula Roth, B. pubescens Ehrh. and Alnus glutinosa (L.) Moench. were exposed to 5 weeks of 10-h SD induction at 9, 15 and 21 degrees C followed by chilling at 5 degrees C for 40, 70, 100 and 130 days and subsequent forcing at 15 degrees C in a 24-h photoperiod for 60 days. In all species and with all chilling periods, high temperature during SD dormancy induction significantly delayed bud burst during subsequent flushing at 15 degrees C. In A. glutinosa, high temperature during SD dormancy induction also significantly increased the chilling requirement for dormancy release. Field experiments at 60 degrees N with a range of latitudinal birch populations revealed a highly significant correlation between autumn temperature and days to bud burst in the subsequent spring. September temperature alone explained 20% of the variation between years in time of bud burst. In birch populations from 69 and 71 degrees N, which ceased growing and shed their leaves in August when the mean temperature was 15 degrees C, bud burst occurred later than expected compared with lower latitude populations (56 degrees N) in which dormancy induction took place more than 2 months later at a mean temperature of about 6 degrees C. It is concluded that this autumn temperature response may be important for counterbalancing the potentially adverse effects of higher winter temperatures on dormancy stability of boreal trees during climate warming.     

Improvement of Betula pendula by clonal and progeny testing of phenotypically selected trees

Abstract

Field tests, each including 39–83 Swedish plus-tree clones of silver birch (Betula pendula Roth) planted on 10 different sites in the southern part of Sweden, were evaluated at up to 10 years of age. The main conclusions can be summarized as follows. Growth traits were mostly under strong genetic influence, and showed substantial genetic variation and high potential genetic gain. Survival did not seem to be a trait of genetic importance. The genetic correlation between growth and growth cessation was weak, suggesting that it should be possible to find clones combining high growth and early autumn growth. Non-significant transfer effects for clones of different origin and weak G×E interaction were found, indicating that southern Sweden can be treated as a single utilization zone and that few sites are needed for genetic tests. Strong genetic age×age correlations were detected, suggesting that short test periods could be used.     

Variability of wood properties in two wild cherry clonal trials

There has been an increase in wild cherry cultivation over recent decades, revealing a need for improvements in planting material to be used for the production of high quality timber. To date, trial tests have been carried out mainly on growth and tree architectural traits, while no investigations have focused on wood properties. The present study investigated the variability of the growth traits and the physical and mechanical properties of wild cherry wood, both within the tree, within and among clones and between sites. The aims of the study are to provide useful information on the genetics of wood traits in valuable hardwoods and to discuss how within-tree variation can affect early selection of genotypes. The results suggest that site was the most important source of variation in growth traits and in the proportion of heartwood, although differences among clones were highly significant. The number of sapwood rings was very homogeneous both between sites and among clones. Concerning the wood traits, most variation was detected within the tree, rather than within or among clones or between sites. Within tree variability seems to be mainly due to heartwood/sapwood presence. However, since there is a high correlation between heartwood and sapwood properties, a quite efficient selection made on the basis of sapwood should provide an efficient means of selecting heartwood traits.     

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.     

杉木干基萌芽能力的遗传变异

分析了两片杉木子代林伐桩和杉木无性系采穗圃萌条数量的遗传变异情况。结果表明：杉木子代林伐桩萌条数量在产地间、产地内家系间及组合间均存在极显著差异，并有中等以上遗传力；遗传变异系数在３７％～３９％之间；产地的遗传方差分量与产地内家系间的相近。伐桩萌条数量与基径粗度的关系随伐桩年龄的变化而表现不一；不同砍伐季节伐桩萌条数量也存在较大差异。杉木无性系采穗圃无性系间有效萌条数量存在极显著差异，无性系的萌芽能力有较高的重复力。     

Genetic variation in fruitfulness in a hinoki (Chamaecyparis obtusa Endl.) seed orchard and its impact on the maintenance of genetic diversity in seedlots

Genetic variation in seed/cone production among clones was studied in a hinoki (Chamaecyparis obtusa Endl.) seed orchard containing 25 plus-trees by analyzing the number of cones, the yield of cones and seeds of individual ramets for 5 successive years (1982 to 1986). There was significant variation among clones each year and parental contribution in the seed orchard. Specifically, in the years 1982, 1983, 1984, 1985, and 1986, 20% of the clones produced 37.2, 60.6, 36.0, 44.3, and 44.8% of the total cones, respectively. The size of the crop greatly influenced the parental balance in the resulting seed/cone crops. The product moment correlation coefficients and Spearman’s coefficients of rank correlation were small and insignificant between consecutive years, but large and highly significant between alternate years, suggesting the presence of carry-over effects in seed/cone production. The broad-sense heritability on a clone mean basis was 0.74 ± 0.15 for the number of cones, 0.72 ± 0.14 for the yield of cones, and 0.68 ± 0.13 for the yield of seeds. The corresponding heritabilities from analyses combined over all years were 0.24, 0.558, and 0.724, respectively. These results indicate that seed/cone production in hinoki is under strong genetic control. Several managerial measures are discussed that maintain the genetic diversity in seedlots used for reforestation, by reducing the variation in seed/cone production among clones and producing seed crops with equal contributions from all parents.     

杉木无性系生长和材性变异及多性状指数选择

对19年生和17年生2块杉木无性系测定林开展生长和材性变异及多性状指数选择研究,其中,19年生测定林包括38个参试无性系,17年生测定林包括102个参试无性系。2块测定林数据分析结果均显示:无性系树高、胸径、单株材积、心材率和木材基本密度存在极显著的遗传差异,无性系重复力达中等偏上至较高水平,遗传变异系数各性状间差异较大;树高、胸径、材积生长性状间存在极显著的表型和遗传相关,与基本密度存在弱负相关。不同选择方法的结果表明,指数选择能整体提高单株材积、心材率和木材基本密度综合的遗传增益,达到比单性状选择更好的选择效果,是多性状改良较理想的选择方法。     

Models of the spring phenology of boreal and temperate trees: Is there something missing?

According to prevailing theory, air temperature is the main environmental factor regulating the timing of bud burst of boreal and temperate trees. Air temperature has a dual role in this regulation. First, after the cessation of growth in autumn, prolonged exposure to chilling causes rest completion, i.e., removes the physiological growth-arresting conditions inside the bud. After rest completion, prolonged exposure to warm conditions causes ontogenetic development leading to bud burst or flowering. During the past three decades, several simulation models based on chilling and forcing have been developed and tested. In recent modeling studies of the timing of bud burst in mature trees, the simpler thermal-time models that assume forcing starts on a fixed date in the spring have outperformed the chilling-forcing models. We hypothesize that this discrepancy may be due to some element missing from the chilling-forcing models. We tested two new model formulations by introducing reversing, temperature-driven elements that precede forcing and by fitting the models to seven historical time series of data of flowering and leaf bud burst of common boreal tree species. In these tests, both of the new models were generally more accurate in predicting the timing of bud burst than a classical chilling-forcing model, but less accurate than the simple thermal-time model. We therefore conclude that besides chilling, other environmental factors are involved in the regulation of the timing of bud burst. Further work is needed to determine if the regulatory factors derive from air temperature or from some other environmental condition such as changes in light conditions, like day length or night length.