Gains from selecting Pinus sylvestris in southern Sweden for volume per hectare

Abstract

An overall objective of the Swedish Scots pine (Pinus sylvestris L.) tree breeding programme is to increase volume production. Selection is based on height at young age as an indicator of future volume production. This study examined the relationship between height at young age and volume production in block-plots at older ages, and the sustainability of increased growth in volume per unit area using parent breeding values. The study included five trials with controlled pollinated progenies from 71 phenotypically selected plus-trees. Height and diameter were measured in the trials at ages ranging from 7 to 36 years. The genetic correlation between height at 9 years and volume growth per unit area at 30 years of age was 0.8. The additive genetic coefficient of variation for growth per area unit was 17% at the age of 30 years, i.e. first thinning. Based on the estimated parameters, selection of the best 25% genetic entries according to their height at age 9 years resulted in 13% volume gain at age of 30 years, while selection of the 10% best resulted in 18% volume gain. The plus-tree progenies had 12% higher volume growth per unit area than the unimproved commercial check lots, and this difference needs to be added to obtain the total gains. This means that selection of the 25% best parents gave a gain of 25% in volume per hectare compared with unimproved check lots.     

Rapid living crown reduction caused by Gremmeniella abietina affects foliar nutrient concentrations of Scots pine

The foliar chemistry of diseased and healthy trees was studied one growing season after severe reduction in living crown caused by Gremmeniella abietina in four young Scots pine (Pinus sylvestris) stands. Sample trees were chosen pairwise on the basis of the living crown length: a diseased tree with about 50% live crown reduction and a healthy tree in each pair. Fifteen elements were determined in the youngest healthy needles on the lateral top shoots of each sample tree. Diseased trees had higher foliar boron, manganese and sodium concentrations and lower magnesium, iron, zinc, copper, potassium, nitrogen and sulphur concentrations compared to the healthy trees. Foliar calcium, aluminium, phosphorus, carbon and hydrogen concentrations did not differ between the diseased and healthy trees, except for P and A1 in two of the stands when the stands were analysed separately. Significant correlations between the needle element concentrations and crown ratio (length of the living crown/tree height) were found especially for B (increasing B with decreasing crown ratio) and for Mg, Fe and Zn (decreasing concentrations with decreasing crown ratio). The effect of G. abietina-induced living crown reduction on tree nutrient status and the role of these mineral nutrients in the susceptibility are discussed.     

Crown transparency, tree mortality and stem growth of Pinus sylvestris, and colonization of Tomicus piniperda after an outbreak of Gremmeniella abietina

In the year 2000, large areas of forest in Sweden, mainly 30-50 year old Pinus sylvestris (L.) stands, were attacked by the fungus Gremmeniella abietina (Lagerb.) Morelet. The aims of this study were to investigate: (i) the relationship between G. abietina-induced tree crown transparency (CT) and P. sylvestris (L.) tree mortality; (ii) the influence of CT levels on stem growth; (iii) the recovery of the crown; and (iv) the association of CT and colonization by Tomicus piniperda (L.). Thirty-five permanent sample plots were established in five P. sylvestris stands (38-46 years old), infested by G. abietina, and 23 plots in four reference stands, not obviously infested.During the 5 years following the attack, the total mortality amounted to 454 trees ha⁻¹ and 7.8 m² ha⁻¹, on average, in the five infested stands, corresponding to 42% of the trees and 34% of the basal area at the time of the attack. Most of the mortality occurred within 2 years of the attack. The mortality of individual trees (2002-2005) was found to be related to the crown transparency (CT), the position of needle loss within the crown and the tree diameter at breast height. Based on our modeling, the probability of mortality was substantially increased if the initial CT-value was higher than 85%.Growth reductions were detected for individual trees with an initial CT of >c. 40%. In contrast, trees with a low initial CT (<c. 40%) were not affected and even exhibited increased growth. In the five infested stands, the reductions in basal area and volume increment were estimated to be 26-58%, and, 42-73%, respectively, during the five growing seasons after the attacks.The trees in the infested stands that were still alive in spring 2005 had started to recover in terms of CT. Breeding of T. piniperda on the P. sylvestris (L.) stems occurred almost exclusively on stems with a CT > 90%.The data from this study suggest that when a P. sylvestris (L.) stand has been attacked by G. abietina, trees with a CT above 80% should be felled; the remaining trees will have a high probability of survival and resistance to successful breeding by the T. piniperda.     

Occurrence of Gremmeniella abietina damage on Scots pine along a pollution gradient from Monchegorsk nickel smelter to western Lapland

Gremmeniella abietina damage was studied at stand and tree level on the Kola Peninsula and in northern Finland. At stand level, damage was recorded in four areas in Finland, while only individual damaged trees were found in Russia. According to the results of branch analysis, there was no sign of endemic epidemics in any of the areas. There was a significant negative correlation between the average G. abietina damage and modelled SO₂ deposition and temperature sum along the gradient. The results suggest that G. abietina is distributed all over northern Finland and the Kola Peninsula in natural stands of Scots pine, and that the direct effect of SO₂ deposition from the Monchegorsk nickel smelter on the damage may be minimal.     

思茅松自由授粉家系遗传参数与早期选择分析

[目的]通过对思茅松自由授粉子代测试林的生长与形质性状遗传参数的估计,进行优良家系选择,为建立思茅松高世代种子园及培育优良无性系苗木提供种质资源。[方法]采用随机区组试验设计,在214个思茅松自由授粉家系子代测试林中选择胸径、地径、树高、枝下高、冠幅、通直度、树干圆满度、树冠圆满度、材积及地上生物量等生长与形质性状,利用线性混合模型进行遗传参数的估计,进行优良家系筛选。[结果]思茅松自由授粉子代测试林生长与形质性状的遗传参数在家系间差异极显著。各性状的家系遗传力较高,其中,地径的家系遗传力最大(1.105)。材积的预期遗传增益高于地上生物量,利用家系/家系内选择方法进行优良家系的筛选,材积的预期遗传增益为60.75%,入选家系为60个;地上生物量的预期遗传增益为44.22%,入选家系为66个。轮盘数、通直度和树干圆满度对材积和地上生物量都有显著影响。[结论]在早期选择中,形质性状的遗传变异对思茅松自由授粉优良家系的选择有重要参考作用,4年生思茅松自由授粉家系子代测试林中各性状存在着丰富的遗传变异,在早期选择中可以筛选出适合不同培育目标的优良家系,最大限度挖掘其遗传潜力。     

Development of fruiting bodies of large tree type of Gremmeniella abietina var. abietina and timing of infection on Scots pine inn northern Finland

The formation and maturing of the large tree type Gremmeniella abietina var. abietina fruiting bodies and their sporulation were investigated for 3 years on Scots pine (Pinus sylvestris) in northern Finland. This was done by monthly assessment of shoots in the field and in the laboratory. Infection caused by G. abietina var. abietina was dated on Scots pine by monthly covering with pollination bags and exposing branches during the growing season. Pycnidia appeared between August and September, 1 year after infection, and they started to release conidia between late June and early July, 2 years after infection. Fresh pycnidia and microconidia were formed during the following August and September in the infected shoots. The causal large tree type of G. abietina var. abietina did not produce apothecia on branches within 3 years of infection. Monthly covering and exposing branches showed that infection took place mainly between June and July.     

Phenology Differs among Norway Spruce Populations in Relation to Local Variation in Altitude of Maternal Stands in the Beskidy Mountains

An investigation of phenological and growth traits was conducted in a lowland plantation, comprised of 23 Norway spruce (Picea abies (L.) Karst.) seed stands from the Istebna region in the Beskidy Mountains of Poland. Significant differences were found among population in growth initiation, growth termination and the length of shoot elongation period. The two latter traits were highly correlated and were attributed to the altitude of the maternal stand. The tested populations were grouped based on phenology. Progeny of lower-alttitude stands ceased growth later, had a longer duration of shoot growth and greater current leader length compared to progeny of higher-altitude stands. Individual populations, however, did not differ in current-year growth and total tree height, implying higher within-population variation in growth traits than in phenology. The similar growth capacity of all tested populations suggests considerable gene flow between maternal stands, although differences in phenology imply the adaptation of progenies to the altitudinal environment of seed origin. These results suggest that within tested populations, selection is possible based on phenology alone, without considerable reduction of early height growth; final decisions, however, should be based on the environmental conditions of the planting site.     

Genetically improved reforestation stock provides simultaneous benefits for growers and a sawmill,a case study in Finland

Recent research has demonstrated benefits of genetically improved seed material (from seed orchards) for growers. In this study, we analyzed whether the profits of genetically faster growing material are also realized as simultaneous economic benefits for a sawmill. We compared three types of improved Scots pine (Pinus sylvestris L.) seedlots to unimproved seedlots on three climatically different sites using stand-simulation software MOTTI. Genetic gains in early-rotation growth traits were incorporated in the growth models as genetic multipliers. Bare land values of the simulated stands were maximized using stand-level optimization. Consequently, removals of each thinning and final harvest were used to estimate the total value of sawing yield. This was subsequently maximized by optimizing the sawing procedure of the logs assuming identical timber quality between improved and unimproved seedlots. The results indicate that simultaneous financial incentives from tree breeding are achievable for both agents (growers and a sawmill). In general, the financial potential increases with the growth potential of the regeneration material, although the form of the relationship is not linear in all cases.     

An outbreak of Gremmeniella abietina in central Finland

G. abietina has damaged P. sylvestris forests in central Finland. The denser the stand, the greater the number of recoverable trees. Artificially regenrated stands were more infected than naturally regenerated ones. Forest fertilization decresed the proportion of recoverable trees.     

Progeny differences of hinoki (Chamaecyparis obtusa) and sawara (C. pisifera) against resinous stem canker disease and spatial distribution of damage (disease severity) in a progeny test

The incidence and severity of resinous stem canker disease were investigated in hinoki (Chamaecyparis obtusa) and sawara (C. pisifera) at a progeny test located in Yamatsuri Town, Fukushima Prefecture, Japan. Symptoms of the disease were observed in 307 trees out of 933 investigated trees (32.9%). The damage was more severe on lower slopes than on upper slopes, indicating that micro-environmental factors are causally associated with the occurrence of the disease. The severity of the disease varied both among nine open-pollinated progenies derived from hinoki plus-tree clones and among 13 progenies derived from pollination between hinoki plus-tree clones and the mixed-pollen of hinoki plus-tree clones. The severity of the disease also varied with height above the ground. This tendency was most obvious where the disease was most severe. An estimation of the narrow sense heritability,h ², of 0.214, was obtained from analysis of the 13 progenies derived from pollination with mixed-pollen. The heritability derived from material that had not been subjected to selection against the disease, indicated that breeding to enhance resistance would be possible to control the disease. The interaction between environments and progenies in relation to severity of the disease is also discussed. Ms. Setsuko Chiba and Mr. Mamoru Ueta gave us vigorous supports, and Mr. Haruki Orita, Dr. Tomiyasu Miyaura, and Dr. Shinichiro Ito gave us many helpful comments. We deeply appreciated their cooperation. The study was supported by a grant from the Ministry of Agriculture, Forestry. and Fisheries of Japan.     

红豆树优树子代遗传多样性及与生长相关性分析

[目的]研究红豆树优树自由授粉子代遗传多样性及其对生长的影响,比较天然居群子代和孤立木子代遗传多样性差异,揭示子代遗传多样性的变化规律及天然居群在子代遗传多样性维持中的作用,为红豆树遗传保育和优异种质挖掘提供科学依据。[方法]以来自浙、闽、赣、川等26个红豆树优树自由授粉家系为研究对象,利用11对SSR引物对765个子代群体进行遗传多样性评价,同时分析子代遗传多样性参数与种子、生长性状的相关性。[结果](1)红豆树优树子代群体具有较高的遗传多样性,有效等位基因数为7.766个,观测杂合度(H_O)和期望杂合度(H_E)分别为0.469和0.865。(2)除SSR8外,其余位点的观测杂合度均小于期望杂合度,表明子代群体绝大多数位点处于杂合子缺失状态。(3)红豆树不同家系的遗传多样性存在明显差异,12号家系的遗传多样性水平最高,8号家系则最低。(4)比较发现,天然居群子代的遗传多样性显著或极显著地高于孤立木子代。(5)F统计量和分子方差分析(AMOVA)均表明,红豆树优树子代群体的遗传变异主要存在于家系内,家系间的遗传分化相对较小。(6)相关性分析发现,子代遗传多样性参数与种子性状、子代年高生长量呈显著正相关(r=0.378~0.527)。[结论]较大的红豆树天然居群在维持其子代较高遗传多样性中发挥了重要作用,子代遗传多样性显著影响苗木生长,这为红豆树遗传保育和优良家系选择提供了理论依据。     

Growth reactions of a Gremmeniella abietina isolate and Scots pine embryogenic tissue cultures differ in a host–parasite in vitro system

Summary The aim of this study was to screen Scots pine embryogenic tissue culture material to Gremmeniella abietina. According to the results Scots pine (Pinus sylvestris) embryogenic tissue lines (27 lines) could be ranked based on significant differences in G. abietina effects estimated by changes in glucosamine concentrations, growth, and dry matter contents of inoculated tissue pieces. The ranking order of the extreme tissue lines stayed basically similar, regardless of the variable measured. These findings imply a possibility to use the ranked embryogenic tissue lines in more detailed investigations of interactions between G. abietina and its host. This is the first report of using embryogenic tissue cultures in studying host–parasite interaction between Scots pine and G. abietina.     

Variation analysis and fine family selection on half-sib progenies from clonal seed orchard of Pinus massoniana

Variation of traits that include height, diameter at breast height (DBH), stem volume, crown width, straightness, wood basic density, tracheid length and width of open-pollinated progenies from clonal seed orchard of Pinus massoniana was analyzed. Differences in traits such as growth, tree form and wood quality among families was significant, which indicated that the improvement potential was notable. Heritability of height and wood basic density reached 0.691 and 0.640, respectively. According to a correlation analysis between traits, stem straightness could be improved indirectly when growth traits are improved; growth traits were significantly correlated with wood basic density; and wood basic density was not significantly correlated with length and width of the tracheid. Under the selection ratio of 10%, genetic gains were 19.74% and 19.32% respectively, and selected families from the progeny test stands at the age of five years and eight years were the same. Genetic correlation of height, DBH and stem volume was also significant at the level of 0.01. These show that selection of the short rotation of P. massoniana for pulp and paper use would be efficient for five-year-old trees. According to the comparison of the heritability of all the traits and taking into account the result of canonical correlation analysis, height could be considered as the most important index for selection. Given wood quality of all the families, and flowering and fruit settings of the clones in a seed orchard, twelve fine families were selected at 10% selection ratio of height growth and stem volume. Translated from Scientia Silvae Sinicae, 2005, 41(6): 43–49 [译自: 林业科学]     

Studies of the life cycle of Gremmeniella abietina on Scots pine in southern Sweden

Aspects of the life cycle of Gremmeniella abietina (Lagerb.) Morelet were studied from 1988 to 1990 in stands of Pinus sylvestris L., 16–32 years old, in southern Sweden, initiated in 1988 with a widespread outbreak of the disease. Pycnidia started to release conidia in late spring and apothecia began to release ascospores in summer. Latent infections could still be detected one year after their establishment by cultivation of healthy looking shoots on agar petri dishes. G. abietina appeared to have a mainly biennal life cycle, as most spores were released two years after infection of the shoot.     

RAPD variation in Gremmeniella abietina attacking Pinus sylvestris and Pinus contorta in northern Sweden

Genomic DNA from 81 isolates of Gremmeniella abietina collected from eleven plantations each of Pinus sylvestris and Pinus contorta in northern Sweden was studied using RAPD markers. The DNA variation between and within populations and the race and type distribution of G. abietina populations, causing symptoms similar to those of the North American race, were studied. The degree of genetic similarity was greater among G. abietina isolates from the same geographical areas than among isolates from different geographical areas, regardless of whether they were isolated from P. sylvestris or P. contorta. RAPD variation was greatest in the central parts of northern Sweden, suggesting that sexual reproduction has been somewhat more important there than further north or south. Only the RAPD fragments characteristic of the EU race of G. abietina were found in the material tested. The RAPD pattern described as characteristic of the northern type within the EU race was identified in 62% of the isolates. Divergence from the expected profile was due to differences in occurrence of fragments OPA12-1400 and 12-1500. This indicates that this part of the RAPD profile cannot be treated as diagnostic for the northern type. A conclusion of practical importance is that there is a considerable risk of G. abietina spreading from infected P. contorta plantations to adjacent areas with indigenous P. sylvestris regeneration, and vice versa, owing to the indicated lack of host-specificity of the pathogen. It is possible, however, that host-specific strains exist, but do not differ in their RAPD profiles.     

马尾松2个世代种子园6年生家系生长的遗传变异与增益比较

【目的】研究与评价马尾松一代、二代无性系种子园的遗传增益及改良效果,指导马尾松良种选育、生产及应用。【方法】利用营建在浙江省淳安县燕山林场和福建省邵武市卫闽林场的6年生马尾松一代和二代自由授粉家系遗传测定林,分析不同世代种子园子代生长性状的遗传变异,并进行家系育种值估算及生长评价。【结果】马尾松不同世代的种子园子代生长差异很大,其树高、胸径和单株材积等生长量显著大于马尾松优良天然林分的混系子代,其中一代种子园子代各家系单株材积表型值和育种值分别大于优良天然林分混系55.78%~84.62%和40.66%~71.39%,二代其值分别大于优良天然林分混系58.96%~108.97%和42.97%~91.89%,二代种子园子代的平均生长量和遗传稳定性总体上高于一代种子园子代。树高、胸径和单株材积等主要生长性状在世代内家系间的生长差异均达极显著水平,在一代种子园子代中最优家系的树高、胸径和单株材积分别大于最差家系21.94%、30.72%和97.75%,在二代种子园子代中最优家系的树高、胸径和单株材积分别大于最差家系23.42%、29.60%和90.09%,说明种子园中对亲本材料的再选择非常重要。依据子代的单株材积育种值在两测定点的位次及其变幅大小,将38个家系分成速生型、较速生稳定型、较速生不稳定型和慢生型4类,各类所占家系数分别有23.68%、26.32%、21.05%和28.95%,其中速生型和较速生稳定型家系所对应的亲本无性系可作为新建马尾松二代无性系种子园的优良建园材料。种子园子代主要生长性状的地点×家系、区组×家系的互作效应极显著,说明各家系在不同立地环境中生长表现不一致。【结论】马尾松无性系种子园良种效益显著,二代增益大于一代。通过选择速生型和较速生稳定型家系将有助于种子园建园亲本的再选择、现有种子园的留优去劣疏伐和重建新的种子园。鉴于子代主要生长性状的基因型与环境互作使各家系在不同立地环境中生长表现不同,建议对子代生长表现突出、结实量较多的无性系进行单系采种利用。     

Magnitude and efficiency of genetic diversity captured from seed stands of <Emphasis Type="Italic">Pinus nigra</Emphasis> (Arnold) subsp. <Emphasis Type="Italic">pallasiana</Emphasis> in established seed orchards and plantations

To test how efficiently plantations and seed orchards captured genetic diversity from natural Anatolian black pine (Pinus nigra Arnold subspecies pallasiana Holmboe) seed stands, seed sources were chosen from 3 different categories (seed stands (SS), seed orchards (SO) and plantations (P)) comprising 4 different breeding zones of the species in Turkey. Twenty-five trees (mother trees) were selected from each SS, SO and P seed sources and were screened with 11 Random Amplified Polymorphic DNA (RAPD) markers. Estimated genetic diversity parameters were found to be generally high in all Anatolian black pine seed sources and the majority of genetic diversity is contained within seed sources (94%). No significant difference in genetic diversity parameters (numbers of effective alleles, % of polymorphic loci and heterozygosity) among seed source categories was found, except for a slight increase in observed heterozygosities in seed orchards. For all seed source categories, observed heterozygosity values were higher (H_o = 0.49 for SS, 0.55 for SO and 0.49 for P) than expected ones (H_e = 0.40 for SS, 0.39 for SO and 0.38 for P) indicating the excess of heterozygotes. In general, genetic diversity in seed stands has been transferred successfully into seed orchards and plantations. However, the use of seeds from seed orchards can increase the amount of genetic diversity in plantations further. The study also demonstrated that number of plus-tree clones (25–38) used in the establishment of seed orchards was adequate to capture the high level of diversity from natural stands.     

Comparison of Gremmeniella abietina isolates from Pinus sylvestris and Pinus contorta in terms of conidial morphology and host colonization

Gremmeniella abietina isolates from Pinus contorta in northern Sweden produced, in vitro, shorter conidia with fewer septa compared with isolates from Pinus sylvestris in the southern part of the country. After mycelial inoculation of shoots with G. abietina isolates from both host species, the resulting necroses were longer in P. sylvestris than in P. contorta. Keeping seedlings in artificial mild winter climate or detaching shoots from the seedling before inoculation caused longer necroses. No host specificity in colonization was found. Isolates from P. sylvestris caused longer necroses than did isolates from P. contorta, and both types of isolates caused longer necroses in P. sylvestris than in P. contorta. The differences found between the two G. abietina populations probably reflect regional variation in the fungus.     

马尾松实生种子园的遗传分析和育种值预测

利用５年生１３９个马尾松天然林优树自由授粉家系的测定材料,研究生长性状和雌、雄球花量的遗传变异,估算家系和家系内个体的育种值。该子代测定林是实生种子园的一部分。研究结果高、径生长和雌、雄球花量在家系间存在着极显著的差异,多数家系的生长最都大于当地的试验对照。性状的家系遗传力较高,而单株遗传力较低、雌、雄球花量所受的遗传控制与生长性状相近,遗传相关分析表明,开花性状似独立于生长性状。利用最佳线性预测     

Incidence of Gremmeniella abietina in planted seedlings of Picea abies and Pinus sylvestris in northern Finland

The ascomycete Gremmeniella abietina causes a disease (scleroderris canker) on conifers. The pathogen kills terminal shoots and branches of Pinus sylvestris and Picea abies, sporulates on host shoots and causes leader changes (i.e. dieback). Damage caused by G. abietina was investigated in young P. abies and P. sylvestris plantations in northern Finland. Side branches from seedlings of both species and the main stems of P. sylvestris were collected from three sites in two locations (Poksa 1 and 2, Kivalo) in northern Finland. The number of cankers and leader changes was counted in branches of each age (i.e. year) in a total of ca. 6300 shoots of P. abies and 1200 shoots of P. sylvestris. Cankers were common on both P. abies and P. sylvestris in Poksa 1 (2002–2008) and on P. abies in Poksa 2 (1998–2001). In Kivalo, cankers occurred only sporadically on P. sylvestris. Leader changes were most frequent in 1999–2009 in Poksa 1, in 2001–2009 in Poksa 2 and in 2003 in Kivalo. Cankers and other symptoms of infection were more obscure on P. abies than on P. sylvestris. According to both conidial morphology and molecular analysis, the strain of G. abietina infecting both P. abies and P. sylvestris was small‐tree type (STT). This is the first report of G. abietina, STT or B type, injuring P. abies in plantations.