Arne Steffenrem
Research Scientist
(+47) 916 70 420
arne.steffenrem@nibio.no
Place
Steinkjer
Visiting address
Statens Hus, Strandvegen 38, 7734 Steinkjer
Authors
Jørn Henrik Sønstebø Mari Mette Tollefsrud Tor Myking Arne Steffenrem Anne Eskild Nilsen Øyvind Meland Edvardsen O. Ragnar Johnskås Yousry A. El-KassabyAbstract
Seed from orchards, established from breeding programs, often dominate the planting stock in economically important tree species, such as Norway spruce. The genetic diversity in seed orchards’ crops depends on effective population size which in turn is affected by many factors such as: number of parents in the orchard, seed orchards’ design, fecundity, and pollen contamination. Even though seed orchards’ seed is extensively used over large regions, very few studies have addressed how well their crops reflect the genetic diversity present in the regions where they are planted. Here we have investigated the genetic diversity (by means of 11 microsatellites) of two Norway spruce seed orchard populations with different number of parents (60 and 25) and compared this with seed crops collected in the semi natural forest and natural unmanaged populations. We found that the ratio between the effective population size (N e ) and actual number of parents (N) varied between 0.60 and 0.76 in the orchards’ seedlots. A reduction in genetic diversity (mainly allelic richness) was detected in a few seedlots, mainly where the number of parents was low. Our results also show that pollen contamination play an important role in maintaining the genetic diversity in orchards’ seedlots, particularly when the number of parents is low. The population genetic structure among seed orhcards and natural populations is shallow suggesting that re- generation with seed from current seed orchards will have limited effect on the overall genetic diversity.
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Arne SteffenremAbstract
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Milan Lstibůrek Yousri A. El-Kassaby Tore Skrøppa Gary R. Hodge Jørn Henrik Sønstebø Arne SteffenremAbstract
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Arne SteffenremAbstract
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Igor A. Yakovlev Thomas Solvin Harald Kvaalen Arne Steffenrem Marcos Viejo Carl Gunnar FossdalAbstract
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The root rot pathogens in Norway spruce (Picea abies) Heterobasidion ssp. cause substantial loss in carbon sequestered in forest and economic revenue for forest owners. To facilitate strategic breeding planning for increased resistance against this pathogen in particular, the blue stain fungus Endoconidiophora polonica, growth and wood quality traits (wood density and spiral grain), we estimated additive genetic parameters, correlations and the potential response from selection. Parameters were estimated from a progeny trial series established at two sites (25 years from planting) and their parents in a seed orchard (43 years from grafting). A standard half-sib analysis based on progenies and a parent–offspring regression was used for estimation of heritabilities. Resistance against the pathogens was measured as lesion length under bark after inoculations in phloem. Heritability values varied with site and estimation procedure from 0.06 to 0.33, whereas the phenotypic variance (as CV P ) is high and fairly stable around 40–50 %. Heritability values for wood density and spiral grain in the same material varied from 0.32 to 0.63. The highest heritability values were generally obtained from parent–offspring regression. There is no evidence of resistance traits being genetically correlated with growth or wood quality traits. Wood density is negatively correlated with stem diameter. Implications for breeding are discussed.
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Plantations of genetically improved forest trees are critical for economic sustainability in forestry. This review summarizes gains in objective traits and the resulting economic impact of tree breeding programmes in Scandinavia and Finland. Genetic improvement of forest trees in these countries began in the late 1940s, when the first phenotypically superior plus-trees were selected from natural environments. The main findings from this review are that (i) tree breeding can increase volume growth in the range 10–25%, and (ii) the bare land value associated with genetically improved trees gives a better return on investment and a shorter rotation period compared to the unimproved forests. As some Nordic countries are quite dependent on the forest industry, breeding programmes that have resulted in economic gains have been beneficial for society. Growth and wood quality traits are often adversely correlated, and the weighting of traits from an economic perspective could provide an index for determining maximum profit from breeding. Tree breeding faces an array of challenges in the future, such as changes in silviculture, climate, new pests and diseases, and demand for wood-based products.
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Venche Talgø Jan-Ole Skage Arne Steffenrem Corina Junker Håvard Eikemo May Bente Brurberg Odd Ragnar JohnskåsAbstract
Delphinella shoot blight (Delphinella abietis) attacks true firs (Abies spp.) in Europe and North America. Especially subalpine fir (A. lasiocarpa), one of the main Christmas tree species in Norway, is prone to the disease. The fungus kills current year needles, and in severe cases entire shoots. Dead needles become covered with black fruiting bodies, both pycnidia and pseudothecia. Delphinella shoot blight has mainly been a problem in humid, coastal regions in the northwestern part of Southern Norway, but, probably due to higher precipitation in inland regions during recent years, heavy attacks were found in 2011 in a field trial with 76 provenances of subalpine fir in Southeastern Norway. However, the amount of precipitation seemed less important once the disease had established in the field. Significant differences in susceptibility between provenances were observed. In general, the more bluish the foliage was, the healthier the trees appeared. The analysis of provenance means indicated that, at least for the southern range, the disease ratings were correlated with foliage color. This study also includes isolation, identification, a pathogenicity test, a seed test and electron microscopy of the wax layer on the needles. The fungus was identified based on the morphology of spores and by sequencing the Internal Transcribed Spacer (ITS) regions of the ribosomal DNA. Koch’s postulates were fulfilled. The fungus was found present on newly harvested seeds and may therefore spread via international seed trade. When comparing the wax layers on green and blue needles, those of the latter were significantly thicker, a factor that may be involved in disease resistance.
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Arne SteffenremAbstract
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Use of genetic materials with a more “southern growth rhythm” has been suggested as one of the measures for adapting our forests to climate change. However, studies on Norway spruce (Picea abies (L.) Karst) provenances and families have shown a possible relationship between phenology (apical growth rhythm) and cambial growth rhythm that might have negative effects on latewood proportion and wood density. We made a detailed study of the xylem formation of four clones during one growth season. The clones were known to express contrasting phenology in terms of timing of bud flush equivalent to two weeks when assessed in 1997. Micro cores from four 20 year old ramets of the four clones, 16 trees in total, were sampled once a week from May to October in 2010. When bud flush were assessed in 2010 there were about one week difference between the most contrasting clones. Temperatures during the spring 2010 were low and flushing started in general late. No relationship was found between the clonal values for timing of bud flush and initiation of xylem formation. Large differences between clones in numbers of formed tracheids were found in later phases of the growing season. Both the rate of cell division and number of formed tracheids varied significantly between clones. Only small differences in latewood percentage were found between the clones. Genetic variation in xylem formation was found, but from this study the genetic variation in xylem formation seems to be independent from the genetic variation in phenology.
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Arne SteffenremAbstract
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Franҫois Lefèvre Jarkko Koskela Jason Hubert Hojka Kraigher Roman Longauer Ditte C. Olrik Silvio Schüler Michele Bozzano Paraskevi Alizoti Remigijus Bakys Cathleen Baldwin Dalibor Ballian Sanna Black-Samuelsson Dagmar Bednarova Sándor Bordács Eric Collin Bart De Cuyper Sven M.G. De Vries Thröstur Eysteinsson Josef Frýdl Michaela Haverkamp Mladen Ivankovic Heino Konrad Czeslaw Koziol Tiit Maaten Eduardo Notivol Paino Hikmet Öztürk Ivanova Denitsa Pandeva Gheorghe Parnuta Andrej Pilipovic Dragos Postolache Cathal Ryan Arne Steffenrem Maria Carolina Varela Federico Vessella Roman T. Volosyanchuk Marjana Westergren Frank Wolter Leena Yrjänä Inga ZarinaAbstract
Dynamic conservation of forest genetic resources (FGR) means maintaining the genetic diversity of trees within an evolutionary process and allowing generation turnover in the forest. We assessed the network of forests areas managed for the dynamic conservation of FGR (conservation units) across Europe (33 countries). On the basis of information available in the European Information System on FGR (EUFGIS Portal), species distribution maps, and environmental stratification of the continent, we developed ecogeographic indicators, a marginality index, and demographic indicators to assess and monitor forest conservation efforts. The pan-European network has 1967 conservation units, 2737 populations of target trees, and 86 species of target trees. We detected a poor coincidence between FGR conservation and other biodiversity conservation objectives within this network. We identified 2 complementary strategies: a species-oriented strategy in which national conservation networks are specifically designed for key target species and a site-oriented strategy in which multiple-target units include so-called secondary species conserved within a few sites. The network is highly unbalanced in terms of species representation, and 7 key target species are conserved in 60% of the conservation units. We performed specific gap analyses for 11 tree species, including assessment of ecogeographic, demographic, and genetic criteria. For each species, we identified gaps, particularly in the marginal parts of their distribution range, and found multiple redundant conservation units in other areas. The Mediterranean forests and to a lesser extent the boreal forests are underrepresented. Monitoring the conservation efficiency of each unit remains challenging; however, <2% of the conserved populations seem to be at risk of extinction. On the basis of our results, we recommend combining species-oriented and site-oriented strategies.
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We have recently found that Norway spruce (Picea abies (L.) Karst.) can rapidly adjust its adaptive performance, probably through an epigenetic mechanism. This appears to employ a kind of long-term memory of temperature sum and (probably) photoperiod from the time of its embryo development. In our research we made identical controlled crosses and produced seed lots under controlled temperature and day-length conditions and later observed phenology, growth and hardiness traits in the progenies. It was repeatedly found that temperature conditions during seed set, in particular, influence the phenotypes of the offspring; seedlings from seeds produced under warm conditions have later terminal bud set and reduced autumn frost hardiness than those from seed produced under colder conditions, and thus perform like a more southern provenance. When embryonic clones were derived from mature zygotic embryos and were cultured at different temperatures, the plants cultured under warm in vitro temperature were the last to set bud and grew taller than those cultured at lower temperatures. Progenies produced in Norway by Central European mother trees had a bud set curve skewed towards that of the local Norwegian performance. A comparison of the performance of seedlings from seeds collected in the same provenance regions in 1970 and 2006 shows that the more recent seed lots consistently produce taller seedlings with a later bud set, probably due to higher temperatures during seed production in 2006. The effect of reproductive environment has been shown to persist for years. It mimics the variation between provenances from different latitudes and altitudes and may explain much of the observed variability in bud set and early height growth between natural populations of Norway spruce. The observed phenomenon suggests an epigenetic mechanism in the developing embryo, either zygotic or somatic, that senses environmental signals such as temperature and influences adaptive traits. Research is underway to understand the molecular basis of this mechanism. We will discuss the implications of this epigenetic phenomenon for the interpretation of provenance differences, for tree breeding and for its possible role in adaptation to climate change.
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Genetic parameters were estimated for wood density and spiral grain in two long-term field trials with families of Picea abies (L.) Karst., and for microfibril angle (MfA) and model-predicted wood stiffness (MOEest) at one of the sites. The trials were located at 600-720 m altitude in Norway and the progenies, which were a sample of 13 half-sib families from plus-trees in a breeding population, were 33 years old from seed when measured. Significant genetic variation (p0.05) was found for all wood quality traits. The narrow-sense heritability was estimated to be 0.50 for density (across two sites), 0.38 for MfA, 0.29 for MOEest and 0.37 for spiral grain (across two sites). No significant genotype by environment interactions were found for density or spiral grain (p0.05). Genetic relationships between ring width and wood quality traits were negative for density and MOEest, and positive for MfA. Site index and competition had major effects on wood density and predicted MOEest but not on MfA and spiral grain.
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Minirhizotrons, transparent acrylic tubes inserted in the soil, are well suited for long term, non destructive, in situ observations of fine roots. In minirhizotrons, the fine roots are regularly photographed and the root images are visually evaluated according to their status as living, dead or disappeared. This evaluation gives the background for further statistical treatment to estimate the fine root longevity. It is inherent in the minirhizotron technique that a large group of roots will be described as “disappeared” due to grazing, overgrowing by other roots, unclear images or other reasons. Because the fraction of disappeared roots is substantial in some cases, this has consequences for the interpretation of the longevity results. We processed three years of minirhizotron images from Norway spruce stands in southeast Norway (30 yr old) and northern Finland (60 yr old). Of all processed fine roots 32 and 23% was evaluated as disappeared in Norway and Finland, respectively. When roots labelled as disappeared were pooled together with dead ones, the fine root longevity estimates, using the Kaplan Meier method, decreased almost by a factor of two (401 and 433 days), as opposed to labeling them as censored observations (770 and 750 days for Norway and Finland, respectively). Here we demonstrate how the early decision making on the fine root status bears consequences on the resulting longevity estimates. The implications will be discussed
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Progenies from a natural stand of Picea abies planted on a forest site with heterogeneous growth conditions were characterized for genetic and environmental control of internodal and whorl branch formation and lammas growth. The progenies studied were 27 years old from seed and planted in a randomized complete block experiment. Internodes and whorls studied were located in the top section of the trees. Significant genetic variation (p<0.01) was found for a number of internodal and whorl branches. Block means for the length of internodes, a strong indicator for the site index in respective blocks, were correlated strongly with branch dimensions (r >= 0.90), intermediately with the frequency of lammas growth (r = 0.81) and number of branches in whorls (r = 0.68), and only weakly with the number of internodal branches per length unit of internode (r = 0.25). A strong environmental effect on branch dimensions was supported by the relative size of variance components. The results show that the environmental effect expressed by site index dominates the control of branch dimensions, while the number of branches is under stronger genetic control in P. abies.
Authors
Arne SteffenremAbstract
Wood based products come from a renewable resource playing an important part of the carbon cycle and are therefore regarded environmentally friendly compared to many other alternatives. We might therefore expect a more intensive use of the timber resource in the future. An increase in the supply and net value of timber products from a given area requires more intensive and cost efficient forest management. However, a faster growth rate in general and an increase in the juvenile wood proportion specifically, might come in conflict with the desirable wood quality. Tree breeding programs for Norway spruce (Picea abies L. Karst.) aim to follow up the general demands by providing forestry with plant materials that have superior growth and high wood quality. It is therefore important to establish a better knowledge of the potential of improving wood quality characters in this species through breeding. Characteristics important for structural strength and dimensional stability in conifers are wood density, spiral grain, microfibril angle (MfA) in the S2 layer in the secondary cell wall, branch characteristics and stem straightness. We have studied the genetic variation and correlation patterns for these traits and the direct end-use performance of clearwood samples in terms of stiffness (MOE) and strength (MOR) in four studies of Norway spruce (Paper I-IV). Wood quality traits of defect free wood were studied in Paper I and II. The materials used were 28 and 29 year old progeny trials planted in South East Norway on fertile soils. Significant genetic variation was found for wood density, MfA and spiral grain in both studies (p < 0.05). MOE and MOR, measured by static bending, were studied in one of the materials and the genetic variation was found to be highly significant (p < 0.01). There was also significant genetic variation (p < 0.05) in MOE estimated (MOEest) from wood density and the x-ray diffractometry pattern from the S2 layer obtained by SilviScan®-3. Genetic parameters were estimated in Paper II with large standard errors. The parameters must therefore be used with caution and preferably with support from other studies. We could confirm earlier studies of high genetic variation and low genotype by environment interactions (GxE) for wood density and spiral grain. Parameter estimates for MfA and MOEest, from one site, suggest that these are under moderate to strong genetic control in Norway spruce. Phenotypic correlations between growth traits (height and diameter growth) and wood quality traits were negative for wood density, positive for MfA and negative for MOEest. Genetic relationships showed the same trends for diameter growth, but not for height growth. The genetic and environmental variation in branch characteristics (Paper III and IV) were studied in three sets of progeny trials that were 22, 28 and 29 years old from planting. Stem straightness was studied in the 22-year old trial only. The branch diameter was highly influenced by site index and spacing. Number of branches formed and stem straightness seems to be under stronger genetic control. Individual tree heritabilities for growth traits, branch diameter and stem straightness were estimated in the 22-year old progeny trial planted at two sites. The trials were planted with different spacing, which allowed us to study genotype by spacing interactions. Heritability for growth traits and branch diameter across sites were moderate (0.14 - 0.19) compared to the higher values found for stem straightness (0.28). GxE, and thereby genotype by spacing interactions were not important for any of the traits. Heritability for number of branches, estimated from one of the sites, was 0.24. There was a strong and positive genetic correlation between growth traits and branch diameter. The presence of genetic variation for most wood quality traits suggests that these can be improved by tree breeding. However, genetic correlations with growth traits indicate that selection solely for growth might have adverse effects on wood quality. It is therefore important to balance the gain in growth and wood quality. Implementation of several adversely correlated traits in a breeding program reduces its efficiency. It is therefore essential that tree breeding, silvicultural practices and forest industrial needs interact when developing tree breeding programs to obtain an optimal strategy for selection and utilization of improved plant materials.
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We investigated whether the stand age affects the life span of tree and understory fine roots (<1mm) in three Norway spruce (Picea abies) stands: 30, 60 and 120-yr-old. In each stand 9 minirhizotrons were installed and images were collected once in a month throughout the growing season during the three years. Norway spruce fine roots in the 30-yr old stand had a life span 401 ± 27 and 341 ± 68 days, and understory 409 ± 162 and 349 ± 142 days, estimated by using the Kaplan Meier survival analysis (KM) and Weibull distribution, respectively...
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We present results from early tests and field trials of offspring from two Norway spruce (Picea abies (L.) Karst.) seed orchards containing clones that have been transferred from high altitudes to sea level and from northern to southern latitudes. Seedlings from seeds produced in the low-altitude seed orchard developed frost hardiness later at the end of the growth season, flushed later in field trials, and grew taller than seedlings from seeds produced in natural stands. They had the lowest mortality rate and the lowest frequency of injuries in the field trials. Similar results were observed in seedlings from seeds produced in the southern seed orchard. We found no adverse effects of the changed growth rhythm. Seedlings from two seed crops in the southern orchard, produced in years with a warm and a cold summer, had different annual growth rhythms. The results are explained mainly by the effects of the climatic conditions during the reproductive phase. Seed crops from different years in the same seed orchard may produce seedlings that perform as if they were from different provenances. It is argued that the effects of the climatic conditions during seed production must contribute to the variation among provenances of Norway spruce.
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Genetic- and environmental variation and correlation patterns were characterized for modulus of elasticity (MOE), modulus of rupture (MOR) and related wood traits: latewood proportion, wood density, spiral grain, microfibril angle and lignin content in five full-sib families of Norway spruce.The families were evaluated on the basis of clearwood specimens from the juvenile -mature wood transition zone of 93 sampled trees at age 30 year from seed. Family-means varied significantly (p 0.05) for all wood traits studied except lignin content. MOE varied between 7.9-14.1 GPa among trees and 9.4-11.0 GPa among families. MOR varied between 47-87 MPa among trees and 61-71 MPa among families.Families remained significantly different in an analysis of specific MOE (MOE/density) and MOR (MOR/density). Hence, solely relying on wood density as a wood quality trait in tree breeding would not fully yield the potential genetic gain for MOE and MOR. Correlations between wood structural traits and specific MOE and MOR are presented and discussed.
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Fine root production, respiration, longevity and mortality are the major processes in carbon dynamics of the forest soils. The objective of the present work was to determine fine root biomass, respiration and root longevity. The study was carried out at a ten year-old stand of planted Norway spruce (Picea abies) (a clearcut, dominated by natural regrowth of Scots pine and birch) and three stands of Norway spruce, approximately 30, 60 and 120 years old, during 2001 and 2002. The stands were located at Nordmoen, a plain of sandy deposits in southeast Norway.Root biomass of both trees and understorey vegetation (0-1, 1-2 and 2-5 mm in diameter) in the humus layer and mineral soil horizons (to depth of 60 cm) was sampled by soil coring. Root respiration was performed in situ, by measuring the CO2 of excavated fine roots by using the CIRAS-I portable gas analyser. For the root turnover study, altogether 60 minirhizotrones were installed and images were processed. Root biomass and necromass (g m-2), specific root length (SRL, m g -1), root length density (RLD, cm cm-3), number of root-tips and mean longevity (y) were estimated.Root biomass was 2-3 times higher in the mineral soil than in the humus horizon. Compared with other stands, root biomass, SRL, RLD and the number of root tips were highest in the 30-year-old stand. At the 10 and 120 year-old stands understorey vegetation roots counted for 70 and 40% of total root biomass, respectively. The amount of necromass at 60 year-old stand was about twice as high (45%) compared to other stands.Root respiration (g C/min./g roots) was significantly lowest at 10-year-old stand. Root respiration among 30, 60 and 120 year-old stands was not significantly different, but it was highest in the 60-year-old stand. The respiration varied seasonally, with high peaks during the summer and lower values during the spring and autumn. Fine root longevity of tree and understorey roots at the 10-year-old stand were 1.2 and 1.4 years, respectively.It is concluded that stand age may influence the dynamics of the fine roots. The complexity of influences will be discussed.
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The minirhizotron technique provides the opportunity to perform in situ measurements of fine root dynamics and obtain accurate estimates of fine root production and turnover. The objective of the present work was to determine the fine root longevity and mycorrhization in a Norway spruce chronosequence. The study was carried out on four stands of planted Norway spruce (Picea abies), approximately 10, 30, 60 and 120 years old, during 2001 and 2002. The stands were located at Nordmoen, a plain of sandy deposits in southeast Norway (60o15 N, 11o06 E). For the root turnover study, altogether 60 minirhizotrones were installed and images were processed.Individual fine roots were identified, their mycorrhization assessed, appearance and possible disappearance dated, and growth in length measured. The data set was subjected to a survival analysis, using a Kaplan-Meier product-limit approach. The minirhizotron samples were stratified according to stand age class, and Coxs F-test was used to analyze differences in survival estimates. The analysis may also be extended to consider other covariates such as tree species (spruce, pine or birch), understory vegetation, or soil depth. Typical survival function estimates will be presented, and the influence of stand age on the mycorrhization and the dynamics of the fine roots will be discussed.