Arne Steffenrem
Research Scientist
(+47) 916 70 420
arne.steffenrem@nibio.no
Place
Steinkjer
Visiting address
Innocamp Steinkjer, Skolegata 22, 7713 Steinkjer
Authors
Arne SteffenremAbstract
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Abstract
The complete diallel cross is the only mating design that provides estimates of variance components of general combining (GCA), specific combining ability (SCA), maternal and reciprocal effects, in addition to heritabilities and genetic correlations. To obtain such estimates, complete diallels were made among 10 trees in each of three natural Norway spruce populations from altitude 300 and 500 m in Norway. Nursery trials were performed with the families from these crosses and families from seeds collected from open pollination. Traits measured and analysed are seed weight, germination rate, germination percentage, terminal bud set, and seedling heights the first and second years. The seedlings from the population at origin 500 m had lower seed weight, lower heights and earlier bud set than those from the two populations from lower altitude. A considerable variation was present among families within each diallel, and the GCA variance components had the highest values and were significant for most traits. Variance components for SCA and maternal effects were also significant for some traits, but with inconsistent values in the three diallels. A strong relationship was present between the weight of the seed lots from the maternal parent and mean family height after one and two growing seasons. The highest estimate of heritability was observed for bud set, with similar values in all three diallels.
Abstract
The complete diallel cross is the only mating design that provides estimates of variance components of general combining (GCA), specific combining ability (SCA), maternal and reciprocal effects, in addition to heritabilities and genetic correlations. To obtain such estimates, complete diallels were made among 10 trees in each of three natural Norway spruce populations from altitude 300 and 500 m in southern Norway. Seedlings from families from the diallels and open pollinations were tested in short-term tests on agricultural soil at one site at altitude 85 m until age 10 years from seed. Tree height at ages 7 and 10 years and diameter at age 10 had strongly significant GCA variance components within each population. The components for SCA and maternal effects were small and not significant, indicating low levels of non-additive genetic variation. For the days of initiation and cessation of the shoot elongation period the GCA components were dominating and had the highest heritability estimates in two of the diallels. Estimates of genetic correlations between traits measured in earlier nursery trials and height and diameter in the short-term trials had low and not consistent values in the three diallels. The duration of the shoot growth period and rate of growth showed positive relationships with height and diameter. Strong relationships were present between half-sib family means from the diallels and open-pollinated families for height, diameter and phenology traits. Progeny trials testing open-pollinated half-sib families from natural populations can be used for selection of candidates for the initial breeding populations.
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Field trials with Norway spruce seedlings from 84 full-sib families from a factorial cross in Opsahl Seed Orchard and 11 provenances were planted at eight sites between altitudes between 600 and 900 m in Oppland County in Norway. Measurements of tree heights and assessments of stem and branch defects were made at regular intervals until 34 years from seed. Data from measurements made in nursery trials and from artificial freezing trials were also available. The families from the seed orchard had on average 12 % better height growth than the provenances. For volume growth per hectare, measured in two of the trials 30 years after planting, the families had a superiority of more than 30 %. A large variation among families was present for height growth and additive genetic variation was the main genetic factor. For the maternal half-sib families, the ranking of families for height was stable after 15 years from seed, and the five best families selected for height at that age were at age 34 years 6 % taller and produced 13 % more volume per hectare compared with the mean of all families. Weak relationships were present between traits measured in the nursery trial, the freezing test and the field trials. Assessments were made of cone production at age 20 years after planting and showed variation among families for the frequency of trees with cones.
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Artificial freezing tests were performed on seedlings from Norway spruce families at the end of the first growing season. Similar tests were made on twigs collected from trees in a progeny test at the end of growing season nine. The 26 families in the early test were included in the short-term progeny test with 100 full-sib families from a 10 x 10 factorial cross. All families were also planted in seven field trials in Norway, Sweden and Finland, from which data on mortality, tree heights and stem damage at age 10 years are available. Significant difference was found among families for freezing test injuries on whole intact seedlings at the end of the first growing season and for lethal temperature of needles on detached twigs collected at the end of growing season nine. However, no relationships were found between the freezing test scores of families in the two types of tests or few between these scores and the traits measured in the short-term and field trials. The results show that frost hardiness testing of families at a young age, grown under artificial temperature and light conditions in nursery, is a weak predictor of their performance under natural conditions in field at older ages.
Authors
Mateusz Liziniewicz Mats Berlin Thomas Solvin Henrik R. Hallingbäck Matti Haapanen Seppo Ruotsalainen Arne SteffenremAbstract
Norway spruce is a major industrial tree species in Fennoscandia and future productivity of the species must be secured by matching the variation in adaptation of the species with suitable sites for optimized performance. An appropriate transfer model for forest reproductive material (FRM) is crucial for regeneration of productive forests in the changing climatic conditions that are predicted to occur in Fennoscandia. We have developed a transfer model for prediction of height of Norway spruce in Norway, Sweden, and Finland, using data acquired from 438 progeny and provenance trials with 1919 genetic entries of local and transferred origins. Transfer of genetic material at a given site was expressed in terms of the difference in daylength (photoperiod) between the site and its origin. This variable best reflected the nonlinear response to transfer that has been commonly reported in previous studies. Apart from the transfer variable, the height prediction model included the age of material when height measurements were acquired, annual temperature sum over 5 °C, precipitation during the vegetation period, and interaction terms between test site and transfer variables. The results show that long northward transfers (4-5° latitude) seem to be optimal for relatively mild sites in southern parts of the countries where growing season is longer, and shorter northward transfers (2-4° latitude) for harsher northern sites with shorter growing seasons. The transfer model also predicts that southward transfers of Norway spruce would result in height growth reductions. The developed model provides foundations for development of common or national recommendations for genetically improving Norway spruce material in Fennoscandia.
Authors
Ishita Ahuja Inger Sundheim Fløistad Per Grande Øyvind Meland Edvardsen Adam O’Toole Arne SteffenremAbstract
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Arne SteffenremAbstract
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Arne SteffenremAbstract
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Norway spruce parents selected from results in progeny trials for testing Svenneby Seed Orchard clones were crossed in a factorial crossing design, and full-sib families were planted in short-term and field trials in southern Norway. Artificial freezing tests with the same families were made the first growing season and at ages 10 and 12 years. Offspring of parents selected for superior growth were taller in the field tests than those from parents with heights lower than the mean in the progeny test. However, the correlations between half-sib family performance in the initial progeny test and in the offspring field tests were only moderate. The variation among families in injury scores in the freezing tests was large, but the relationships with field trial performance were weak, indicating that frost hardiness testing at a young age is not valuable for predicting later field performance. Families with a late timing of bud flush had the highest frequency of injuries after a frost event at mid-summer. These families were also the tallest in the field tests. The families from crosses among Svenneby Seed Orchard clones had better height growth than commercial provenances. Foreldretrær av gran, valgt ut fra resultater i avkomforsøk for å teste kloner i Svenneby frøplantasje, ble krysset etter en faktoriell krysningsplan, og full-søsken familiene ble plantet i kortidsforsøk og i langsiktige feltforsøk. Fryseforsøk med familiene ble gjort etter første vekstsesong og ved alder 10 og 12 år. Avkom fra foreldre valgt ut for god høydevekst var høyere i feltforsøkene enn de fra foreldre som ble valgt ut med midlere høyde. Det var bare moderate korrelasjoner mellom høyder til halv-søsken i de første avkomforsøkene og i feltforsøkene. Det var betydelig variasjon mellom familiene i skader etter frysforsøkene, men sammenhengene var svake mellom frostskader og vekst og skader i feltforsøkene. Dette indikerer at resultater fra testing av frostherdighet i ung alder sier lite om hvordan trær fra familiene senere vokser i felt. Familier med sen skuddskyting fikk mest skader etter frostnetter 22.-23. juni. De sent skytende familiene hadde best høydevekst i feltforsøkene. Trærne fra familiene fra Svenneby frøplantasje var høyere enn de fra hadde handelsprovenienser.
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Arne SteffenremAbstract
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Genetic variation and phenotypic stability in Norway spruce were studied based on provenances, families, and clones planted in trials at 12 sites in four Nordic countries. The families were generated in a factorial cross between 10 parents of Norwegian origin and 10 parents of Eastern European origin, and the clones were propagated from seedlings within 20 of the same families. Traits analyzed were survival, proportion of trees with stem defects, and tree heights. Stability was analyzed by regression analyses with the genetic entries’ annual shoot increment as the dependent variable and the total site mean as an environmental index. Information about growth and phenology traits were available from short-term tests. For tree heights, significant variance components were present both among female and male parents, but not for their interactions, indicating that non-additive genetic effects are small. Genotype × environment interactions were significant at all three genetic levels, but their variance components had considerably lower values than the variance components estimated for the effects of families and clones. For the set of families of Norwegian origin, strong relationships were observed between the timing of annual shot elongation, mortality, and height growth. Large variation was found at all three genetic levels for phenotypic stability measured by regression coefficients. A positive relationship was present between the regression coefficient and the timing of annual shoot growth for families, indicating that later flushing families responded more to a high site index. The regression coefficient can be a useful supplement to the breeding value when selecting for superior and stable genotypes.
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Arne SteffenremAbstract
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Arne SteffenremAbstract
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Seedlings from seeds collected in three natural populations of Norway spruce (Picea abies (L. Karst)) in each of 21 provenances distributed between latitudes 63ºN to 66º20’N and altitudes from 25 to 500 m in Trøndelag og Nordland counties were tested in nursery trials and one long-term field trial. Large provenance variation was found for phenology traits and early height growth in the nursery trials. A strong clinal variation was found for these traits relative to latitude and altitude. These relationships were weaker for height and diameter at ages 26 and 40 years. The timing of bud flush was strongly related to the temperature conditions at the seed collection sites, whilst terminal bud set and lammas shoot percentages showed high correlation coefficients with the provenance latitude. Provenances in the same geographic region showed large differences in height and diameter Growth in the field trial. The long-term experimental site Spelrem is situated within the northern Natural range of Norway spruce and the general trend in this material is that provenances from the Northern part of the range perform better compared with provenances from more southern areas. Hence, the gain from provenance transfer seems to be limited under the present climatic conditions in this region.
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Temperature during seed maturation can induce an epigenetic memory effect in growth phenology of Norway spruce (Picea abies (L.) Karst.) that lasts for several years. To quantify the epigenetic modifications induced by natural climatic variation, common garden experiments with plants originating from different provenances and seed years were performed. Plants from warmer seed years showed delayed phenology with later bud flush, bud set and growth cessation. This effect was quantified by linear models of phenology traits as a function of climate indices for the origin and seed year of the plants. Significant effects of the temperature during seed production (seed year) was found for the bud set in seedlings in their first growing season and for bud flush and growth cessation in the 7th-8th growing season from seed. The models suggest that growth start and growth cessation are delayed 0.7–1.8 days per 100 additional degree days experienced by the seed during embryo development and seed maturation. Models that include factors that are known to induce epigenetic effects could be used to better predict future performance of forest reproductive material.
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Knowledge about spatiotemporal variability of climate change effect on tree-ring width (TRW) and crown condition is essential to optimize the modelling of future forest ecosystem responses to the changing climate. Geographical differences in the climate–growth relationship are a reflection of the regional climatic conditions mainly. In this study, 175 Picea abies trees from the north-western edge of its geographical distribution in Central Norway were evaluated with respect to geographical and age-dependent differences during the common period of 1950–2015. The results showed that the most significant positive correlations between TRW and the current June temperature were unstable although the temperature increased. The correlations suddenly started to decrease (regardless of the site placement and tree age) at the beginning of the 1990s, but subsequently unexpectedly increased in the 2010s. The superposed epoch analysis revealed longer TRW regeneration of the southern plots (except over-mature trees) after negative pointer years compared to the northern plots. Previous summer temperature and related physiological processes (cone crops, storage of nutrients, etc.) significantly negatively affected P. abies growth in the current year. Additionally, our results showed that the selection of the chronology version (standard or residual) significantly affects the resulting correlations and thus must be carefully considered in dendroclimatological studies. Our main outputs can contribute to better understanding of the climate–growth relationship variability and general prediction of the radial growth.
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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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There have been intense discussions about reducing fossil energy dependence for many years, indeed there is a vision to have a carbon-neutral energy system in Scandinavia by the year 2050. One way to address this ambition is to increase the use of woody biomass. This places a focus on forest tree breeding, since it is considered to be one of the most effective and environmentally friendly options to increase sustainable biomass production in our forests. In this report we have summarised information about forest tree breeding in Denmark, Finland, Norway and Sweden, covering historic, current and future activities. It includes estimates of realised genetic gain in volume and dry matter production on a regional basis for regeneration materials of different improvement levels, which are available today and will be available in the year 2050. Genetic gains in economic terms are also discussed, and basic breeding and mass propagation principles are described. The report includes the most relevant commercial forest tree species in Scandinavia: Scots pine, Norway spruce, Sitka spruce, Douglas fir, grand fir, birch, poplars, alder, oak and beech. Intensive and long-term breeding is carried out continuously in Sweden and Finland for Scots pine, Norway spruce and to some extent birch, while other species are worked with intermittently. In Norway the interest in breeding has, after a break of some years, now resumed, resulting in the initiation of a new breeding cycle for Norway spruce. The present resources put into breeding in Denmark are small, due to a major change in silvicultural management towards natural regeneration in the State Forests. However, as in all the other Scandinavian countries, there is great potential to increase growth through breeding. Seed from seed orchards is, and will within coming decades, be the main way to supply the forestry with genetically improved plant material. The extra gain in yield that is obtained by using material from existing seed orchards varies among species, but in general it is, on average, 10–15% when compared to local unimproved material. In 2050 it is estimated that this gain will be 20–25% for many species, which is a substantial increase in productivity. The time elapsed from establishment of a seed orchard to the first seed harvest for many species is quite long (10–20 years) and delays the realisation of the genetic improvement efforts. An alternative is to use plants obtained from the vegetative propagation of genetically well-performing seed sources or individuals, which makes it possible to capture the progress from breeding immediately. Utilisation of clones is also a way to reduce the consequences of limited amounts of seed from seed orchards, which for instance, is the case for Norway spruce in southern Sweden. For Norway spruce, using clonal material can immediately deliver a gain of around 25–35% in yield, and it is estimated that by 2050 this figure will have increased to 40%. It should be noted, however, that the use of clonal material may be limited by forest regulations. Future climate change will probably alter the growing conditions in Scandinavia in a way that makes forestry with high productivity exotic species such as Douglas fir, grand fir, Sitka spruce and poplars more attractive. Using several species, instead of the few traditional ones, is a way to spread the risk of an unknown future. In Denmark, several exotic species have been an integrated part of forestry for more than 100 years. Limited breeding work has been performed for these species and the gain in yield is, for instance, estimated to amount to 40% for existing seed orchard material of Sitka spruce compared to unimproved material.
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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.
Authors
Arne SteffenremAbstract
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Authors
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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Gunnhild Søgaard Inger Sundheim Fløistad Aksel Granhus Kjersti Holt Hanssen Harald Kvaalen Tore Skrøppa Arne SteffenremAbstract
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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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Authors
Ari Hietala Nina Elisabeth Nagy Arne Steffenrem Harald Kvaalen Carl Gunnar Fossdal Halvor SolheimAbstract
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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.
Abstract
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Abstract
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...
Abstract
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Abstract
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.
Abstract
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Abstract
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.
Abstract
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.
Abstract
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.
