Optimum contribution selection in large general tree breeding populations with an application to Scots pine

Development of selection methods that optimises selection differential subject to a constraint on the increase of inbreeding (or coancestry) in a population is an important part of breeding programmes. One such method that has received much attention in animal breeding is the optimum contribution (OC) dynamic selection method. We implemented the OC algorithm and applied it to a diallel progeny trial of Pinus sylvestris L. (Scots pine) focussing on two traits (total tree height and stem diameter). The OC method resulted in a higher increase in genetic gain (8–30%) compared to the genetic gain achieved using standard restricted selection method at the same level of coancestry constraint. Genetic merit obtained at two different levels of restriction on coancestry showed that the benefit of OC was highest when restriction was strict. At the same level of genetic merit, OC decreased coancestry with 56 and 39% for diameter and height, respectively, compared to the level of coancestry obtained using unrestricted truncation selection. Inclusion of a dominance term in the statistical model resulted in changes in contribution rank of trees with 7 and 13% for diameter and height, respectively, compared to results achieved by using a pure additive model. However, the genetic gain was higher for the pure additive model than for the model including dominance for both traits.     

Biomass functions applicable to Scots pine

This study describes parameterization of biomass functions applicable to Scots pine (Pinus sylvestris, L.) in the conditions of Central Europe. Fifty-two sample trees from seven sites in different regions of the Czech Republic were used for destructive measurements. The observed aboveground biomass (B _AB) and its individual components were examined by different types of non-linear regression models using one to five independent variables including stem diameter (D), tree height (H), tree age (A), crown length (C) and altitude (Z). The best single-equation approximation of B _AB was a three-parameter model using D, H and Z, which was also best suited for bark. Including altitude into classical multiplicative exponential function with three independent variables (DHZ) and four parameters yielded the best model for stem over or under bark. Age was important for assessment of dead branches within DHA model, whereas living branches were best approximated using four variables including C (DHAC model). The most complex model (DHACZ) worked best for needle biomass. The comparison of B _AB assessment by single regression equation and from sum of individual components showed a negligible difference for full-grown trees, whereas the additive assessment yielded considerably higher B _AB for small or young trees. This corresponds to the assessed confidence intervals for individual trees that were larger for smaller trees. The paper also discusses the issue of commonly used linearization of biomass equations and application of linear regression, and provides comparative examples with nonlinear approach used here. The paper presents the parameter sets for the tested equations for B _AB and individual biomass components.     

Increased contribution of wheat nocturnal transpiration to daily water use under drought

Increasing evidence suggests that in crops, nocturnal water use could represent 30% of daytime water consumption, particularly in semi‐arid and arid areas. This raises the questions of whether nocturnal transpiration rates (TR_N) are (1) less influenced by drought than daytime TR (TR_D), (2) increased by higher nocturnal vapor pressure deficit (VPD_N), which prevails in such environments and (3) involved in crop drought tolerance. In this investigation, we addressed those questions by subjecting two wheat genotypes differing in drought tolerance to progressive soil drying under two long‐term VPD_N regimes imposed under naturally fluctuating conditions. A first goal was to characterize the response curves of whole‐plant TR_N and TR_N/TR_D ratios to progressive soil drying. A second goal was to examine the effect of VPD_N increase on TR_N response to soil drying and on 13 other developmental traits. The study revealed that under drought, TR_N was not responsive to progressive soil drying and – intriguingly – that TR_N seemingly increased with drought under high VPD_N consistently for the drought‐sensitive genotype. Because TR_D was concomitantly decreasing with progressive drought, this resulted in TR_N representing up to 70% of TR_D at the end of the drydown. In addition, under drought, VPD_N increase was found not to influence traits such as leaf area or stomata density. Overall, those findings indicate that TR_N contribution to daily water use under drought might be much higher than previously thought, that it is controlled by specific mechanisms and that decreasing TR_N under drought might be a valuable trait for improving drought tolerance.     

Field studies of water relations and photosynthesis in Scots pine.

Needle water potential at noon and diurnal variation in needle water potential was measured with a pressure chamber during the growth seasons (1974, 1975 and 1976) in a 20-year-old stand of Scots pine (Pinus sylvestris L.), subjected to daily irrigation and nutrient treatments at Jädraås, in central Sweden (60°50'N).
In unstressed conditions there was little difference in water potential amongst the treatments. However, during a dry spell in 1976 the diurnal curves of water potential in the control and combined irrigated and fertilized treatment diverged by up to 6 bar in the middle of the day. Irrigation alone resulted in higher midday water potentials (Ψ_n) than in the controls only during this period. At other times in 1974, 1975 and early 1976, the values of Ψ_n in the irrigated treatment were similar to the controls. Fertilization alone resulted in higher Ψ_n than in the controls in both 1975 and 1976 during both wet and dry periods. However, the largest difference between treatment and control occurred in the combined, irrigated and fertilized treatment during the dry spell in 1976.
Several possible explanations for the effect of fertilization on leaf water potential are discussed, including changes in hydraulic conductivity and water storage. The most likely explanation is a reduced transpiration rate as a result of more effective stomatal control of water loss.     

Recovery of photosynthesis in winter-stressed Scots pine

Abstract. . Winter-induced inhibition of photosynthesis in Scots pine (Pinns sylvestris L.) is caused by the combined effects of light and freezing temperatures; light causes photoinhibition of photosystem II (Strand & Oquist, 1985b, Physiologic Plantarum, 65 , 117–123), whereas frost causes inhibition of enzymatic steps of photosynthesis (Strand & Öquist, 1988, Plant, Cell & Environment, 11 , 231–238). To reveal limiting steps during recovery from winter stress, the potential of photosynthesis to recover and the actual recovery outdoors during spring, were studied in Scots pine. Studies of light dependent O₂-evolution under saturating CO₂ and recordings of room temperature fluorescence induction kinetics were used. When branches of pine, in February and March, were brought into the laboratory and kept at 18°Cand 100μmol m^?2 s^?1, light saturated rates and apparent quantum yields of photo-synthetic O₂-evolution recovered fully within approximately 48h. The photochemical efficiency of photosystem II, as measured by Fv/Fm ratios, recovered fully within 24h after an initial lag-phase of 2-3 h. Under natural winter conditions, the Fv/Fm ratio decreased more in exposed than in shaded pine, whereas the efficiency of photosynthesis was similarly inhibited in exposed and shadedpine. However, when recovery from winter stress occurred during spring, the Fv/Fm ratios of both shaded and exposed pine recovered well before photosynthesis. It is concluded that the light-induced photoinhibition component of winter stress in photosynthesis of pine recovers well before the frost induced component(s) of winter stress. In this context, reversible photoinhibition of photosynthesis in evergreen conifers is considered as a dynamic down-regulation of photosystem II to prevent more severe photodynamic damage of the thylakoid membrane when photosynthesis is inhibited by frost.     

Effects of water and nutrient applications in a Scots pine stand to tree growth and nutrient cycling

In a Scots pine forest stand, demineralized water and a complete set of nutrients with water were applied to the soil by means of frequent irrigation for four years in order to eliminate water and nutrient shortage of the trees. Apart from this optimization, dissolved (NH₄)₂SO₄ was irrigated at a rate of 120 kg N ha^-1 y^-1 to create a situation of N excess. Effect of treatments on tree growth and chemical composition of soil water and vegetation were monitored. From the first treatment year onwards basal area growth increased by ca. 35% as a result of the increased water supply. Nutrient applications increased K and P concentrations in pine needles immediately, but growth was enhanced only in the fourth treatment year and coincided with an improved K supply. Most of the applied P and K was retained in the soil, and only 6% was recovered in the vegetation. Tree nutrient status did not respond on Ca and Mg applications, whereas Ca and Mg seepage losses were increased with ca. 5 kg ha^-1 y^-1. The applied NH₄ was mostly retained in the 0–20 cm surface soil and caused a drastic increase of Al in soil solution. Tree growth was stimulated initially by extra NH₄, but was hampered after three years obviously because of a decreased P nutrition. The applied base cations were absorped to the soil and the accompanying anions were leached, thus temporarily increasing the acidification of the soil solution.     

Mating dynamics of Scots pine in isolation tents

Seed orchards are forest tree production populations for supplying the forest industry with consistent and abundant seed crops of superior genetic quality. However, genetic quality can be severely affected by non-random mating among parents and the occurrence of background pollination. This study analyzed mating structure and background pollination in six large isolation tents established in a clonal Scots pine seed orchard in northern Sweden. The isolation tents were intended to form a physical barrier against background pollen and induce earlier flowering relative to the surrounding trees. We scored flowering phenology inside and outside the tents and tracked airborne pollen density inside and outside the seed orchard in three consecutive pollination seasons. We genotyped 5683 offspring collected from the tents and open controls using nine microsatellite loci, and assigned paternity using simple exclusion method. We found that tent trees shed pollen and exhibited maximum female receptivity approximately 1 week earlier than trees in open control. The majority of matings in tents (78.3 %) occurred at distances within two trees apart (about 5 m). Self-fertilization was relatively high (average 21.8 %) in tents without supplemental pollination (SP), but it was substantially reduced in tents with SP (average 7.7 %). Pollen contamination was low in open controls (4.8–7.1 %), and all tents remained entirely free of foreign pollen. Our study demonstrates that tent isolation is effective in blocking pollen immigration and in manipulating flowering phenology. When complimented with supplemental pollination, it could become a useful seed orchard management practice to optimize the gain and diversity of seed orchard crops.     

Quantitative characterization of clumping in Scots pine crowns

Background and Aims

Proper characterization of the clumped structure of forests is needed for calculation of the absorbed radiation and photosynthetic production by a canopy. This study examined the dependency of crown-level clumping on tree size and growth conditions in Scots pine (Pinus sylvestris), and determined the ability of statistical canopy radiation models to quantify the degree of self-shading within crowns as a result of the clumping effect.

Methods

Twelve 3-D Scots pine trees were generated using an application of the LIGNUM model, and the crown-level clumping as quantified by the crown silhouette to total needle area ratio (STAR_crown) was calculated. The results were compared with those produced by the stochastic approach of modelling tree crowns as geometric shapes filled with a random medium.

Key Results

Crown clumping was independent of tree height, needle area and growth conditions. The results supported the capability of the stochastic approach in characterizing clumping in crowns given that the outer shell of the tree crown is well represented.

Conclusions

Variation in the whole-stand clumping index is induced by differences in the spatial pattern of trees as a function of, for example, stand age rather than by changes in the degree of self-shading within individual crowns as they grow bigger.     

Abiotic factors modulate post-drought growth resilience of Scots pine plantations and rear-edge Scots pine and oak forests

The proportion of planted forests in the Mediterranean Basin is one of the largest in the world. These plantations are dominated by pine species and present a series of characteristics such as low elevation, high competition or small tree size that make them more vulnerable to droughts. However, quantitative assessments of their post-drought growth resilience in accordance with species, site factors and tree characteristics are lacking. In this study we sampled 164 trees at four forest sites located in the drought-prone Sierra Nevada, southeastern Spain. We compared growth responsiveness to drought in rear-edge planted vs. relic natural Scots pine (Pinus sylvestris) and coexisting Pyrenean oak (Quercus pyrenaica) stands. Our objective was to characterize and compare the different growth responses to drought between species and sites and the effect of the main physiographic factors (altitude, aspect, and slope) on these responses since the influence of these factors on post-drought resistance and resilience has received little attention to date. Our results reveal that the planted pine sites with the lowest mean growth rates displayed greater resistance during drought, and that higher altitude was associated with improved resistance and/or resilience for all species and sites. Natural pine and Pyrenean oak stands were better adapted to the dry climatic conditions of the Mediterranean region where the study was undertaken, displaying greater resistance and/or resilience and lower influence of drought on growth in comparison to stands of planted pines. These results suggest that promoting the conservation of high-elevation pine plantations and enhancing the regeneration of natural pine and oak may improve the resistance and resilience of these drought-prone forest ecosystems.     

Sapwood water storage: its contribution to transpiration and effect upon water conductance through the stems of old-growth Douglas-fir

Abstract Enough water is stored in the sapwood of large Douglas-fir to significantly contribute to transpiration. Sapwood water content falls through the season, causing the wood's conductivity to fall. This leads to low leafwater potentials, stomatal closure, and reduced photosynthesis by the trees. The amount of water stored in the sapwood of Douglasfir 50-60 m tall, growing in the Cascade Mountains of Oregon, was estimated periodically over two seasons from measurements of sapwood relative water content (R_s). The relationship between R_s and volume of water contained in the sapwood was determined in the laboratory, and an equation describing the variation of relative conductivity (K) with R_s was derived from the literature. Stomatal conductance (k_s) and leaf water potentials were measured in the field. The relative conductivity of the sapwood was calculated from estimates of the flow rate through the tree and differences in water potential between dawn and the time of comparison. Flow rate was assumed to equal transpiration rate, calculated from the Penman-Monteith equation using measured k_s values. A sixfold decrease in K during the summer was attributed to changes in R_s. The maximum observed diurnal variation in K would require a change in R_S estimated at 25%. About 270 m³ ha^?1 (27 mm) of water were stored in sapwood, and 75% of that was in the stemwood. Withdrawal from this store reached 1.7 mm day^?1 on clear days after cloudy or rainy weather. Recharge could be almost as fast (up to 1.6 mm day^?1) after rain, but was very slow if the foliage remained wet.     

Seasonal development of phloem in Scots pine stems

The formation of phloem was studied for two years in stems of 50 to 60 year old trees of Scots pine (Pinus sylvestris L.) growing in nature. The development of phloem of the current year begins 10 to 20 days before the xylem formation and is completed with the termination of shoot growth in the end of June. Observations over the seasonal activity of cambium producing sieve-like cells of phloem and duration of their differentiation as compared to the xylem derivatives of cambium have shown that the maxima of formation of phloem and xylem cells could coincide or not coincide by season, while the activities of their differentiation were always at antiphase. The sieve-like cells of early phloem were separated from those of late phloem by a layer of tannin-containing cells, which are formed simultaneously with the formation of late xylem cells by the cambium. Seasonal dynamics of accumulation of starch grain in structural elements of the phloem is related to the xylem development. The content of metabolites in differentiating and mature phloem elements, in the cambium zone, and in the xylem cells growing in the radial direction depended on cell specificity, stage of their development, and type of forming wood, early or late, which differ in the cell wall parameters and, hence, requirement of assimilates. Significant differences were described between the content of low molecular weigh carbohydrates, amino acids, organic acids, and phenol compounds using two methods of calculation: per dry weight and per cell.     

Evaporative demand determines branchiness of Scots pine

Analysis of the branch area/stem area ratio of Scots pine growing in different climatic conditions in Europe and Siberia indicates that the branch area supported by a stem increases in warmer and drier conditions. The ratio was significantly correlated with several climatic variables, especially with potential evapotranspiration (E _p). The ratio was negatively correlated with stand density (d _s). A regression model combining E _p and d _s accounted for 85% of the total variation. These trends are believed to reflect hydraulic segmentation of trees and may represent a strategy to avoid cavitation in the tree, especially in the branches.     

Cryopreservation of embryogenic cultures of Scots pine

The aim of the study was to develop an effective cryopreservation method for Scots pine (Pinus sylvestris L.) embryogenic cultures. Altogether nine cell lines derived from three mother trees were cryopreserved after cold hardening using dimethylsulfoxide or two different mixtures of polyethyleneglycol 6000, glucose and dimethylsulfoxide as cryoprotectants. Seventy-eight percent of the cell lines remained viable after cryostorage, the best cryoprotectant treatment being 10% polyethyleneglycol 6000, 10% glucose, and 10% dimethylsulfoxide in water. This treatment resulted in significantly better regrowth of the embryogenic cultures than with the other cryoprotectants or with the controls. According to microscopical observations, the cells that retained their viability and regrowth ability after cryopreservation were the embryonal head cells, as well as some elliptic suspensor cells close to the embryonal head cell area. When proliferation growth of the frozen cultures had started, their morphological appearance was the same as the non-frozen cultures. In addition, the RAPD assays suggested that the cryostorage treatment used here preserved the genetic fidelity of the Scots pine embryogenic cultures. This revised version was published online in June 2006 with corrections to the Cover Date.     

A hydrological tracer study of water uptake depth in a Scots pine forest under two different water regimes

Little is known about the vertical distribution of water uptake by trees under different water supply regimes, the subject of this study, conducted in a Scots pine stand on sandy loam in northern Sweden. The objective was to determine the water uptake distribution in pines under two different water regimes, desiccation (no precipitation) and irrigation (2?mm day^?1 in July and 1?mm day^?1 in August), and to relate the uptake to water content, root and soil texture distributions. The natural ¹⁸O gradient in soil water was exploited, in combination with two added tracers, ²H at 10?cm and ³H at 20?cm depth. Extraction of xylem sap and water from the soil profile then enabled evaluation of relative water uptake from four different soil depths (humus layer, 0–10, 10–25 and 25–55?cm) in each of two 50-m² plots per treatment. In addition, water content, root biomass and soil texture were determined. There were differences in vertical water uptake distribution between treatments. In July, the pines at the irrigated and desiccated plots took up 50% and 30%, respectively, of their water from the upper layers, down to 25?cm depth. In August, the pines on the irrigated plots took up a greater proportion of their water from layers below 25?cm deep than they did in July. In a linear regression, the mean hydraulic conductivity for each mineral soil horizon explained a large part of the variation in relative water uptake. No systematic variation in the residual water uptake correlated to the root distribution. It was therefore concluded that the distribution of water uptake by the pines at Åheden was not a function of root density in the mineral soil, but was largely determined by the unsaturated hydraulic conductivity.     

Freeze-preservation of buds from Scots pine trees

A procedure has been developed for freeze-preservation of buds of the Scots pine (Pinus sylvestris L.). Instead of liquid nitrogen, cold storage in –80°C was used. The partly dormant material used in the experiments was obtained directly from a natural stand in Northern Finland and no prefreezing or cryoprotectants for preconditioning were used. Cooling velocity was 1°C/min up to a terminal freezing temperature of –39°C, after which the buds were immersed in liquid nitrogen at –196°C for 10 minutes. The material was then transferred to a deepfreezer at –80°C and stored up to 6 months. After rapid thawing, the buds were sterilized and their viability was tested by FDA staining and by culturing meristems on 1/2 MS medium for at least two weeks. All the freezing experiments were performed during March and April. The best survival of buds (90–100%) was achieved at the beginning of April, after which a pronounced decline in survival occurred obviously due to a rise in the water content of the buds.     

Structural adaptation rather than water conservation was observed in Scots pine over a range of wet to dry climates

We investigated the functional and structural responses of Scots pine to climate and estimated the importance of the genotype on the traits studied. We analysed ¹³C isotope discrimination (Δ¹³C) of various provenances in a common garden experiment and gas exchange characteristics for provenances growing in their natural environment. No clear climatic trend was found in the foliar Δ¹³C values of common garden trees. Similar results were obtained from estimation of λ (a largely VPD, temperature and light independent measure of intrinsic water use efficiency) from the gas exchange data. The ratio of needle mass to unit stem area and branch area to stem area increased towards south in both experiments and hence, seemed to be genetic. Trees from drier and warmer conditions seemed not to have either lower needle mass or higher intrinsic water use efficiency compared to northern latitudes. Received: 6 November 1998 / Accepted: 1 July 1999     

Stress reactions of Scots pine trees to injuring by ground fire

Controlled burning has been carried out to stimulate moderate ground fire in the Scots pine (Pinus sуlvestris L.) stand (age class I) in the Krasnoyarsk forest steppe at the beginning of the growing season. The structure of annual rings and trophic supply, as well as the cambial zone antioxidant system condition and the content of photosynthetic pigments in the needles of dominant and suppressed trees in the experimental and control plots, were studied 3 h, 5 days, and 2.5 months after the fire. The fire narrowed the annual ring width, since the number and size of early and late tracheids decreased. The primary reaction to pyrogenic heat shock (sharp reduction of chlorophyll content in needles and manifestation of oxidative stress in the stem cambial zone) was an elevated level of hydrogen peroxide and activity of most antioxidant enzymes and a reduction in the nonstructural carbohydrates pool. Restoration of the cambium function after the fire was promoted by an increased activity of amylase, invertase, and peroxidase.     

The effects of ozone and acid mist on Scots pine saplings

Summary It has been suggested that the forest decline (Neuartige Waldschäden) seen recently in parts of West Germany is due to the direct effects of ozone combined with acid mists, rather than soil-mediated effects of acid deposition. It has been proposed that ozone (a) makes the needles of affected conifers more susceptible to leaching by acid mist and (b) damages the photosynthetic apparatus, giving rise to diminished carbohydrate reserves which reduce the ability of affected trees to replace the leached nutrients. This nutrient deficiency (especially of Ca and Mg) is a characteristic symptom of the Waldschäden, which progresses through growth decline, needle loss, and eventually death. Parts of this hypothesis were tested in a preliminary experiment in which 3-year old Pinus sylvestris (Scots pine) saplings were exposed to 4 different O₃ levels, with and without acid mist (pH 3) treatment, for 56 days between July and September, 1983 in outdoor solardome fumigation chambers. The visual symptoms observed at >100 g m^-3 were more characteristic of the chlorotic mottle seen on O₃-affected trees in the USA than the general chlorosis of affected stands in Germany. O₃ at mean concentrations of >200 g m^-3 for 56 days reduced the fine root biomass and accelerated the senescence of older needles, in keeping with field effects observed in Germany. However, these O₃ levels increased, rather than decreased, the concentrations of most elements in the needles. Acid mist had no effect on needle concentrations, and there was no O₃-acid mist interaction. O₃ up to 300 g m^-3 also had no effect on the amount of ions leached from the needles, whereas acid mist increased the leaching of some ions, and again there was no interaction. The only nutritional effect of O₃ was to reduce the foliar uptake of NO _- ³ from the acid mist solution. An aphid infection part way through the experiment caused a large increase in leaching, particularly of K, and affected the intermediate O₃ and watersprayed plants most. Caution is needed in extrapolating these results to the field, as the experiments were of short duration on young trees with fully-formed needles, growing in a soil better supplied with nutrients than field soils. Nevertheless, these preliminary results do not support the hypothesis of an O₃-mediated increase in foliar leaching as the major cause of forest decline nor were the symptoms of O₃-injury on Scots pine comparable with those reported in the field.     

Plasticity in hydraulic architecture of Scots pine across Eurasia

Widespread tree species must show physiological and structural plasticity to deal with contrasting water balance conditions. To investigate these plasticity mechanisms, a meta-analysis of Pinus sylvestris L. sap flow and its response to environmental variables was conducted using datasets from across its whole geographical range. For each site, a Jarvis-type, multiplicative model was used to fit the relationship between sap flow and photosynthetically active radiation, vapour pressure deficit (D) and soil moisture deficit (SMD); and a logarithmic function was used to characterize the response of stomatal conductance (G _s) to D. The fitted parameters of those models were regressed against climatic variables to study the acclimation of Scots pine to dry/warm conditions. The absolute value of sap flow and its sensitivity to D and SMD increased with the average summer evaporative demand. However, relative sensitivity of G _s to D (m/G _s,ref, where m is the slope and G _s,ref is reference G _s at D = 1 kPa) did not increase with evaporative demand across populations, and transpiration per unit leaf area at a given D increased accordingly in drier/warmer climates. This physiological plasticity was linked to the previously reported climate- and size-related structural acclimation of leaf to sapwood area ratios. G _s,ref, and its absolute sensitivity to D (m), tended to decrease with age/height of the trees as previously reported for other pine species. It is unclear why Scots pines have higher transpiration rates at drier/warmer sites, at the expense of lower water-use efficiency. In any case, our results suggest that these structural adjustments may not be enough to prevent lower xylem tensions at the driest sites.