Genetic parameters for early wood and latewood densities and development with increasing age in Scots pine

? Each annual ring in pines consists of earlywood and latewood with considerable difference in density and width. To get a better determination of the genetic regulation of total wood density in Scots pine (Pinus sylvestris L.), density and width of those ring sections were measured in annual ring numbers 12 to 21 of Scots pines in a full-sib progeny test. Tree height and stem diameter were also measured.

? Heritabilities for the annual ring sections increased with age for earlywood density from 0.08 to approximately 0.25; latewood density showed similar reductions. Heritability over all 10 annual rings was 0.25 for earlywood density, 0.22 for latewood density, 0.29 for height and 0.10 for stem diameter. Genetic correlations between earlywood and latewood density and growth traits were negative, while they were strongly positive between densities of adjacent annual rings (0.70–1.0).

? Despite the higher heritability of earlywood density, the strong positive genetic correlation between those traits indicates little benefit from focusing solely on earlywood density when selecting for wood density. Analysing earlywood and latewood separately does not benefit from including the width of the corresponding ring section as a covariate. Juvenile wood may possibly turn into mature wood 15–20 y from the pith.

     

Influence of drought on tree rings and tracheid features of Pinus nigra and Pinus sylvestris in a mesic Mediterranean forest

We investigated the influence of climate on the ring width and xylem anatomy of two co-occurring pines (Pinus nigra Arn. and P. sylvestris L.) in the mountains of east-central Spain in order to test their utility for dendroclimatic reconstructions. We developed chronologies of ring width, mean lumen diameter and mean cell-wall thickness (in the earlywood, latewood, and the total annual ring) and the number of cells between 1960 and 2006. Drought, expressed as the standardized precipitation-evapotranspiration index (SPEI), was the main climatic driver of tree radial growth, although trees were also sensitive to temperature (negative effect in previous autumn and current summer) and precipitation (with a general positive effect). P. sylvestris response was stronger to climate of the current year, whereas the effect of previous-year climate was more important for P. nigra. Warm and dry summers reduced ring width, tracheid lumen, and wall thickness in both species, whereas warm winter-spring temperatures had the opposite effect, primarily for P. sylvestris. Previous-year or early-season conditions mainly affected earlywood features, whereas latewood was more responsive to summer climate. Overall, climate appeared to be a stronger limiting factor for P. sylvestris. During periods of drought, cell-wall thickness was reduced while lumen width increased in the latewood of P. sylvestris. This could compromise its hydraulic safety against drought-induced cavitation as our site was close to the southern and dry edge of the species distribution area. Our results suggest that anatomical variables record different and stronger climate information than ring width variables, especially in P. sylvestris. Reconstruction models for SPEI at the 3-month scale were developed for July–August and September–October using principal components regression. The best models included anatomical and width variables of both pine species suggesting that tracheid chronologies can be useful for drought reconstructions especially at mesic sites or with species that encode a mixed drought and temperature-precipitation signal.     

The effect of weed control and fertilization on survival and growth of four pine species in the Virginia Piedmont

The growth response of loblolly pine (Pinus taeda L.), shortleaf pine (Pinus echinata Mill.), Virginia pine (Pinus virginiana Mill.), and white pine (Pinus strobus L.) to weed control and fertilization in the Piedmont of Virginia was assessed. Four different silvicultural treatments were evaluated: (1) check (no treatment); (2) weed control; (3) fertilization; (4) weed control plus fertilization. The weed control treatment included a series of herbicide and mechanical treatments to eliminate competing hardwoods. The fertilizer treatments added N, P, K, and S. Survival and growth was measured annually through age 5. There were significant differences in survival and growth among species. Survival was greatest for loblolly pine, lower in shortleaf and Virginia pine, and lowest in white pine. Fertilization without controlling the competing hardwoods decreased survival in all planted pines due to the increased hardwood competition. Loblolly pine was tallest through the 5-year period, shortleaf and Virginia pine were shorter and white pine was shortest. Silvicultural treatments had no impact on tree height but significantly affected DBH. Weed control increased DBH while fertilization did not. When applied in combination with weed control, there was no additional increase in growth of the pines due to fertilization beyond that from weed control only. Fertilization stimulated the growth of the competing hardwoods which were significantly taller in the fertilized plots.     

Influence of silvicultural practices on twisting rust infection and damage in maritime pine,as related to growth

A three-year study was carried out on the influence of silvicultural intensification on infection and damage caused by Melampsora pinitorqua, the twisting rust fungus, in maritime pine, Pinus pinaster Ait. The silvicultural practices studied were phosphorus fertilisation, the mode of stand establishment (either seeds or plants), and pine density. Growth and nutrition variables were also measured. Infection and damage were the most important in the first two years, which could reflect higher resistance with aging. At the end of the experiment, when pines were 4 years old, the planted, fertilised pines turned out to be the most damaged and the tallest, while the sown, non-fertilised pines were the least damaged and the smallest. Final damage was mainly explained by the damage observed in the earlier years. Annual damage was significantly correlated with infection percentage, more closely in the first year of observation. Higher susceptibility of the planted pines was associated with better growth and was explained by a greater amount of susceptible tissues, Under our experimental conditions, this explanation could not be retained for the fertilisation effect. However, phosphorus fertilisation induced changes in the nutritional status of pines. A significant positive correlation was observed between the phosphorus and potassium content on one hand and the susceptibility and damage on the other hand. Density effects were seldom observed in the experiment, either on growth or susceptibility variables. The positive relation between pine growth and rust damage, as confirmed in this study, emphasises the need for an integrated management of intensively managed pine stands, taking into account the disease hazard.     

Specific gravity responses of slash and loblolly pine following mid-rotation fertilization

Wood quality attributes were examined in six stands of slash pine (Pinus elliottii Engelm. var. elliottii) and loblolly pine (P. taeda L.) in the lower Coastal Plain of Georgia and Florida. Several plots comprised each stand, and each plot was divided so that it received three fertilizer treatments: a control treatment with herbaceous weed control at planting and brush control at mid-rotation only (control); 45 kg ha⁻¹ N + 56 kg ha⁻¹ P + herbaceous weed control at planting and 224 kg ha⁻¹ N + 45 kg ha⁻¹ P + brush control at mid-rotation (fertilizer with N at planting); and 56 kg ha⁻¹ P + herbaceous weed control at planting and 224 kg ha⁻¹ N + 45 kg ha⁻¹ P + brush control at mid-rotation (fertilizer without N at planting). Ring width, ring earlywood specific gravity (SG), ring latewood SG, whole ring SG, and ring percent latewood were measured on each of seven trees. Of these measurements, this study focused mainly on the properties related to SG. Examination of the rings showed that latewood SG was significantly lower in trees treated with fertilizers with and without N at planting in the two to three years following fertilization, but that latewood SG gradually returned to a level similar to the control. Fertilizer without N at planting may also have had a brief negative effect on earlywood SG following mid-rotation fertilization, but it was not as clear or lasting as the effect on latewood SG. Additionally, although slash and loblolly pine appear to differ in the developmental patterns of these SG properties, there were no significant differences in how these patterns interacted with treatment. This study demonstrated that fertilization treatments have similar short-term effects on the SG of slash and loblolly pines, particularly in latewood, but the trees will return to a SG pattern consistent with unfertilized trees within two or three years.     

Seasonal variation in gene expression for loblolly pines (Pinus taeda) from different geographical regions

In developing xylem, gene expression levels vary in different genotypes, at different stages of development, throughout a growing season, and in response to stresses. Commercially important characteristics such as wood-specific gravity are known to differ with seed source. For example, when grown on a common site, the specific gravity of Arkansas loblolly pine (Pinus taeda L.) trees is greater than that of Louisiana loblolly pine, and Texas loblolly pines have a greater specific gravity than loblolly pines from the Atlantic coast. A microarray analysis was performed to examine variation in gene expression among trees from different geographical sources when grown on a common site, and seasonal variation in gene expression in each seed source. We used microarrays containing 2171 expressed sequence tags (ESTs) with putative functions of interest, selected from several loblolly pine xylem partial cDNA libraries and a shoot tip library. Genes with significant variation in expression for each factor were identified. Many genes preferentially expressed in latewood compared with earlywood were for proteins involved in cell wall biosynthesis. Variation in gene expression among trees from the two seed sources in each growing season suggests that there may be more differences between South Arkansas trees and South Louisiana trees in latewood than in earlywood. Variation in gene expression among trees from different regions may reflect adaptation to different environments.     

Pinus pinaster Ait. tree mortality following wildfire in Spain

Maritime pine (Pinus pinaster Ait.) is the tree species most affected by wildfire in the Iberian Peninsula. Prediction of the probability of fire-injured tree mortality is critical for management of burned areas, evaluation of the ecological and economic impact of wildfire and prescribed fire planning and application. Pine bark beetles (Scolytidae) frequently attack burned maritime pine stands and cause extensive post-fire mortality throughout the Iberian Peninsula. In the present study, maritime pine trees were monitored for three years following 14 wildfires in four ecotypes in Spain (11 fires in Galicia (Galician ecotype - NW Spain), one fire in Portillo (Meseta-Castellana ecotype - Central Spain), one fire in Rodenal (Rodenal ecotype - Central Spain), and one fire in Genalguacil (Sierra Bermeja ecotype - SW Spain)). Data on tree attributes, crown and bole injury, ground fire severity, Ips sp. presence and tree survival were obtained by examining 3085 trees. Logistic regression models for predicting the probability of delayed maritime pine mortality were developed by use of generalized estimated equations (GEE). An ample range of response to fire damage in mortality was evident among the four ecotypes and different models were fitted for each. The most important variables for predicting tree mortality were total crown volume damaged, presence of Ips sp. attack and cambium kill rating. The results highlight the extensive presence of Ips sp. in burned maritime pine forests and its importance in tree mortality process, the ample range of response of P. pinaster, in terms of post-fire mortality, as well as the need to develop site specific mortality models for the different ecotypes of this species following fire.     

木材材性株内径向变异模式初探──Ⅲ.种植密度对湿地松、火炬松木材密度径向变异模式的影响

对两个地区不同种植密度的湿地松、火炬松进行了微密度分析，研究了种植密度对木材密度径向变异模式的影响。主要结果是：种植密度对湿地松和火炬松各密度特征值径向变异模式的影响，主要表现在变异曲线平均水平高低的变动，而对变异曲线形状的影响并不显著。种植密度对湿地松和火炬松各年轮宽特征值径向变异模式的影响，主要表现在年轮宽度RW和早材宽度EW变异曲线平均水平高低的变动和RW、EW与晚材宽度LW变异曲线形状的变动。种植密度对年轮密度RD的影响从年轮内宽度和年轮密度变异曲线平均水平高低两个方面起作用，出现较复杂的情况，并非所有情况下都是单调的正相关。     

Pathogenicity of canadian isolates of the Bursaphelenchus xylophilus (pinewood nematode) to provenances of Pinus sylvestris and Pinus contorta as grown in Finland: A greenhouse study

Seedlings of Scots pine (Pinus sylvestris L.) and lodgepole pine (P. contorta Dougl.) provenances, as grown in Finland, were inoculated with “m”; and “r”; “forms”; of the pinewood nematode (Bursaphelenchus xylophilus) from Canada, an m form isolate from France and an r form isolate from Missouri, USA. Scots pine was highly susceptible to an Alberta r form and a British Columbia m form isolate and moderately susceptible to two Quebec m form isolates. Lodgepole pine was higly susceptible to the two r form (Alberta and Missouri) isolates and moderately susceptible to the British Columbia m form and the two Quebec m form isolates. Mortality of both pines after inoculation with the French isolate was inconsistent. Mortality of both pines occurred more rapidly following inoculation with r form than with m form nematodes. Large numbers of nematodes were generally found in the tissues of both pines. Our results with seedlings need to be corroborated by inoculating larger field‐grown trees.     

Pine growth response to management of the subterranean clover understory

Management of subterranean clover (Trifolium subterraneum L.) understory grown with loblolly pine (Pinus taeda L.) and slash pine (Pinus elliottii Engelm.) resulted in differences in both clover yield and pine diameter. Loblolly and slash pines were planted into a subterranean clover pasture in 1984. Suppression of warm season herbaceous vegetation by applying herbicides or disking the site in late summer resulted in significantly greater subterranean clover production than on the control site where no additional understory management was applied. Pine species had no effects on clover yield. The N content of the pine foliage on the blocks managed with herbicide application or disking generally was significantly greater than the control from the 5th through 7th growing seasons (1988–1990). The dbh of the pine trees in the disked and herbicide treatments was greater than the control after the 7th growing season (1990).     

Development of oaks (Quercus petraea (Matt.) Liebl.) emerged from bird-dispersed seeds under old-growth pine (Pinus silvestris L.) stands

In East Germany, there are a lot of areas covered by old pine stands. They are growing on soils, on which under natural conditions without anthropogenic impact mixed stands of oak (Quercus petraea (Matt.) Liebl.) and pine (Pinus silvestris L.), would form a natural forest. An important objective of ecological silviculture in these areas is to convert the pure pine stands into mixed oak–pine stands by using natural regeneration methods. A highly appreciated assistant in this connection is the European jay (Garrulus glandarius L.). The remarkable results of its work have been found by analyzing the natural regeneration in a few old pine stands in the forest district of Weißwasser in Saxony. Although mother trees of oak are very scarce there, natural regeneration of oak was found everywhere in the research area. The oak regeneration, undoubtedly created by the jay, amounted to at least 2000 oaks/ha. They were mainly distributed at random, while pine regeneration was aggregated in places where gaps in the pine canopy occurred. Oak regeneration was established much earlier than pine regeneration. Thus, oaks exceeded pines in age, height and diameter. The oaks were also superior to pines concerning height increment for the last three years. Thus, there is a good chance for oak to defend its prevailing role in the regeneration in the future. Probably, the next forest generation will be composed of oak and pine trees. The analysis of the quality of the oak regeneration shows that there is no substantial difference to artificially sown oak stands. This indicates that the European jay creates oak stands sufficiently both in number and quality.     

Influence of the fertilisation method in controlled ectomycorrhizal inoculation of two Mediterranean pines

The influence of the fertilisation method: soluble (SF) vs. slow-release fertiliser (SRF) and of inoculation with Laccaria laccata (Scop.) Fr., Pisolithus tinctorius (Pers.) Coker & Couch and Melanogaster ambiguus (Vittad.) Tul & C. Tul. on ectomycorrhizal colonization and growth of Pinus pinea L. and Pinus pinaster Ait. was evaluated. For both pines, mycorrhization with L. laccata was not affected by the fertilisation method. Percentages of ectomycorrhizas (ECM) formed by P. tinctorius were dependent on the fertilisation method, the inoculum type (vegetative or spores) and the pine species involved. ECM formed by M. ambiguus were increased with fertilisation in both pines. Inoculation significantly improved P. pinea biomass when seedlings were fertilised with SRF whereas no effect was found in non-fertilised ones. For non-fertilised P. pinaster, inoculation with L. laccata and both inocula of P. tinctorius increased seedling biomass whereas fertilisation neutralised the fungal effect. Fertilisation increased P. pinea and P. pinaster biomass, independently of the inoculation treatment.     

Response of planted Pinus taeda L. to brush control in Northern Louisana

Eradication of brush, defined as hardwood trees, shrubs, blackberries, vines, and volunteer pines, was not necessary to maximize juvenile pine growth in northern Louisiana Pinus taeda L. plantations. Pine height and diameter gains were similar on treatments when either (1) only the above ground portion of the brush was cut yearly or (2) both the above ground stems and root systems of brush species were controlled each year. At age 10, planted pines on both treatments averaged 0.9 m taller with a 2.5 cm greater stem diameter at breast height (1.4 m) than those on untreated checks. Total outside bark volume per pine averaged 51.9 dm³ on checks vs 78.5 dm³ on treated plots.     

Susceptibility of Coulter pine (Pinus coulteri) to western dwarf mistletoe (Arceuthobium campylopodum) in southern California

The susceptibility of Coulter pine (Pinus coulteri) to western dwarf mistletoe (Arceuthobium campylopodum) was evaluated in southern California. Eight stands infested with western dwarf mistletoe and dominated by Coulter and Jeffrey pines (Pinus jeffreyi) were sampled. Infection of Coulter pine was evaluated using circular plots established around large, severely infected Jeffrey pines and with rectangular transects. A total of 25 plots and 10 transects were sampled. In each plot and transect, live trees > 5 cm in diameter at breast height (1.4 m above the ground) were assigned a dwarf mistletoe rating (6‐class system). Nearly 600 Jeffrey pines and more than 400 Coulter pines were evaluated. Infection of Jeffrey and Coulter pines in the plots and transects averaged 96% and 97%, respectively. These data indicate that Coulter pine should be classified as a principal host of WDM and not as a secondary host as previously reported.     

Aggressiveness of Fusarium oxysporum and F. verticillioides isolates on stone and scots pine under greenhouse conditions

Scots pine (Pinus sylvestris) and Stone pine (Pinus pinea) are two important species used in re-forestation that are subject to damage by damping-off fungi in forest nurseries. Twenty-two isolates of Fusarium oxysporum and F. verticillioides from diseased seedlings of eight different hosts were tested for aggressiveness on seeds and seedlings of both pine species, including the effects on seedling emergence and mortality. Scots pine was more susceptible to damping-off than Stone pine, as indicated both by reduced seedling emergence and elevated seedling mortality. The impact of F. oxysporum and F. verticillioides on seeds and seedlings did not differ significantly for either pine species. Our findings support previous studies that found that these are damping-off pathogens on the studied pines. Whereas most isolates proved to be pathogenic, some isolates of both Fusarium species showed to be non-pathogenic.     

Young conifer stands form a deer browsing refuge for an oak admixture: silvicultural implications for forest regeneration under herbivore pressure

There is little knowledge how ungulate pressure on forest regeneration may be mitigated by silvicultural methods. The knowledge is especially needed for artificially regenerated, deciduous tree species. We studied factors affecting browsing incidence by deer in the Pisz Forest District in Poland, an area where 10,000 ha of forest was damaged by a 2002 hurricane. In 2006, we established three experimental plots (in total, 22.6 ha), in which the main species was Scots pine (Pinus sylvestris) admixed with pedunculate oak (Quercus robur). The data on browsing were collected in 2008–2015. In general, oak browsing incidence was unrelated to oak planting density. On a plantation scale, it was significantly affected by the pine age. Although in each variant all the oaks were browsed for four consecutive years (2009–2012), in 2013 browsing incidence began to decrease. When the pines grew higher and formed a physical barrier, it was harder for deer—roe deer (Capreolus capreolus), red deer (Cervus elaphus) and moose (Alces alces)—to move through and locate the oaks. Moreover, within plantations, oak browsing incidence was higher in the patches with shorter pines. Browsing of individual saplings or small groups of saplings was also negatively affected by the height of neighbouring pine saplings. Oak density influenced deer selectivity depending on the tree height. In a low oak tree density, browsing incidence was unrelated to oak height, while in higher tree density, deer selected oaks of the height between 40 and 100 cm. We postulate that deciduous admixture in a coniferous (unattractive) stand can be planted with a few year delay. Older coniferous trees should impede locating of attractive tree species by deer and the browsing incidence.     

Effect of cleaning methods in mixed pine–deciduous stands on moose damage to Scots pines in southern Finland

Abstract

The effect of different cleaning treatments on moose (Alces alces L.) damage to Scots pines (Pinus sylvestris L.) was examined using a generalized linear mixed model (GLMM) with a binomial response (logit-link function). The treatments were the following: no cleaning, point-cleaning of broadleaves (mainly birch, Betula pendula Roth and B. pubescens Ehrh.) within a radius of 1 m from the pine, point-cleaning delayed 2 years, topping of competing broadleaves (main stem cut at a height of 1.3 m) and total cleaning of broadleaves. A randomized complete block design with three replicates was established in three planted pine stands (mean height of pines 1.5 m). Measurements taken four growing seasons later (mean height of pines 3.5 m) showed that all cleaning treatments reduced moose damage, although the difference was not significant (p=0.078) in the case of topping. The proportion of damaged pines was predicted as a function of the characteristics of pines and broadleaves in two untreated, planted pine stands with a height of 3 m. Within these untreated stands the number of damaged pines increased in plots with a higher number of broadleaves and taller broadleaves. In conclusion, the results show that on relatively fertile growing sites of pine cleaning treatments reduced moose damage on pine. It is recommended that cleaning is performed before broadleaved trees occur as overgrowth above pines.     

Nutrient and water availability to and growth of young radiata pine plantations intercropped with lupins

Because the growth of Pinus radiata (D. Don) plantations on podzolized sandy soils can be increased by improving the content of organic matter and nitrogen in soil, the potential for improving these by intercropping with annual lupins (Lupinus angustifolius) in young plantations on first and second rotation sites was examined.Lupins can be grown as an intercrop along with transplanted pine seedlings without causing detrimental effects on the water relations of pines. Because the lupins self-seed, two to three crops can be grown before being shaded out by the pines. Lupins increased the amount of organic matter and concentrations of total nitrogen and mineral nitrogen in the soil, and increased the concentrations of nitrogen in pine needles. The benefit of lupins to tree growth was substantial (16–32% increase in stem growth) and nearly equaled that achieved by repeated additions of nitrogen fertilizer.The choice and management of legume intercrop should be based on an understandig of how site resources (especially water) are shared between young trees and the leguminous plant.     

Effects of light/darkness, thermal shocks and inhibitory components on germination of Pinus canariensis, Pinus halepensis and Pinus pinea

Knowledge of the germination characteristics of Pinus species can help in the understanding, prediction and management of the regeneration of pine forests. In the Canary Islands, several exotic pines (Pinus halepensis L. and Pinus pinea L.) were planted with Pinus canariensis Chr. Sm. Ex DC, the only native pine species, and there is now an interest in controlling these exotic species to restore the original forest. The main objective of the present study is to determine the germination response of P. canariensis, P. halepensis and P. pinea to different light regimes (darkness and light/darkness), thermal shocks and the presence of inhibitory substances from the leaves and litter of P. canariensis. P. halepensis seeds showed the highest viability and germination rate. Darkness accelerated P. halepensis germination, while exudates accelerated P. canariensis germination. Only treatments of 200 °C for 5 min and 300 °C for 5 min significantly decreased the germination of all three pine species. In the absence of strong differences in germination among species after treatments, the key for fire management or prescribed burning in this case may be the sprouting ability of P. canariensis. A possible management strategy to control the two obligate seeder exotic species could be to use one intense prescribed fire followed by a low-intensity prescribed fire after seed bank germination in the field, so in this way, only P. canariensis could resprout after fire and it would eliminate only the exotic pines in a mixed stand.     

Effective rainfall seasons for interannual variation in δ13C and tree-ring width in early and late wood of Chinese pine and black locust on the Loess Plateau, China

We studied the importance of effective rainfall for interannual variation in water use efficiency (WUE) and tree-ring growth of Chinese pine (Pinus tabulaeformis Carr.) and black locust (Robinia pseudoacacia L.) by examining correlations of seasonal precipitation with annual values of stable carbon isotope ratio (δ¹³C) and tree-ring width in early and late wood. The correlations with precipitation were examined for each month and for periods of all possible lengths from 2 to 22 months starting from January of the previous year to October of the current year. The period with the highest correlation was adopted as the most effective rainfall season for interannual variations in WUE and tree-ring width. In early wood, precipitation during the dry season (October to May) before the growing season was negatively correlated with δ¹³C in pine trees and positively correlated with ring width in pine and locust trees. In late wood, rainfall during the growing season in the current year was negatively correlated with δ¹³C in pine and locust trees, and positively correlated with ring width in locust trees. Our results demonstrated the differences in the water use strategies of pine and locust trees. The δ¹³C in pines indicated higher WUE and more conservative water use than in locust trees. Precipitation during the dry season affected the interannual variation in WUE and tree-ring growth in pine and locust trees, indicating that rainfall during the dry season is important for carbon gain and tree-ring growth during the following growing season.