Growth of Young Sitka Spruce (Picea sitchensis (Bong.) Carr.) and the Effect of Simulated Browsing, Staking and Treeshelters

Sitka spruce (Picea sitchensis (Bong.) Carr.) 1 + 1 transplantswere grown at 1 x 1 m for 3 years with 100 per cent chemicalweed control on a very fertile site. Treatments were control,firm staking, staked with 15 cm diameter netting with simulatedbrowsing, staked with simulated browsing to 15 cm diameter crown,1.2 m treeshelter, 1.2 treeshelter with holes and no browsing.No leaders were browsed. Trees were assessed for annual heightgrowth, stem diameter at 0.5 m and 1.0 m, total above and belowground biomass, root:shoot ratio, root plate diameter, rootdepth and root volume. No 3-year heights were significantlydifferent but third-year increments were; the greatest finalheight was in the control at 200.7 cm. All other tree parametershad significant treatment differences demonstrating the needto select carefully the criteria of success in experiments.Treatments which restricted the horizontal development of thecanopy either physically or by simulated browsing seriouslyreduced stem diameter and root development. The controls hada root:shoot ratio of 0.418; treeshelter treatments had thesmallest root:shoot ratio of 0.238; staked trees were 0.379while staked browsed trees had the highest value at 0.447. Acceptanceof browsing or use of shelters or guards to protect trees frombrowsing may seriously reduce growth.     

Genetic Variation in Seedling Development of Sitka Spruce, Picea sitchensis (Bong). Carr.

The development of the terminal bud in one-year seedlings ofSitka spruce, Picea sitchensis (Bong.) Carr., is illustratedand described as a continuous process. The same sequence ofmorphological events occurred in forty-seven provenances examinedbut the timing was different. Buds formed in response to decreasingdaylength, and an essentially continuous relationship was demonstratedbetween time of bud formation and latitude of seed origin. Flushingwas controlled largely by temperature, and time of flushingreflected the nature of the temperature r?gime in the nativehabitat. The height attained at the end of the growing season was demonstratedto be related to date of bud formation and latitude of seedorigin, but the relationship was modified by temperature andphotoperiod. There was little genetic variation in rate of heightgrowth, and variability in time of bud formation was consideredto be the major factor causing provenance variation in totalheight growth. Bud size was correlated with the number of needle primordia,but the effect of bud size on height growth was secondary, aswas the effect of the number of lateral branches. The reactions of two provenances suggested that ecotypic differentiationhas occurred in response to specific, local, environmental selectionpressures. The results supplement the conclusions of earlier reports, namelythat there is a broad interaction of genotype with environmentthat should be evaluated before planting the species at anygiven site.     

Branch thickness in unthinned stands of Sitka spruce (Picea sitchensis (Bong.) Carr.)

Results of branch measurings in unthinned stands of Sitka spruce in Denmark are presented. Three different measures of branch thickness are discussed in relation to visual grading of structural timber. On basis of the average thickest branch 5.0 m above ground it is recommended not to let spacing exceed 1.8x1.8 m‐2.0x2.0 m at establishment. Since Sitka spruce branches react profusely when spacing widens, this is the upper limit, presupposing a no‐thinning regime. With reference to the literature, the same recommendation can be given on basis of machine stress gradings.     

Early Growth of Picea sitchensis (Bong.) Carr. on Deep Oligotrophic Peat in Northern Ireland

Relationships are examined between growth of young Sitka spruce,planted on deep oligotrophic blanket peat, and uptake of N,P, and K by the trees under various fertilizer and herbicideregimes. Despite the provision of adequate P, tree growth eventuallybecomes limited by a shortage of available N. This can be overcomeat present only by repeated applications of fertilizer N. Theuse of herbicides to kill heather in checked crops increasesthe availability of N temporarily, and fertilizer K in all situationsimproves growth, though often only slightly. It is suggested that following the initial application of phosphatea finite fraction of the total N content of the peat becomesavailable to the trees. Once this has been used the releaseof mineral nitrogen is so slow that crops become N-deficientand growth is limited. There is no evidence to suggest thatonce canopy closes the situation will change, so that on oligotrophicpeat sites managers are faced with the possibility of havingto apply nitrogenous fertilizers every 3 or 4 years to maintaingrowth at a satisfactory level. Although there is as yet nodirect evidence on oligotrophic peat that there will not bean increase in the rate of N mineralization once crops reach5–7 m in height, work on other poorly drained soils inNorthern Ireland indicates that it is unlikely to happen.     

Effects of elevated carbon dioxide and drought on the growth and physiology of clonal Sitka spruce plants (Picea sitchensis (Bong.) Carr.)

Two-year-old Sitka spruce (Picea sitchensis (Bong.) Carr.) plants from four clones were grown in naturally lit growth chambers for 6 months at either ambient (350 ppm) or ambient + 250 ppm (600 ppm) CO(2) concentration. Plants were grown in large boxes filled with peat, in a system that allowed the roots of individual plants to be harvested easily at the end of the growing season. Half of the boxes were kept well watered and half were allowed to dry out slowly over the summer. Plants growing in elevated CO(2) showed a 6.9% increase in mean relative growth rate compared to controls in the drought treatment and a 9.8% increase compared to controls in the well-watered treatment, though there was considerable variation in response among the different clones and water treatments. Rates of net CO(2) assimilation were higher and stomatal conductances were lower in plants grown in elevated CO(2) than in ambient CO(2) in both the well-watered and drought treatments. Both of these factors contributed to the doubling of instantaneous water use efficiency. The partitioning of biomass to roots was unaffected by elevated CO(2), but the ratio of needle mass/stems + branches mass decreased. Together with reduced stomatal conductance, this probably caused the observed increases in xylem pressure potentials with elevated CO(2).     

The effect of cultivation technique on root architecture of young Sitka spruce (Picea sitchensis (Bong.) Carr.) trees on surface water gleys

The research on the impact of soil cultivation/drainage practices upon root architecture of young Sitka spruce (Picea sitchensis (Bong.) Carr.) trees reported in this article was carried out in the period from 1995 to 1997. A total of 279 trees between five and eight years of age were excavated on three sites in the mid-west of Ireland. The general soil type was surface water gley. Horizontal and vertical root spread was found to be related to the size of the tree. Larger trees in the machine mound and mole and mound treated plots tended to have greater root spread compared to the smaller trees in the moled and uncultivated plots. Between the two sampling periods in 1995 (at the age of 5 years) and 1997 (at the age of 7 or 8 years) the percentage root distribution decreased close to the root stock and increased further out. However, at all three sites very little of the root cross sectional area was found beyond 60 cm from the root stock in either the vertical or the horizontal directions, even after 7 growing seasons. Generally the location of the centre of gravity of the tree root systems was not affected by cultivation treatment, nor was it found to differ significantly from the ideal co-ordinates of (0,0). Therefore, most of the root systems appeared to be symmetrical.     

An outbreak of Vapourer Moth (Orgyiaantiqua L.: Lepidoptera Lymantridae) on Sitka spruce (Picea sitchensis (Bong.) Carr.) in Central Scotland

Highly significant differences were observed in the intensityof defoliation on Sitka spruce of different heights at CarronValley Forest, Central Region. Most severe defoliation occurredon the smallest trees. Significant differences also occurredbetween degree of defoliation and foliar potassium and phosphorusconcentrations. Winter and summer rainfall patterns immediatelyprior to the outbreak showed higher than average winter andlower than average summer rainfall compared with the 30-yearmean. Experience with this pest suggests that it will continueto have local nuisance value.     

Relationships between Growth of Sitka Spruce (Picea sitchensis), Soil Factors and Mycorrhizal Activity on Basaltic Soils in Western Scotland

A survey was made of soil factors likely to be responsible forchecked growth of Sitka spruce on soils derived from basaltin western coastal areas of Scotland. Despite the high organicP contents of these soils, availability of P appears to be themain factor limiting growth; growth indices were significantlycorrelated with foliar P concentrations and inorganic P levelsof the soil. The low availability of soil P appears to be associatedwith high levels of oxalate-extractable Al (amorphous) in thesoils, a strong correlation existing between the latter andthe P sorption index of Bache and Williams. CaCl₂-extractableAl (‘available’) values were also high and althoughthey exhibited no significant correlation with growth therewas a negative correlation between growth and foliar Al concentrations. High mycorrhizal counts were associated with tree roots on siteswith better growth while beaded roots were mainly associatedwith poor growth. There is a suggestion that high CaCl₂-extractableAl in these soils may affect mycorrhizal development, as a strongcorrelation was found between the occurrence of beaded rootsand foliar Al concentration. Factor analysis confirmed that low availability of P in thesesoils may be ameliorated by mycorrhizal development and thata negative association exists between growth, the occurrenceof beaded roots and foliar Al concentration.     

Resistance to the green spruce aphid (Elatobium abietinum Walker) in progenies of Sitka spruce (Picea sitchensis (Bong) Carr.)

Damage caused by Elatobium abietinum was estimated as needle loss in a progeny trial of Sitka spruce in Denmark in 1990 and 1992. The parents of the progeny had been selected for putative aphid resistance during a massive attack of aphids in 1957. Needle loss was significantly less in the progeny than in a standard provenance. Family heritability was estimated as 0.72 (P>0.0001). Aphid damage had an impact on growth and vigour of trees following warm winters in 1989 and since, but no correlation could be found between growth parameters and damage of different families. Due to high heritability and large variation, high gains are expected by breeding. Practical and theoretical consequences of including aphid resistance in breeding programmes of Sitka spruce are discussed.     

The effect of seed pretreatment and sowing date on the nursery emergence of Sitka spruce (Picea sitchensis [Bong.] Carr.) seedlings

Sitka spruce seeds which had been: (1) untreated (U), (2) prechilled(P), or (3) prechilled and then redried and stored for 16 months(R), were sown into progressively warmer seed beds on four datesat monthly intervals from March to June. At each sowing date,samples of seed were germinated in the laboratory at optimum(20°C) and sub-optimum (10°C) temperatures. Prechillingand prechilling + redrying both increased the rate and totalamount of germination compared with untreated seed at each testtemperature. However in the nursery, the beneficial effectsof pretreating Sitka spruce seed were restricted to sowing inMarch and April when soil temperatures were sub-optimal forthe germination of untreated seeds. Pretreatment resulted inrapid seedling emergence at soil temperatures between 10°Cand 15°C, whereas untreated seed germinated slowly and seedlingemergence continued until high soil temperatures in June killedungerminated seeds. The pattern of seedling emergence from theredried seed was similar to that from the prechilled seed, exceptfor a lag in the start of seedling emergence probably due tothe extra time needed for redried seed to imbibe water in theseed beds. Thus redrying caused insignificant loss of seed performanceon the nursery. Seed sown in May and June experienced average temperatures closeto the optimum for germination of 20°C, however daily maximaregularly exceeded 35°C resulting in the cessation of seedlingemergence in all three seed treatments, and the loss of newlyemerged seedlings. Periods of warm weather from June onwardswere the main factors limiting the final number and size ofseedlings.     

The use of isozyme genetic markers to estimate the rate of outcrossing in a Sitka spruce (Picea sitchensis (Bong.) Carr.) seed orchard in Scotland

Effects of genotype, collection date, auxin treatment, crown position and artificial chilling on rooting and subsequent vegetative growth of Cupressus sempervirens L. softwood cuttings taken from 13-year-old plants were evaluated. Three different clones (318, 296 and 22) previously selected for resistance to cypress cortical cancer caused by Seiridium cardinale were used. In all these clones, both the IBA treated and the untreated cuttings showed seasonal periodicity of rooting. Untreated cuttings rooted best when collected in April (up to 49% for the best clone, i.e. 296) and rooted very poorly in July and October. Auxin treatments appeared unable to overcome this periodicity, as the cuttings taken in July and October did not respond to IBA stimulation. However, in January and April, IBA treatments enhanced adventitious root formation of poorly rooting clones (318 and 22), as well as of clone 296, which performed best (89%) when cuttings were treated in April with 1.5% IBA. Cuttings of each clone collected in the lower third of the crown of the donor trees showed a significantly higher rooting percentage, when compared with those from the upper third. A 4-week post-severance chilling treatment (4°C) significantly enhanced the rooting capacity of cuttings taken in January, while a longer chilling treatment (8 weeks) had a negative effect. The subsequent growth of the rooted cuttings was not affected by either the crown position of the severed cuttings, or the genotype.     

Variation in the levels of the antifungal stilbene glucosides astringin and isorhapontin in the bark of Sitka spruce [Picea sitchensis (Bong.) Carr.

The concentrations of the constitutive antifungal stilbene glucosides astringin and isorhapontin were measured in the bark of a number of Sitka spruce [Picea sitcbensis (Bong.) Carr.] genotypes. These included trees of different provenances, trees grown at different sites, and young clonal plants. The stilbene contents of individual trees or clones varied widely. In general this was attributable to variation between individual genotypes, and was not related to provenance or site.     

Growth Changes in 20-year-old Sitka Spruce Picea sitchensis after Attack by the Green Spruce Aphid Elatobium abietinum

Following a severe attack by Elatobium abietinum (Walker) in1979–80, annual observations were made for 5 years on320 pole-size Sitka spruce in a nitrogen and phosphate experimentlaid out on oligotrophic blanket bog. Leader length was reduced by 25.5 per cent in 1980–81compared with the 3 previous years. This reduction was highlycorrelated with defoliation, being 39.4 per cent for trees thatlost all 1977–79 needles and 8.4 per cent in trees thatlost no needles. There were strong interactions with fertilizertreatment. Defoliated trees continued to grow less well duringthe 5 years of observations and were more likely to go intogrowth check. Early flushing trees suffered significantly less growth reductionfollowing defoliation. The same trees consistently broke budearly over 4 years of observations. Early bud burst was notcorrelated with height, girth or defoliation. No reattack by aphids was observed in the experiment until 1985–86and in this minor infestation the trees previously colonizedwere found to be significantly more likely to be recolonized. A novel measuring-rod for tree height is described.     

Grass and legume seeding on a scarified coastal alluvial site in northwestern British Columbia: response of native non-crop vegetation and planted Sitka spruce (Picea sitchensis (Bong.) Carr.) seedlings

The success of various grass-legume mixtures in controlling competing vegetation, and their effect on subsequent survival and growth of Sitka spruce seedlings was studied on a coastal alluvial site in northwestern British Columbia. Mechanically scarified (bladed) strips were hand seeded to pure and combined mixtures of legumes, bunchgrasses, and sodforming grasses. An unseeded control (bladed but not seeded) was also established.Alsike clover (Trifolium hybridum L.) was the most successful legume species. Orchardgrass (Dactylis glomerata L.), big bluegrass (Poa ampla Merr.), and creeping red fescue (Festuca rubra L.) were the most successful grass species. Red alder (Alnus rubra Bong.) density and height were lower in legume-seeded treatments. Red elderberry (Sambucus racemosa L.) and salmonberry (Rubus spectabilis Pursh.) were effectively reduced by blading and have reestablished slowly. A combination of blading and early establishment of either sodforming grasses or bunchgrasses effectively decreased reinvasion by thimbleberry (Rubus parviflorus Nutt.).Growth of Sitka spruce was best in the unseeded control treatment or in the treatments with legumes but no sodforming grasses. Presence of sodforming grasses decreased both diameter and height growth. Sitka spruce diameter decreased with increasing red alder density. After 5 growing seasons, spruce has outgrown all competitors except red alder.     

The Growth of Sitka Spruce on Poorly Drained Soils in Northern Ireland

A soil survey of Lisnaskea Forest, Co. Fermanagh, showed thatgrowth of Sitka spruce was far from uniform on areas of apparentlyuniform soil. The relation between soil factors, tree growth,and foliar nutrient status on two blocks of trees growing ongleyed soils was therefore studied. Simple regression showed that tree growth was poor when thelevels of foliar nutrients were low and when there was a matof undecomposed needles on the forest floor. It is thereforeconsidered that lack of organic matter breakdown in the soilis causing poor tree nutrition and growth. Multivariate statistical analysis also showed highly significantregressions between tree growth, level of foliar nutrients,and depth of litter layer. The correlations between soil factorsalone and tree growth were barely significant and the multivariatetechnique did not give an acceptable method of predicting treegrowth from soil measurements. Methods of improving soil conditions are discussed.     

Biomass expansion factors for Sitka spruce (<Emphasis Type="Italic">Picea sitchensis</Emphasis> (Bong.) Carr.) in Ireland

The assessment of a forest resource in national inventories provides a firm basis for the calculation of biomass and carbon (C) stocks of forests. Biomass expansion factors (BEFs) and conversion factors provide a robust and simple method of converting from forest tree stem volume to total forest biomass. These factors should be constructed on the basis of nationally specific data in order to take account of regional differences in growth rates, management practices, etc. The objective of this study is to improve the accuracy of biomass estimation by calculating a range of age-dependant BEFs from representative data that more accurately describe the allometry of present forests. The results from this study show that the allocation of biomass to compartments in forest stands and throughout a rotation varies considerably, and that the use of BEFs for the calculation of C stocks in forests of sub-timber dimensions is highly impractical.