Relationships between gas exchange adaptation of Sitka x interior spruce genotypes and ribosomal DNA markers

Adaptive physiological changes were investigated in seven populations of Sitka (Picea sitchensis (Bong.) Carr.) x interior spruce (P. glauca (Moench) Voss x P. engelmannii Parry ex Engelm.) spanning the Nass-Skeena transition zone in British Columbia, Canada. Each population was represented by an Si rDNA index that was calculated from the relative optical densities on a gel autoradiogram of five ribosomal DNA bands characteristic of Sitka spruce and interior spruce. This index estimates the proportion of the genome contributed by interior spruce. Physiological adaptations were assessed by gas exchange parameters measured under both well-watered and drought conditions. Under well-watered conditions, Sitka spruce populations had higher maximal photosynthesis at saturating light and ambient CO(2), higher quantum yield at the light compensation point, and higher dark respiration than interior spruce populations. Sitka spruce populations also reached maximal photosynthesis at lower photosynthetically active radiation and higher CO(2) concentrations, and had higher stomatal densities that resulted in lower stomatal limitations to photosynthesis than interior spruce populations. In contrast, interior spruce populations exhibited greater drought tolerance than Sitka spruce populations. Their gas exchange rates declined at a slower rate in response to drought. They maintained higher gas exchange rates in response to moderate to severe drought (predawn plant water potentials = -1.5 MPa), and their photosynthetic rates recovered faster when they were rewatered after exposure to drought. Comparison of the seven populations indicated that physiological parameters were significantly related to the Si rDNA index. An increase in Si rDNA index was associated with proportional changes in physiological measurements, suggesting that genetic interchange among species with contrasting ecological adaptations can enhance the environmental adaptation of natural populations.     

Relationships between gas exchange and carbon isotope discrimination of Sitka x interior spruce introgressive genotypes and ribosomal DNA markers

Effects of hybridization on physiological performance were investigated in seven natural introgressive populations and one F(1) population of Sitka (Picea sitchensis (Bong.) Carr.) x interior spruce (P. glauca (Moench) Voss. x P. engelmannii Parry ex Engelm.). Each population was represented by a Sitka-interior spruce ribosomal DNA (Si rDNA) index that was calculated from the relative abundance of species-specific DNA polymorphisms. Gas exchange parameters were measured under well-watered conditions on current-year needle tissues, which were also analyzed for carbon isotope discrimination. Sitka spruce populations had higher gas exchange rates (net photosynthesis, stomatal conductance to water vapor), but lower carbon isotope discrimination values than interior spruce populations. The natural hybrid populations had intermediate values of these parameters consistent with their Si rDNA index. The F(1) population had gas exchange parameters resembling those of Sitka spruce populations, but its carbon isotope discrimination was skewed toward that of interior spruce populations. These results confirmed previous findings that physiological performance of introgressive hybrid spruce populations varied as their DNA constitution changed.     

Temporal changes in functional diversity of culturable bacteria populations in Sitka spruce stumps

Changes in culturable aerobic bacterial communities of Sitka spruce stumps between 1 and 10 years old were investigated by isolating from the top 55 mm of the stumps. High numbers of bacteria were recovered from 15 mm below the surface of 1‐ and 2‐year‐old stumps, particularly in the sapwood, after which numbers dropped by an order of magnitude. Proportions of colony‐forming units (cfu) capable of degrading cellulose and chitin increased with stump age, whereas siderophore producers decreased. Colony‐forming units were grouped on the basis of 13 phenotypic characteristics, including colony form and colour, Gram stain, heat tolerance, siderophore production and production of various enzymes. Four groups of bacteria were identified; the proportions of the three Gram‐negative groups varied with stump age. A quadratic relationship between bacterial diversity and stump age was detected using the Shannon–Weaver diversity index. Bacterial diversity was higher 15 mm below the stump surface than in deeper sampling points. The results are discussed in terms of the possible influence of prokaryotic organisms on the development of decay communities in Sitka spruce stumps.     

In vitro interactions between bacteria isolated from Sitka spruce stumps and Heterobasidion annosum

Culture medium composition affected antagonism by bacterial isolates from Sitka spruce (Picea sitchensis) stumps against Heterobasidion annosum. Fifty percent of bacterial isolates inhibited H. annosum growth on sporulation agar or yeast–dextrose–peptone agar; only 10% of isolates caused inhibition on both media. Proportions of isolates inhibiting H. annosum varied with stump age; fewer isolates from 4‐ or 6‐year‐old stumps exhibited antagonism than isolates from older or younger stumps. Fifteen isolates showing antagonism on sporulation agar were tested against H. annosum in spruce wood cubes. None of the bacterial isolates alone caused a significant weight reduction in inoculated cubes. Relative inoculation times of bacterial isolates and H. annosum had an effect on weight loss in interactions; simultaneous inoculation with isolates and H. annosum inhibited weight loss caused by H. annosum compared with bacteria‐free controls. Inoculation with bacterial isolates 10 days before H. annosum had no effect on the decay rate. In contrast, inoculation with H. annosum 10 days before bacteria increased weight loss of cubes by 200% relative to cultures lacking bacteria. The effect of a mixed bacterial inoculum on weight change in 0.2‐mm spruce wood slips co‐inoculated with H. annosum, Resinicium bicolor, Hypholoma fasciculare, Stereum sanguinolentum or Melanotus proteus differed between different fungi.     

Interaction between moisture, nutrients and growth of white spruce in interior Alaska

Two thinning and fertilization studies, the first in 1969 and the second in 1971, were established to evaluate the question of nutrient limitation to tree growth and the consequences of stand manipulation of soil moisture supply. Fertilizer was applied yearly for the first 5 years in both studies; growth response has been measured through 1987. Results indicate that thinning is necessary to obtain a growth response to fertilizer applied at the rate of 111 kg nitrogen ha⁻¹. The response to fertilization after fertilization ended lasted for 4 years in plots thinned to 800 stems ha⁻¹, while a significant response continued for only 2 years in plots thinned to 1600 stems ha⁻¹.

A soil water-balance model was calibrated for the control and treatment plots of these two studies. Soil water-deficits were estimated and correlated with yearly average basal-area growth per tree. Results indicated that there is a correlation between seasonal soil-moisture deficit and growth during the years when soil moisture was measured for the unthinned control plots (r² = −0.787, P = 0.002) but not for the thinned and fertilized plots (r² = −0.652, P = 0.057).     

Relationships between nutrient supply, nitrogen partitioning and growth in young Sitka spruce (Picea sitchensis)

Budget studies have shown that internal cycling may contribute a large proportion of the annual nutrient supply required to support new growth in trees. Use of budgets to quantify internal cycling only quantifies the net transfer of nutrients within the plant. Differential partitioning of remobilized nutrients and current nutrient uptake could lead to errors in the interpretation of results from these studies. We have quantified the dynamic relationships among tree growth, nutrient uptake and internal cycling by labeling the current uptake of N in trees that received contrasting amounts of nutrient. Two-year-old seedlings of Sitka spruce (Picea sitchensis (Bong.) Carr.) were grown in sand culture in a greenhouse for one year. The trees received nutrients in a balanced solution at either a high (high-RAR) or a low (low-RAR) relative addition rate throughout the experiment. Current N uptake was labeled with (15)N from April 13 to July 25. Thereafter, trees were re-potted in clean sand and unlabeled N applied until November 13. Overall growth was sustained for approximately 10 weeks before treatment effects were observed. Initially, no differences in the partition of growth or remobilized N occurred, although partition of current uptake favored the roots of plants in the low-RAR treatment. After 6 weeks, the partition of both growth and remobilized N altered in favor of roots of plants in the low-RAR treatment. Nutrient supply had no effect on the amount or rate of N remobilization. No evidence was found to suggest that N taken up in the current season and partitioned to preexisting shoots or roots is remobilized late in the season to support growth of new shoots. However, some trees in the high-RAR treatment exhibited a second flush of growth later in the season that was partially sustained by remobilization of (15)N from current shoots formed earlier in the season. Use of (15)N demonstrated differential partitioning of current uptake and remobilized N. The results highlight the limitations of simple budget studies for quantifying internal cycling.     

Trade-offs between seedling growth,thinning and stand stability in Sitka spruce stands: a modelling analysis

The forest industry is increasingly adopting alternative silvicultural systems, involving regeneration beneath an existing forest canopy, rather than clear-felling and replanting. To apply these silvicultural systems in windy regions such as Britain and Ireland, it is essential that the interactions between thinning intensity, stand stability and seedling growth are properly understood. Here, we present a modelling analysis of the three key relationships between: (i) stand density and the proportion of incident radiation transmitted through a forest canopy as a stand is thinned; (ii) transmitted radiation and seedling growth, and (iii) stand density and stand stability. These relationships were examined using separate models of radiative transfer (MAESTRO), seedling growth, and stand stability/wind risk (ForestGALES). Output from the three models was synthesised to calculate whether a given stand thinned to a pre-defined stability limit would allow sufficient light to penetrate the canopy for seedling growth. A minimum transmittance of 20% was identified as a requirement for seedling growth, which corresponds to removing 45% of stand basal area. A thinning of this intensity left some stands susceptible to unacceptable wind damage, especially in old or previously thinned stands on soils where rooting is impeded. The results emphasised the fact that rooting conditions, thinning history and age of intervention are major constraints on the silvicultural options. In general, older stands are not suitable for conversion to continuous cover forestry (CCF) systems, and the transformation process should begin at pole stage, when heavy thinning does not leave the stand unstable. The analysis approach used here illustrates the potential for combining models to address complex forest management issues.     

Internal cycling of nitrogen, potassium and magnesium in young Sitka spruce

Potassium (K) and magnesium (Mg) are essential macro-nutrients, but little is known about how they are cycled within plants. Stable isotope studies have shown that the internal cycling of nitrogen (N) is independent of current nutrient supply in temperate tree species. This is ecologically significant because it allows trees to produce rapid shoot growth in spring independent of current soil N uptake. We used stable isotopes to quantify N, K and Mg in new shoots of Sitka spruce (Picea sitchensis (Bong.) Carr.) seedlings and to compare the relative contributions from current uptake and internal cycling. Two-year-old Sitka spruce seedlings were labeled with (15)N, (41)K and (26)Mg in an abundant or a limited supply for one growing season. The trees were repotted in the subsequent dormant season to prevent further root uptake of enriched isotopes and provided with an abundant or a limited supply of unlabeled nutrients until they were harvested in early summer of the following year. The supply was switched for half the trees in the second year to create four nutrient regimes. Enrichment of (15)N, (41)K and (26)Mg in current-year growth was attributed to internally cycled N, K and Mg uptake from the previous year. The internal cycling of N, K and Mg in new growth was significantly affected by the first-year nutrient treatments. The second-year nutrient supply affected the growth rates of the trees, but had no effect on the amounts of N, K or Mg contributed from internal cycling. Thus, internal cycling of K and Mg in Sitka spruce are, like that of N, independent of current nutrient supply.     

Mycorrhizal and associated fungi of Sitka spruce in Irish forest mixed stands

Identity of mycorrhizas and isolation of symbionts and associated fungi from Sitka spruce growing in pure and mixed stands with either Japanese larch or Lodgepole pine are described and compared. More mycorrhizal types and sporocarps of the Agaricales were collected from mixed stands. Mycorrhizas of Lactarius rufus, Paxillus involutus and Suillus spp. were more prevalent on roots from mixed stands. The most common unidentified mycorrhizal type (type B) had features similar to synthesized mycorrhizas of two Basidiomycete isolates. Suillus grevillei and three un-identified types were associated specifically with Japanese larch. The main associating fungi were Oidiodendron sp. and Mycelium radicis atrovirens. The association of a “nurse” tree with Sitka spruce provides a more diverse mycorrhizal flora, the majority of which are shared between the tree species.     

In vitro interactions in artificial and wood‐based media between fungi colonizing stumps of Sitka spruce

Interspecific interactions between fungi that colonize stumps of Picea sitchensis in Scotland were tested in dual cultures on Norkrans agar, spruce sawdust and in autoclaved blocks cut from roots. Isolates were ranked according to competitive ability on the different media, based on their ability to overgrow competitors or to form deadlock interactions. On the defined medium, Phaeolus schweinitzii was the species most able to overgrow competitors, followed by Stereum sanguinolentum and Heterobasidion annosum; Resinicium bicolor was the least able to overgrow competitors on this medium. By contrast, R. bicolor was the most competitive on spruce sawdust medium. Deadlocking interactions were formed most often in dual cultures on Norkrans agar. Observation of hyphal interactions on Norkrans agar under the microscope identified several different response types including growth of thin hyphae compared to control cultures, hyphal coiling, vacuolation of hyphae, hyphal lysis of one competitor and deposition of crystals in the agar. Hymenomycetes caused varying amounts of decay in autoclaved root blocks. Resinicium bicolor was able to replace other species in most co‐inoculations. Stereum sanguinolentum appeared to be the least competitive species in root block inoculations, being replaced by Melanotus proteus and R. bicolor, although interactions with H. annosum varied widely. These results indicate that substrate has a marked effect on interspecific fungal interactions, with wood‐based, particularly intact woody tissues closely matching competitive behaviour displayed in the field.     

Capercaillie habitat, diet and management in a Sitka spruce plantation in central Scotland

The ancestral habitat of capercaillie in most of its range isScots pine forest with an understorey of blaeberry. Keillourforest in Perthshire is dominated by Sitka spruce, has littleblaeberry, yet had a high density of capercaillie. Food andhabitat use by capercaillie were studied there from 1991 to1994. Seven cocks and two hens were caught, and marked withsmall radio transmitters. The habitats they occupied and foodremains in their droppings provided information on home range,habitat utilization and diet through the year. Sitka sprucewas the main food in autumn, winter and spring but was equalledby the fruiting heads of sedge in summer. Larch was the secondmost abundant item in spring and autumn diets. Sedge was plentifulon the wet restocks associated with the gleyed soils at Keillourand may provide a partial substitute for blaeberry in summer.Capercaillie showed a clear preference for the oldest blocksof trees, especially in winter. Sitka spruce, Norway spruceand Scots pine were used roughly in proportion to their availabilityin the birds' home range, but larch stands were used more thanexpected from availability. In summer, all birds used restocksto some extent. Management recommendations to encourage capercailliein commercial plantations largely comprising exotics are thatmature stands over 45 years and well-vegetated restocks shouldeach occupy at least 20 per cent of the forest, larch and blocksof mixed conifers each at least 15 per cent of the ground. Blaeberryshould be encouraged and, if possible, patches of windthrowinvolving living trees with the main root intact should be retained.     

Storage and internal cycling of nitrogen in relation to seasonal growth of Sitka spruce

Three-year-old clonal cuttings of Picea sitchensis (Bong.) Carr. were grown for two years (1988-1989) in sand irrigated with a nutrient solution containing either 1.0 mol N m(-3) (low N) or 6.0 mol N m(-3) (high N) NH(4)NO(3). In 1988, all the N provided was enriched with (15)N to 4.95 atom % (labeled N). In 1989, N was supplied with (15)N at natural abundance (unlabeled N). The recovery of unlabeled and labeled N in new foliage was used to quantify the internal cycling of N. In the high-N treatment, trees had two flushes of shoot growth and a period of rapid root growth, which coincided with the second flush of shoot growth in August. The timing of root growth and the first flush of shoot growth was similar in the low-N treatment, but there was no second flush of shoot growth and a greater proportion of biomass was recovered in roots. By November 1989, the root/needle dry matter ratio was 1.95 for the low-N trees and 1.36 for the high-N trees. Nitrogen was stored overwinter in roots and current-year needles. During the first six weeks of growth in the spring of 1989, stored N was remobilized for new foliage growth. Subsequent growth depended on root uptake of N. Remobilization of stored N was apparently not affected by the current N supply, because the amount of unlabeled N recovered in foliage produced in 1988 was the same for both N treatments. During 1989, the proportion of (15)N remobilized from roots relative to that from leaves produced in 1988 was greater in low-N trees than in high-N trees. In the autumn of both years, there was rapid uptake of N into roots and current-year needles. The effects of N supply on tree growth and nitrogen use efficiency are discussed in terms of the capacity for both N storage and internal cycling.     

Long-term growth and yield effects of respacing natural regeneration of Sitka spruce in Britain

Stands of Sitka spruce (Picea sitchensis (Bong.) Carr.) originating from natural regeneration can be extremely dense with high mortality, poor growth form and low volume production. Although costly, respacing (pre-commercial thinning) can reduce resource competition for the remaining trees, altering the final crop produced. Three experiments were established to examine the effect of respacing 1–2 m tall trees to different target densities. The impact on stand growth and yield was measured 11–17 years after respacing, and the longer-term impact was modelled to year 50. Unrespaced control treatments self-thinned at a similar and constant rate at two sites. At the third, extremely high initial density resulted in a higher rate of self-thinning and lower density after 11 years. Wider respacing treatments resulted in larger mean tree diameter, but there was no significant effect of respacing on stand volume 11–17 years after respacing; greater diameter growth did not compensate for low tree number. Results indicate a volume penalty associated with delaying respacing until trees were 4 m tall, but this treatment was unreplicated. Modelled stand volume in year 50 was higher for the 2.6 m × 2.6 m respacing treatment than for the 1.8 m × 1.8 m, 2.1 m × 2.1 m or 3.3 m × 3.3 m treatments. However, open-grown conditions may reduce timber quality compared to closer spacing treatments. The results are presented using a stand density management diagram for Sitka spruce growing in Canada and support recent suggestions that British stands have a shallower size–density relationship than Canadian stands.     

Dynamic soil processes on heathland due to changes in vegetation to oak and Sitka spruce

Soil and soil water chemistry under three different vegetation-types on a former heath area were investigated in Jutland, Denmark. An unmanaged heathland left undisturbed since the start of this century is now invaded by the oaks. Together with a small Sitka plantation (age 60 years), soil development due to vegetation changes was investigated. The typical heathland podzol soil had changed within decades under the oak trees towards an acid brown soil. The soil under the Sitka spruce had changed towards a more strongly podzolized soil compared to the former heath soil. Present processes occurring within the soil were investigated by analysis of soil solution chemistry. The leaching of inorganic ions and soluble organic matter had increased manifold under Sitka spruce. Nitrification increased markedly with soil depth probably due to effects of higher pH. The increase of nitrate with soil depth was followed by a similar increase in dissolved aluminium. The equivalence of the dissolved organic carbon varies under oak and heath between 6 and 10 m eqv g⁻¹ TOC and under Sitka the equivalence of the dissolved carbon was between 10 and 15 m eqv g⁻¹. No nitrate and only traces of ammonium were found in soil water from the upper 30 cm of the soil under heath. Soil nitrogen was almost exclusively (more than 90%) connected to the carbon cycle. Under Sitka spruce generally less than 20% of soil water nitrogen was coupled to the carbon cycle, i.e. most of the nitrogen in the soil water leaving the rooting zone under Sitka spruce was nitrate. In the oak forest, nitrate only appeared as traces in the soil water in the winter period and in the summer period in relatively higher amounts. Generally, between 80% and 90% of the nitrogen was coupled to the carbon cycle in the winter and in the summer only 50%.     

Root growth in Sitka spruce and Douglas-fir transplants: dependence on the shoot and stored carbohydrates

When dormant Sitka spruce (Picea sitchensis (Bong.) Carr.) transplants were lifted from the nursery in March and transferred to favorable growing conditions in a controlled environment chamber, new roots emerged and elongated even when the plants were bark-ringed so that the roots could not receive metabolites from the shoot by phloem translocation. Carbohydrate analysis showed that starch reserves in the roots of the bark-ringed plants were depleted, indicating that these reserves provided the energy source for new root growth. The shoot also had a role in new root growth as excised root systems showed greatly reduced root growth. This response is compared to that of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) in which bark-ringing prevented root growth indicating that current photosynthate is needed for new root growth. The findings are discussed in relation to the problems of establishing these species in forest plantations.     

The effect of nitrogen supply on root growth and development in sycamore and Sitka spruce trees

Sycamore (Acer pseudoplatanus L.) and Sitka spruce (Picea sitchensis(Bong.) Carr.) were grown in sand culture in glasshouses withtwo levels of N supplied with irrigation 1.0 (deficient) and6.0 (sufficient) molNm^–3, all as NH₄NO₃. Providing trees with a sufficient N supply increased both theirabove ground growth and N content compared with the N deficientplants. Provision of the generous N supply also altered thedistribution of growth between leaves and roots, with the N-deficientplants having a higher root:leaf or root:needle ratio than thewell fertilized trees. Provision of a generous N supply stimulatedroot growth, especially at the later harvests, which was reflectedmainly in growth of the tap root in sycamore. There was an increasein both root mass and root diameter with N supply. When thegrowth of white roots was monitored using a borescope, it wasshown that there were significantly more new roots producedby the well-fertilized trees. The borescope data also showeddetail of the bimodal seasonal pattern of root growth underhigh N in Sitka spruce which related to trends in nitrogen uptake.     

Physical and chemical responses of Sitka spruce (Picea sitchensis) clones to colonization by Heterobasidion annosum as potential markers for relative host susceptibility

Forty-one 2-year-old clones of Picea sitchensis (Bong.) Carr. from three full-sib families (14 clones from each of two families and 13 clones from a third family) were either wounded and inoculated with an isolate of Heterobasidion annosum (Fr.) Bref. or wounded without inoculation. Lesion lengths on the inner bark from the point of inoculation varied among clones 35 days after treatment. There was no relationship between lesion length and relatedness within families. Two clones (21342 and 25202) with the shortest lesions, tentatively designated as less susceptible to H. annosum, and two clones (21176 and 27166) with the longest lesions, designated more susceptible, were selected for comparison of host anatomical and chemical responses to infection. The position and structure of the ligno-suberized boundary zone (LSZ) in the bark of the clones suggested that the less susceptible clones formed thicker layers of suberized cells in the LSZ following wounding plus inoculation. No LSZ was observed in two ramets of the more susceptible Clone 27166 following wounding and inoculation with H. annosum. Compared with more susceptible genotypes, clones of P. sitchensis with low susceptibility to H. annosum had high relative proportions of (+)-alpha-pinene, (-)-beta-pinene and one unidentified terpene constituent (Unknown-15) in cortical resin sampled 25 cm from the lesions. In contrast, more susceptible clones had higher relative proportions of (-)-limonene, Unknown-16, Unknown-18 and Unknown-19. In the secondary resin produced in bark tissues surrounding the lesions, proportions of several monoterpene constituents varied; these changes included a decrease in the relative amount of beta-phellandrene and corresponding small increases in some minor constituents. The concentrations of the monoterpenes, except a few minor constituents, increased in the infected tissues. Wounding plus inoculation with H. annosum resulted in varied monoterpene responses, with distinct differences between less susceptible and more susceptible clones. In less susceptible clones, Unknown-19 increased following wounding plus inoculation, whereas in more susceptible clones, concentrations of delta-3-carene and Unknown-13 and Unknown-16 increased. Differences in both constitutive and induced resin monoterpene profiles may provide useful markers for resistance to H. annosum in selection and breeding programs.