Classifying seedlots of Picea sitchensis and P. glauca in zones of introgression using restriction analysis of chloroplast DNA

Summary Chloroplast DNA (cpDNA) restriction analysis was used to classify five reforestation seedlots as to species. The material included two Sitka spruce (Picea sitchensis (Bong.) Carr.), one white spruce (P. glauca (Moench) Voss) from interior British Columbia, and two putative hybrid seedlots from the coast-interior introgression zone in British Columbia. The cpDNA patterns generated by Bam-HI and Bc1-I from individual trees of Sitka spruce, white spruce, western white spruce (P. glauca var. albertiana (S. Brown)), and Engelmann spruce (P. engelmanni (Parry)) were species-specific. They were used as reference patterns for comparisons. In addition, two controlled crosses between white and Sitka spruce were analyzed to demonstrate the paternal inheritance of cpDNA in spruces. The cpDNA restriction patterns for the five seedlots were obtained from composite samples of seedlings from each lot and compared to the typical cpDNA patterns of each species. Restriction patterns for the two Sitka spruce seedlots agreed with those from the Sitka spruce tree, while patterns for the white spruce seedlots from British Columbia agreed with those from the white spruce tree, lacking evidence of any Engelmann spruce component in the sample. On the other hand, one putative hybrid seedlot showed cpDNA patterns similar to white spruce while the other showed fragments unique to both Sitka and white spruce, indicating that this was a hybrid seedlot. The analysis of cpDNA restriction polymorphism has proven to be an effective tool for classifying seedlots in regions of introgression. To our knowledge, these results provide the first demonstration of the use of cpDNA analysis for solving practical forestry problems.     

Effect of open-air fumigation with sulphur dioxide and ozone on phyllosphere and endophytic fungi of conifer needles

The phyllosphere microbial populations inhabiting the needles of three conifer species, Scots pine (Pinus sylvestris L.), Sitka spruce (Picea sitchensis L.) and Norway spruce (Picea abies (L.) Karst.), exposed to SO₂ and O₃, in an open-air fumigation experiment were analysed over a 3 year period using serial dilution after washing, direct plating and a fluorescein diacetate (FDA) enzyme assay. Total fungal populations ranged from 10² to 10⁵ colonyforming units (CPU) g^?1 fresh weight of needles. The dominant fungi isolated from needles varied with tree species and isolation technique; Aureobasidium pullulans (de Bary) Arnaud was most common on Scots pine and Norway spruce and white yeasts on Sitka spruce using the dilution plating method. However, direct plating of needle segments onto culture media indicated that Sclerophoma pythiophila (Corda) Hohnel was dominant on Scots pine and A. pullulans on Sitka and Norway spruce. Green needles of Sitka spruce were found to be endophytically colonized by Rhizosphaera kalkhoffii Bubak, but seldom by Lophodermium piceae (Fuckel) Hohn during extensive sampling in 1990. Statistical analyses revealed significant differences (P<0.05) between plots in the 3 year mean of the total fungal populations or the fungal biomass (FDA assay) on all three tree species. Differences between plots were also observed for a number of dominant component species. Data were also analysed for treatment effects. A significant effect of SO₂ treatment was observed on the total fungal populations on Sitka spruce (P<0.05) which were reduced markedly by the low-SO₂ treatment, while the O₃ treatment caused a significant increase in total fungal numbers on Scots pine (P<0.05). The FDA activity on needles of both Scots pine and Sitka spruce was noticeably higher in the 0₃-only treatment plot, but the overall O₃ effect was not significant. Treatment effects were also detected on the occurrence of component species. The serial dilution method revealed an SO₂ effect (P<0.05) of a reduction in the occurrence of pink yeasts on Sitka spruce and an O₃ effect (P<0.05) of an increase in the occurrence of S. pythiophila on Sitka spruce (P<0.01) but a decrease of Epicoccum nigrum Link and Cladosporium spp. on Scots pine. The direct-plating method revealed an SO₂ effect of an increase in S. pythiophila on Norway spruce (P<0.05). Ozone treatment caused a significant increase in the isolation of a black strain of A. pullulans on Norway spruce (P<0.05). Endophytic colonization of Sitka spruce needles by R. kalkhoffii was found to be increased on two occasions by O₃ exposure.     

Interactions between green spruce aphid (Elatobium abietinum (Walker)) and Norway and Sitka spruce under high and low nutrient conditions

1 Green spruce aphid (Elatobium abietinum) is a serious pest of spruce (Picea spp.) in north‐west Europe, causing defoliation of one‐year‐old and older needles. 2 Relationships between population development of E. abietinum, needle loss and tree growth were compared for five pure genotypes of Sitka spruce and mixed‐genotype material of Sitka and Norway spruce, grown under high and low nutrient conditions. 3 Despite wide differences in flushing date between spruce genotypes, E. abietinum populations peaked on the same date on each genotype and on the mixed‐genotype material, irrespective of nutrient supply. 4 Larger aphid populations developed on trees grown under high nutrient conditions than under low nutrients. However, more needles were lost per aphid in the low nutrient treatment and overall defoliation rates in the two nutrient treatments were similar. 5 Total aphid numbers differed significantly between Sitka spruce genotypes within nutrient treatments, but not in relation to bud‐burst or needle terpene content. Reductions in height growth caused by infestation were greater (15–44%), and related to mean aphid densities and defoliation, in the low nutrient treatment, but were smaller (11–27%) and not related to aphid density and defoliation in the high nutrient treatment. 6 Development of E. abietinum populations was similar on Norway and Sitka spruce, but Norway spruce lost fewer needles. However, the effects of infestation on tree growth were more closely related to aphid density and were similar for Norway and Sitka spruce. 7 Infestation caused a decrease in total root dry weight of Norway and Sitka spruce in proportion to the reductions observed in above‐ground growth.     

Microsatellites as DNA markers in Sitka spruce

Nine microsatellite loci were found by screening a genomic DNA library of Sitka spruce (Picea sitchensis) with the four oligonucleotide probes (TG), (CAC), (GATA) and (AT). Pairs of flanking primers were generated for seven microsatellites. Five primer pairs were used to screen up to 58 Sitka spruce clones. The five loci SStg3a, SStg4, SStg4a, SStg4c and SSgataS were found to have 15, 13, 4, 3 and 6 different length alleles respectively, and in using a combination of them almost all 58 Sitka spruce genotypes could be identified. The five primer pairs were successful in amplifying DNA from two other spruce species (Picea albutilia and Picea smithiana), while only one primer pair could amplify DNA from the pine species, Pinus sylvestris and Pinus latifolia. The inheritance of microsatellites in Sitka spruce was co-dominant Mendelian.     

Investigating the abundance and flight period of bark beetles (Coleoptera: Curculionidae: Scolytinae) over elevational gradients in Sitka spruce forests

1. A warming climate, as predicted under current climate change projections, is likely to influence the population dynamics of many forest insect species. Numerous bark beetle species in both Europe and North America have already responded to a warming climate by significantly expanding their geographical ranges.
2. The aim of the current study was to investigate how populations of bark beetles within stands of Sitka spruce, a widely planted non-native commercial plantation tree species in the U.K., were likely to respond to a warming climate. Experimental plots were established in stands of Sitka spruce over elevational gradients in two commercial forest plantations, and the abundance and emergence times of key bark beetle species were assessed over a 3-year period using flight interception traps. The air temperature difference between the lowest and highest experimental plot in each forest was consistently >1°C throughout the 3-year period.
3. In general, the abundance of the most dominant bark beetle species (e.g. Trypodendron, Dryocoetes, Hylastes spp.) was higher, and emergence times tended to be earlier in the year at the lower elevation plots, where temperatures were higher, although not all bark beetle species responded in the same manner.
4. The results of the study indicated that, under the projected future climate warming scenarios, monoculture Sitka spruce stands at low elevations may potentially be more vulnerable to significant outbreak events from existing or invasive bark beetle species. Hence, consideration of establishing more resilient forests of Sitka spruce by diversifying the species composition and structure of Sitka spruce stands is discussed.

     

Effect of P supply upon seasonal growth and internal cycling of P in Sitka spruce (Picea sitchensis(Bong.)Carr.) seedlings

The availability of phosphorus in many UK forest soils limits growth of Sitka spruce (Picea sitchensis (Bong.) Carr.). Efficient cycling of P within such systems is therefore necessary for sustained tree growth. Internal cycling of P is an important component of the overall P cycle in forests and the current work aims to quantify the impact of P nutrition on internal cycling and seasonal growth of Sitka spruce.Two-year old seedlings of Sitka spruce were grown in sand culture in the glasshouse for one year. Two treatments were imposed in which trees received either a complete nutrient solution from which P was excluded (-P) or one in which P was applied as labelled ³²P (+P). Internal cycling of P was measured directly in plants which had received no P and by difference in those which received ³²P.The contrasting P treatments produced an eight-fold difference in P content and a three-fold difference in tree growth between May and October. Root:shoot ratios increased during the growing season from 0.29 to 0.38 and from 0.29 to 0.52 in +P and-P treatments, respectively. In both treatments P was translocated from old shoots to support new shoot growth. P supply did not affect the amount of P remobilised but there was evidence that the rate of remobilisation may have been affected. The partition of remobilised P was affected by current P supply and differed from the partition of current P uptake.Results are compared to those from studies of growth and internal cycling of nitrogen in Sitka spruce.     

Do vascular plants and bryophytes respond differently to coniferous invasion of coastal heathlands?

Invasion by non-native conifers may pose a threat to local biodiversity, but knowledge about introduced conifer effects on Northern Hemisphere ecosystems is scarce. The coastal heathlands of north-west Europe are threatened by invasion of native and introduced tree species. We assess how spread of the introduced conifer Sitka spruce (Picea sitchensis (Bong.) Carr.) into European coastal heathlands affect two major functional groups; vascular plants and bryophytes, and how these effects relate to the environmental changes imposed by the developing tree canopies. We compared the impact of introduced Sitka spruce and native Scots pine (Pinus sylvestris L.) by analysing effects on species richness and turnover of vascular plants and bryophytes along fine-scale transects from individual tree stems into open heathland vegetation. Environmental impacts were assessed by measured environmental variables, and the responses of the two species groups were assessed by calculating changes in their respective mean Ellenberg indicator values. Species richness decreased beneath both conifers, related to decreased light and increased nitrogen and pH. Whereas vascular plants responded negatively to poor light conditions beneath dense and low Sitka spruce canopies, bryophytes were more negatively affected by the warmer and drier microclimates beneath Scots pine. Introduced Sitka spruce impacts the sub-canopy environment differently from the native Scots pine, and the two functional plant groups responded differently to these impacts. This suggests that future forests are likely to differ in species richness and composition, depending on whether succession is based on native or introduced coniferous trees.     

Differences in the probing behaviour of Elatobium abietinum feeding on Sitka and Norway spruces

The probing behaviour of Elatobium abietinum (Walk.) on the needles of Norway spruce and Sitka spruce was directly related to the stomatal frequency accompanied by heavy deposition of saliva, and multiple branching and divergence of the salivary sheath, all of which were associated with the impenetrability of the endodermis. On Norway spruce these features were very greatly reduced where there was a higher rate of probing coupled with a higher rate of penetration into the phloem, which was not accompanied by early uptake of phloem sap when compared with aphids on Sitka spruce. These differences in probing behaviour are related to damage symptoms exhibited by the host trees.     

Morphogenesis in Sitka spruce (<Emphasis Type="Italic">Picea sitchensis</Emphasis> (Bong.) Carr.) bud cultures – tree maturation and explants from epicormic shoots

The adventitious bud forming ability of Sitka spruce (Picea sitchensis (Bong.) Carr.) buds in vitro was found to be dependent upon the age of tree from which the explants were taken. Bud formation declined exponentially with increasing tree age when 1.0 and 10 μM 6-benzylaminopurine (BA) were used to induce adventitious buds. When less BA was used (0.1 μM) bud production was much lower with all ages of tree and no mathematical relationship between declining bud production and tree age was found. By a tree age of 6 years bud-forming ability had declined severely. Even the few buds that developed on older tree tissues failed to elongate into shoots, became necrotic and eventually died, indicating that adventitious bud induction in this species is not a rejuvenative process. Callusing of bud explants also declined with increasing tree age when 0.1 μM BA was used whilst very little callusing occurred at the higher cytokinin concentrations (1.0 and 10 μM BA). Tissue necrosis in vitro increased with tree age, whilst the ability of BA to retard necrosis declined with increasing tree age. Buds from epicormic shoots, formed on the lower trunks of 20-year-old trees when these were exposed to light, were not significantly better at forming adventitious buds in vitro than buds taken from the lower branches of the crown.     

Successful reproduction and pheromone production by the spruce bark beetle in evolutionary naïve spruce hosts with familiar terpenoid defences

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Development of a set of highly polymorphic genomic microsatellites (gSSRs) in Sitka spruce (Picea sitchensis (Bong.) Carr.)

Robust, polymorphic microsatellite DNA markers (simple sequence repeats—SSRs) are valuable tools for a range of tree conservation and breeding applications. SSRs are routinely used in the study of population genetic structure and diversity, pedigree reconstruction and genetic linkage mapping. Their abundance in the genome, co-dominant inheritance and potential for cross-species amplification make microsatellites highly prized markers. This paper characterises 22 novel genomic polymorphic microsatellite loci for Sitka spruce (Picea sitchensis (Bong.) Carr.). Amplification of DNA from Sitka spruce material was carried out both with a set of unrelated trees to obtain diversity statistics for each locus, and with the progeny of a full-sib family to test simple Mendelian inheritance. Observed heterozygosity ranged from 0.38 to 0.91 and allele number per locus ranged from 6 to 21, with a mean of 12.2. In addition, the primer pairs were tested with DNA from Norway spruce (P. abies) and white spruce (P. glauca) to investigate their potential for cross-species amplification and ten loci amplified in all three species. The results from these genomic microsatellites are compared to data generated from microsatellites derived from Picea EST libraries. In summary, this novel, highly polymorphic markers represent a significant addition to the rapidly expanding Picea genomics tool-box. Electronic supplementary material  The online version of this article (doi:) contains supplementary material, which is available to authorized users.     

The effect of light on the endogenous levels of cytokinins and gibberellins in seeds of sitka spruce (Picea sitchensis Carriere)

Abstract Maximum germination of Sitka spruce seed was obtained at 22°C under continuous fluorescent light or following a single 10 min exposure to red light (RL) given after 48 h of dark imbibition. The levels of endogenous cytokinins and gibberellins were examined in seeds which were exposed to different light regimes. Cytokinin levels in Sitka spruce seeds increased approximately 1.7–fold during exposure to continuous white light, during dark stratification, or within 1 h or a 30 min exposure to RL. A single peak of activity could be eluted from the LH^?20 Sephadex column which appeared in the same volume as authentic zeatin riboside. Gibber-ellin activity in Sitka spruce seeds increased rapidly during exposure to 30 min of RL and then declined during subsequent dark incubation. Similarly, under continuous white light, gibberellin activity was at least twice as high as in the dark. Two gibberellm-like substances were found in Sitka spruce seeds which appeared to be similar to GA₇ and GA₉ on the basis of their chromato-graphic properties. The zone of gibberellin activity directly associated with photo-excitation of Sitka spruce seeds contains conjugate gibberellins, possibly glucosyl esters, which when hydrolysed with β-glucosidase form two products which are similar to the free gibberellins which were detected. The light response in Sitka spruce seed is compared to a similar response in Scots pine (Pinus sylvestris L.) seeds.     

Growth of coniferous trees exposed to SO2 and O3 using an open-air fumigation system

Scots pine (Pinus sylvestris L.), Norway spruce (Picea abies (L.) Karst.) and Sitka spruce (Picea sitchensis Bong. Carr.) were planted as 2-year-old seedlings in an open-air fumigation facility at Liphook in southern England in March 1985. The soil was a humoferric podzol of pH 4. SO₂ fumigation began in May 1987 and continued until December 1990. Long-term mean SO₂ concentrations were 4,13 and 22 nmol mo^?1. Three plots, one at each SO₂ level, were also exposed to O₃ at an average of 1–3.times the ambient level. O₃ fumigation ran from March to December 1988, May to December 1989 and February to December 1990. Each species reacted differently to treatment. Scots pine showed no growth response to either pollutant, although other work on the site demonstrated a number of deleterious effects of SO₂ on this species, including increased leaf loss and foliar injury. Stem basal diameter growth of Norway spruce was depressed in SO₂-treated plots. In contrast, extension growth of shoots of Sitka spruce increased in SO₂-treated plots, in apparent response to codeposition of NH₃-N. However, diameter growth of Sitka spruce main stems did not increase. No effects of O₃ on growth were recorded for any species.     

Influence of ectomycorrhizal fungi on the response of Sitka spruce and Japanese larch to forms of phosphorus

Ectomycorrhizal fungi (Paxillus involutus, Suillus grevillei and two unidentified basidiomycetes from excised Sitka spruce mycorrhizas) were isolated from stands of Sitka spruce either in monoculture or in a mixture with Japanese larch in an Irish conifer plantation. The growth of these fungi and their mycorrhizal formation in Sitka spruce and Japanese larch were examined after incubation in modified Melin-Norkrans medium containing either KH₂PO₄, Ca₃(PO₄)₂ or Fe phytate as the phosphorus (P) source. P. involutus and S. grevillei utilized all three P sources. The unidentified basidiomycetes had limited ability to utilize Fe phytate. Basidiomycete 1 showed poor growth on KH₂PO₄ whereas growth of basidiomycete 2 was low on Ca₃(PO₄)₂. Pure culture synthesis studies confirmed that P. involutus and the two basidiomycetes formed mycorrhizas with both tree species but S. grevillei was mycorrhizal only on Japanese larch. P. involutus formed more mycorrhizas in both conifers than the other fungi. Following inoculation with each of the four fungi, shoot and root dry mass of both Sitka spruce and Japanese larch seedlings was enhanced compared with uninoculated/nonmycorrhizal controls. On Fe phytate, Paxillus-inoculated Sitka spruce seedlings had the lowest primary root length and on KH₂PO₄, Suillus-inoculated Japanese larch had the greatest number of short roots. The only differences when Sitka spruce seedlings were grown in either monoculture or in a mixture with Japanese larch mycorrhizal with S. grevillei were primary root length and number of short roots after growth on media containing Fe phytate.     

Damage caused by the Green Spruce Aphid to Norway and Sitka spruce needles

Three-month-old needles of Sitka spruce were less susceptible to Elatobium abietinutn than 15-month-old needles. Symptoms appeared after longer aphid feeding times but only a proportion of damaged needles fell. After short feeding periods symptoms appeared in more Norway spruce needles than in Sitka spruce, whereas longer feeding periods resulted in more needles producing symptoms in the Sitka spruce. The symptoms took 4—6 days longer to appear in Norway spruce, and needle fall followed a longer feeding period than on Sitka spruce. Following 72 h feeding, needle fall occurred more quickly on Sitka spruce than on Norway spruce. The time taken for needle fall to occur was inversely related to the feeding time in Sitka spruce but such a response was not evident in Norway spruce. The results are discussed in relation to the differences exhibited in the probing behaviour of the aphid on the two spruces.     

Drought intensity and frequency have contrasting effects on development time and survival of the green spruce aphid

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Are bird density,species richness and community structure similar between native woodlands and non-native plantations in an area with a generalist bird fauna?

This study compared the bird assemblages of native semi-natural woodlands and non-native Sitka spruce (Picea sitchensis) plantations in Ireland to identify what vegetation variables most influenced birds and to identify management targets in plantations to maximise future bird conservation. Point counts were conducted in 10 Oak (Quercus spp.) and 10 Ash (Fraxinus excelsior) native woodlands and in five Mid-rotation (20–30 years old) and five Mature (30–50 years old) Sitka spruce plantations. Ordination was used to characterise woodland types according to their constituent bird species. Total bird density (calculated using Distance software) and species richness were assessed for the different woodland types. Oak and Ash woodland bird assemblages were separated from Mid-rotation and Mature plantations by the ordination. There was no difference in total bird density between any of the woodland types. Oak woodlands had significantly higher species richness than either Mid-rotation or Mature Sitka spruce plantations. Ash had higher species richness than Mature Sitka spruce plantations. Understorey vegetation was negatively associated with total bird density, which also varied with survey year. Understorey vegetation was positively associated with species richness. Reasons for the relationships between vegetation and bird assemblages are discussed. Management should seek to increase shrub and understorey vegetation in the Mid-rotation phase to improve the contribution of plantations to bird conservation.     

Insects associated with the Sitka spruce weevil,Pissodes Strobi [Col.: Curculionidae] on Sitka spruce,Picea sitchensis in British Columbia,Canada

BioControl - Insects associated with the Sitka spruce weevil,Pissodes strobi (Peck), on Sitka spruce,Picea sitchensis (Bong.) Carr. were sampled at 9 locations in British Columbia. Fourteen species...     

Host shifting by Operophtera brumata into novel environments leads to population differentiation in life-history traits

Abstract.  1. Operophtera brumata L. (Lepidoptera: Geometridae), a polyphagous herbivore usually associated with deciduous trees such as oak Quercus robur L . , has expanded its host range to include the evergreen species heather Calluna vulgaris (L.) Hull and, most recently, Sitka spruce Picea sitchensis (Bong.) Carrière.
2. Phenology, morphology, and survival of O. brumata were measured at several life-history stages in populations from the three different host plant communities sampled from a range of geographical locations. The data were used to test for population differences, reflecting the marked differences in host-plant secondary chemistry, growth form, and site factors such as climate. The hypothesis that spruce-feeding populations originated from populations feeding on moorland, commonly sites of coniferous afforestation, was also tested.
3. Altitude, not host plant species, was the major influence on the timing of adult emergence. An effect of insect population independent of altitude was found, implying that additional unidentified factors contribute to this phenological variation. Larval survival and adult size varied between populations reared on different host plant species. Survival of larvae was affected negatively when reared on the novel host plant, Sitka spruce, versus the natal plant (oak or heather) but oak and heather-sourced insects did not differ in survivorship on Sitka spruce.
4. Host range extension into novel environments has resulted in population differentiation to the local climate, demonstrating that host shifts pose challenges to the herbivore population greater than those offered by the host plant alone. The hypothesis that Sitka spruce feeding populations have arisen predominantly from moorland feeding populations was not supported.     

Ecology of streams draining forested and non-forested catchments in an area of central Scotland subject to acid precipitation

A study of 12 streams draining forested and non-forested catchments was made in an area of central Scotland where slow-weathering bedrock was predominantly quartzite, schists and slates. Sitka spruce (Picea sitchensis Carriere) was the most common tree species. Precipitation in the area had an annual mean pH in the range 4.3–4.5. Streams within the planted zone were always more acid than those outside and had higher concentrations of aluminium and manganese. With one exception, trout were absent from streams within long-established forests and planted salmon eggs (Salmo salar L.) died within a few weeks. A high proportion of such eggs survived in streams outside the forest. Siphlonurus lacustris Eaton was the only mayfly nymph found in the most acid streams in summer collections. In winter samples, mayfly nymphs, Heptagenia lateralis (Curtis) were found in only one forest stream but several species were present in the non-forested catchments. It is suggested that spruce forests can effectively collect acid pollutants which are subsequently washed out, thus accelerating the acidification of the soil. Streams therefore become increasingly acid as the neutralisation capacities of their catchments decrease.