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PubMed2006

Sensitivity of Mediterranean woody seedlings to copper, nickel and zinc.

The restoration of heavy metal contaminated areas requires information on the response of native plant species to these contaminants. The sensitivity of most Mediterranean woody species to heavy metals has not been established, and little is known about phytotoxic thresholds and environmental risks. We have evaluated the response of four plant species commonly used in ecological restoration, Pinus halepensis, Pistacia lentiscus, Juniperus oxycedrus, and Rhamnus alaternus, grown in nutrient solutions containing a range of copper, nickel and zinc concentrations. Seedlings of these species were exposed to 0.048, 1 and 4 microM of Cu; 0, 25 and 50 microM of Ni; and 0.073, 25 and 100 microM of Zn in a hydroponic silica sand culture for 12 weeks. For all four species, the heavy metal concentration increased in plants as the solution concentration increased and was always higher in roots than in shoots. Pinus halepensis and P. lentiscus showed a higher capacity to accumulate metals in roots than J. oxycedrus and R. alaternus, while the allocation to shoots was considerably higher in the latter two. Intermediate heavy-metal doses enhanced biomass accumulation, whereas the highest doses resulted in reductions in biomass. Decreases in shoot biomass occurred at internal concentrations ranging from 25 to 128 microg g-1 of Zn, and 1.7 to 4.1 microg g( -1) of Cu. Nickel phytoxicity could not be established within the range of doses used. Rhamnus alaternus and J. oxycedrus showed higher sensitivity to Cu and Zn than P. halepensis and, especially, P. lentiscus. Contrasted responses to heavy metals must be taken into account when using Mediterranean woody species for the restoration of heavy metal contaminated sites.

Anacardiaceae↗

PubMed2004

Cumulative impact of 40 years of industrial sulfur emissions on a forest soil in west-central Alberta (Canada).

The impact of 40 years of sulfur (S) emissions from a sour gas processing plant in Alberta (Canada) on soil development, soil S pools, soil acidification, and stand nutrition at a pine (Pinus contorta x Pinus banksiana) ecosystem was assessed by comparing ecologically analogous areas subjected to different S deposition levels. Sulfur isotope ratios showed that most deposited S was derived from the sour gas processing plant. The soil subjected to the highest S deposition contained 25.9 kmol S ha(-1) (uppermost 60 cm) compared to 12.5 kmol S ha(-1) or less at the analogues receiving low S deposition. The increase in soil S pools was caused by accumulation of organic S in the forest floor and accumulation of inorganic sulfate in the mineral soil. High S inputs resulted in topsoil acidification, depletion of exchangeable soil Ca2+ and Mg2+ pools by 50%, podzolization, and deterioration of N nutrition of the pine trees.

Alberta↗

PubMed2005

Case study: inorganic pollutants associated with particulate matter from an area near a petrochemical plant.

The area of Gela (Sicily, Italy) contains one of the largest petroleum refineries in Europe and also has several oil fields both on land and offshore. This paper discusses how the oil refinery and traffic-related air pollution affect the chemical composition of airborne particulate matter over the town of Gela, using pine needles and urban road dust as the means of survey. Forty-one samples of pine needles from Pinus halepensis (Mill.) and two composite samples of roadway dust, each subdivided into six size fractions, were analyzed for major and trace elements. Information on the natural or anthropogenic origin of the observed heavy metals was deduced from factor analysis and element distribution maps. Factor analysis was applied to a data set of 20 element concentrations in pine needles and identified three main sources of metals: soil, vehicle traffic, and industrial emissions. The petrochemical plant appears to be associated with raised levels of As, Mo, Ni, S, Se, V, and Zn. Similarly, enhanced Cu, Pb, Pt, Pd, Sb, and partly Zn concentrations are closely associated with traffic. High correlations between Ni and V, As and Se, and Pb and Sb were observed. Element distribution maps, showing a decrease in heavy metal contents immediately farther inland, confirm that local sources play a considerable role in heavy metal pollution. Morphological alterations and accumulation of phenols were observed in sections of Pinus halepensis needles collected from sites with high traffic density and industrial emissions.

Air Pollutants↗

PubMed2006

Antimicrobial and cytotoxic knotwood extracts and related pure compounds and their effects on food-associated microorganisms.

Knotwood or bark extracts prepared from 30 species of hard and soft wood trees as well as selected pure compounds (lignans, stilbenes and flavonoids) were assayed for their antimicrobial activity against a battery of both gram positive and negative bacteria, yeasts, and filamentous fungi (Bacillus cereus, Staphylococcus aureus, Listeria monocytogenes, Lactobacillus plantarum, Escherichia coli, Salmonella infantis, Pseudomonas fluorescens, Candida albicans, Saccharomyces cerevisiae, Aspergillus fumigatus and Penicillium brevicompactum). By far the most consistent antibacterial and antifungal properties were associated with extracts of Pinus species. These extracts showed also cytotoxicity against a mouse hepatoma cell line. Both antimicrobial and cytotoxic properties correlated with the stilbene content of the extracts. Purified stilbenes showed the most consistent antimicrobial and cytotoxic activities, while purified lignans had marginal effects, only. The results suggest that stilbenes account both for the antimicrobial and cytotoxic properties of Pinus knotwood extracts.

Anti-Bacterial Agents↗

PubMed2006

Evaluation of some plants used in Turkish folk medicine against parasitic infections for their in vivo anthelmintic activity.

Ethanolic and aqueous extracts obtained from nine plant species from seven families selected depending on their use in Turkish folk medicine, including Citrillus lanatus (Thunb.) Matsum. (seed), Jasminum fruticans L. (branches), Juniperus drupacea Labill. (fruits), Juniperus nana L. (fruit and leaves), Juniperus oxcycedrus L (fruit and leaves), Mentha longifolia L. (herba), Pinus nigra ssp. pallasiana (Lamb.) Richt. (fruits), Plantago lanceolata L. (leaves), and Zea mays L. (seed) were evaluated for their in vivo anthelmintic activity. Among the plant extracts studied, both ethanolic and aqueous extracts of Jasminum fruticans, Mentha longifolia and Pinus nigra ssp. pallasiana, the aqueous extracts of Zea mays, the ethanolic extracts of Citrillus lanatus, Juniperus drupacea (fruit), Juniperus oxcycedrus and Plantago lanceolata displayed significant anthelmintic activity against pinworms, Syphacia obvelata and Aspiculuris tetraptera, in mice. Rest of the extracts from plants did not show any remarkable anthelmintic activity. The results were considered significant at p<0.05.

Animals↗

PubMed2003

Quantifying ozone uptake at the canopy level of spruce, pine and larch trees at the alpine timberline: an approach based on sap flow measurement.

Micro-climatic and ambient ozone data were combined with measurements of sap flow through tree trunks in order to estimate whole-tree ozone uptake of adult Norway spruce (Picea abies), cembran pine (Pinus cembra), and European larch (Larix decidua) trees. Sap flow was monitored by means of the heat balance approach in two trees of each species during the growing season of 1998. In trees making up the stand canopy, the ozone uptake by evergreen foliages was significantly higher than by deciduous ones, when scaled to the ground area. However, if expressed per unit of whole-tree foliage area, ozone flux through the stomata into the needle mesophyll was 1.09, 1.18 and 1.40 nmol m(-2) s(-1) in Picea abies, Pinus cembra and Larix decidua, respectively. These fluxes are consistent with findings from measurements of needle gas exchange, published from the same species at the study site. It is concluded that the sap flow-based approach offers an inexpensive, spatially and temporally integrating way for estimating ozone uptake at the whole-tree and stand level, intrinsicly covering the effect of boundary layers on ozone flux.

Climate↗

PubMed2003

Development of enzyme-linked immunosorbent assays for isocupressic acid and serum metabolites of isocupressic acid.

The consumption of ponderosa pine (Pinus ponderosa), lodgepole pine (Pinus contorta), common juniper (Juniperus communis), and Monterey cypress (Cupressus macrocarpa) causes abortions in pregnant cattle. Recent studies have identified isocupressic acid (1) as the primary abortificient compound in these plants. In vitro and in vivo studies using rumen and blood have shown isocupressic acid (1) is rapidly metabolized to agathic acid (3), dihydroagathic acid (4), and tetrahydroagathic acid (5). Rapid and sensitive diagnostic techniques are needed to identify poisoned animals, to study toxicokinetics, and to elucidate the mechanism of isocupressic acid-induced abortion in cattle. In this study, four competitive inhibition enzyme-linked immunosorbent assays for isocupressic acid and its sera metabolites were developed using polyclonal antibodies. One assay is specific to 1, whereas the other three assays show cross-reactivity to 3-5 in addition to 1. The assay specific to 1 had a limit of detection of 44.1 pg. The other assays which demonstrated cross-reactivity to the isocupressic acid blood metabolites also had comparably low limits of detection. One assay was used to follow the absorption and elimination profile of isocupressic acid metabolites in both cow serum and urine after oral dosage of a cow with common juniper.

Abortifacient Agents↗

PubMed2003

Characterisation of Pt MYB1, an R2R3-MYB from pine xylem.

A cDNA encoding a member of the R2R3-MYB family of transcription factors was cloned from a library constructed from differentiating Pinus taeda (loblolly pine) xylem RNA. This MYB family member, Pinus taeda MYB1 (PtMYB1), was most abundantly expressed in differentiating xylem, as assessed by both ribonuclease protection assays, and by northern blot analysis with poly(A)-enriched RNA. Similar to other plant R2R3-MYB family members, recombinant Pt MYB1 protein was able to bind to AC elements in electrophoretic mobility shift assays (EMSAs). AC elements are DNA motifs rich in adenosine and cytosine that have been implicated in the xylem-localised regulation of genes encoding lignin biosynthetic enzymes. Pt MYB1 not only bound to AC elements, but was also able to induce AC-element-dependent shifts in the electrophoretic mobility of a plant promoter that contains three AC elements, the minimal PHENYLALANINE AMMONIA-LYASE 2 (PAL2) promoter from Phaseolus vulgaris. Transcriptional activation assays conducted using yeast showed that Pt MYB1 also activated transcription, and that it did so in an AC-element-dependent fashion. Pt MYB1 also activated transcription from the minimal PAL2 promoter in plant cells in an AC-element-dependent fashion, as revealed by transient transcriptional activation assays with microprojectile-bombarded tobacco NT-1 cells. Taken together, these finding are consistent with the hypothesis that Pt MYB1 may regulate transcription from cis -acting AC elements in pine xylem.

Adenosine Triphosphate↗

PubMed2004

Automated SNP detection in expressed sequence tags: statistical considerations and application to maritime pine sequences.

We developed an automated pipeline for the detection of single nucleotide polymorphisms (SNPs) in expressed sequence tag (EST) data sets, by combining three DNA sequence analysis programs: Phred, Phrap and PolyBayes. This application requires access to the individual electrophoregram traces. First, a reference set of 65 SNPs was obtained from the sequencing of 30 gametes in 13 maritime pine (Pinus pinaster Ait.) gene fragments (6671 bp), resulting in a frequency of 1 SNP every 102.6 bp. Second, parameters of the three programs were optimized in order to retrieve as many true SNPs, while keeping the rate of false positive as low as possible. Overall, the efficiency of detection of true SNPs was 83.1%. However, this rate varied largely as a function of the rare SNP allele frequency: down to 41% for rare SNP alleles (frequency < 10%), up to 98% for allele frequencies above 10%. Third, the detection method was applied to the 18498 assembled maritime pine (Pinus pinaster Ait.) ESTs, allowing to identify a total of 1400 candidate SNPs, in contigs containing between 4 and 20 sequence reads. These genetic resources, described for the first time in a forest tree species, were made available at http://www.pierroton.inra/genetics/Pinesnps. We also derived an analytical expression for the SNP detection probability as a function of the SNP allele frequency, the number of haploid genomes used to generate the EST sequence database, and the sample size of the contigs considered for SNP detection. The frequency of the SNP allele was shown to be the main factor influencing the probability of SNP detection.

Algorithms↗

PubMed2004

A community of unknown, endophytic fungi in western white pine.

The endophytic fungi of woody plants may be diverse as often claimed, and likewise, they may be functionally novel as demonstrated in a few studies. However, the endophyte taxa that are most frequently reported tend to belong to fungal groups composed of morphologically similar endophytes and parasites. Thus, it is plausible that endophytes are known (i.e., described) parasites in a latent phase within the host. If this null hypothesis were true, endophytes would represent neither additional fungal diversity distinct from parasite diversity nor a symbiont community likely to be novel ecologically. To be synonymous with parasites of the host, endophytes should at least be most closely related to those same parasites. Here we report that seven distinct parasites of Pinus monticola do not occur as endophytes. The majority of endophytes of P. monticola (90% of 2,019 cultures) belonged to one fungal family, the Rhytismataceae. However, not a single rhytismataceous endophyte was found to be most closely related by sequence homology to the three known rhytismataceous parasites of P. monticola. Similarly, neither endophytic Mycosphaerella nor endophytic Rhizosphaera isolates were most closely related to known parasites of P. monticola. Morphologically, the endophytes of P. monticola can be confounded with the parasites of the same host. However, they are actually most closely related to, but distinct from, parasites of other species of Pinus. If endophytes are generally unknown species, then estimates of 1 million endophytes (i.e., approximately 1 in 14 of all species of life) seem reasonable.

Base Sequence↗

PubMed2004

Nucleotide diversity and linkage disequilibrium in loblolly pine.

Outbreeding species with large, stable population sizes, such as widely distributed conifers, are expected to harbor relatively more DNA sequence polymorphism. Under the neutral theory of molecular evolution, the expected heterozygosity is a function of the product 4N(e)mu, where N(e) is the effective population size and mu is the per-generation mutation rate, and the genomic scale of linkage disequilibrium is determined by 4N(e)r, where r is the per-generation recombination rate between adjacent sites. These parameters were estimated in the long-lived, outcrossing gymnosperm loblolly pine (Pinus taeda L.) from a survey of single nucleotide polymorphisms across approximately 18 kb of DNA distributed among 19 loci from a common set of 32 haploid genomes. Estimates of 4N(e)mu at silent and nonsynonymous sites were 0.00658 and 0.00108, respectively, and both were statistically heterogeneous among loci. By Tajima's D statistic, the site frequency spectrum of no locus was observed to deviate from that predicted by neutral theory. Substantial recombination in the history of the sampled alleles was observed and linkage disequilibrium declined within several kilobases. The composite likelihood estimate of 4N(e)r based on all two-site sample configurations equaled 0.00175. When geological dating, an assumed generation time (25 years), and an estimated divergence from Pinus pinaster Ait. are used, the effective population size of loblolly pine should be 5.6 x 10(5). The emerging narrow range of estimated silent site heterozygosities (relative to the vast range of population sizes) for humans, Drosophila, maize, and pine parallels the paradox described earlier for allozyme polymorphism and challenges simple equilibrium models of molecular evolution.

Animals↗

PubMed2003

Photosynthetic functions of cembran pines and dwarf pines during winter at timberline as regulated by different temperatures, snowcover and light.

Trees at timberline in the high Alps are exposed to a variety of climatic conditions. Most climatic stresses occur during winter and spring, when frost, occasionally low snow cover, and high irradiation interact. In this study, we follow reactions of photosynthesis from high winter to spring in two dominating tree species of the alpine timberline, which may indicate the status of stress response to a changing environment. The results indicate a level of physiological stability in trees, which are important for stabilising natural high mountain ecosystems. Trees of Pinus cembra and of Pinus mugo were selected at altitudes between 1850 m a.s.l. and 1950 m a.s.l. near innsbruck, Austria. At six sampling times from January to May, fast chlorophyll fluorescence was measured in the field and twigs were collected for further investigation in the laboratory. The following measurements were taken: photosynthetic oxygen formation, needle chlorophyll and carotenoid determination, and kinetic studies of the xanthophyll cycle. In general, both tree species showed similar results in most parameters studied. P. mugo seems to have some advantages if winter precipitation is high, when, because of its growth habitus, most needles will be snow covered. Primary photochemistry (trapping per reaction centre) in PS II does not change with sampling dates despite the fact that temperature and light are changing. However, first events in electron transport and whole needle photosynthesis are strongly affected by light and temperature conditions during the days before sampling. The kinetics of the xanthophyll cycle indicate not only light, but also strong temperature effects. P. mugo photosynthesis seems to have a higher stability under changing weather. Both tree species are well prepared to start with photosynthesis in winter, if favourable conditions, like foehn events, occur.

Adaptation, Physiological↗

PubMed2003

Diffusion barriers of tripartite sporopollenin microcapsules prepared from pine pollen.

Tripartite sporopollenin microcapsules prepared from pine pollen (Pinus sylvestris L. and Pinus nigra Arnold) were analysed with respect to the permeability of the different strata of the exine which surround the gametophyte and form the sacci. The sexine at the surface of the sacci is highly permeable for polymer molecules and latex particles with a diameter of up to 200 nm, whereas the nexine covering the gametophyte is impermeable for dextran molecules, with a Stokes' radius > or =4 nm (Dextran T 70), and for the tetravalent anionic dye Evans Blue (Stokes' radius = 1.3 nm). The central capsules obtained by dissolution of the sporoplasts showed strictly membrane-controlled exchange of non-electrolytes, with half-equilibration times in the range of minutes (monosaccharides, oligosaccharides) to hours (dextran molecules with Stokes' radii up to 2.5 nm). The dependence of the permeability coefficients of the nexine for non-electrolytes on Stokes' radius or molecular weight shows that the aqueous pores through the nexine are inhomogeneous with respect to their size, and that most pores are too narrow for free diffusion of sugar molecules. To explain the barrier function of the nexine for Evans Blue, it is assumed that at least the larger pores, which enable slow permeation of dextran molecules, contain negative charges.

Carbohydrate Metabolism↗

PubMed2006

Intra-annual radial growth and water relations of trees: implications towards a growth mechanism.

There is a missing link between tree physiological and wood-anatomical knowledge which makes it impossible mechanistically to explain and predict the radial growth of individual trees from climate data. Empirical data of microclimatic factors, intra-annual growth rates, and tree-specific ratios between actual and potential transpiration (T PET(-1)) of trees of three species (Quercus pubescens, Pinus sylvestris, and Picea abies) at two dry sites in the central Wallis, Switzerland, were recorded from 2002 to 2004 at a 10 min resolution. This included the exceptionally hot and dry summer of 2003. These data were analysed in terms of direct (current conditions) and indirect impacts (predispositions of the past year) on growth. Rain was found to be the only factor which, to a large extent, consistently explained the radial increment for all three tree species at both sites and in the short term as well. Other factors had some explanatory power on the seasonal time-scale only. Quercus pubescens built up much of its tree ring before bud break. Pinus sylvestris and Picea abies started radial growth 1-2 weeks after Quercus pubescens and this was despite the fact that they had a high T PET(-1) before budburst and radial growth started. A high T PET(-1) was assumed to be related to open stomata, a very high net CO2 assimilation rate, and thus a potential carbon (C)-income for the tree. The main period of radial growth covered about 30-70% of the productive days of a year. In terms of C-allocation, these results mean that Quercus pubescens depended entirely on internal C-stores in the early phase of radial growth and that for all three species there was a long time period of C-assimilation which was not used for radial growth in above-ground wood. The results further suggest a strong dependence of radial growth on the current tree water relations and only secondarily on the C-balance. A concept is discussed which links radial growth over a feedback loop to actual tree water-relations and long-term affected C-storage to microclimate.

Carbon↗

PubMed2001

Family differences in height growth and photosynthetic traits in three conifers.

We investigated variation in height growth, gas exchange, chlorophyll fluorescence and leaf stable carbon isotope ratio among wind-pollinated progenies of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco var. glauca), ponderosa pine (Pinus ponderosa Dougl. ex Laws.) and western white pine (Pinus monticola Dougl. ex D. Don) from a small group of contiguous stands on the Priest River Experimental Forest in northern Idaho. Photosynthetic variables differed between height classes in the pines, but not in Douglas-fir. Among species and families, tall families of ponderosa pine regained photosynthetic capacity earliest in the spring and maintained it latest in the growing season. Tall families of western white pine had higher instantaneous water-use efficiencies and lower photosynthetic rates than short families on warm days in August.

Chlorophyll↗

PubMed2001

Time-series analysis of delta13C from tree rings. I. Time trends and autocorrelation.

Univariate time-series analyses were conducted on stable carbon isotope ratios obtained from tree-ring cellulose. We looked for the presence and structure of autocorrelation. Significant autocorrelation violates the statistical independence assumption and biases hypothesis tests. Its presence would indicate the existence of lagged physiological effects that persist for longer than the current year. We analyzed data from 28 trees (60-85 years old; mean = 73 years) of western white pine (Pinus monticola Dougl.), ponderosa pine (Pinus ponderosa Laws.), and Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco var. glauca) growing in northern Idaho. Material was obtained by the stem analysis method from rings laid down in the upper portion of the crown throughout each tree's life. The sampling protocol minimized variation caused by changing light regimes within each tree. Autoregressive moving average (ARMA) models were used to describe the autocorrelation structure over time. Three time series were analyzed for each tree: the stable carbon isotope ratio (delta(13)C); discrimination (delta); and the difference between ambient and internal CO(2) concentrations (c(a) - c(i)). The effect of converting from ring cellulose to whole-leaf tissue did not affect the analysis because it was almost completely removed by the detrending that precedes time-series analysis. A simple linear or quadratic model adequately described the time trend. The residuals from the trend had a constant mean and variance, thus ensuring stationarity, a requirement for autocorrelation analysis. The trend over time for c(a) - c(i) was particularly strong (R(2) = 0.29-0.84). Autoregressive moving average analyses of the residuals from these trends indicated that two-thirds of the individual tree series contained significant autocorrelation, whereas the remaining third were random (white noise) over time. We were unable to distinguish between individuals with and without significant autocorrelation beforehand. Significant ARMA models were all of low order, with either first- or second-order (i.e., lagged 1 or 2 years, respectively) models performing well. A simple autoregressive (AR(1)), model was the most common. The most useful generalization was that the same ARMA model holds for each of the three series (delta(13)C, delta, c(a) - c(i)) for an individual tree, if the time trend has been properly removed for each series. The mean series for the two pine species were described by first-order ARMA models (1-year lags), whereas the Douglas-fir mean series were described by second-order models (2-year lags) with negligible first-order effects. Apparently, the process of constructing a mean time series for a species preserves an underlying signal related to delta(13)C while canceling some of the random individual tree variation. Furthermore, the best model for the overall mean series (e.g., for a species) cannot be inferred from a consensus of the individual tree model forms, nor can its parameters be estimated reliably from the mean of the individual tree parameters. Because two-thirds of the individual tree time series contained significant autocorrelation, the normal assumption of a random structure over time is unwarranted, even after accounting for the time trend. The residuals of an appropriate ARMA model satisfy the independence assumption, and can be used to make hypothesis tests.

Carbon Isotopes↗

PubMed2002

Response of old-growth conifers to reduction in stand density in western Oregon forests.

The positive growth response of healthy young trees to density reduction is well known. In contrast, large old trees are usually thought to be intrinsically limited in their ability to respond to increased growing space; therefore, density reduction is seldom used in stands of old-growth trees. We tested the null hypothesis that old-growth trees are incapable of responding with increased growth following density reduction. The diameter growth response of 271 Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), ponderosa pine (Pinus ponderosa Dougl. ex Laws) and sugar pine (Pinus lambertiana Dougl.) trees ranging in age from 158 to 650 years was examined 20 to 50 years after density reduction. Density reduction involved either light thinning with removal of less vigorous trees, or shelterwood treatments in which overstory trees were not removed. Ratios of basal area growth after treatment to basal area growth before treatment, and several other measures of growth, all indicated that the old trees sometimes benefited and were not harmed by density reduction. Growth increased by 10% or more for 68% of the trees in treated stands, and nearly 30% of trees increased growth by over 50%. This growth response persisted for at least 20 years. During this 20-year period, only three trees in treated stands (1.5%) exhibited a rapid decrease in growth, whereas growth decreased in 64% of trees in untreated stands. The length of time before a growth response to density reduction occurred varied from 5 to 25 years, with the greatest growth response often occurring 20 to 25 years after treatment. These results have important implications both for the basic biology of aging in woody plants as well as for silvicultural practices in forests with old-growth trees.

Forestry↗

PubMed2002

Effects of phloem girdling in conifers on apical control of branches, growth allocation and air in wood.

We investigated effects of stem phloem girdles on apical control of branch angle, stem and branch growth and stem air content in six conifer species. A stem girdle 2 cm above a branch caused the branch to bend upward in all six species. Upward bending was associated with increased formation and action of compression wood (CW) in the lower portion of the branch. Compression wood also formed in the main stem below the branch, suggesting increased auxin production in the branch. A stem girdle 2 cm below a branch (the branch remained directly connected to the apex and distal branches) released the branch from apical control in Tsuga canadensis (L.) Carr., Pinus contorta Dougl. ex Loud. and Pseudotsuga menziesii (Mirb.) Franco. The branch bent up, but there was no CW formation in the stem. In Pinus rigida Mill., the branch exhibited increased cambial activity but did not bend up. A stem girdle > 20 cm below a branch did not release the branch from apical control in any of the species. These results support the hypothesis that branches compete with the subjacent stem for branch-produced photosynthate and that when the branch lacks this competitive sink it is released from apical control. A stem girdle 2 cm below a branch did not cause release of apical control in either Juniperus virginiana L. or Picea abies (L.) Karst. In these species, decreased shoot elongation and cambial activity above the girdle probably prevented release. A stem girdle 2 cm below a branch increased air content in the stem below the girdle in four of five species, whereas the other girdle treatments had no significant effect on stem air content. Although growth was inhibited above the girdle in the two species with the largest increase in air content, growth was not inhibited in the other species. High air content in stem segments isolated from distal auxin and carbohydrate sources is consistent with the hypothesis that a carbohydrate supply is required to refill embolized cells.

Juniperus↗