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A large body of literature suggests that asymmetric competition, where large individuals suppress the growth of smaller individuals by intercepting a disproportionate share of incoming light, is a dominant process in tree population development. This has not been examined extensively for long-lived tree species that accumulate growth over many years under varying growing conditions. Using dendrochronological techniques, we reconstructed annual growth and mortality rates at ten stands of jack pine (Pinus banksiana Lamb.) in Western Canada. We used these data to calculate an annual index of the size asymmetry of growth for each stand for the last 50 years. Jack pine is a shade-intolerant species found in even-aged monoculture stands, so the simple hypothesis is that large trees should consistently perform relatively better than small trees. Inter-annual variation in the index of size-asymmetric growth was positively associated with interannual variation in stand productivity at eight of ten sites. The size asymmetry of growth also showed a positive trend with age at eight of ten sites, even though all sites were in a period of declining leaf area. This should have reduced the intensity of asymmetric competition for light and reduced the size asymmetry of growth over time. Alternate hypotheses for this trend are (1) that physical collisions between crowns result in asymmetric competition for growing space because they are more damaging to small trees, or (2) that a differential build up of diseases in susceptible trees suppresses their growth, even in the absence of competition.

Background and Aims: Changes in size inequality in tree populations are often attributed to changes in the mode of competition over time. The mode of competition may also fluctuate annually in response to variation in growing conditions. Factors causing growth rate to vary can also influence competition processes, and thus influence how size hierarchies develop., Methods: Detailed data obtained by tree-ring reconstruction were used to study annual changes in size and size increment inequality in several even-aged, fire-origin jack pine (Pinus banksiana) stands in the boreal shield and boreal plains ecozones in Saskatchewan and Manitoba, Canada, by using the Gini and Lorenz asymmetry coefficients., Key Results: The inequality of size was related to variables reflecting long-term stand dynamics (e.g. stand density, mean tree size and average competition, as quantified using a distance-weighted absolute size index). The inequality of size increment was greater and more variable than the inequality of size. Inequality of size increment was significantly related to annual growth rate at the stand level, and was higher when growth rate was low. Inequality of size increment was usually due primarily to large numbers of trees with low growth rates, except during years with low growth rate when it was often due to small numbers of trees with high growth rates. The amount of competition to which individual trees were subject was not strongly related to the inequality of size increment., Conclusions: Differences in growth rate among trees during years of poor growth may form the basis for development of size hierarchies on which asymmetric competition can act. A complete understanding of the dynamics of these forests requires further evaluation of the way in which factors that influence variation in annual growth rate also affect the mode of competition and the development of size hierarchies.

We investigated whether root grafts between lodgepole pine (Pinus contorta var. latifolia Dougl. ex. Loud.) trees can transfer sufficient carbohydrate reserves from a source tree to a grafted sink tree to affect the vigor of trees growing in a light-limited environment. Eleven plots were established in early spring and two grafted tree pairs and two independent non-grafted trees were selected at each plot. One tree in a grafted pair and one non-grafted tree were shaded at each plot, whereas the remaining trees were non-shaded during the experimental period. Shaded trees had significantly lower carbohydrate reserves and smaller crowns than non-shaded trees following one growing season. Grafted shaded trees had significantly higher root total nonstructural carbohydrate concentrations than non-grafted shaded trees, indicating that root grafts partially offset the effects of shading. Also, large root grafts transferred proportionately more carbohydrates to the shaded trees than small root grafts. Carbohydrates transferred through root grafts could allow grafted trees to persist under conditions where non-grafted trees would be removed by competition.

We determined the effects of removal of leaves, stem axillary buds, or the entire shoot on root suckering (adventitious shoot formation by roots) and basal stem sprouts in 3- and 4-year-old potted seedlings of aspen (Populus tremuloides Michx.). The greatest number of root suckers (67.9 +/- 8.5 per plant) emerged after excision of the entire shoot. Defoliated and debudded stems were the major source of inhibitory agents for root suckering, although axillary buds and developing new leaves also exerted a significant inhibitory effect. Removal of mature leaves had only a minor effect on root suckering. Removal of a continuous band of bark (girdling) at the base of the stem consistently stimulated growth of adventitious shoots from the stem below the girdle and occasionally promoted root suckering. Exogenous application of indole-3-acetic acid to excised stumps inhibited root suckering and basal stem sprouting. Naphthylphthalamic acid (NPA), an auxin polar transport inhibitor, had no effect on root suckering or stem sprouting when it was applied to the bark of the basal stem. However, NPA significantly increased root suckering when it was applied to the exposed surface of xylem after girdling. These results suggest that polar transport of auxin in the xylem parenchyma is an important inhibitor of root suckering. On decapitated stems, vacuum extraction of xylem sap from the root system lowered the frequency of root suckering compared with decapitation alone, indicating that substance(s) originating in the root system also play a significant role in controlling root suckering.

The literature dealing with the water conducting properties of sapwood xylem in trees is inconsistent in terminology, symbols and units. This has resulted from confusion in the use of either an analogy to Ohm's law or Darcy's law as the basis for nomenclature. Ohm's law describes movement of electricity through a conductor, whereas Darcy's law describes movement of a fluid (liquid or gas) through a porous medium. However, it is generally not realized that, in their full notation, these laws are mathematically equivalent. Despite this, plant physiologists have failed to agree on a convention for nomenclature. As a result, the study of water movement through sapwood xylem is confusing, especially for scientists entering the field. To improve clarity, we suggest the adoption of a single nomenclature that can be used by all plant physiologists when describing water movement in xylem. Darcy's law is an explicit hydraulic relationship and the basis for established theories that describe three-dimensional saturated and unsaturated flow in porous media. We suggest, therefore, that Darcy's law is the more appropriate theoretical framework on which to base nomenclature describing sapwood hydraulics. Our proposed nomenclature is summarized in a table that describes conventional terms, with their formulae, dimensions, units and symbols; the table also lists the many synonyms found in recent literature that describe the same concepts. Adoption of this proposal will require some changes in the use of terminology, but a common rigorous nomenclature is needed for efficient and clear communication among scientists.

We examined how tree growth and hydraulic properties of branches and boles are influenced by periodic (about 6 years) and annual fertilization in two juvenile lodgepole pine (Pinus contorta Dougl. var. latifolia Engelm.) stands in the interior of British Columbia, Canada. Mean basal area (BA), diameter at breast height (DBH) and height increments and percent earlywood and sapwood hydraulic parameters of branches and boles were measured 7 or 8 years after the initial treatments at Sheridan Creek and Kenneth Creek. At Sheridan Creek, fertilization significantly increased BA and DBH increments, but had no effect on height increment. At Kenneth Creek, fertilization increased BA, but fertilized trees had significantly lower height increments than control trees. Sapwood permeability was greater in lower branches of repeatedly fertilized trees than in those of control trees. Sapwood permeabilities of the lower branches of trees in the control, periodic and annual treatments were 0.24 x 10(-12), 0.35 x 10(-12) and 0.45 x 10(-12) m2 at Kenneth Creek; and 0.41 x 10(-12), 0.54 x 10(-12) and 0.65 x 10(-12) m2 at Sheridan Creek, respectively. Annual fertilization tended to increase leaf specific conductivities and Huber values of the lower branches of trees at both study sites. We conclude that, in trees fertilized annually, the higher flow capacity of lower branches may reduce the availability of water to support annual growth of the leader and upper branches.

Stem sapwood hydraulic permeability, tree leaf area, sapwood basal area, earlywood to latewood ratio of annual rings, radial variation in hydraulic permeability and stem hydraulic capacity were examined in dominant (D), codominant (CD) and suppressed (SP) lodgepole pine (Pinus contorta Dougl. ex Loud.) trees growing on medium and poor sites. Hydraulic permeability on a sapwood area basis (ks) was lower in suppressed trees (0.71 x 10(-12) m2) compared to dominants (1.97 x 10(-12) m2) and codominants (1.79 x 10(-12) m2), and higher on medium than on poor sites. The leaf/sapwood area ratio (S) varied with crown dominance position (D > CD > SP) but not by site type. Leaf specific conductivity (kL) did not vary between crown classes or site types. The relationship between leaf area and stem hydraulic supply capacity (Q*) was strong, but differed among crown classes. Dominant trees and trees from the medium sites had a greater proportion of earlywood in outer rings of sapwood than suppressed trees. Sapwood permeability declined from the cambium to the sapwood-heartwood boundary in all samples, but the decline was more gradual in dominant trees compared to codominant and suppressed trees; differences in the radial variation in sapwood permeability may be related to differences in S. Sapwood permeability is positively related to crown dominance, whereas subdominant (CD and SP) trees have greater Q* in relation to leaf area, leading us to propose that this may give subdominant trees a survival advantage, slowing self-thinning.

Global warming and the resultant increase in evapotranspiration might lead to lowered water tables in peatlands and an increase in fire frequency. The objective of this study was to investigate some of the potential effects of these changes on peat decomposition. Dry mass losses and emissions of CO 2 and CH 4 from peat samples taken from three depth layers (0-10, 10-20, and 30-40 cm) of a black spruce peatland were measured in the laboratory at 8°, 16°, and 24°C under two moisture treatments. Effects of deep peat fire on decomposition were also simulated by burning the upper layer (0-10 cm) of peat and adding the ash to peat samples from the 10-20 cm layer. CH 4 release averaged <1% of total carbon loss in flooded samples. Release of CO 2 was 4-9 times greater from the 0-10 cm layer than from the 30-40 cm layer. After 120 d, the 30-40 cm layer had lost <1% of its original dry mass in all treatments. Higher temperatures strongly promoted decomposition of samples exposed to drying cycles but had little effect on decomposition of continuously flooded samples. Ash addition had variable effects on CO 2 emissions but may have promoted CH 4 production. It is suggested that in certain situations, global warming may not cause appreciable increases in carbon loss from peat deposits. The results indicate that some deeper peats are resistant to decay even when exposed to warm, aerobic conditions. However, further experimental work is needed to predict the long-term response of peat deposits to changes in water levels in different peatland types., (© 1992 by the Ecological Society of America.)