Freezing tolerance of conifer seeds and germinants

Survival after freezing was measured for seeds and germinants of four seedlots each of interior spruce (Picea glauca x engelmannii complex), lodgepole pine (Pinus contorta Dougl. ex Loud.), Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) and western red cedar (Thuja plicata Donn ex D. Donn). Effects of eight seed treatments on post-freezing survival of seeds and germinants were tested: dry, imbibed and stratified seed, and seed placed in a growth chamber for 2, 5, 10, 15, 20 or 30 days in a 16-h photoperiod and a 22/17 degrees C thermoperiod. Survival was related to the water content of seeds and germinants, germination rate and seedlot origin. After freezing for 3 h at -196 degrees C, dry seed of most seedlots of interior spruce, Douglas-fir and western red cedar had 84-96% germination, whereas lodgepole pine seedlots had 53-82% germination. Freezing tolerance declined significantly after imbibition in lodgepole pine, Douglas-fir and interior spruce seed (western red cedar was not tested), and mean LT50 of imbibed seed of these species was -30, -24.5 and -20 degrees C, respectively. Freezing tolerance continued to decline to a minimum LT50 of -4 to -7 degrees C after 10 days in a growth chamber for interior spruce, Douglas-fir and lodgepole pine, or after 15 days for western red cedar. Minimum freezing tolerance was reached at the stage of rapid hypocotyl elongation. In all species, a slight increase in freezing tolerance of germinants was observed once cotyledons emerged from the seed coat. The decrease in freezing tolerance during the transition from dry to germinating seed correlated with increases in seed water content. Changes in freezing tolerance between 10 and 30 days in the growth chamber were not correlated with seedling water content. Within a species, seedlots differed significantly in freezing tolerance after 2 or 5 days in the growth chamber. Because all seedlots of interior spruce and lodgepole pine germinated quickly, there was no correlation between seedlot hardiness and rate of germination. Germination rate and freezing tolerance of Douglas-fir and western red cedar seedlots was negatively correlated. There was a significant correlation between LT50 after 10 days in the growth chamber and minimum spring temperature at the location of seedlot origin for interior spruce and three seedlots of western red cedar, but no relationship was apparent for lodgepole pine and Douglas-fir.     

Supercritical fluid impregnation of selected wood species with tebuconazole

The effects of pressure and temperature on supercritical fluid impregnation of tebuconazole were evaluated on Douglas-fir, western red cedar, red alder, white spruce, and white oak. Higher pressure markedly enhanced both the retention and distribution of tebuconazole in these species. When the rate of pressure release was altered at the ends of treatments of Douglas-fir, results varied. Generally, a higher rate of venting increased the steepness of the preservative gradient inward from the surface. Elevated pressures also affected some wood properties. Western red cedar and white spruce showed collapse, while the other three species were free of such defects. Modulus of Elasticity (MOE) and Modulus of Rupture (MOR) tended to decline with higher pressure in western red cedar and white spruce, but the differences were rarely significant. No significant changes in MOE/MOR occurred with the other 3 species. Received 9 November 1998     

Responses of transpiration and photosynthesis to reversible changes in photosynthetic foliage area in western red cedar (Thuja plicata) seedlings

Experiments were conducted on 1-year-old western red cedar (Thuja plicata Donn.) seedlings to determine the response of illuminated foliage to reversible changes in total photosynthetic foliage area (L(A)). Reductions in L(A) were brought about by either shading the lower foliage or by reducing the ambient CO2 concentration (c(a)) of the air surrounding the lower part of the seedling. In the latter case, the vapor pressure was also changed so that transpiration rates (E) could be manipulated independently of photosynthetic rates (A). We hypothesized that following such treatments, short-term compensatory changes would occur in stomatal conductance (g(s)) and A of the remaining foliage. These changes would occur in response to hydraulic signals generated by changes in the water potential gradient rather than changes in the distribution of sources and sinks of carbon within the seedling. When a portion of the foliage was shaded, there was an immediate reduction in whole-seedling E and a concomitant increase in g(s), A and E in the remaining illuminated foliage. However, the intercellular CO2 concentration did not change. These compensatory effects were fully reversed after the shade was removed. When the lower foliage A was reduced to < 0 micromol m-2 s-1, by shading or lowering c(a), and E was either unchanged or increased (by adjusting the vapor pressure deficit), there was no significant increase in g(s) and A in the remaining foliage. We conclude that compensatory responses in illuminated foliage occur only when reductions in L(A) are accompanied by a reduction in whole-plant E. The relationship between the reduction in whole-seedling E and the increase in A is highly linear (r2 = 0.68) and confirms our hypothesis of the strong regulation of g(s) by hydraulic signals generated within the seedling. We suggest that the mechanism of the compensatory effects is a combination of both increased CO2 supply, resulting from increased g(s), and a response of the rate of carboxylation, possibly related to the activity of Rubisco.     

Nutrition and bud removal affect biomass and nutrient allocation in Douglas-fir and western red cedar

Seedlings of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco) and western red cedar (Thuja plicata J. Donn ex D. Don) were grown at high (250 mg l(-1)) and low (20 mg l(-1)) nitrogen (N) supply for a year. Before the second growing season, half of the seedlings in each nutrient treatment were allocated to the other treatment. Half of the seedlings in each nutrient treatment then had all growing points removed. Biomass and N, phosphorus (P) and potassium (K) concentrations of old and new shoots and roots were measured three times in the second year to test the interaction of current-year and previous-year nutrient supply on biomass and nutrient allocation in these two species with different growth habits. Pruned seedlings served as controls. Unpruned seedlings of both species increased in height throughout the second growing season, except for Douglas-fir in the N250 --> N20 treatment. Repeated pruning did not prevent new shoot growth, but resulted in a 12 to 52% reduction in biomass of new shoots and new and old roots. Seedlings receiving a low N supply in the first growing season were more severely affected by pruning than seedings receiving a high N supply. Growth was reduced more by pruning in western red cedar than in Douglas-fir. Concentrations of N, P and K were higher in pruned seedlings than in unpruned seedlings. Although dry weights of all plant parts in all treatments increased throughout the second growing season, some retranslocation of N, P and K was observed from old shoots of both species in the N250 --> N20 and N20 --> N20 treatments after August. Quantities of N, P and K retranslocated were greatest in seedlings grown the previous year in the high-N treatment.     

Dynamic variation in sapwood specific conductivity in six woody species

Our goals were to quantify how non-embolism-inducing pressure gradients influence trunk sapwood specific conductivity (k(s)) and to compare the impacts of constant and varying pressure gradients on k(s) with KCl and H2O as the perfusion solutions. We studied six woody species (three conifers and three angiosperms) which varied in pit membrane structure, pit size and frequency of axial water transport across pits (long versus short conduits). Both stepwise ("steady") and nonlinear continuous ("non-steady") decreases in the pressure gradient led to decreased k(s) in all species but white oak (Quercus garryana Dougl. ex Hook), a ring-porous and long-vesseled angiosperm. In one diffuse-porous angiosperm (red alder, Alnus rubra Bong.) and two conifers (western red cedar, Thuja plicata Donn. ex D. Don, and Douglas-fir, Pseudotsuga menziesii (Mirb.) Franco), k(s) was 10-30% higher under steady pressure gradients than under non-steady pressure gradients, and a decrease in the pressure gradient from 0.15 to 0.01 MPa m(-1) caused a 20-42% decrease in k(s). In another diffuse-porous angiosperm (maple, Acer macrophyllum Pursh) and in a third coniferous species (western hemlock, Tsuga heterophylla (Raf.) Sarg), there was no difference between k(s) measured under steady and non-steady pressure gradients. With the exception of western red cedar, a conifer with simple pit membranes, the differences in k(s) between low and high pressure gradients tended to be lower in the conifers than in the diffuse-porous angiosperms. In Douglas-fir, western red cedar and the diffuse-porous angiosperms, k(s) was higher when measured with KCl than with H2O. In white oak, there were no differences in k(s) whether measured under steady or non-steady pressure gradients, or when xylem was perfused with KCl or H2O. The species differences in the behavior of k(s) suggest that elasticity of the pit membrane was the main factor causing k(s) to be disproportionate to the pressure gradient and to the different pressure regimes. The results imply that, if nonlinearities in pressure-flux relationships are ignored when modeling tree water relations in vivo, large errors will result in the predictions of tree water status and its impact on stomatal control of transpiration and photosynthesis.     

Effects of moisture content and specific gravity on static bending properties and hardness of six wood species

This study was designed to investigate the effects of moisture content (MC) and specific gravity (SG) on the bending strength and hardness of six wood species including Japanese cedar (Cryptomeria japonica D. Don), China fir (Cunninghamia lanceolata), western hemlock (Tsuga heterophylla), red meranti (Shorea spp.), Selangan batu (Shorea spp.), and red oak (Quercus spp.). The experimental results are summarized as follows: Effects of MC and SG on the strength (MOR), stiffness (MOE), and hardness (H _B) could be represented by a multiregression formulas. A negative correlation existed between these properties and MC, whereas a postive correlation showed between them and the SG. The changing rate of these properties induced by 1% MC changes varied with the wood species: 2.6% change in MOR was observed in Japanese cedar, China fir, western hemlock, red meranti, and Selangan batu; and 3.9% was found in red oak. For MOE, a 0.58% change was observed in Japanese cedar, China fir, and red meranti; western hemlock and Selangan batu exhibited 1.2% and red oak 2.5%. For hardness, a 1.1% change was observed in Japanese cedar, western hemlock, and red oak; red meranti and China fir exhibited 3.3%; and Selangan batu 1.8%.A part of this report was presented at the 48th annual meeting of the Japan Wood Research Society in Shizuoka, Japan, April 3-5, 1998     

Leaf longevity of western red cedar (Thuja plicata) increases with depth in the canopy

Leaf longevity determines the annual cost of replacing foliage biomass and influences water and nutrient budgets. Longevity is readily estimated in most evergreen species by annual bud scars along the shoot. However, some species with indeterminate growth do not show these annual markers, making estimation of longevity difficult. One of these species is the widespread and economically valuable western red cedar (Thuja plicata J. Donn ex D. Don), for which no dependable estimates of leaf longevity exist. In this study, we estimated leaf longevity for western red cedar by counting growth rings in shoots at the point of leaf abscission. Estimates were obtained on 26 dominant or codominant trees growing in natural stands in a montane forest in northern Idaho, USA. Leaf longevity averaged 8.9 (SE = 0.2) years, but it strongly increased with depth in the canopy (0.3 year m(-1); mean crown depth was 15 m), increasing from a mean of 6.8 years in the upper third of the canopy to 10.6 years in the lower third. The increase in longevity with depth in the crown is consistent with many reports showing that longevity increases in resource-limiting environments. Longevity did not vary significantly with altitude or solar insolation in these montane forests. Among stand-level variables, longevity was correlated only with leaf area index: it increased slightly in stands with high leaf area indices. This approach to longevity estimation may be useful for any species that produces annual rings but no obvious bud scars, including many Cupressaceae species.     

Growth and transpiration of Japanese cedar (Cryptomeria japonica) and Hinoki cypress (Chamaecyparis obtusa) seedlings in response to soil water content

To investigate the effects of soil water content on growth and transpiration of Japanese cedar (Cryptomeria japonica D. Don) and Hinoki cypress (Chamaecyparis obtusa (Siebold et Zucc.) Endl.), potted seedlings were grown in well-watered soil (wet treatment) or in drying soil (dry treatment) for 12 weeks. Seedlings in the wet treatment were watered once every 2 or 3 days, whereas seedlings in the dry treatment were watered when soil water content (Theta; m3 m(-3)) reached 0.30, equivalent to a soil matric potential of -0.06 MPa. From Weeks 7 to 12 after the onset of the treatments, seedling transpiration was measured by weighing the potted seedlings. After the last watering, changes in transpiration rate during soil drying were monitored intensely. The dry treatment restricted aboveground growth but increased biomass allocation to the roots in both species, resulting in no significant treatment difference in whole-plant biomass production. The species showed similar responses in relative growth rate (RGR), net assimilation rate (NAR) and shoot mass ratio (SMR) to the dry treatment. Although NAR did not change significantly in either C. japonica or C. obtusa as the soil dried, the two species responded differently to the dry treatment in terms of mean transpiration rate (E) and water-use efficiency (WUE), which are parameters that relate to NAR. In the dry treatment, both E and WUE of C. japonica were stable, whereas in C. obtusa, E decreased and WUE increased (E and WUE counterbalanced to maintain a constant NAR). Transpiration rates were lower in C. obtusa seedlings than in C. japonica seedlings, even in well-watered conditions. During soil drying, the transpiration rate decreased after Theta reached about 0.38 (-0.003 MPa) in C. obtusa and 0.32 (-0.028 MPa) in C. japonica. We conclude that C. obtusa has more water-saving characteristics than C. japonica, particularly when water supply is limited.     

Branch and foliage morphological plasticity in old-growth Thuja plicata

At the Wind River Canopy Crane Facility in southeastern Washington State, USA, we examined phenotypic variation between upper- and lower-canopy branches of old-growth Thuja plicata J. Donn ex D. Don (western red cedar). Lower-canopy branches were longer, sprouted fewer daughter branches per unit stem length and were more horizontal than upper-canopy branches. Thuja plicata holds its foliage in fronds, and these had less projected area per unit mass, measured by specific frond area, and less overlap, measured by silhouette to projected area ratio (SPARmax), in the lower canopy than in the upper canopy. The value of SPARmax, used as an indicator of sun and shade foliage in needle-bearing species, did not differ greatly between upper- and lower-canopy branches. We suggest that branching patterns, as well as frond structure, are important components of morphological plasticity in T. plicata. Our results imply that branches of old-growth T. plicata trees have a guerilla growth pattern, responding to changes in solar irradiance in a localized manner.     

Relationship between freezing tolerance and shoot water relations of western red cedar

Freezing tolerance and shoot water relations parameters of western red cedar (Thuja plicata Donn) seedlings were measured every 2 weeks from October 1989 to April 1990. Freezing tolerance, measured by freeze-induced electrolyte leakage, showed seasonal shifts in the temperature causing 50% foliage electrolyte leakage (LT(50)). The LT(50) value was -4 degrees C in October, it decreased to -20 degrees C in February and then increased to -6 degrees C in April. The foliage index of injury at -10 degrees C (II(-10)) also showed seasonal shifts from a high of 98% in October to a low of 18% in February followed by an increase to 82% in April. Osmotic potentials at saturation (Psi(s(sat))) and turgor loss point (Psi(s(tlp))) were, respectively, -1.07 and -1.26 MPa in October, -1.57 and -2.43 MPa in January, and -1.04 and -1.86 MPa in April. Dry weight fraction (DWF) increased and symplastic volume at full turgor (V(o)) decreased during the fall-winter acclimation phase, whereas DWF decreased and V(o) increased during the late winter-spring deacclimation phase. Relationships between seasonal patterns of freezing tolerance and shoot water relations parameters showed that LT(50) and II(-10) decreased linearly as Psi(s(tlp)) and V(o) decreased and DWF increased. There was no discernible difference in the relationship during fall acclimation or spring deacclimation. The freezing dehydration index at -10 degrees C (FDI(-10)) declined from 0.69 in November to 0.41 in February and increased to 0.56 in April. The value of II(-10) decreased linearly as FDI(-10) decreased, although a measurement made on actively growing spring foliage did not fit this relationship. The results indicate that seasonal changes in freezing tolerance of western red cedar are partially due to changes in tissue water content, symplastic volume, passive osmotic adjustment and FDI(-10).     

The impact of black cottonwood on soil fertility in coastal western hemlock forest

Black cottonwood (Populus trichocarpa Torr. and Gray) is a deciduous tree species that extends from Alaska through coastal regions of western Canada into the northwestern United States and as far south as Baja California. We examined the influence of black cottonwood on soil fertility within a forest dominated by Douglas-fir [Pseudotsuga menziessi (Mirb.) Franco], western hemlock [Tsuga heterophylla (Raf.) Sarg], and western red cedar (Thuja plicata Donn ex. D. Don.). Six circular 0.008 ha plots with a single cottonwood tree in the center of conifers were paired with six conifer plots (of the same size) without cottonwood. Litterfall, litter decomposition, properties of forest floor and mineral soil, and N mineralization were compared between plot types. Cottonwood litter had higher concentrations of almost all elements relative to conifer litter. Mass loss did not differ between cottonwood and fir/hemlock litter on cottonwood sites. Twice the amount of mull-like humus form (vermimull and mullmoder, 56%) was found in cottonwood plots compared to 28% in conifer plots. Higher pH (4.4) was found in the forest floor under cottonwood compared to conifer (3.9). Total N concentration (3.33 g/kg) and base saturation (68%) were higher in the mineral soil under cottonwood compared to conifers (2.98 g/kg total N and 50% base saturation). Net ammonification and net mineralization were both lower under cottonwood. These results suggest a variable effect of cottonwood on soil fertility within coastal western hemlock forests with some soil variables changed in a favourable direction and some in an unfavourable direction.     

Blight of conifer seedlings caused by Colletotrichum gloeosporioides

In a growth chamber experiment, 70-day-old seedlings of 10 conifer species were inoculated with Colletotrichum gloeosporioides conidia to determine the host range of the fungus. Based on the percentage of seedlings affected and the disease severity on individual seedlings, the order of most to least susceptibility was: western hemlock (WH), mountain hemlock, western larch, Sitka spruce, Engelmann spruce, Douglas-fir (coastal form, then interior form), white spruce and ponderosa pine; lodgepole pine and western red cedar remained unaffected. Inoculation of WH needles showed that within 24 h C. gloeosporioides conidia germinate and appressoria (penetration structures) form. A growth chamber study demonstrated that the pathogen can infect WH at needle wetness periods as short as 15 min; number of needles affected was higher at 0.5 h, but did not increase further even when wetness was extended up to 8 h. The results are discussed in relation to blight management of greenhouse-grown conifer seedlings.Portion of a Bachelor of Science (Honors) thesis, Department of Biology, University of Victoria, Victoria, British Columbia.     

Ecosystem production in Douglas-fir plantations: Interaction of red alder and site fertility

Pure stands of Douglas-fir (Pseudotsuga menziesii [Mirb.] Franco) and mixed stands of Douglas-fir and naturally established red alder (Alnus rubra Bong.) were examined on two different sites for soil properties, tree growth and nutrition, and aboveground ecosystem biomass and net primary production. One site (Mt. Benson, Nanaimo, B.C.) was nitrogen (N)-deficient and had a low site index (expected Douglas-fir height of 24 m at 50 years). The other site (Skykomish, western Washington) was N-rich and had a site index of 45 m at 50 years. Soil N accretion on the red alder units was estimated at 65 (Mt. Benson) and 42 (Skykomish) kg ha^?1 year^?1 for 23 years to a soil depth of 50 cm. At the current stage of plantation development, presence of red alder at the infertile Mt. Benson site increased average Douglas-fir diameter but did not affect its basal area and basal area growth rate; including alder stem biomass increased total stand basal area and basal area growth 2.5 fold. Presence of red alder at the fertile Skykomish site decreased average diameter, basal area, and basal area growth of Douglas-fir; including alder biomass left total stand basal area and basal area growth unchanged. Douglas-fir foliar N concentrations on Mt. Benson increased from 0.93 without alder to 1.41% on the red alder unit but were 1.55% for both units at Skykomish. Although alder did not affect Douglas-fir aboveground biomass and net primary production on Mt. Benson, total ecosystem biomass doubled and production tripled when alder biomass was included. Conversely, at Skykomish, Douglas-fir biomass and production decreased, and total ecosystem values were essentially unchanged. Mixing red alder and Douglas-fir seems to have great potential for increasing Douglas-fir growth and ecosystem production on infertile, N-deficient sites but probably has limited value on fertile, N-rich sites.     

A comparison of annual transpiration and productivity in monoculture and mixed-species Douglas-fir and red alder stands

Although much is known about drivers of productivity in Douglas-fir and red alder stands, less is known about how productivity may relate to stand transpiration and water use efficiency. We took advantage of a 15-year-old experiment involving Douglas-fir (Pseudotsuga menziesii) and red alder (Alnus rubra) in the western Cascade Range of western Oregon to test the following hypotheses: (a) more productive stands transpire more water, (b) the relationship between productivity and transpiration differs between species, and (c) the relationship between productivity and transpiration differs between sites varying in soil moisture and fertility. Furthermore, the experimental design included alder, a facultative nitrogen-fixing species, which could also affect fertility. Fixed area plots (20 × 20 m) were planted as monocultures of each species or in mixtures at a common density (1100 trees ha⁻¹) in a randomized-block design. Transpiration of Douglas-fir and red alder was measured using heat dissipation sensors installed in eight trees per plot and scaled to the plot level based on sapwood basal area for each species. Although up to 53% of the variability in tree transpiration was explained by basal area, irrespective of species or site conditions, the two stands with the highest biomass and sapwood basal area did not transpire the most. Instead of more productive stands transpiring more water, the greatest variability in both productivity and transpiration was determined by site conditions and to a lesser degree, species composition. For example, 70% of the variation in tree biomass increment (TBI) was determined by leaf area index, which was much higher at the site with higher fertility and soil moisture (p < 0.05). Despite marked phenological and physiological differences, Douglas-fir and red alder performed similarly. Only 19% of annual water use of Douglas-fir occurred between October and March when alder was leafless. Also, there was no evidence of a fertilization effect of the nitrogen-fixing red alder on the Douglas-fir: the nitrogen concentration and N-isotopic ratio of Douglas-fir needles did not differ whether trees were grown in monoculture or in mixtures with red alder. We conclude that lower soil fertility and contrasting microclimate at one site relative to the other suppressed NPP while maintaining higher transpiration, thus reducing water use efficiency.     

Utility pole decay

Summary The Basidiomycetes associated with decay in pine, Douglas-fir, and cedar utility poles within various geographic regions of North America were investigated. On the basis of 313 isolations from these poles, 9 fungi appeared to be of major importance in internal pole decay in the United States. These fungi were the following: Lentinus lepideus, Lenzites saepiaria, L. trabea, Peniophora A., P. gigantea, and Poria radiculosa in pine, and L. lepideus, Poria carbonica, P. monticola, and P. xantha in Douglas-fir poles. Lentinus lepideus was overwhelmingly predominant in pine poles, whereas Poria carbonica was similarly predominant in Douglas-fir poles. Lenzites trabea was the fungus most often isolated from cedar poles; however, it is believed to be associated primarily with shell rot of cedar poles and to be of little significance, therefore, in causing internal decay of cedar. The fungi associated with western red-cedar and red and jack pine poles in Canada are listed, although the frequency of their occurrence is not included.This work was in cooperation with the Navy Department, Naval Facilities Engineering Command.The author is indebted to the following for furnishing cultures and pole sections or both for culturing or for information on the identities of the species of decay fungi found in Canadian poles: Joe Clark and John Kulp, U. S. Forest Products Laboratory, Madison, Wisconsin; Robert Graham and John Mothershead, Forest Research Laboratory, Oregon State University, Corvallis, Oregon; O. Floyd Hand, Bonneville Power Administration, Vancouver, Washington; John Shields, Canadian Forest Products Laboratory, Ottawa, Ontario; and John Roff, Canadian Forest Products Laboratory, Vancouver, British Columbia. The author is especially indebted to members of the Forest Disease Laboratory, Laurel, Maryland, for their invaluable aid in identifying representative cultures of many of the fungi discussed in this investigation.The Laboratory is maintained at Madison in cooperation with the University of Wisconsin.     

Titratable Acids and Bases in Tree and Shrub Leaf Litters

Acidity of aqueous extracts of several tree and shrub leaf litterswas determined by titration to pH 7 with 0.01N NaOH. Bases weredetermined by back titration of 0.05N HCl extracts of groundlitters and after ashing. Lowest acidity was found in the broadleavedspecies (except sycamore) and western red cedar; greatest aciditywas found in western hemlock, grand fir, and one sample of Douglasfir. Largest contents of ash bases were found in some broadleavesplus western red cedar; the smallest contents were in most ofthe conifers. Excess ash bases (ash bases minus acidity) weregreatest in elm, hawthorn, western red cedar, hazel, willow,ash, and southern beech, and were smallest in western hemlock,Douglas fir, lodgepole pile, Sitka spruce, grand fir, hybridlarch, Scots pine, and Norway spruce. Acidity, directly titratableand ash bases, and excess ash bases showed significant, andin some cases quite large, variation between sites for somespecies. There is some support for the suggestion that excessash bases are greater in mull-forming than in mor-forming litters.Acidity was significantly greater for litters collected drythan for those collected wet. Litters which were stored airdry for several months showed increased acidity compared withtheir initial values.     

Vegetation competition effects on aboveground biomass and macronutrients,leaf area,and crown structure in 5-year old Douglas-fir

Vegetation control (VC) in forest plantations often increases growth of crop trees but can also affect biomass and nutrient partitioning to tree components. We examined above-ground biomass and macronutrients, leaf area and crown structure in 5-year old Douglas-fir (Pseudotsuga menziessi (Mirb.) Franco) growing with VC and with no vegetation control (NVC) in coastal Washington, United States of America. Trees in VC had larger stem, branch, foliage and total biomass than trees of equal stem diameter at 1.3 m above ground in NVC. The difference in component biomass between treatments was in the order: branch > foliage > stem. Trees in the VC regime did not differ in macronutrient concentration in stem, branch and foliage except for branch N which was greater in NVC than in VC. Differences in tree macronutrient stores between VC and NVC ranged from 2.2 times for Mg to 2.6 times for N and K. The relationship between stem diameter and leaf area was linear in both VC treatments. The relationship between stem diameter and foliage biomass was curvilinear for both VC treatments. Results of this study support the need for separated allometric equations to estimate biomass for young Douglas-fir growing in areas with and without VC.     

Atmospheric and soil drought reduce nocturnal conductance in live oaks

Nocturnal and daytime whole-canopy transpiration rate (E) and conductance (g = E/VPD, where VPD is leaf to air vapor pressure difference) were assessed gravimetrically in drought-treated and well-watered 3-year-old saplings of live oak species (Quercus series Virentes Nixon) from the southeastern USA (Quercus virginiana Mill.) and Central America (Q. oleoides Cham. and Schlecter). Our objectives were to: (1) quantify nocturnal and daytime E and g in a controlled environment; (2) determine the impact of severe drought on nocturnal E and g; and (3) examine whether unavoidable water loss through the epidermis could account for nocturnal water loss. We calculated daytime E during peak daylight hours (between 0930 and 1330 h) and nocturnal E during complete darkness (between 2200 and 0500 h). In addition to reducing E and g during the daytime, drought-treated plants reduced nocturnal E and g on a whole-canopy basis by 62-64% and 59-61%, respectively, and on a leaf-level basis by 27-28% and 19-26%, respectively. In well-watered plants, nocturnal g declined with increasing VPD, providing evidence for stomatal regulation of nocturnal transpiration. In drought-treated plants, g was low and there was no relationship between nocturnal g and VPD, indicating that water loss could not be reduced further through stomatal regulation. Both daytime and nocturnal g declined curvilinearly with predawn water potential for all plants, but nocturnal g was unrelated to predawn water potentials below -1 MPa. The reductions in daytime and nocturnal E and g during drought were associated with decreases in whole-plant and leaf hydraulic conductances. Observed nocturnal g was within the same range as epidermal conductance for oak species determined in previous studies under a range of conditions. Nocturnal E rose from 6-8% of daytime E for well watered plants to 19-20% of daytime E for drought-treated plants. These results indicate that, during drought, saplings of live oak species reduce g to a minimum through stomatal closure, and experience unavoidable water loss through the epidermis.     

An investigation of hydraulic limitation and compensation in large,old Douglas-fir trees

The hydraulic limitation hypothesis (Ryan and Yoder 1997) proposes that leaf-specific hydraulic conductance (kl) and stomatal conductance (gs) decline as trees grow taller, resulting in decreased carbon assimilation. We tested the hydraulic limitation hypothesis by comparison of canopy-dominant Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirb.) Franco) trees in stands that were approximately 15 m (20 years old), 32 m (40 years old) and 60 m (> 450 years old) tall in Wind River, Washington, USA. Carbon isotope discrimination (Delta) declined with tree height (18.6, 17.6 and 15.9 per thousand for stands 15, 32 and 60 m tall, respectively) indicating that gs may have declined proportionally with tree height in the spring months, when carbon used in the construction of new foliage is assimilated. Hydraulic conductance decreased by 44% as tree height increased from 15 to > 32 m, and showed a further decline of 6% with increasing height. The general nonlinear pattern of kl versus height was predicted by a model based on Darcy's Law. Stemwood growth efficiency also declined nonlinearly with height (60, 35 and 28 g C m-2 leaf area for the 15-, 32- and 60-m stands, respectively). Unlike kl and growth efficiency, gs and photosynthesis (A) during summer drought did not decrease with height. The lack of decline in cuvette-based A indicates that reduced A, at least during summer months, is not responsible for the decline in growth efficiency. The difference between the trend in gs and A and that in kl and D may indicate temporal changes (spring versus summer) in the response of gas exchange to height-related changes in kl or it may be a result of measurement inadequacies. The formal hydraulic limitation hypothesis was not supported by our mid-summer gs and A data. Future tests of the hydraulic limitation hypothesis in this forest should be conducted in the spring months, when carbon uptake is greatest. We used a model based on Darcy's Law to quantify the extent to which compensating mechanisms buffer hydraulic limitations to gas exchange. Sensitivity analyses indicated that without the observed increases in the soil-to-leaf water potential differential (DeltaPsi) and decreases in the leaf area/sapwood area ratio, kl would have been reduced by more than 70% in the 60-m trees compared with the 15-m trees, instead of the observed decrease of 44%. However, compensation may have a cost; for example, the greater DeltaPsi of the largest trees was associated with smaller tracheid diameters and increased sapwood cavitation, which may have a negative feedback on kl and gs.