Interacting influence of light and size on aboveground biomass distribution in sub-boreal conifer saplings with contrasting shade tolerance

Plant size often influences shade tolerance but relatively few studies have considered the functional response of taller plants to contrasting light environments. Several boreal and sub-boreal Abies, Picea and Pinus species were studied along a light (0-90% full sunlight) and size (30-400-cm high) gradient to examine the interactive influence of tree size and light availability on aboveground biomass distribution. Sampling was conducted in two regions of Canada: (A) British Columbia, for Abies lasiocarpa (Hook.) Nutt., the Picea glauca (Moench.) Voss x P. engelmannii Parry ex. Engelm. complex and Pinus contorta Dougl. var. latifolia Engelm.; and (B) Quebec, for Abies balsamea (L.) Mill., Picea glauca (Moench. Voss) and Pinus banksiana (Lamb.). All biomass distribution traits investigated varied with size, and most showed a significant interaction with both size and light, which resulted in increasing divergences among light classes as size increased. For example, the proportion of needle mass decreased as size increased but the rate of decrease was much greater in saplings growing at below 10% full sunlight. Needle area ratio (total needle area:aboveground mass) followed a similar pattern, but decreased more rapidly with increasing tree size for small trees up to 1 m tall. The proportion of needle biomass (needle mass ratio) was always lower in taller trees (i.e., > 1 m tall) than in small trees (< 1 m tall) and increasingly so at the lowest solar irradiances (0-10% full sunlight). Thus, extrapolating the functional response to light from small seedling to taller individuals is not always appropriate.     

Shoot structure,light interception,and distribution of nitrogen in an Abies amabilis canopy

We studied the effects of variation in shoot structure and needle morphology on the distributions of light and nitrogen within a Pacific silver fir (Abies amabilis (Dougl.) Forbes) canopy. Specifically, we investigated the role of morphological shade acclimation in the determination of resource use efficiency, which is claimed to be optimal when the distribution of nitrogen within the canopy is directly proportional to the distribution of intercepted photosynthetically active radiation (PAR). Shoots were collected from different heights in the crowns of trees representing four different size classes. A new method was developed to estimate seasonal light interceptance (SLI, intercepted PAR per unit needle area) of the shoots using a model for the directional distribution of above-canopy PAR, measurements of shoot silhouette area and canopy gap fraction in different directions. The ratio SLI/SLI(o), where the reference value SLI(o) represents the seasonal light interceptance of a spherical surface at the shoot location, was used to quantify the efficiency of light capture by a shoot. The ratio SLI/SLI(o) doubled from the top to the bottom of the canopy, mainly as a result of smaller internal shading in shade shoots than in sun shoots. Increased light-capturing efficiency of shade shoots implies that the difference in intercepted light by sun shoots versus shade shoots is much less than the decrease in available light from the upper to the lower canopy. For example, SLI of the five most sunlit shoots was only about 20 times greater than the SLI of the five most shaded shoots, whereas SLI(o) was 40 times greater for sun shoots than for shade shoots. Nitrogen content per unit needle area was about three times higher in sun needles than in shade needles. This variation, however, was not enough to produce proportionality between the amounts of nitrogen and intercepted PAR throughout the canopy.     

Foliar morphological and physiological plasticity in Picea abies and Abies alba saplings along a natural light gradient

The role of morphological versus physiological foliar plasticity in the capacity for, and mechanisms of, photosynthetic acclimation was assessed in Picea abies (L.) Karst. and Abies alba Mill. saplings in a forest gap-understory light gradient (relative irradiance, RI, ranging from 0.02 to 0.32). The species investigated showed a similar foliar morphological plasticity along the light gradient, at both the needle level (through alteration in leaf dry mass per area) and the shoot level (through alteration in the silhouette area ratio, e.g., shoot silhouette to projected needle area ratio). In both species chlorophyll (Chl) concentration on a mass basis decreased at increasing RI, but was independent of RI when expressed on an area basis. In contrast, leaf N concentration on a mass basis was independent of RI, but was positively influenced by RI when expressed on an area basis. The parameters describing photosynthetic performance at low light (dark respiration rate, apparent quantum yield and light compensation point) suggest that Abies alba was better suited to maintain a positive carbon balance in shaded conditions. By contrast, parameters describing biochemical capacity at high light (maximum electron transport rate, Jmax and maximum ribulose-1,5-biphosphate carboxylation capacity, Vcmax) indicate that only Picea abies was capable of acclimating physiologically to high photosynthetic photon flux densities (PPFDs) by increasing nitrogen partitioning to Rubisco and Vcmax/mass by increasing RI. These results support the hypothesis that interspecific differences in nitrogen partitioning within the photosynthetic apparatus may provide a mechanistic basis for species separation along a light gradient. The differences in photosynthetic plasticity observed are likely to influence regeneration patterns and habitat breadth of the species investigated. The limited ability of Abies alba saplings to exploit high-light conditions may be a competitive disadvantage in large canopy gaps and thus limit recruitment of this species to small gaps.     

Differences in biomass allocation patterns between saplings of two co-occurring Mediterranean oaks as reflecting different strategies in the use of light and water

Production and biomass allocation patterns, the growth rates of aboveground biomass, and crown traits were examined in saplings of the deciduous Quercus faginea and the evergreen Q. ilex to determine whether differences in these traits might account for the greater mortality during periods of drought undergone by Q. faginea. Strong differences were observed in almost all the traits analyzed, which suggests that the two species use different strategies to cope with the main limiting factors for woody seedling establishment in Mediterranean environments: excess light and low water availability. In Q. faginea, sapling design seems to be oriented to maximize light capture and, hence, leaf productivity during the short life span of the leaf biomass. Thus, the seedlings of Q. faginea showed crown traits that permit self-shading to be minimized: longer shoots with more spaced leaves that result in lower leaf area index than in Q. ilex. In addition, the larger area per unit leaf biomass in Q. faginea leads to a larger interceptive leaf area per unit plant mass and to higher light capture. These characteristics imply higher investments in woody tissues (SWR) that permit the plants to support a wide canopy and facilitate water transport to meet the strong transpiratory demands of a canopy with such characteristics. By contrast, in Q. ilex, saplings are apparently designed to guarantee leaf survival against temperature extremes and photoinhibition through avoidance of excessive radiation.     

Generalized biomass equations for the main aboveground biomass components of maritime pine across contrasting environments

• Introduction   

In order to predict the biomass of aerial components of maritime pine stands (Pinus pinaster Ait.), generalized allometric equations were developed using data collected from the southern and northern margins of its geographical area.     

Timing, light availability and vigour determine the response of Abies alba saplings to leader shoot browsing

Herbivore browsing on tree saplings is a common phenomenon that can cause damage particularly on preferred species. In this study, the combined effects of light availability and timing of browsing on the response of 9-year-old Abies alba saplings were tested experimentally. Leader shoot clipping was applied before budburst, shortly after budburst or in autumn on saplings grown in full light or under artificial shade. Timing of clipping, light availability and tree vigour (expressed as height and tree ring width before clipping) had an effect on the height after clipping. After clipping in autumn or before budburst, fast-growing fir saplings bent up twigs to form new leader shoots and overcompensated height loss; saplings characterised by intermediate growth rates formed new shoots out of regular visible lateral buds; and slow-growing saplings had no new shoot in the first year after clipping, such that the clipping-induced height difference even increased over time. Saplings clipped shortly after budburst elongated the remaining part of the shoot in the first year and developed shoots out of the most distal lateral buds in the second growing season, leading to complete height compensation. Multi-trunking was typical for all clipped trees. We conclude that the microscale conditions under which a tree is growing (i.e. which affect tree vigour) are highly important for determining whether the height reduction imposed by browsing is offset by overcompensation or increases over time relative to unclipped trees. This response can partly be influenced by forest management via enhancing tree vigour via the light regime.     

Ammonium and nitrate uptake, nitrogen productivity and biomass allocation in interior spruce families with contrasting growth rates and mineral nutrient preconditioning

Four full-sib families of interior spruce (Picea glauca (Moench) Voss) x Picea engelmanii Parry ex Engelm.) with contrasting growth rates (two fast-growing and two slow-growing families) were grown aeroponically with either a 2% relative nitrogen addition rate or free access to nitrogen. Fast-growing families showed greater plasticity in allocating biomass to shoots at high nitrogen supply and to roots at low nitrogen supply than slow-growing families. Compared with the slow-growing families, short-term net ammonium uptake rate measured with an ion selective electrode was significantly greater in fast-growing families at high ammonium supply, but not at low supply. Net nitrate uptake showed the same trend, but differences among families were not significant. Results indicate that differences in seedling growth rate are partly a result of physiological differences in net nitrogen uptake efficiency and nitrogen productivity.     

Boundary layer conductance,leaf temperature and transpiration of Abies amabilis branches

We used three methods to measure boundary layer conductance to heat transfer (g(bH)) and water vapor transfer (g(bV)) in foliated branches of Abies amabilis Dougl. ex J. Forbes, a subalpine forest tree that produces clumped shoot morphology on sun-formed branches. Boundary layer conductances estimated in the field from energy balance measurements increased linearly from approximately 10 mm s(-1) at low wind speeds (< 0.1 m s(-1)) to over 150 mm s(-1) at wind speeds of 2.0 m s(-1). Boundary layer conductances measured on shoot models in a wind tunnel were consistently higher than field measurements. The difference between wind tunnel values and field measurements was attributable to variation in path length between the two experimental environments. Boundary layer conductance estimated by subtracting stomatal resistance (r(sV)) measured with a porometer from the total branch vapor phase resistance were unusually small. Sensitivity analysis demonstrated that this method is not suitable for coniferous foliage or when stomatal conductance (g(sV)) is small compared with g(bV). Analysis of the relative magnitudes of g(sV) and g(bV) revealed that, under most conditions, A. amabilis branches are well coupled (i.e., g(sV) is the dominant controller of transpiration). The boundary layer conductance to heat transfer is small enough that leaf temperature can become substantially higher than air temperature when radiation is high and wind speed is low. Over a two-month period, the maximum difference between leaf and air temperatures exceeded 6 degrees C. Leaf temperature exceeded air temperature by more than 2 degrees C on 10% of the daylight hours during this period. Consideration of both the photosynthetic temperature response of A. amabilis foliage as well as the summer air temperature conditions in its habitat suggests that these elevated leaf temperatures do not have a significant impact on carbon gain during the growing season.     

Relative importance of photosynthetic physiology and biomass allocation for tree seedling growth across a broad light gradient

Studies of tree seedling physiology and growth under field conditions provide information on the mechanisms underlying inter- and intraspecific differences in growth and survival at a critical period during forest regeneration. I compared photosynthetic physiology, growth and biomass allocation in seedlings of three shade-tolerant tree species, Virola koschynii Warb., Dipteryx panamensis (Pittier) Record & Mell and Brosimum alicastrum Swartz., growing across a light gradient created by a forest-pasture edge (0.5 to 67% diffuse transmittance (%T)). Most growth and physiological traits showed nonlinear responses to light availability, with the greatest changes occurring between 0.5 and 20 %T. Specific leaf area (SLA) and nitrogen per unit leaf mass (N mass) decreased, maximum assimilation per unit leaf area (A area) and area-based leaf N concentration (N area) increased, and maximum assimilation per unit leaf mass (A mass) did not change with increasing irradiance. Plastic responses in SLA were important determinants of leaf N and A area across the gradient. Species differed in magnitude and plasticity of growth; B. alicastrum had the lowest relative growth rates (RGR) and low plasticity. Its final biomass varied only 10-fold across the light gradient. In contrast, the final biomass of D. panamensis and V. koschynii varied by 100- and 50-fold, respectively, and both had higher RGR than B. alicastrum. As light availability increased, all species decreased biomass allocation to leaf tissue (mass and area) and showed a trade-off between allocation to leaf area at a given plant mass (LAR) and net gain in mass per unit leaf area (net assimilation rate, NAR). This trade-off largely reflected declines in SLA with increasing light. Finally, A area was correlated with NAR and both were major determinants of intraspecific variation in RGR. These data indicate the importance of plasticity in photosynthetic physiology and allocation for variation in tree seedling growth among habitats that vary in light availability.     

Morphological acclimation to understory environments in Abies amabilis, a shade- and snow-tolerant conifer species of the Cascade Mountains, Washington, USA

Light-related plasticity in a variety of crown morphology and within-tree characteristics was examined in sun and shade saplings of Abies amabilis Dougl. ex J. Forbes growing in two late-successional forests with different snow regimes in the Cascade Mountains of Washington, USA. Compared with sun saplings, shade saplings typically had broad flat crowns as a result of acclimation at several scales (needle, shoot, branch, crown and whole sapling). Shoots of shade saplings had a smaller needle mass per unit of stem length than shoots of sun saplings, a feature that enhances light-interception efficiency by reducing among-needle shading. The low annual rate of needle production by shade saplings was associated with a longer needle lifespan and slower needle turnover. Reduced needle production within a shoot was reflected at the branch level, with lateral branches of shade saplings having a smaller needle mass than branches of the same length of sun saplings. Reduced allocation to needles permits greater investment in branches and stems, which is necessary to support the horizontally expanding branch system characteristic of shade saplings. Mean branch age of shade saplings was significantly higher than that of sun saplings. Shade saplings had lower needle mass per unit of trunk biomass or total biomass, reflecting greater investment in the trunk as a support organ. Increased investment in support organs in shade was more evident in the snowier habitat. The observed morphological acclimation makes A. amabilis highly shade and snow-tolerant and thus able to dominate in many late-successional forests in snowy coastal mountain regions.     

Biomass and nutrient allocation in Douglas-fir and amabilis fir seedlings: influence of growth rate and temperature

Allocation of biomass and nutrients to current-year and one-year-old shoots and roots of two-year-old conifer seedlings with differing rates of growth was studied. Differences in growth rate were achieved by selecting fast- and slow-growing populations of the relatively fast- and slow-growing conifer species, Pseudotsuga menziesii (Mirb.) Franco and Abies amabilis Dougl. ex Loud, respectively. Environmentally controlled differences in growth rate were induced by placing half of the seedlings in a 10 degrees C growth chamber and half in a 20 degrees C growth chamber in their second growing season. Seedling samples were harvested in May, before the temperature treatment, and in July and November of the second growing season, and biomass and nutrient concentrations of current-year and one-year-old shoots and roots were determined. Seedling biomass and nutrient allocation differed among the high-growth treatments. Seedlings exhibiting high growth in response to the 20 degrees C treatment and faster growing populations within species both showed increased allocation to new shoots, whereas seedlings of the fast-growing species showed greater allocation to old shoots than to new shoots. Increased growth increased nutrient uptake, but nutrient concentration decreased with growth rate as a result of dilution, so that faster-growing seedlings had greater nutrient-use efficiency than slower-growing seedlings. Retranslocation of P and K was seen in the second year only in slow-growing populations at 10 degrees C. Nutrient concentrations of one-year-old plant parts decreased in the second year, indicating new growth was a stronger sink for nutrients than second-year growth.     

Biomass and nutrient allocation in Douglas-fir and amabilis fir seedlings: influence of growth rate and nutrition

Allocation of biomass and nutrients to shoots and roots was followed for three years in fast and slow growing populations of Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco), a fast growing pioneer species, and amabilis fir (Abies amabilis Dougl. ex J. Forbes), a slow growing shade-tolerant species. Seedlings were grown for three seasons in five nutrient treatments containing varying proportions of nitrogen and phosphorus (N:P). In both species, growth was greatest in the 250:20 N:P treatment followed by the 100:60 and 100:20 treatments. Vector analysis showed that, in both species, relative to the 100:20 treatment, seedlings in the 20:20 treatment were N deficient and seedlings in the 100:4 treatment were P deficient, i.e., where deficiency is defined to mean that an increase in nutrient supply increases nutrient content, nutrient concentration and plant dry weight. Seedlings in the 100:60 treatment had a higher P content than seedlings in the 100:20 treatment but the same dry weight, indicative of what Timmer and Armstrong (1987) termed luxury consumption. No nutrient retranslocation was observed in either species until the third growing season. In Douglas-fir, the greatest percentage of nutrients was exported from one-year-old shoots between May and July of the third growing season to support new growth. The total amount and percent of nutrients retranslocated was higher in Douglas-fir than in amabilis fir. Amabilis fir seedlings also exported N and P from older shoots, but this was later partially replenished. In both species, P retranslocation was greatest in treatments with a high N:P ratio. Nitrogen retranslocation was greatest in amabilis fir seedlings in treatments with a low N:P ratio, and greatest in Douglas-fir seedlings in the 250:20 and 100:60 treatments. Potassium retranslocation was correlated with seedling size. Douglas-fir retranslocated more of its shoot N reserves into new growth at the expense of older needles when soil fertility was high and sinks were strong. Otherwise, both species preferentially translocated the elements in short supply. Thus, retranslocation varied with the ecological characteristics of species, the relative availability of soil nutrients and sink strength.     

Shoot growth patterns in saplings of Cleyera japonica in relation to light and architectural position

To gain further insight into crown development, the influences of shoot architectural position (branch order) and light environment on patterns of shoot growth of Cleyera japonica Thunberg (Theaceae) were investigated. Annual shoot length and light environment were positively correlated within same-order branches. Shoot length differed significantly among branch orders: shoot length was greater for the lower-order branches when light environments were comparable. Lower-order branches lengthened to a certain extent even if light availability was relatively low, whereas higher-order branches did not grow vigorously even when light availability was relatively high. Within same-order branches, branching was independent of the light environment of the shoot. Sylleptic shoot production differed significantly among branch orders, with most sylleptic shoots being produced on second-order branches. It is concluded that both light condition and architectural position of shoots must be considered when examining the mechanisms underlying crown development.     

Photosynthesis and biomass allocation of beech (Fagus crenata) and dwarf-bamboo (Sasa kurilensis) in response to contrasting light regimes in a Japan Sea-Type beech forest

Regeneration of beech (Fagus crenata) forests depends on the formation of canopy gaps. However, in Japan Sea-type beech forests, a dwarf bamboo (Sasa kurilensis) conspicuously occupies sunny gaps. Therefore,F. crenata seedlings must escape the severe interference ofS. kurilensis in the gaps and persist beneath a closed canopy of the beech forest. We hypothesized that the growth ofF. crenata seedlings in the understory would be favored by their being more plastic thanS. kurilensis in photosynthetic and morphological traits, which would support the matter production ofF. crenata seedlings in a wide range of light availabilities. To examine this hypothesis, the photosynthetic-light response of individual leaves and the biomass allocation in aboveground parts (i.e., the culm/foliage ratio) were surveyed at sites with contrasting light availabilities in a Japan Sea-type beech forest in central Japan. InF. crenata, photosynthetic light utilization efficiency at relatively low light was greater, and the dark respiration rate was smaller in the leaves of seedlings (10 cm in height) beneath the closed canopy than in the leaves of saplings at the sunny forest edge. The culm/foliage (C/F) ratio of theF. crenata seedlings at the shady site was small, suggesting effective matter-production beneath the beech canopy. On the other hand,S. kurilensis both in the gap and beneath the beech canopy showed low plasticity in photosynthesis and the culm/foliage ratio. Because the shoot density ofS. kurilensis was smaller beneath the beech canopy than in the gap, the light availability at the bottom of theS. kurilensis layer was greater beneath the beech canopy. These results suggest that the photosynthetic productivity of theF. crenata seedlings would be enough for the seedlings to survive in the understory with a low density ofS. kurilensis shoots beneath the closed beech canopy.     

植物激素对黄檗幼树生长性状的影响及生物量模型的构建

以黄檗5 a生幼树为材料,对叶面喷施不同组合的植物激素,利用正交设计,研究其对生长性状的影响,并建立生物量模型,结果表明:栽植当年和翌年秋季幼树的成活率与保存率分别为97%和95%,平茬与未平茬在定植2年后未发现生长性状有显著差异;植物激素不同组合处理对幼树树高、地径生长量影响较为明显,3种植物激素对树高和地径生长量影响大小依次为NAA6-BAGA3;有效促进树高生长的最佳植物激素配比为NAA200 mg·L-1+6-BA400 mg·L-1+GA3400 mg·L-1,有效促进地径生长的最佳植物激素配比为NAA200 mg·L-1+6-BA400 mg·L-1+GA3200 mg·L-1。地径是影响生物量的主因子,拟合程度最高的一元生物量方程为Y=0.061-0.083X+0.047X2。     

Five-year vegetation control effects on aboveground biomass and nitrogen content and allocation in Douglas-fir plantations on three contrasting sites

Despite widespread use of intensive vegetation control (VC) in forest management, the effects of VC on allocation of biomass and nutrients between young trees and competing vegetation are not well understood. On three Pacific Northwest sites differing in productivity, soil parent material, and understory vegetation community, we evaluated year-5 effects of presence/absence of 5 years of VC on allocation of aboveground biomass and nitrogen (N) between planted Douglas-fir (Pseudotsuga menziesii var. menziesii) and competing vegetation. Equations for predicting bole, branch, foliar, and total dry weights based on stem diameter at a height of 15 cm and total tree height did not differ significantly among sites or by presence or absence of VC. This contrasts with previous research, using diameter at breast height rather than at 15 cm, which found that separate equations were warranted for trees with and without competing vegetation. Estimated whole-tree biomass among the six site/VC combinations ranged from 0.8 to 7.5 Mg ha⁻¹, and increases in tree biomass associated with VC ranged from 62% to 173% among sites. Among the three sites, there were positive, linear relationships between soil total N content to a depth of 60 cm and both N content of aboveground vegetation (trees plus competing vegetation) and Douglas-fir foliar N concentration. Tree N content increased by 8.4, 8.2, and 40.0 kg N ha⁻¹ with VC at the three sites, whereas competing vegetation N content decreased with VC by 0.9, 18.8, and 32.0 kg N ha⁻¹, respectively, at the same sites. Thus, VC did not lead to a direct compensatory tradeoff between aboveground N content of trees and other vegetation. However, soil N content was linearly related to N accumulation and plant growth across the three sites. In addition to differences in N availability among sites, the effect of VC on the redistribution of resources among trees and competing vegetation also was influenced by vegetation community composition and efficacy of VC treatments.     

Warming effect on growth and physiology of seedlings of Betula albo-sinensis and Abies faxoniana under two contrasting light conditions in subalpine coniferous forests of western Sichuan, China

The subalpine coniferous forests on the eastern Qinghai-Tibet Plateau provide a natural laboratory for studying the effect of climate warming on terrestrial ecosystems. Research on differences between tree species in their responses to experimental warming can provide insights into their regeneration behavior and community composition under a future warmer climate. We used open-top chamber (OTC) to determine the short-term effect of two levels of air temperature (ambient and warmed) and light (full light and ca. 10% of full-light regimes) on the early growth and physiology of Betula albo-sinensis and Abies faxoniana seedlings. The OTC manipulation increased mean air temperature and soil surface temperature by 0.51 and 0.34°C, respectively, in a 60-year-old plantation and 0.69 and 0.41°C in forest openings, respectively. Warming generally increased plant growth, biomass accumulation, and advanced physiological processes for seedlings of both species. In response to warming, both tree species allocated relatively more biomass to foliage and had significantly decreased root/shoot ratios (R/S), which might provide the two species with an adaptive advantage when other environmental factors were not limiting. Warming may enhance photosynthesis in the two seedlings by increasing efficiency of PSII in terms of increases in F _v/F _m, photosynthetic pigment concentrations, and apparent quantum yield (Φ). However, the effects of warming on seedling growth and physiological performance varied by light conditions and species. For B. albo-sinensis seedlings, the effects of warming were pronounced only under full-light conditions, while the growth and physiological responses of A. faxoniana seedlings to warming were found only under low-light conditions. Competitive and adaptive relationships between the two species may be altered as a result of response differences to warming manipulation. The shortterm beneficial impact of warming on the early growth and development of the two species suggests that global warming may lead to changes in regeneration dynamics and species composition in subalpine coniferous forest ecosystems.     

Structural characteristics of Abies mariesii saplings in a snowy subalpine parkland in central Japan

Structural characteristics of Abies mariesii M.T. Mast. saplings growing in sun and shade in a snowy subalpine parkland in central Japan were assessed to infer how saplings acclimate to suppression by larger individuals in a conifer clump and to extremely snowy conditions. Sun and shade saplings produced structurally different current-year shoots, and allocated biomass to needles and stem differently. Compared with sun saplings, shoots of shade saplings had lower needle mass per unit shoot size, which indicates less dense needle packing and more effective use of the limited available light by avoiding mutual shading among needles. Biomass allocation within lateral branches also differed between sun and shade saplings. Compared with sun saplings, needle mass was a smaller proportion of total branch mass in shade saplings although shade saplings retained needles for longer, thereby compensating, in part, for their lower annual production of needles. Thus shade saplings incur a high mechanical cost to support their low-light acclimated, conspicuously flattened crowns in this snowy habitat. Suppressed saplings are an important component of the persistent conifer clumps in snowy subalpine parklands. The observed structural characteristics of A. mariesii saplings, which ensure high shade- and snow-tolerance, contribute to the dominance of the species in snowy subalpine regions in Honshu, Japan.     

Foliage dark respiration in Abies amabilis (Dougl.) Forbes: variation within the canopy

Dark respiration of foliage was measured in a 30-year-old stand of Abies amabilis in western Washington from June to November. Both laboratory and field measurements were used to study the effect of environmental and tree variables on respiration. Foliage respiration rates were most strongly influenced by needle temperature. After accounting for leaf temperature differences, foliage respiration decreased with depth in the canopy for all age classes of foliage. Respiration differences attributed to location within the canopy were greatest early in the growing season, but were still significant in November. Younger foliage respired more than older foliage in the upper canopy, but not in the lower canopy. Respiration differences due to foliage age were highly significant in the early growing season, but were not detectable by mid-October.