Branch growth and biomass allocation in Abies amabilis saplings in contrasting light environments

Aboveground biomass allocation, and height and branch growth were studied in saplings of the shade-tolerant conifer, Abies amabilis Dougl. ex Forbes growing in large openings and in the understory of an old-growth forest in western Oregon. The presence of annual overwintering budscale scars was used to infer extension growth histories; annual growth rings in branches and stems were used in combination with extension histories to compute partitioning of new biomass among leaves, branches and stems. Saplings growing in large gaps had conical crowns, whereas understory saplings had umbrella shaped crowns as a result of much greater rates of branch extension than stem extension. Understory saplings grew slowly in height because of low rates of biomass production and low allocation of biomass to stem extension. About 40% of new biomass was allocated to foliage in both groups, but understory saplings allocated more of the remaining growth increment to branches and less to stem than did saplings growing in large gaps. These results differ from the patterns observed in shade-tolerant saplings of tropical forests, where allocation to foliage increases with shading and branch allocation is much lower than observed here. This difference in allocation may reflect mechanical constraints imposed by snow loads on the evergreen A. amabilis crowns, particularly on flat-crowned understory saplings.     

Evaluation of leaf display of evergreen broadleaved tree species and deciduous tree species in warm temperate conifer plantations

We investigated the sapling leaf display in the shade among trees of various leaf lifespans co-occurring under the canopy of a warm-temperate conifer plantation. We measured leaf-area ratio (aLAR) and morphological traits of saplings of evergreen broadleaved tree species and a deciduous tree species. Although we found large interspecific and intraspecific differences in aLAR even among saplings of similar size in the homogeneous light environment, we did not find a consistent trend in aLAR with leaf lifespan among the species. While deciduous trees annually produced a large leaf area, some evergreen broadleaved trees retained their leaves across years and had aLAR values as high as those of deciduous trees. Among leaf-level, shoot-level, and individual-level morphological traits, aLAR was positively correlated with current-year shoots mass per aboveground biomass in deciduous trees, and with the area of old leaves per aboveground mass in evergreen broadleaved trees. Thus, tree-to-tree variation in the degrees of annual shoot production and the accumulation of old leaves were responsible for the interspecific and intraspecific variations in aLAR.     

Enhancement of understory productivity by asynchronous phenology with overstory competitors in a temperate deciduous forest

Some saplings and shrubs growing in the understory of temperate deciduous forests extend their periods of leaf display beyond that of the overstory, resulting in periods when understory radiation, and hence productivity, are not limited by the overstory canopy. To assess the importance of the duration of leaf display on the productivity of understory and overstory trees of deciduous forests in the north eastern United States, we applied the simulation model, BIOME-BGC with climate data for Hubbard Brook Experimental Forest, New Hampshire, USA and mean ecophysiological data for species of deciduous, temperate forests. Extension of the overstory leaf display period increased overstory leaf area index (LAI) by only 3 to 4% and productivity by only 2 to 4%. In contrast, extending the growing season of the understory relative to the overstory by one week in both spring and fall, increased understory LAI by 35% and productivity by 32%. A 2-week extension of the growing period in both spring and fall increased understory LAI by 53% and productivity by 55%.     

Photosynthetic differences between saplings and adult trees: an integration of field results by meta-analysis

Ontogenetic changes in gas exchange parameters provide both insight into mechanisms underlying tree growth patterns, and data necessary to scale environmental impacts on young trees to predict responses of older trees. We present a quantitative review and meta-analysis of field measurements of gas exchange parameters in saplings and mature trees of 35 tree species (seven conifers, seven temperate deciduous trees, and 21 tropical evergreen trees). Data for saplings were obtained in both understory environments and open areas or large gaps. We also present data on ontogenetic changes in photosynthesis for Pseudotsuga menziesii (Mirb.) Franco and Tsuga heterophylla (Raf.) Sarg., species of particular interest because of their large maximal heights and long life-spans. Among tree species, there is evidence for both ontogenetic increases and ontogenetic decreases in photosynthetic capacity on a leaf area basis (A(area)). Overall, A(area) is generally higher for upper-canopy leaves of adult trees than for saplings, especially in temperate deciduous trees. However, the pattern for photosynthetic capacity on a leaf mass basis (A(mass)) is the reverse of that observed for A(area). Saplings of both conifers and broad-leaved trees, even when acclimated to low-light conditions, characteristically have a higher A(mass) than adult trees. This pattern is driven largely by an ontogenetic increase in leaf mass per unit area (LMA), as found in 100% of studies reviewed. Data for Pacific Northwest conifers, although including measurements on some of the tallest trees studied, did not differ greatly from patterns found in other tree species. We conclude that ontogenetic changes in LMA are the single most consistent difference between saplings and adult trees, and that changes in LMA and related aspects of leaf morphology may be critical to understanding both variation in gas exchange during tree growth, and stage-dependent responses of trees to environmental change.     

Functional relationships between crown morphology and within-crown characteristics of understory saplings of three codominant conifers in a subalpine forest in central Japan

Light-related plasticity of crown morphology and within-crown characteristics were investigated in understory sun and shade saplings of three codominant subalpine conifers, Abies mariesii M.T. Mast., Abies veitchii Lindl. and Picea jezoensis var. hondoensis (Mayr) Rehd. Compared with those of sun saplings, current-year shoots of shade saplings allocated less biomass to needles, resulting in less dense needle packing and hence less mutual needle shading. The proportion of lateral branch biomass in foliage was either similar in sun and shade saplings or greater in shade saplings, depending on the species, suggesting that, over the lifetime of a branch, greater needle longevity in shade compensates for reduced biomass investment in needles of current-year shoots of shade saplings. Saplings with slower-growing branches tended to have greater needle life spans, suggesting that plasticity of branch growth rate and plasticity of needle life span are interdependent. Both Abies species showed greater light-related plasticity of needle life span and branch growth than P. jezoensis. The greater shade tolerance of the Abies species derives from their broad flattened crowns with slow-growing branches. This type of crown development incurs substantial support costs, but the long needle life span of shade saplings of the Abies species compensates, at least in part, for their low annual investment in foliage, especially in the case of A. mariesii, which has a longer needle life span and slower-growing and stouter branches than A. veitchii. Compared with the Abies species, P. jezoensis had a less plastic crown morphology, and less variability of needle life span and branch growth in response to light, resulting in lower shade tolerance. However, compared with the flattened crown of Abies shade saplings, the conical crown of P. jezoensis saplings imposes a smaller support cost, making this species better adapted to rapid height growth than to survival in shade.     

Crown architecture of understory and open-grown white pine (Pinus strobus L.) saplings

Crown architecture and growth allocation were studied in saplings of eastern white pine (Pinus strobus L.), a species classified as intermediate in shade tolerance. A comparison was made of 15 understory saplings and 15 open-grown saplings that were selected to have comparable heights (mean of 211 cm, range of 180-250 cm). Mean ages of understory and open-grown trees were 25 and 8 years, respectively. Understory trees had a lower degree of apical control, shorter crown length, and more horizontal branch angle, resulting in a broader crown shape than that of open-grown trees. Total leaf area was greater in open-grown saplings than in understory saplings, but the ratio of whole-crown silhouette (projected) leaf area to total leaf area was significantly greater in understory pine (0.154) than in open-grown pine (0.128), indicating that the crown and shoot structure of understory trees exposed a greater percentage of leaf area to direct overhead light. Current-year production of understory white pine was significantly less than that of open-grown white pine, but a higher percentage of current-year production was allocated to foliage in shoots of understory saplings. These modifications in crown structure and allocation between open-grown and understory white pine saplings are similar to those reported for more shade-tolerant fir (Abies) and spruce (Picea) species, but the modifications were generally smaller in white pine. As a result, white pine did not develop the flat-topped "umbrella" crown structure observed in understory fir and spruce, which approaches the idealized monolayer form that maximizes light interception. The overall change to a broader crown shape in understory white pine was qualitatively similar, but much more limited than the changes that occurred in fir and spruce. This may prevent white pine from persisting in understory shade as long as fir and spruce saplings.     

Physiological, morphological and allocational plasticity in understory deciduous trees: importance of plant size and light availability

In a 4-year study, we investigated changes in leaf physiology, crown morphology and whole-tree biomass allocation in seedlings and saplings of shade-tolerant sugar maple (Acer saccharum Marsh.) and intermediate shade-tolerant yellow birch (Betula alleghaniensis Britt.) growing in natural understory light (0.5 to 35% of full sunlight) or in understory light reduced by 50% with shade nets to simulate the effect of gap closure. Leaf physiological parameters were mainly influenced by the light gradient, whereas crown morphological and whole-tree allocational parameters were mainly influenced by tree size. No single physiological, morphological or allocational trait was identified that could explain the difference in shade tolerance between the species. Yellow birch had higher growth rates, biomass allocation to branches and leaf physiological plasticity and lower crown morphological plasticity in unmodified understory light than sugar maple. Sugar maple did not display significant physiological plasticity, but showed variation with tree size in both crown morphology and whole-tree biomass allocation. When sugar maple was small, a greater proportion of whole-tree biomass was allocated to roots. However, physiological differences between the species decreased with decreasing light and most morphological and allocational differences tended to disappear with increasing tree size, suggesting that many species differences in shade-tolerance are expressed mainly during the seedling stage. Understory trees of both species survived for 4 years under shade nets, possibly because of higher plasticity when small and the use of stored reserves when taller.     

Spatial pattern and regeneration dynamics in a temperate Abies–Tsuga forest in southwestern Japan

This article reports the regeneration dynamics of a temperate Abies–Tsuga forest in Kirishima Yaku National Park, southwestern Japan, and examines the influence of species coexistence mediated by gap disturbances on biomass production. All trees taller than 2 m in a 1-ha plot were monitored over four growing seasons. Three growth-form groups occupied different vertical layers. Evergreen conifers and deciduous broad-leaved trees tended to be spatially segregated from evergreen broad-leaved trees, which formed thickets in the understorey. The regeneration of understorey evergreen broad-leaved trees was affected by canopy gaps. The recruitment of conifers and deciduous broad-leaved species was not observed during the four growing seasons. This suggests that regeneration is sporadic and the present environmental conditions are not favorable for these canopy species. The mortality and unsuccessful recruitment of conifers and deciduous trees appeared to cause fluctuations in the productivity of the stand. However, an abundance of canopy gaps accelerates the regrowth of shorter species, and the fluctuation of productivity resulting from the population dynamics of canopy species would be partly mitigated by the regeneration of evergreen understorey species. The horizontal and vertical heterogeneity of the temperate mixed forest was a result of the patch structures of the three growth-form groups. The different regeneration patterns among the three groups, which were driven by interactions of species-specific regeneration niches and disturbance regimes, might be an important factor in maintaining the aboveground productivity in a transitional mixed forest between warm-temperate and cool-temperate zones.     

Crown architecture of stand-grown sugar maple (Acer saccharum Marsh.) in the Adirondack Mountains

Leaf and crown morphology of shade-tolerant sugar maple (Acer saccharum Marsh.) were examined to test the hypotheses (1) that leaf area exhibits significant plasticity both within and between crown classes and individual tree crowns and (2) that leaf area is accurately predicted from estimates of crown volume. A total of 18 trees, ranging from 3.3 to 43.4 cm dbh, were felled and dissected into upper, middle, lower, and below-crown layers, for measurements of leaf, bark, and xylem dimensions. For dominant trees only, bark thickness and xylem radii were higher within the crown than below the crown. Cumulative leaf area index increased with decreasing stratum height at similar rates in all trees, except for two trees that were located in the understory. Area leaf weight declined with decreasing stratum height within the crown of all except four overstory trees. These four trees showed an increase with decreasing stratum height, i.e., leaves were heavier per unit area in the lower crown stratum and below the crown than they were at mid-crown. Within-tree leaf area density was usually higher in the upper crown of overstory trees and in the lower crown of understory trees. Total crown volume was the best predictor of whole-tree leaf area, but it was only slightly better than dbh.     

Deciduous trees affect small-scale floristic diversity and tree regeneration in conifer forests

Abstract

There is a growing interest in the effects of deciduous trees on biodiversity, soil processes and long-term productivity in boreal, conifer-dominated forests. This study investigated whether individual birch trees allowed to grow to maturity in the coniferous forest can have a local effect on floristic richness and regeneration of tree saplings. The ground vegetation was compared in 2?m radius plots around the stem under the canopies of matched conifer–deciduous trees in a mature, conifer-dominated forest, and included in the analysis variables that could potentially mediate the tree effect (soil pH, cover of lichens, bryophytes, leaf and needle litter). The field layer vegetation was more species rich under birch (Betula pendula and B. pubescens) than under conifers (Picea abies and Pinus sylvestris), and several vascular plant species (including saplings of tree species) occurred more often under birch than under conifers. However, when the effect of the number of subordinate trees was taken into account the difference between birch and pine was not significant. The number of tree regenerations (saplings) was lowest under pines, but did not differ between spruce and birch. There were no effects of the canopy species on soil pH or on cover of lichens and bryophytes. The difference in diversity may be caused by the different effects of leaf and needle litter, and it is also likely that canopy structure has an influence via interception and throughfall and by affecting the light and microclimate.     

Belowground competition from overstory trees influences Douglas-fir sapling morphology in thinned stands

We evaluated effects of belowground competition on morphology of naturally established coast Douglas-fir (Pseudotsuga menziesii var. menziesii (Mirb.) Franco) saplings in 60- to 80-year-old thinned Douglas-fir stands in southwestern Washington. We separately quantified belowground competition from overstory and understory sources using trenching and understory removal. In this light-limited environment of 26 ± 16% (std. dev.) full sunlight, 2-year exclusion of tree root competition by trenching increased sapling stem biomass by 18%, total aboveground biomass by 21%, number of interwhorl buds by 68%, total foliar biomass by 33%, and foliar biomass on branch components over 4 years old by 143%. Belowground competition did not influence shoot:root ratio or foliar efficiency (i.e., stem growth per unit foliage biomass). Sapling needle size, specific leaf area, and internodal distance also were not affected by belowground competition; these variables were apparently a function of the low-light environment. The principal source of belowground competition was roots of overstory trees; effects of belowground competition from understory vegetation were minor. Thus, under a partial overstory, morphology of Douglas-fir regeneration was influenced by both belowground and aboveground competition from overstory trees. In this environment, understory vegetation control would not likely influence belowground competition to an extent that would affect sapling morphology.     

Osmotic potential of several hardwood species as affected by manipulation of throughfall precipitation in an upland oak forest during a dry year

Components of dehydration tolerance, including osmotic potential at full turgor (Psi(pio)) and osmotic adjustment (lowering of Psi(pio)), of several deciduous species were investigated in a mature, upland oak forest in eastern Tennessee. Beginning July 1993, the trees were subjected to one of three throughfall precipitation treatments: ambient, ambient minus 33% (dry treatment), and ambient plus 33% (wet treatment). During the dry 1995 growing season, leaf water potentials of all species declined to between -2.5 and -3.1 MPa in the dry treatment. There was considerable variation in Psi(pio) among species (-1.0 to -2.0 MPa). Based on Psi(pio) values, American beech (Fagus grandifolia Ehrh.), dogwood (Cornus florida L.), and sugar maple (Acer saccharum Marsh.) were least dehydration tolerant, red maple (A. rubrum L.) was intermediate in tolerance, and white oak (Quercus alba L.) and chestnut oak (Quercus prinus L.) were most tolerant. During severe drought, overstory chestnut oak and understory dogwood, red maple and chestnut oak displayed osmotic adjustment (-0.12 to -0.20 MPa) in the dry treatment relative to the wet treatment. (No osmotic adjustment was evident in understory red maple and chestnut oak during the previous wet year.) Osmotic potential at full turgor was generally correlated with leaf water potential, with both declining over the growing season, especially in species that displayed osmotic adjustment. However, osmotic adjustment was not restricted to species considered dehydration tolerant; for example, dogwood typically maintained high Psi(pio) and displayed osmotic adjustment to drought, but had the highest mortality rates of the species studied. Understory saplings tended to have higher Psi(pio) than overstory trees when water availability was high, but Psi(pio) of understory trees declined to values observed for overstory trees during severe drought. We conclude that Psi(pio) varies among deciduous hardwood species and is dependent on canopy position and soil water potential in the rooting zone.     

Within-crown structural variability of dwarfed mature Abies mariesii in snowy subalpine parkland in central Japan

We investigated mature dwarf Abies mariesii trees growing in conifer thicket–meadow parklands on a snowy subalpine plateau, where these dwarf trees are buried in the accumulated snow in winter. We focused on structural variation in the needles, shoots, and branchlets within different crown positions (leader crown vs lower crown) of the dwarf trees. In the leader crown, which appears above the snow surface earlier than the lower crown, current-year shoots and branchlets had greater total biomass, and foliage was more closely packed along the stem axis than in the lower crown, whereas current-year shoots in the leader crown had a lower needle mass ratio than in the lower crown. These results suggest that current-year shoots and branchlets in the leader crown have a specific structure that allows them to harvest more light, although construction and maintenance costs would be higher. In contrast, the structural characteristics of current-year shoots and branchlets in the lower crown efficiently concentrate incoming light by avoiding mutual shading within foliage, thus leading to increased biomass of photosynthetic needles within shoot and branchlet biomass. Such within-crown variability at various hierarchical levels from needles to branches in mature, but very dwarf, A. mariesii trees maintains the crown and allows survival within conifer clumps in areas of subalpine parklands that receive heavy snowfall.     

The effect of stand age on the accumulation of nutrients in the aboveground components of an Aleppo pine ecosystem

Nutrient dynamics of an Aleppo pine (Pinus halepensis, Mill.) ecosystem located in the Kassandra peninsula, Central Macedonia, Northern Greece, were studied using a chronosequence approach. The nutrient composition of the Aleppo pine trees, the understory evergreen broadleaves and forest floor in adjacent stands of 23, 48, 70 and over 100 years old was determined to estimate postfire nutrient losses. The concentration of nutrients in the Aleppo pine trees, except of Ca, was reduced with increasing stand age. Ca was the most abundant nutrient in the aboveground vegetation and in forest litter, followed by N, K, Mg and P. The accumulation of nutrients in the aboveground biomass was positively related to stand age. For younger stands nutrient accumulation was considerably larger in the understory vegetation as compared to the pines, due to substantial enhancement of the understory biomass and the number of understory species present. In middle-aged stands, however, nutrient accumulation in the understory and overstory vegetation reached a balance. In addition, considerable quantities of nutrients have been accumulated in the forest floor particularly in stands of 48 years old. Therefore, any destruction during the period of maximum nutrient accumulation in the forest floor will cause degradation of the ecosystem. It is postulated that the competition for nutrients between overstory and understory vegetation may be as important as competition in soil. Forest management practices leading to the direct conversion of the understory biomass into littermass would be of great significance for the sustainability of the Aleppo pine ecosystem.     

Light-related competitive effects of overstory trees on the understory conifer saplings in a subalpine forest

To clarify the mechanism by which overstory trees shade understory saplings, we investigated the relationships among light conditions of the saplings (measured as indirect site factor; ISF and direct site factor; DSF), the calculated competition effects of overstory trees on the saplings (W), and relative height growth rate of the saplings (RHGR). We calculated several W values in order to find a W value which can express the light conditions as appropriately as possible, and the results indicated that W explained only 21.9%–24.7% of the total variance of light conditions in the cases where W gave the best fit. In this study, W was calculated based on the basal areas of overstory trees. However, it is known that canopy structure also affects the light regimes in the forest understory, and this might yield the possible errors even within W representing the shading effects most adequately. Therefore, although W significantly represents the shading effect from overstory trees, a great proportion of the variance remained without being explained by W. RHGR was negatively correlated with W, and the W value which had the most adequate explanation of the shading effect also showed the best negative correlation with RHGR. This provides the evidence that the competitive effect of overstory trees on sapling growth is mediated by the shading effect, indicating that competition for light clearly exists within this forest. Such competition for light may closely relate to the well-known phenomenon of gap regeneration in subalpine forests in central Japan.     

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.     

Fuel buildup and potential fire behavior after stand-replacing fires,logging fire-killed trees and herbicide shrub removal in Sierra Nevada forests

Typically, after large stand-replacing fires in mid-elevation Sierra Nevada forests, dense shrub fields occupy sites formerly occupied by mature conifers, until eventually conifers overtop and shade out shrubs. Attempting to reduce fuel loads and expedite forest regeneration in these areas, the USDA Forest Service often disrupts this cycle by the logging of fire-killed trees, replanting of conifers and killing of shrubs. We measured the effects of these treatments on live and dead fuel loads and alien species and modeled potential fire behavior and fire effects on regenerating forests. Sampling occurred in untreated, logged and herbicide-treated stands throughout the Sierra Nevada in four large fire areas 4–21 years after stand-replacing fires. Logging fire-killed trees significantly increased total available dead fuel loads in the short term but did not affect shrub cover, grass and forb cover, alien species cover or alien species richness. Despite the greater available dead fuel loads, fire behavior was not modeled to be different between logged and untreated stands, due to abundant shrub fuels in both logged and untreated stands. In contrast, the herbicide treatment directed at shrubs resulted in extremely low shrub cover, significantly greater alien species richness and significantly greater alien grass and forb cover. Grass and forb cover was strongly correlated with solar radiation on the ground, which may be the primary reason that grass and forb cover was higher in herbicide treated stands with low shrub and tree cover. Repeat burning exacerbated the alien grass problem in some stands. Although modeled surface fire flame lengths and rates of spread were found to be greater in stands dominated by shrubs, compared to low shrub cover conifer plantations, surface fire would still be intense enough to kill most trees, given their small size and low crown heights in the first two decades after planting.     

Determinants of grassland primary production in seasonally-dry silvopastoral systems in Central America

Grassland primary productivity is the function that underpins the majority of the fodder production in cattle-rearing silvopastoral farms. Hence, understanding the factors that determine grassland productivity is critical for the design and management of silvpastoral systems. We studied the effect of two factors with documented impact on grassland productivity in seasonally dry silvopastures of Nicaragua, rainfall and trees. We assessed the effects of three species that differed in crown size and phenology, one evergreen, Cassia grandis, and two deciduous species, Guazuma ulmifolia and Tabebuia rosea. Overall, grassland ANPP had a quadratic response to rainfall, with a decline at high rainfall that coincided with peak standing biomass and grassland cover. Trees had a predominately negative effect on grassland productivity, and the effect was concentrated in the rainy season at peak productivity. The effect of the trees corresponded with the tree crown area, but not with crown density. Trees reduced the standing biomass of graminoids and increased forb biomass; thus, the effect of trees on grassland ANPP appears in part to respond to changes in grassland composition. We also found higher levels of soil moisture content below the tree canopy, particularly at the peak of the rainy season when soils tend to become waterlogged. The evergreen species, C. grandis, affected grassland ANPP more strongly than the deciduous species.     

Overstory–understory biomass changes over a 35-year period in southcentral Oregon

Forest canopy development is known to influence understory biomass relationships. An 85,268 ha area in southcentral Oregon was examined for changes in overstory canopy closure using 1953 and 1988 aerial and satellite imagery and a geographic information system. A negative exponential curve predicted a loss of approximately half of understory biomass over the 35-year interval. Reductions in understory biomass were most pronounced at higher elevations where growing conditions for conifers and canopy closure changes were most pronounced. The loss of understory biomass was related to declines in mule deer (Odocoileus hemionus) populations in the area.