Structural variation in current-year shoots of broad-leaved evergreen tree saplings under forest canopies in warm temperate Japan

Stem length and leaf area of current-year shoots were measured in saplings of eight broad-leaved evergreen tree species growing under a forest canopy. Stem length varied over a range of one to two orders of magnitude within each species. In all species, both the number of leaves and the mean stem length between successive leaves were greater in longer shoots. Mean leaf size and stem length were not correlated in six of eight species, and only weakly positively correlated in the other two species. Thus, total leaf area per stem increased with stem length, but not in direct proportion: leaf area per stem length was smaller in shoots with long stems and larger in shoots with short stems. I conclude that the within-species variation in the leaf-stem balance of current-year shoots is related to variation in shoot functional roles, as has been observed for long and short shoots in many deciduous tree species: shoots with long stems are extension oriented and contribute to the framework of the crown, whereas shoots with short stems serve mainly for leaf display. Among species, large differences were found in the leaf area per stem length ratio. In the species with larger leaf area per stem length ratios, leaves had narrower blades or longer petioles, or both, resulting in a reduction of mutual shading among the leaves on the shoot.     

Changes in shoot properties in relation to vertical positions within the crown of mature canopy trees of Abies mariesii and Abies veitchii

We investigated current shoot properties in two contrasting vertical positions (leader crown; LC, and lower branch; LB) within the crowns of mature trees of two subalpine conifer species, Abies mariesii and A. veitchii. For both LCs and LBs, shoot length decreased with increasing branching order. However, shoot properties were different between LCs and LBs. Shoots in LCs had more needle biomass per unit of shoot length. Shoots in sunny conditions pack needles closer along the shoot and intercept incoming light more completely. This causes the shoots in the LCs to have more needles. In contrast, less needle packing per unit shoot length in LBs results in the avoidance of mutual shading among needles in order to intercept limited light more effectively. Because branch systems in lower layers tend to be more shaded, the quantity of irradiance received by the shoots in LBs is smaller. Thus, reduced needle amounts on the shoots in LBs reflect the needle arrangement acclimating to the lower light availability. This study suggests the importance of changes in the properties of individual shoot as a component of a branch system and accordingly a whole-crown system in mature canopy trees of A. mariesii and A. veitchii.     

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.     

Leaf traits in relation to crown development, light interception and growth of elite families of loblolly and slash pine

Crown architecture and size influence leaf area distribution within tree crowns and have large effects on the light environment in forest canopies. The use of selected genotypes in combination with silvicultural treatments that optimize site conditions in forest plantations provide both a challenge and an opportunity to study the biological and environmental determinants of forest growth. We investigated tree growth, crown development and leaf traits of two elite families of loblolly pine (Pinus taeda L.) and one family of slash pine (P. elliottii Mill.) at canopy closure. Two contrasting silvicultural treatments -- repeated fertilization and control of competing vegetation (MI treatment), and a single fertilization and control of competing vegetation treatment (C treatment) -- were applied at two experimental sites in the West Gulf Coastal Plain in Texas and Louisiana. At a common tree size (diameter at breast height), loblolly pine trees had longer and wider crowns, and at the plot-level, intercepted a greater fraction of photosynthetic photon flux than slash pine trees. Leaf-level, light-saturated assimilation rates (A(max)) and both mass- and area-based leaf nitrogen (N) decreased, and specific leaf area (SLA) increased with increasing canopy depth. Leaf-trait gradients were steeper in crowns of loblolly pine trees than of slash pine trees for SLA and leaf N, but not for A(max). There were no species differences in A(max), except in mass-based photosynthesis in upper crowns, but the effect of silvicultural treatment on A(max) differed between sites. Across all crown positions, A(max) was correlated with leaf N, but the relationship differed between sites and treatments. Observed patterns of variation in leaf properties within crowns reflected acclimation to developing light gradients in stands with closing canopies. Tree growth was not directly related to A(max), but there was a strong correlation between tree growth and plot-level light interception in both species. Growth efficiency was unaffected by silvicultural treatment. Thus, when coupled with leaf area and light interception at the crown and canopy levels, A(max) provides insight into family and silvicultural effects on tree growth.     

Analysis of the morphology and structure of crowns in a young sugi (Cryptomeria japonica) stand

Crown form, vertical changes in branch inclination and vertical distribution of foliage density in a young sugi (Crytomeria japonica D. Don) stand were analyzed using allometric equations. Tall trees had deeper crowns than short trees, whereas the crown diameters of both tall and short trees were similar. Apical roundness was more pronounced in the lower-story trees, which were characterized by umbrella-shaped crowns. The vertical distribution of foliage density was approximated by a nearly symmetrical curve. Tall trees had higher spatial densities of foliage than short trees. Branch inclination also varied significantly with tree height. The middle-story trees had the largest branch inclinations and the lower-story trees had the smallest branch inclinations. Even in young uniform stands, crown morphology and structure were modified in response to the light environment.     

Structural scaling of light interception efficiency in Picea engelmannii and Abies lasiocarpa

Sunlight interception efficiency was compared at the leaf, shoot, branch and crown levels for Picea engelmannii (Parry) and Abies lasiocarpa ((Hook.) Nutt.), dominant tree species of the central Rocky Mountains, USA. The ratio of silhouette to total leaf area (STAR) was used to quantify the efficiency of direct-beam sunlight interception at each structural scale. Total mean reductions in STAR from the leaf to the crown level were 0.39 to 0.06 in P. engelmannii and 0.46 to 0.02 in A. lasiocarpa. These reductions in STAR occurred for both species as structural scale increased due to a more upright leaf inclination, increased leaf twisting and curvature, or greater mutual shading among plant structures. A steeper leaf inclination between the leaf and shoot level accounted for 26 +/- 19% (95% C.I.) of the total leaf-to-crown STAR reduction; mutual shading among leaves on shoots caused a 14 +/- 7% reduction, whereas leaf curvature and twisting accounted for 22 +/- 3% for a total reduction of 62 +/- 8%. The STAR varied slightly from the shoot to the branch level (+/- 7%) except for a 26% reduction in shade shoots of A. lasiocarpa as a result of increased mutual shading among leaves at lateral nodes. Another substantial reduction in STAR occurred from the branch to the crown level (35 +/- 3% of total) as a result of shading of one branch layer by another within the crown. Thus, light interception efficiency decreased as structural scale increased in both species, especially from the leaf to the shoot level and from the branch to the crown level.     

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.     

Simulation of competition for light in even-aged stands of Douglas Fir

Three versions of an explanatory model to simulate competition between trees within forest stands are presented, based on the distribution of photosynthetically active radiation (PAR) over the trees in the stand. From the amount of PAR absorbed, the rates of assimilation and volume increment are calculated for the trees, which are presented in size classes. Volume increments calculated this way for periods of 35–45 years are compared with measurements from permanent field plots to evaluate different versions of the model.In the first elementary version of the model, it is assumed that the foliage of all trees is distributed uniformly over the field area, neglecting any differences in individual tree height. In this case, the growth of each tree is proportional to its share of the total foliage area of the stand and approximately proportional to its size. This model version slightly underestimated the variation in diameters observed in the field plots.The second version of the model takes into account differences in tree height. As a result of the prior assumption that the foliage of all classes is distributed uniformly over the field area, this model version severely overestimated the shading of short trees by tall neighbours, and the variation in diameters was overestimated as compared to the field data.The third version of the model accounts for clustering of needles within individual crowns in addition to height differences between the classes. This approach gave the better fit to the field data for normally stocked and for dense stands.     

Tree allometry, leaf size and adult tree size in old-growth forests of western Oregon

Relationships between tree height and crown dimensions and trunk diameter were determined for shade-tolerant species of old-growth forests of western Oregon. The study included both understory and overstory species, deciduous and evergreen angiosperms and evergreen conifers. A comparison of adult understory species with sapling overstory species of similar height showed greater crown width and trunk diameter in the former, whether the comparison is made among conifers or deciduous trees. Conifer saplings had wider crowns than deciduous saplings, but the crown widths of the two groups converged with increase in tree height. Conifer saplings had thicker trunks than deciduous saplings of similar crown width, possibly because of selection for resistance to stem bending under snow loads. The results suggest that understory species have morphologies that increase light interception and persistence in the understory, whereas overstory species allocate their biomass for efficient height growth, thereby attaining the high-light environment of the canopy. The greater crown widths and the additional strength requirements imposed by snow loads on conifer saplings result in less height growth per biomass increment in conifer saplings than in deciduous saplings. However, the convergence in crown width of the two groups at heights greater than 20 m, and the proportionately smaller effect of snow loads on large trees, may result in older conifers equalling or surpassing deciduous trees in biomass allocation to height growth.     

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.     

人工整形对观赏果树生长发育的影响

为了科学而有效地利用苹果树资源,以促进休闲观光农业的发展,采用不同的整形处理方法,就苹果树的树体造型及其造型后的生长发育状况进行了试验与观测,结果表明:双环处理的致矮效应最好;"S"树形最为均衡,其平均干粗明显地高于其它处理,其长、中、短枝的比例适宜;双环处理的高效叶最多,其新梢能够及时停止生长,树体积累的养分多,对形成花芽有利;双环、单环和"S"形处理能够促使苹果树形成优美独特的景观。     

Patterns of shoot mortality in Betula platyphylla in northern Japan

To understand the development of crown structure in Betula platyphylla Sukatch., mortality patterns of long shoots were analyzed quantitatively. We selected 25 saplings growing under various light conditions and measured the relative photosynthetically active radiation (rPAR) at, and the three-dimensional position of, first-order branches. A long shoot was assigned "no buds" (NB) status if it lacked buds at the end of the growing season, including at the tips of short shoots. A long shoot was classified as dead if it was NB and all the offspring long shoots issuing from it were NB. The probability that a leafy long shoot (a current-year long shoot with leaves or an older long shoot with short shoots with leaves) would become NB by the end of the season was positively dependent on shoot age and branch age, and negatively dependent on shoot length, centripetal shoot order, branch height and rPAR at the branch. Randomization tests revealed that shoots became NB and dead in clusters of connected shoots. In particular, shoot clusters originating from 3-year-old shoots were more likely to die than expected if each shoot was assumed to become NB regardless of the connection. Stepwise logistic regression revealed that the maximum rPAR within the crown of an individual tree had a significant effect on the mortality rate of 3-year-old shoot clusters, together with the rPAR at the level of the branch and other structural entities. Correlative inhibition is an important mechanism for determining shoot mortality patterns.     

Variation in the ratio of shoot silhouette area to needle area in fertilized and unfertilized Norway spruce trees

We compared the range and variation in shoot silhouette area to projected leaf area ratio (SPAR) in fertilized and unfertilized (control) Norway spruce (Picea abies (L.) Karst.) trees. We measured SPAR for several view directions of 169 shoots at different depths in the crown of fertilized and control trees. There was an increase in SPAR with depth in the crown in both control and fertilized trees. In the fertilized trees, however, mean SPAR was larger overall, the increase with depth in the crown was steeper, and there was a larger variation in SPAR with inclination and rotation angle of the shoot (relative to the view direction). In particular, shoots in the lower crown of fertilized trees were rotationally asymmetrical ("flat") and had high values of the maximum ratio of shoot silhouette area to projected leaf area (SPAR(max)). Differences in SPAR between fertilized and control trees were explained by changes in shoot structure in response to fertilization and shading. Shoots of fertilized trees were larger and had more needle area than shoots of control trees. However, the ratio of needle area to shoot size was smaller in fertilized trees than in control trees, implying less within-shoot shading and, consequently, a larger SPAR. Also, the increase in SPAR with increased shading (depth in the crown) could be explained by a decrease in the ratio of needle area to shoot size. In addition, because fertilized trees had more needle area than control trees, the effect of shading at a given depth in the crown was more pronounced in fertilized trees than in control trees.     

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.     

Height growth of young larch (Larix kaempferi) in relation to the frequency of deer browsing damage in Hokkaido, Japan

The effect of browsing frequency by Sika deer (Cervus nippon yezoensis) on the height growth of Japanese larch (Larix kaempferi) in Hokkaido, Japan, was analyzed. Tree heights at the time of the census and at the start of the current growing season were determined for 100 trees selected randomly at each site. The tallest shoot on each tree was identified and the number of browsing occurrences on the current-year shoot was determined. Young larches browsed by deer repeatedly develop new shoots during a growing season. Larch is a browsing-tolerant species, and the average height growth of the most heavily browsed trees was >15 cm. Although resprouting partly compensated for decreased height growth caused by browsing, the annual increase in height decreased with increased browsing frequency. Thus, browsing frequency during a growing season best reflects the intensity of browsing damage to larch trees.     

Aboveground growth interactions of paired conifer seedlings in close proximity

Where belowground resources are relatively abundant, naturally established trees sometimes occur in very close proximity to one another. We conducted a two-year study to assess the aboveground interactions between Douglas-fir (Pseudotsuga menziesii), grand fir (Abies grandis) and noble fir (Abies procera) seedlings planted in closely spaced (stems 10 cm apart) conspecific and heterospecific pairs. Paired seedling growth also was compared to seedlings planted with no neighbor. Stem height growth was not affected by the presence of a neighbor seedling, although diameter growth was slightly reduced. Branch diameter growth and weight were reduced where seedling crowns overlapped; branch morphological data suggested that this was caused by shading rather than mechanical interactions. Light measurements showed the potential for significant shading, particularly by the relatively large, dense crowns of Douglas-fir. Heterospecific pairs including Douglas-fir demonstrated the competitive production principle in that their mean growth was greater than the average of conspecific pairs of both species. Neighbor seedling height significantly affected subject seedling growth; neighbor effects were similar whether the neighbor seedling was growing on the north or south side of the subject seedling. Light reflected from Douglas-fir crowns had a lower red: far-red ratio than that of noble fir, although there was no evidence of a phytochrome-mediated growth response to the neighbor seedling. While heterospecific seedling pairs showed reduced competition, we found no evidence of facilitation for seedlings growing in very close proximity.     

Variations in leaf functional traits among plant species grouped by growth and leaf types in Zhenjiang,China

Leaf functional traits are adaptations that enable plants to live under various environmental conditions. This study aims to determine the differences in leaf functional traits among plants grouped by growth habit, leaf life span, leaf lifestyle, leaf form, and origin. Specific leaf area (SLA) of perennial or evergreen species was lower than that of annual or deciduous species because longer-lived leaves of perennial or evergreen species require more investment in structural integrity and/or defense against disturbances, especially with any resource constraint. SLA of large individuals was lower than that of small individuals. The low SLA in large individuals can improve their response to changing light and water conditions because increasing plant height is advantageous for light competition, but it can also impose a cost in terms of structural support and water transport. Petioles of plants with compound leaves were significantly longer than those of simple leaves because branching is expensive in terms of gaining height. SLA of plants increased with increasing invasiveness accordingly, and SLA of invasive plants was higher than that of their native congeners because invasive plants should invest more biomass on leaf growth rather than leaf structures per unit area to achieve a higher growth rate. Overall, variation in leaf functional traits among different groups may play an adaptive role in the successful survival of plants under diverse environments because leaf functional traits can lead to pronounced effects on leaf function, especially the acquisition and use of light. Plant species with different growth and leaf traits balance resource acquisition and leaf construction to minimize trade-offs and achieve fitness advantages in their natural habitat.     

Extent of vegetation-free zone necessary for silvopasture establishment of eastern black walnut seedlings in tall fescue

Silvopasture—the integration of trees, forage, and livestock can be established by planting trees in existing pastures. Successful tree establishment and acceptable tree growth in existing tall fescue [Lolium arundinaceum (Schreb.) Darbysh.] pastures requires a vegetation-free zone near the tree base. This study was conducted to determine how large a vegetation-free zone was necessary for the establishment of black walnut (Juglans nigra L.) in tall fescue pastures. Half-sib black walnut seeds were planted in seven different-sized vegetation-free zones [0.0 (control), 0.3, 0.6, 0.9, 1.2, 1.5, and 1.8 m radii] that were created and maintained by applying glyphosate. The study included two consecutive planting years at two locations, one in central Missouri and another in north-central Missouri. The central Missouri site was underlain with well drained, deep soil while the other site was underlain with a poorly drained soil with a defined argillic horizon. Tree height growth was greatest in 0.9-m or larger vegetation-free zones. Diameter growth was greatest in 1.2-m vegetation-free zones and larger. The results suggest that a minimum of a 1.2-m vegetation-free zone in tall fescue pastures should be used to maximize black walnut height and diameter growth in the critical first years of tree establishment.     

Characterization of a type II chlorophyll a/b-binding protein gene (Lhcb2*Pp1) in peach. II. mRNA abundance in developing leaves exposed to sun or shade

Leaf development of shoots exposed to full sunlight and shoots shaded by the canopy was followed in field-grown, mature peach trees (Prunus persica (L.) Batsch, cv. Loring) during the first half of the 1995 growing season. The architecture and size of shaded shoots and sun-exposed shoots differed significantly. Total number of leaves produced on shaded shoots was significantly less than on sun-exposed shoots throughout the season, and differences in leaf number between light conditions increased as the season progressed. The overall patterns of leaf development along sun-exposed and shaded shoots were qualitatively similar. The expression pattern of the type II chlorophyll a/b-binding protein gene, Lhcb2*Pp1, determined by RNA abundance in leaves at different positions along the shoot, was also similar between the two light conditions. The major difference between sun-exposed and shaded leaves was a lower abundance of Lhcb2*Pp1 RNA in mature, shaded leaves compared with sun-exposed leaves. Although the number of fruit per shoot was significantly lower on shaded shoots than on sun-exposed shoots, the rate of fruit drop was not substantially different during the growing season, indicating that quantitative differences in leaf initiation and growth caused by differences in light exposure did not adversely affect fruit retention. However, based on comparison with a previous study of leaf development in non-fruiting trees, reproductive development slowed the rate of vegetative growth without affecting the overall pattern of leaf development along the shoots.     

Influence of tree shelters,irrigation and branch pruning on early field performance of micropropagated wild cherry cv. F12/1

The growth potential of micropropagated wild cherry cv. F12/1 under field conditions was assessed. Relatively short trees (19–22 cm tall) were planted in the field just seven months after transfer from tissue culture to soil. Irrigation and shelters promoted shoot heights in the first (establishment) year. If planting techniques can be developed which reduce branch production, then the cost of corrective pruning to achieve clean stems is also reduced. Shelters, but not irrigation almost totally inhibited outgrowth of branches produced in spring. Outgrowth of spring branches by unsheltered trees was strongly influenced by tree height at planting, with shorter trees producing fewer branches. In the second and third years of the trial, pruning branches from trees that had been planted in shelters, resulted in taller trees. By the end of the third year, pruned trees had smaller stem diameters than unpruned trees. These results are discussed with respect to using cv. F12/1 as a productive timber tree.