Elevated CO2 concentration, fertilization and their interaction: growth stimulation in a short-rotation poplar coppice (EUROFACE)

We investigated the individual and combined effects of elevated CO2 concentration and fertilization on aboveground growth of three poplar species (Populus alba L. Clone 2AS-11, P. nigra L. Clone Jean Pourtet and P. x euramericana Clone I-214) growing in a short-rotation coppice culture for two growing seasons after coppicing. Free-air carbon dioxide enrichment (FACE) stimulated the number of shoots per stool, leaf area index measured with a fish-eye-type plant canopy analyzer (LAIoptical), and annual leaf production, but did not affect dominant shoot height or canopy productivity index. Comparison of LAIoptical with LAI estimates from litter collections and from allometric relationships showed considerable differences. The increase in biomass in response to FACE was caused by an initial stimulation of absolute and relative growth rates, which disappeared after the first growing season following coppicing. An ontogenetic decline in growth in the FACE treatment, together with strong competition inside the dense plantation, may have caused this decrease. Fertilization did not influence aboveground growth, although some FACE responses were more pronounced in fertilized trees. A species effect was observed for most parameters.     

Spatial distribution of leaf morphological and physiological characteristics in relation to local radiation regime within the canopies of 3-year-old Populus clones in coppice culture

Spatial distributions of leaf characteristics relevant to photosynthesis were compared within high-density coppice canopies of Populus spp. of contrasting genetic origin. We studied three clones representative of the range in growth potential, leaf morphology, coppice and canopy structure: Clone Hoogvorst (Hoo) (Populus trichocarpa Torr. & Gray x Populus deltoides Bartr. & Marsh), Clone Fritzi Pauley (Fri) (Populus trichocarpa Torr. & Gray) and Clone Wolterson (Wol) (Populus nigra L.). Leaf area index ranged from 2.7 (Fri and Wol) to 3.8 (Hoo). The clones exhibited large vertical variation in leaf area density (0.02-1.42 m2 m-3). Leaf dry mass per unit leaf area (DM(A)) increased with increasing light in Clones Hoo and Fri, from about 56 g m-2 at the bottom of the canopy to 162 g m-2 at the top. In Clone Wol, DM(A) varied only from 65 to 100 g m-2, with no consistent relationship with respect to light. Conversely, nitrogen concentration on a mass basis was nearly constant (around 1.3-2.1%) within the canopies of Clones Hoo and Fri, but increased strongly with light in Clone Wol, from 1.4% at the bottom of the canopy to 4.1% at the top. As a result, nitrogen per unit leaf area (N(A)) increased with light in the canopies of all clones, from 0.9 g m-2 at the bottom to 2.9 g m-2 at the top. Although a single linear relationship described the dependence of maximum carboxylation rate (17-93 micromol CO2 m-2 s-1) or electron transport capacity (45-186 micromol electrons m-2 s-1) on N(A), for all clones, Clone Wol differed from Clones Hoo and Fri by exhibiting a higher dark respiration rate at low N(A) (1.8 versus 0.8 micromol CO2 m-2 s-1).     

Shoot culture dynamics of six Populus clones

Shoot tips of five genotypically diverse Populus clones, P. alba x P. grandidentata 'Crandon,' P. nigra 'Betulifolia' x P. trichocarpa, P. nigra x P. laurifolia 'Strathglass,' P. maximowiczii x P. trichocarpa 'Androscoggin' and P. deltoides x P. nigra 'Eugenei,' were collected from hardwood cuttings, sterilized,and established in vitro. Stable shoot cultures were obtained from all clones except P. deltoides x P. nigra 'Eugenei'. The four poplar clones that formed stable shoot cultures together with a previously established P. tremula 'Erecta' clone were placed as two-node explants on either Murashige and Skoog medium or Woody Plant Medium containing benzyladenine to determine the rate of shoot multiplication, shoot growth and other responses of the clones. All five poplar clones showed rapid shoot multiplication when cultured in the presence of 0.4-1.0 microM benzyladenine on Murashige and Skoog medium, although P. tremula 'Erecta' produced a greater number of healthy shoots when grown on Woody Plant Medium. Individual shoot growth of all clones was more vigorous when the medium contained 0-0.1 microM benzyladenine, and 100% of such shoots rooted ex vitro.     

Coppicing ability of dry miombo woodland species harvested for traditional charcoal production in Zambia:a win–win strategy for sustaining rural livelihoods and recovering a woodland ecosystem

The miombo woodland is one of the most extensive woodlands in Africa, supporting livelihoods based on biomass fuel for millions of rural people. However, there are growing concerns about the sustainability of harvesting for biomass fuel(mainly charcoal). Thus, the aim of this study was to examine whether regeneration by coppice is a viable option for sustainably managing miombo woodlands for biomass fuel production. We tested the hypotheses that(1) species, stump diameter, stump height and time since cutting significantly affect the number of sprouts per cut stump(coppice density) and mean sprout height(shoot vigour) and(2) higher coppice density reduces shoot vigour due to competition among coppice shoots in a given stump. In an inventory in areas that were harvested for charcoal production by the local people, 369 stumps of 11 species were recorded with mean coppice stumps ranged from 6 to 84. The mean coppice density ranged from 5 to 8 shoots per stump while the mean height of coppice shoots ranged from 46 to 118 cm with marked interspecific variations. Stump size was significantly and positively correlated with coppice density for some of the species, but not with shoot vigour for the majority of the species. However, shoot vigour was significantly positively correlated to time since cutting of trees for nearly half of the species. Coppice density had a significant negative correlation with shoot vigour for two species, and a positive correlation for one species. In conclusion, the results provide evidence about the importance of coppice management as a win–win strategy for sustaining charcoal-based rural livelihoods and recovering the miombo woodland ecosystem.     

Shoot structure and growth along a vertical profile within a Populus-Tilia canopy

We investigated shoot growth patterns and their relationship to the canopy radiation environment and the distribution of leaf photosynthetic production in a 27-m-tall stand of light-demanding Populus tremula L. and shade-tolerant Tilia cordata Mill. The species formed two distinct layers in the leaf canopy and showed different responses in branch architecture to the canopy light gradient. In P. tremula, shoot bifurcation decreased rapidly with decreasing light, and leaf display allowed capture of multidirectional light. In contrast, leaf display in T. cordata was limited to efficient interception of unidirectional light, and shoot growth and branching pattern facilitated relatively rapid expansion into potentially unoccupied space even in the low light of the lower canopy. At the canopy level, T. cordata had higher photosynthetic light-use efficiency than P. tremula, whereas P. tremula had higher nitrogen-use efficiency than T. cordata. However, at the individual leaf level, both species had similar efficiencies under comparable light conditions. Production of new leaf area in the canopy followed the pattern of photosynthetic production. However, the species differed substantially in extension growth and space-filling strategy. Light-demanding P. tremula expanded into new space with a few long shoots, with shoot length strongly dependent on photosynthetic photon flux density (PPFD). Production of new leaf area and extension growth were largely uncoupled in this species because short shoots, which do not contribute to extension growth, produced many new leaves. Thus, in P. tremula, the growth pattern was strongly directed toward the top of the canopy. In contrast, in shade-tolerant T. cordata, shoot growth was weakly related to PPFD and more was invested in long shoot growth on a leaf area basis compared with P. tremula. However, this extension growth was not directed and may serve as a passive means of avoiding self-shading. This study supports the hypothesis that, for a particular species, allocation patterns and crown architecture contribute as much to shade tolerance as leaf-level photosynthetic acclimation.     

Assessment of Populus wood chemistry following the introduction of a Bt toxin gene

Unintended changes in plant physiology, anatomy and metabolism as a result of genetic engineering are a concern as more transgenic plants are commercially deployed in the ecosystem. We compared the cell wall chemical composition of three Populus lines (Populus trichocarpa Torr. & A. Gray x Populus trichocarpa Bartr. ex Marsh., Populus trichocarpa x Populus nigra L. and Populus deltoides x Populus nigra) genetically modified to express the Cry3A or Cry3B2 protein of Bacillus thuringiensis (Bt) with the cell wall chemistry of non-transformed isogenic control lines. Three genetically modified clones, each represented by 10 independent transgenic lines, were analyzed by pyrolysis molecular beam mass spectrometry, gas chromatography/mass spectrometry and traditional wet chemical analytical methods to assess changes in cell wall composition. Based on the outcome of these techniques, there were no comprehensive differences in chemical composition between the transgenic and control lines for any of the studied clones.     

Phytohormones and shoot growth in a three-generation hybrid poplar family

Hybrid vigor for secondary growth in poplar has been linked with increased gibberellin (GA) concentration in cambial tissue, but the relationship between concentrations of GAs and hybrid vigor of primary growth has not been investigated. We explored associations between concentrations of GAs, abscisic acid (ABA) and indoleacetic acid (IAA) and shoot extension in a hybrid family of Populus deltoides Bartr. ex Marsh. (Clone ILL-129) and P. trichocarpa Torr. & Gray (Clone 93-968) and two F1 and 67 F2 genotypes. Rapidly elongating subapical internodes from shoots of 4-year-old clonal saplings were selected for hormone analysis. The F1 hybrids displayed heterosis (hybrid vigor) for primary growth as a result of the complementation of dominance for increased internode length from the P. trichocarpa parent and dominance for increased diameter from the P. deltoides parent. Internodes from the faster-elongating shoots of the P. trichocarpa parent had a fourfold higher concentration of bioactive GA1 and higher concentrations of GA20, GA44, GA29 and GA8 than the P. deltoides parent. However, the two fast-growing F1 hybrids had low concentrations of all five GAs, with concentrations similar to those of the slower-elongating P. deltoides parent. Concentrations of ABA and IAA were correlated with GA concentrations and there was thus no evidence of a consistent promoting or inhibiting effect of ABA or IAA within the F1 family. These results indicate that heterosis for internode growth was not primarily regulated by hormone concentration. The segregating population of F2 hybrids was analyzed to assess the possible utility of hormone screening as a selection tool. The internodes of most of the F2 hybrids were smaller than those of their F1 parents and the larger P. trichocarpa parent. Among the F2 hybrids, mature internode length varied fourfold, and was correlated with lengths of young, subapical internodes from which hormones were analyzed. In these internodes, GA1 concentrations were negatively correlated with length (r = 0.41), diameter (0.33) and mass (0.50). Indoleacetic acid concentration was negatively correlated only with diameter (r = 0.37) and positively correlated with GA1 (r = 0.46), whereas ABA concentration was not correlated with any parameter. Thus, in the F2 population, variation in hormone concentration was not primarily responsible for the variation in shoot growth, indicating that selection for individuals with high GA concentration may not be an effective strategy for identifying vigorous hybrid genotypes.     

Lateral bud outgrowth on decapitated shoots of low-pruned mulberry (Morus alba L.)

After decapitation, lateral shoot growth of mulberry coppice (Morus alba L. cv. Shin-ichinose) from 10-year-old stumps had an extremely acrotonic form compared with that of intact one-year-old stems of trees during the spring. When one dominant shoot from each stump was decapitated, only a few upper lateral buds grew out and elongated. Defoliation of the decapitated shoots resulted in an increase in the number of shoots sprouting and a decrease in the rate of elongation of the laterals. Further enhancement of laterals on decapitated dominant shoots resulted from the removal of competing intact coppice shoots from the stumps. 1-Naphthaleneacetic acid (NAA) applied as a spray reduced the number of lateral buds sprouting from both defoliated, decapitated erect shoots and intact horizontally trained shoots. The results suggest that management of mulberry coppice could increase the supply of leaves for commercial silk production.     

Root produced DHZR-, ZR- and IPA-like cytokinins in xylem sap in relation to coppice shoot initiation and growth in cloned trees of Betula pubescens

Six-year-old cloned Betula pubescens Ehrh. trees, grown outdoors at 65 degrees 01' N, were cut on six dates during the growing season to study coppice shoot development in relation to root-produced cytokinin-like compounds. Bleeding sap was collected over timed intervals for two days after cutting, and endogenous cytokinin-like compounds were measured by ELISA assay in HPLC-purified fractions of xylem sap. Initiation and development of coppice shoots on the clonally propagated plants were comparable to those in seedlings. Coppice shoot initiation was affected by the time of cutting, diminishing significantly after June. Of the cytokinin-like compounds detected in the xylem sap, zeatin riboside-like (ZR) compounds were present in the highest concentrations, and the concentrations of dihydrozeatin riboside-like (DHZR) and isopentenyladenoside-like (IPA) compounds were approximately one third and one eighth of the ZR concentrations, respectively. The concentration of cytokinin-like compounds was positively correlated with xylem sap flow rate. The export of cytokinin-like compounds, especially DHZR- and ZR-types, was positively correlated with the initiation and elongation rate of coppice shoots, the number of lateral branches, and the radial growth of the more slowly growing coppice shoots. The export of cytokinin-like compounds collected immediately after cutting may represent the basal value for each tree. This value is probably affected by the size and activity of the root system and may be a relevant estimate for predicting the success of coppicing.     

Growth and carbohydrate status of coppice shoots of hybrid poplar following shoot pruning

Fifteen, 1-year-old Populus maximowiczii Henry x P. nigra L. 'MN9' trees were decapitated and allowed to sprout. After 8 weeks, all had 6 to 10 coppice shoots. All shoots, except the tallest (dominant) shoot, were removed from five of the trees (pruned treatment), and shoot growth, gas exchange and carbohydrate status were compared in the pruned and unpruned trees. Although photosynthetic rate of recently mature leaves of pruned trees was approximately 50% greater than that of leaves on the dominant shoot of unpruned trees, and the dry weight of leaves of pruned trees was 37% greater than that of the leaves on the dominant shoot of unpruned trees, the shoot dry matter relative growth rate did not differ between treatments. Concentrations of water-soluble carbohydrates and starch in the uppper stem and leaves of the dominant shoot were similar in pruned and unpruned trees. However, relative to that of the dominant shoot in unpruned trees, the lower stem in pruned trees was depleted in both soluble carbohydrates and starch. Starch deposition, assessed as the quantity of (14)C-starch in tissues 24 h after a fully expanded source leaf was labeled with (14)CO(2), was 3.9 times greater in roots of pruned trees than in roots of unpruned trees. We conclude that early removal of all but the dominant shoot reduces the carbohydrate status of the roots and the lower portion of the stem by eliminating the excised shoots as a source of photosynthate.     

Coppice regrowth in a disturbed tropical dry limestone forest in Jamaica

The Hellshire Hills are an extensive area of tropical dry forest over limestone, located on the south coast of Jamaica. Despite being a protected area, they are subject to encroachment and clearance by charcoal burners, leaving a relatively pristine core within more disturbed areas of forest. Future management requires a consideration of the forest’s resilience to such disturbance, and this study reports on coppice regrowth following clearance. The number of coppice shoots and the length and diameter at breast height (DBH) of the leading shoots were measured on tree stumps 14 months after they were cut in an experiment. Coppice regrowth after cutting was measured on a total of 476 stumps, representing 51 species. Coppice production and growth was vigorous as there was an extremely high incidence of coppice regrowth among and within the species in this forest after it had been cut. Initial regrowth was high with the diameter recovered by the leading shoots after 14 months approximating to 29% per year. Of the 51 species sampled three did not coppice at all and the average percent of individuals found with shoots within each species was 81%. The height and diameter of the leading shoots was significantly different among the species and the average height recorded of the leading shoots was 114 cm while the average diameter recorded was 10 mm. Larger sized stems (>10 cm DBH) on average produced more coppice shoots and larger sized shoots than smaller sized stems (<10 cm DBH) while smaller sized stems recovered more biomass than larger sized stems irrespective of species. The average diameter of the leading shoots decreased significantly with increasing shoot densities. At the species level there was a significant relationship between stem diameter and the number of shoots produced in seven of the 27 species analysed and three species returned significant relationships between stem diameter and average height of leading shoots. Coppice regrowth clearly offers considerable resilience to disturbance in this dry forest where successful regeneration by seed is highly susceptible to rainfall seasonality. However, the different abilities of species to produce coppice shoots will likely affect long-term species diversity if the present rate of clearance continues.     

Ecophysiology of riparian cottonwoods: stream flow dependency, water relations and restoration

Cottonwoods (Populus spp.) are adapted to riparian or floodplain zones throughout the Northern Hemisphere; they are also used as parents for fast-growing hybrid poplars. We review recent ecophysiological studies of the native cottonwoods Populus angustifolia James, P. balsamifera L., P. deltoides Marsh., P. fremontii S. Watson and P. trichocarpa T. & G. in North America, and P. nigra L. in Europe. Variation exists within and across species and hybrids; however, all riparian cottonwoods are dependent on shallow alluvial groundwater that is linked to stream water, particularly in semi-arid regions. This conclusion is based on studies of their natural occurrence, decline following river damming and dewatering (water removal), water relations, isotopic composition of xylem water, and by the establishment of cottonwoods along formerly barren natural channels after flow augmentation in response to the conveyance of irrigation water. When alluvial groundwater is depleted as a result of river dewatering or groundwater pumping, riparian cottonwoods exhibit drought-stress responses including stomatal closure and reduced transpiration and photosynthesis, altered 13C composition, reduced predawn and midday water potentials, and xylem cavitation. These physiological responses are accompanied by morphological responses including reduced shoot growth, altered root growth, branch sacrifice and crown die-back. In severe cases, mortality occurs. For example, severe dewatering of channels of the braided Big Lost River in Idaho led to mortality of the narrowleaf cottonwood, P. angustifolia, and adjacent sandbar willows, Salix exigua Nutt., within 5 years, whereas riparian woodlands thrived along flowing channels nearby. The conservation and restoration of cottonwoods will rely on the provision of river flow regimes that satisfy these ecophysiological requirements for survival, growth and reproduction.     

Light interception and partitioning between shoots in apple cultivars influenced by training

The effect of two training systems (Central Leader with branch pruning versus Centrifugal Training with minimal pruning, i.e., removal of fruiting laterals only) on canopy structure and light interception was analyzed in three architecturally contrasting apple (Malus domestica Borkh.) cultivars: 'Scarletspur Delicious' (Type II); 'Golden Delicious' (Type III); and 'Granny Smith' (Type IV). Trees were 3D-digitized at the shoot scale at the 2004 and 2005 harvests. Shoots were separated according to length (short versus long) and type (fruiting versus vegetative). Leaf area density (LAD) and its relative variance (xi), total leaf area (TLA) and crown volume (V) varied consistently with cultivar. 'Scarletspur Delicious' had higher LAD and xi and lower TLA and V compared with the other cultivars with more open canopies. At the whole-tree scale, training had no effect on structure and light interception parameters (silhouette to total area ratio, STAR; projected leaf area, PLA). At the shoot scale, Centrifugal Training increased STAR values compared with Central Leader. In both training systems, vegetative shoots had higher STAR values than fruiting shoots. However, vegetative and fruiting shoots had similar TLA and PLA in Centrifugal Trained trees, whereas vegetative shoots had higher TLA and PLA than fruiting shoots in Central Leader trees. This unbalanced distribution of leaf area and light interception between shoot types in Central Leader trees partly resulted from the high proportion of long vegetative shoots that developed from latent buds. These shoots developed in the interior shaded zone of the canopy and therefore had low STAR and PLA. In conclusion, training may greatly affect the development and spatial positioning of shoots, which in turn significantly affects light interception by fruiting shoots.     

Shoot dynamics of Pinus pumila in relation to altitudinal and wind exposure gradients on the Kiso mountain range, central Japan

Seasonal shoot dynamics of Japanese mountain stone pine (Pinus pumila Regel.) growing at six sites with different altitudes and slope aspects on the Kiso mountain range in central Japan were investigated. The shoots followed a similar growth pattern at all sites; current shoots elongated rapidly during June and July, and the current needles grew during July and August. Final lengths of the current shoots and needles decreased with increasing altitude. Current-shoot lengths were also shorter at the windward sites than at the leeward sites. At all sites, leaf senescence occurred intensively between mid-August and mid-September. Needle longevity, i.e., the age of the oldest living needle attached to a shoot, increased with increasing altitude. Fascicle density, i.e., the number of current needle fascicles on a unit shoot length, was greater at higher altitudes and windward sites than at lower altitudes and leeward sites. Seasonal dynamics of the P. pumila shoots were characterized by simultaneous replacement of old needles with new needles in the early autumn, thus avoiding any loss of canopy photosynthetic production during the growing season. Increases in needle longevity and fascicle density were associated with declining air temperature and increasing wind exposure. Needle longevity and fascicle density were characteristics of adaptive plasticity in P. pumila that prevent a reduction in growth potential in the stressful conditions of alpine regions.     

Hydraulic function contributes to the variation in shoot morphology within the crown in Quercus crispula

Hydraulic and light environments have variation within the crown in well-grown trees. Shoot morphology and shoot hydraulics were compared between the upper and lower crown or among branching patterns in well-grown Quercus crispula Blume. Shoots in the upper crown had longer and thicker axes and lower water potential than did shoots in the lower crown. Hydraulic conductance from the soil to the shoot did not differ between the upper crown and the lower crown. Shoots in the upper crown are exposed to hydraulic stress, and shoots in the lower crown are under shade stress. Shoot morphology and shoot hydraulic traits (i.e., higher Huber value and higher hydraulic conductivity) in the upper crown affected the hydraulic conductance of shoots. Shoots in the lower crown showed larger light-receiving leaf area per leaf biomass investment, which is an adaptive morphology under shaded environments. Shoot morphology and shoot hydraulics were not correlated to branching pattern significantly, but shoots with higher branching intensity in the upper crown represented trends for higher hydraulic conductivity. These results reveal that shoot morphological and physiological characteristics in the upper crown reduce hydraulic stress, and those in the lower crown reduce shade stress. I conclude that vertical position within a crown affects both morphological and physiological acclimation for light acquisition and hydraulic conductance, and that hydraulic architecture is associated with crown architecture.     

Shoot growth and crown development: effect of crown position in three-dimensional simulations

Trees have been increasingly considered as modular organisms, with individual shoots forming autonomous units that respond semi-independently to their surrounding environment. However, there is evidence for fairly strict hormonal control of tree crown development. Studies on the hydraulic architecture of trees suggest a closer functional connection between shoots and crown development than is postulated by the theory of branch autonomy. We studied how shoot growth pattern influences growth and crown architecture in young Scots pine trees simulated by the LIGNUM model assuming that (a) the growth of a shoot mainly depends on its light climate and (b) the growth of a shoot is influenced by its position within the crown. We determined shoot position within the crown based on a recently developed vigor index. The vigor index compares the relative axis cross-sectional area from the base of the tree to each shoot and gives a value of 1 to the pathway of the greatest cross-sectional area. All other shoots attain values between 0 and 1 depending on their cross-sectional areas and the cross-sectional areas of the branches leading there from the main axis. The shoot light climate is characterized by annually intercepted photosynthetically active radiation. We compared the results from simulations (a) and (b) against an independent data set. The addition of a within-shoot position index (the vigor index) to our simulation (simulation b) resulted in a more realistic tree form than that obtained with simulation (a) alone. We discuss the functional significance of the results as well as the possibilities of using an index of shoot position in simulations of crown architecture.     

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.     

Altered light conditions following thinning affect xylem structure and potential hydraulic conductivity of Norway spruce shoots

Trees must respond to many environmental factors during their development, and light is one of the main stimuli regulating tree growth. Thinning of forest stands by selective tree removal is a common tool in forest management that increases light intensity. However, morphological and anatomical adaptations of individual shoots to the new environmental conditions created by thinning are still poorly understood. In this study, we evaluated shoot morphology (shoot length, needle number, projected leaf area) and anatomy (tracheid lumen area, tracheid number, tracheid dimensions, xylem area, potential hydraulic conductivity) in three Norway spruce (Picea abies/L./Karst.) families exposed to different thinning regimes. We compared shoot characteristics of upper-canopy (i.e. sun-exposed) and lower-canopy (i.e. shaded) current-year shoots in a control plot and a plot thinned to 50 % stand density the previous year. One tree per family was chosen in each treatment, and five shoots were taken per canopy position. We found that upper-canopy shoots in both plots had higher values than lower-canopy shoots for all studied parameters, except lumen roundness and tracheid frequency (i.e. tracheid number per xylem area). Thinning had little effect on shoot morphology and anatomy 1 year after thinning, except for small but significant changes in tracheid dimensions. Needles were more sensitive to altered light conditions, as projected leaf area of shoot, needle number and leaf hydraulic conductivity changed after thinning. Differences between upper- and lower-canopy shoots did not seem to be influenced by thinning and were almost the same in both plots. Our results suggest that lower-canopy shoots require several years to modify their morphology and anatomy to new light conditions following thinning. The slow light adaptation of the lower canopy may be of practical importance in forest management: thinned stands may be predisposed to drought stress because newly exposed shoots experience increased illumination and transpiration after thinning.     

整形技术对马尾松种子园结实层的影响

本实验研究表明：截除马尾松高生长主梢和各级侧枝的主梢能有效地控制马尾松的树高生长和树冠增长,缩小树冠内膛空间,提高树冠有效采光面,从而可提高种子园采种母树密度;控制结实层上移,可有效降低种子园采种难度;增大采种母树结实层厚度,可提高采种母树的单株产量.     

Changes in shoot allometry with increasing tree height in a tropical canopy species,Elateriospermum tapos

Allometry of shoot extension units (hereafter termed "current shoots") was analyzed in a Malaysian canopy species, Elateriospermum tapos Bl. (Euphorbiaceae). Changes in current shoot allometry with increasing tree height were related to growth and maintenance of tree crowns. Total biomass, biomass allocation ratio of non-photosynthetic to photosynthetic organs, and wood density of current shoots were unrelated to tree height. However, shoot structure changed with tree height. Compared with short trees, tall trees produced current shoots of the same mass but with thicker and shorter stems. Current shoots with thin and long stems enhanced height growth in short trees, whereas in tall trees, thick and short current shoots may reduce mechanical and hydraulic stresses. Furthermore, compared with short trees, tall trees produced current shoots with more leaves of lower dry mass, smaller area, and smaller specific leaf area (SLA). Short trees adapted to low light flux density by reducing mutual shading with large leaves having a large SLA. In contrast, tall trees reduced mutual shading within a shoot by producing more small leaves in distal than in proximal parts of the shoot stem. The production of a large number of small leaves promoted light penetration into the dense crowns of tall trees. All of these characteristics suggest that the change in current shoot structure with increasing tree height is adaptive in E. tapos, enabling short trees to maximize height growth and tall trees to maximize light capture.