Reduced photosynthesis in old oak (Quercus robur): the impact of crown and hydraulic architecture

We tested the hypothesis that changes in crown architecture of old pedunculate oak trees (Quercus robur L. ssp. robur Kl. et Kr. et Rol.) reduce leaf specific hydraulic conductance of shoots, thereby limiting stomatal conductance and assimilation of affected shoots. At the end of summer 1999, hydraulic conductance and leaf specific hydraulic conductance, measured with a high-pressure flow meter in 0.5- to 1.5-m long shoots, were 27 and 39% lower, respectively, in shoots of low vigor compared with vigorously growing shoots in a 165-year- old stand in southeastern Germany. Two types of bottlenecks to water transport can be identified in shoots of old oak trees, namely nodes and abscission zones. The reduction in hydraulic conductance was especially severe in shoots with diameters of less than 2 mm. Maximum stomatal conductance and maximum net assimilation rate increased significantly with hydraulic conductance and leaf specific hydraulic conductance. Our data support the hypothesis that changes in shoot and consequently crown architecture observed in aging trees can limit photosynthesis by reducing shoot hydraulic conductance. Thus, in addition to increasing pathway length and lower conductivity of xylem in old trees, structural changes in shoot and crown architecture need to be considered when analyzing water relations and photosynthesis in mature and declining trees.     

Influence of shoot architectural position on shoot growth and branching patterns in Cleyera japonica

The influence of shoot architectural position on shoot growth and branching patterns was examined in saplings of Cleyera japonica Thunb. (Theaceae), an understory, broad-leaf evergreen woody species. Shoot length varied with branching order and the vertical position of the branch in the crown. In the upper crown, shoot length decreased with increasing branching order, whereas in the lower crown, differences in shoot length among branching orders were not significant. These results demonstrate that it is important to consider not only individual shoots, but also the relationships between shoots in terms of their architectural positions when studying the development of crown architecture in trees. Shoot branching patterns also varied with branching order and the vertical position of the branch in the crown. In the upper crown, branching was mainly sylleptic. In the middle of the crown, mainly proleptic branches were produced. In the lower crown, there was little branching. The importance of these trends in shoot growth and shoot branching patterns in terms of carbon production efficiency is discussed.     

Is rootstock-induced dwarfing in olive an effect of reduced plant hydraulic efficiency?

We investigated the hydraulic architecture of young olive trees either self-rooted or grafted on rootstocks with contrasting size-controlling potential. Clones of Olea europea L. (Olive) cv 'Leccino' inducing vigorous scion growth (Leccino 'Minerva', LM) or scion dwarfing (Leccino 'Dwarf', LD) were studied in different scion/rootstock combinations (LD, LM, LD/LD, LM/LM, LD/LM and LM/LD). Shoots growing on LD root systems developed about 50% less leaf surface area than shoots growing on LM root systems. Root systems accounted for 60-70% of plant hydraulic resistance (R), whereas hydraulic resistance of the graft union was negligible. Hydraulic conductance (K = 1/R) of LD root systems was up to 2.5 times less than that of LM root systems. Total leaf surface area (A(L)) was closely and positively related to root hydraulic conductance so that whole-plant hydraulic conductance scaled by A(L) did not differ between experimental groups. Accordingly, maximum transpiration rate and minimum leaf water potential did not differ significantly among experimental groups. We conclude that reduced root hydraulic conductance may explain rootstock-induced dwarfing in olive.     

Irradiance-induced plasticity in the hydraulic properties of saplings of different temperate broad-leaved forest tree species

We assessed the irradiance-related plasticity of hydraulic architecture in saplings of Betula pendula Roth., a pioneer species; Acer pseudoplatanus L., Fraxinus excelsior L. and Quercus robur L., which are post-pioneer light-requiring species; and Quercus petraea Matt. Liebl. and Fagus sylvatica L. Plants were grown in pots in 36%, 16% and 4% of full sunlight. Hydraulic conductance was measured with a high-pressure flow-meter in entire, in situ root systems and in excised shoots. Leaf-specific whole-plant conductance (LSC) increased with irradiance, due, in part, to an effect of irradiance on plant size. In addition, there was a size-independent effect of irradiance on LSC due, in part, to an increase in root hydraulic conductance paralleled by an increase in root biomass scaled to leaf area. Changes in shoot conductivity also contributed to the size-independent plasticity of LSC. Vulnerability to cavitation measured in current-year twigs was much larger in shade-grown plants. Betula pendula had the highest whole-plant, root and shoot conductances and also the greatest vulnerability to cavitation. The other species were similar in LSC, but showed some variation in root conductance scaled to biomass, with Q. robur, Q. petraea and F. sylvatica having the lowest root conductance and susceptibility to cavitation. All species showed a similar irradiance-related plasticity in LSC.     

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.     

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

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

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.     

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.     

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.     

Seasonal shoot and needle growth of loblolly pine responds to thinning, fertilization, and crown position

The impacts of thinning, fertilization and crown position on seasonal growth of current-year shoots and foliage were studied in a 13-year-old loblolly pine (Pinus taeda L.) plantation in the sixth post-treatment year (1994). Length of new flushes, and their needle length, leaf area, and oven-dry weight were measured in the upper and lower crown from March through November. Total shoot length was the cumulative length of all flushes on a given shoot and total leaf area per shoot was the sum of leaf areas of the flushes.

By the end of June, first-flush foliage reached 70% of the November needle length (14.3 cm) and 65% of the final leaf area (15.0 cm²). Cumulative shoot length of first- and second-flush shoots achieved 95% of the annual length (30.3 cm), whereas total leaf area per shoot was 55% of the final value (75.3 dm²). Fertilization consistently stimulated fascicle needle length, dry weight, and leaf area in the upper crown. Mean leaf area of upper-crown shoots was increased by 64% six years after fertilization. A significant thinning effect was found to decrease mean leaf area per shoot in the crown. For most of the growing season, the thinned-fertilized trees produced substantially more leaf area per shoot throughout the crown than the thinned-nonfertilized trees. These thinned-fertilized trees also had greater needle length and dry weight, longer first flush shoots, and more leaf area per flush than trees in the thinned-nonfertilized plots. Needle length and leaf area of first flush shoots between April and July were linearly related to previous-month canopy air temperature (T_a). Total shoot length strongly depended on vertical light gradient (PPFD) within the canopy, whereas shoot leaf area was a function of both PPFD and T_a. Thus, trees produced larger and heavier fascicles, more and longer flush shoots, and more leaf area per shoot in the upper crown than the lower crown. We conclude that thinning, fertilization, and crown position regulate annual leaf area production of current-year shoots largely by affecting the expansion of first flush shoots and their foliage during the first half of the growing season.     

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.     

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.     

A dynamic model for water flow in a single tree: evidence that models must account for hydraulic architecture

A model is presented for the dynamics of water flow in a single eastern white cedar tree (Thuja occidentalis L.). The model takes into account the spatial and temporal dependence of the evaporative flux from leaves in the crown. It also accounts for the quantitative hydraulic architecture of the tree, i.e., the model characterizes the tree as a branched catena of > 4000 stem segments in which account is taken of the segment length, diameter, hydraulic resistance, and the total area of leaves attached to the segment. Input values needed to run the model are measurements of evaporative flux, hydraulic conductance of stems versus stem diameter, and leaf and stem water storage capacitances. Output parameters are the spatial and temporal characterization of stem and leaf water potentials, stem and leaf water deficits, sap flow rate, and relative sap velocity. The input and output values of the branched catena model are compared and contrasted to that of an unbranched catena model. It is shown that the branched catena model fits independently measured field parameters better than an unbranched catena model. Close correspondence is found between model predictions and field measurements of shoot water potential, pressure gradients in stems, hysteresis in sap velocity between the lower and upper parts of the tree, and diurnal changes in stem and leaf water deficits. This model is discussed in terms of both the hydraulic architecture of trees and the potential application of the model to questions of tree morphology, ecology, physiology and evolution.     

Water utilization,plant hydraulic properties and xylem vulnerability in three contrasting coffee (Coffea arabica) cultivars

Water use, hydraulic properties and xylem vulnerability to cavitation were studied in the coffee (Coffea arabica L.) cultivars San Ramon, Yellow Caturra and Typica growing in the field under similar environmental conditions. The cultivars differed in growth habit, crown morphology and total leaf surface area. Sap flow, stomatal conductance (g(s)), crown conductance (g(c)), apparent hydraulic conductance of the soil-leaf pathway (G(t)), leaf water potential (Psi(L)) and xylem vulnerability to loss of hydraulic conductivity were assessed under well-watered conditions and during a 21-day period when irrigation was withheld. Sap flow, g(c), and G(t) were greatest in Typica both with and without irrigation, lowest in San Ramon, which was relatively unresponsive to the withholding of irrigation, and intermediate in Yellow Caturra. The cultivars had similar g(s) when well watered, but withholding water decreased g(s) more in Typica and Yellow Caturra than in San Ramon. Typica had substantially lower Psi(L) near the end of the unirrigated period than the other cultivars (-2.5 versus -1.8 MPa), consistent with the relatively high sap flow in this cultivar. Xylem vulnerability curves indicated that Typica was less susceptible to loss of hydraulic conductivity than the other cultivars, consistent with the more negative Psi(L) values of Typica in the field during the period of low soil water availability. During soil drying, water use declined linearly with relative conductivity loss predicted from vulnerability curves. However, cultivar-specific relationships between water use and predicted conductivity loss were not observed because of pronounced hysteresis during recovery of water use following soil water recharge. All cultivars shared the same functional relationship between integrated daily sap flow and G(t), but they had different operating ranges. The three cultivars also shared common functional relationships between hydraulic architecture and water use despite consistent differences in water use under irrigated and dry soil conditions. We conclude that hydraulic architectural traits, rate of water use per plant and crown architecture are important determinants of short- and long-term variations in the water balance of Coffea arabica.     

Hydraulic differences along the water transport system of South American Nothofagus species: do leaves protect the stem functionality?

Hydraulic traits were studied for six Nothofagus species from South America (Argentina and Chile), and for three of these species two populations were studied. The main goal was to determine if properties of the water conductive pathway in stems and leaves are functionally coordinated and to assess if leaves are more vulnerable to cavitation than stems, consistent with the theory of hydraulic segmentation along the vascular system of trees in ecosystems subject to seasonal drought. Vulnerability to cavitation, hydraulic conductivity of stems and leaves, leaf water potential, wood density and leaf water relations were examined. Large variations in vulnerability to cavitation of stems and leaves were observed across populations and species, but leaves were consistently more vulnerable than stems. Water potential at 50% loss of maximum hydraulic efficiency (P(50)) ranged from -0.94 to -2.44 MPa in leaves and from -2.6 to -5.3 MPa in stems across species and populations. Populations in the driest sites had sapwood and leaves more vulnerable to cavitation than those grown in the wettest sites. Stronger diurnal down-regulation in leaf hydraulic conductance compared with stem hydraulic conductivity apparently has the function to slow down potential water loss in stems and protect stem hydraulics from cavitation. Species-specific differences in wood density and leaf hydraulic conductance (K(Leaf)) were observed. Both traits were functionally related: species with higher wood density had lower K(Leaf). Other stem and leaf hydraulic traits were functionally coordinated, resulting in Nothofagus species with an efficient delivery of water to the leaves. The integrity of the more expensive woody portion of the water transport pathway can thus be maintained at the expense of the replaceable portion (leaves) of the stem-leaf continuum under prolonged drought. Compensatory adjustments between hydraulic traits may help to decrease the rate of embolism formation in the trees more vulnerable to cavitation.     

Experimental evidence supporting the concept of light-mediated modulation of stem hydraulic conductance

It is a well-described phenomenon that plant leaves respond to changes in light intensity and duration by adjusting leaf hydraulic efficiency, and there is current consensus that up- or down-regulation of water channels (aquaporins) in the plasma membrane of the bundle sheath and mesophyll cells play a central role in the underlying mechanisms. Recently, experimental evidence has been provided also for light-mediated changes of stem hydraulic conductance (K(stem)) in field-grown laurel plants. This effect was attributed to differences in potassium ion concentration of xylem sap as a function of light conditions. In the present article, we report evidence obtained in silver birch (Betula pendula Roth), supporting the concept of light-mediated modulation of K(stem). Both canopy position (long-term effect) and current photosynthetic photon flux density (PPFD; short-term effect) had a significant impact (P < 0.001) on K(stem) measured in shoots taken from the lower (shade shoots) and upper (sun shoots) third of the crowns of ～25-year-old trees growing in a natural forest stand. The shade shoots responded more sensitively to light manipulation: K(stem) increased by 51% in shade shoots and 26% in sun shoots when PPFD increased from 70 to 330 μmol m?2 s?1. In 4-year-old trees growing in a dense experimental plantation, K(stem), specific conductivity of branch-wood (k(bw)) and potassium ion concentration ([K(+)]) in xylem sap varied in accordance with canopy position (P < 0.001). Both K(stem) and k(bw) increased considerably with light availability, increasing within the tree crowns from bottom to top; there was a strong relationship between mean values of K(stem) and [K(+)] in hydraulically sampled branches.     

Effects of branch height on leaf gas exchange,branch hydraulic conductance and branch sap flux in open-grown ponderosa pine

Recent studies have shown that stomata respond to changes in hydraulic conductance of the flow path from soil to leaf. In open-grown tall trees, branches of different heights may have different hydraulic conductances because of differences in path length and growth. We determined if leaf gas exchange, branch sap flux, leaf specific hydraulic conductance, foliar carbon isotope composition (delta13C) and ratios of leaf area to sapwood area within branches were dependent on branch height (10 and 25 m) within the crowns of four open-grown ponderosa pine (Pinus ponderosa Laws.) trees. We found no difference in leaf gas exchange or leaf specific hydraulic conductance from soil to leaf between the upper and lower canopy of our study trees. Branch sap flux per unit leaf area and per unit sapwood area did not differ between the 10- and 25-m canopy positions; however, branch sap flux per unit sapwood area at the 25-m position had consistently lower values. Branches at the 25-m canopy position had lower leaf to sapwood area ratios (0.17 m2 cm-2) compared with branches at the 10-m position (0.27 m2 cm-2) (P = 0.03). Leaf specific conductance of branches in the upper crown did not differ from that in the lower crown. Other studies at our site indicate lower hydraulic conductance, sap flux, whole-tree canopy conductance and photosynthesis in old trees compared with young trees. This study suggests that height alone may not explain these differences.     

Hydraulic architecture and water relations of a flood-tolerant tropical tree, Annona glabra

Hydraulic architecture parameters, water relation parameters and wood anatomy were studied in roots and shoots of the flood-tolerant tree Annona glabra L. on Barro Colorado Island, Panama. Hydraulic conductivity, leaf specific conductivity, and Huber value were similar to the corresponding values for tree species living in non-flooded habitats. The vulnerability of stems to loss of hydraulic conductivity resulting from embolism was low (50% loss of conductivity at -3.3 MPa). The lowest leaf water potential measured in the field was about -1.0 MPa, indicating that A. glabra has a large margin of safety from embolism, which may provide protection against rare drought events, or may be an adaptation to brackish mangrove habitats. Low absolute conductivity of roots was compensated for by an increase in the number of roots. More than two-thirds of whole-plant resistance to water flow was located in the roots.     

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.     

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.