Response of branch growth and mortality to silvicultural treatments in coastal Douglas-fir plantations: Implications for predicting tree growth

Static models of individual tree crown attributes such as height to crown base and maximum branch diameter profile have been developed for several commercially important species. Dynamic models of individual branch growth and mortality have received less attention, but have generally been developed retrospectively by dissecting felled trees; however, this approach is limited by the lack of historic stand data and the difficulty in determining the exact timing of branch death. This study monitored the development of individual branches on 103 stems located on a variety of silvicultural trials in the Pacific Northwest, USA. The results indicated that branch growth and mortality were significantly influenced by precommercial thinning (PCT), commercial thinning, fertilization, vegetation management, and a foliar disease known as Swiss needle cast [caused by Phaeocryptopus gaeumannii (T. Rohde) Petr.]. Models developed across these datasets accounted for treatment effects through variables such as tree basal area growth and the size of the crown. Insertion of the branch growth and mortality equations into an individual-tree modeling framework, significantly improved short-term predictions of crown recession on an independent series of silvicultural trials, which increased mean accuracy of diameter growth prediction (reduction in mean bias). However, the static height to crown base equation resulted in a lower mean square error for the tree diameter and height growth predictions. Overall, individual branches were found to be highly responsive to changes in stand conditions imposed by silvicultural treatments, and therefore represent an important mechanism explaining tree and stand growth responses.     

Applicability of non-destructive substitutes for leaf area in different stands of Norway spruce (Picea abies L. Karst.) focusing on traditional forest crown measures

Since individual tree leaf area is an important measure for productivity as well as for site occupancy, it is of high interest in many studies about forest growth. The exact determination of leaf area is nearly impossible. Thus, a common way to get information about leaf area is to use substitutes. These substitutes are often variables which are collected in a destructive way which is not feasible for long term studies. Therefore, this study aimed at testing the applicability of using substitutes for leaf area which could be collected in a non-destructive way, namely crown surface area and crown projection area. In 8 stands of Norway spruce (Picea abies L. Karst.), divided into three age classes and two thinning treatments, a total of 156 trees were felled in order to test the relationship between leaf area and crown surface area and crown projection area, respectively. Individual tree leaf area of the felled sample trees was estimated by 3P-branch sampling with an accuracy of ±10%. Crown projection area and crown surface area were compared with other, more commonly used, but destructive predictors of leaf area, namely sapwood area at different heights on the bole. Our investigations confirmed findings of several studies that sapwood area is the most precise measure for leaf area because of the high correlation between sapwood area and the leaf area. But behind sapwood area at crown base and sapwood area at three tenth of the tree height the predictive ability of crown surface area was ranked third and even better than that of sapwood area at breast height (R² = 0.656 compared with 0.600). Within the stands leaf area is proportional to crown surface area. Using the pooled data of all stands a mixed model approach showed that additionally to crown surface area dominant height and diameter at breast height (dbh) improved the leaf area estimates. Thus, taking dominant height and dbh into account, crown surface area can be recommended for estimating the leaf area of individual trees. The resulting model was in line with many other findings on the leaf area and leaf mass relationships with crown size. From the additional influence of dominant height and dbh in the leaf area model we conclude that the used crown model could be improved by estimating the position of the maximum crown width and the crown width at the base of the crown depending on these two variables.     

Quantifying branch,crown and bole development in Populus tremuloides Michx. from north-eastern British Columbia

Trembling aspen (Populus tremuloides Michx.) from 14 aspen stands ranging in age from 5 to 142 years were destructively sampled to provide branch, crown and bole growth information. Analysis of crown development processes such as branch angle, branch extensional growth, branch crookedness and annual height growth suggested a significant increase in relative branch growth with height and age which was linked to the temporal decline of height growth. Crown class dependent crown length (%) and crown area relationships were observed indicating crown size followed the sequence: open grown trees > dominants > suppressed trees. A significant shift in crown development from vertical to lateral crown expansion was observed as aspen mature.     

Canopy density in stands of Picea abies and Pinus sylvestris after different thinning methods

Altogether 82 plots (261 estimations) of Picea abies (L.) Karst, and 193 plots (360 estimations) of Pinus sylvestris (L.) stands were estimated by a vertical tube. The “crown free projection”, CFP, of stands thinned in three methods with different thinning grades was measured: unthinned, heavily and very heavily thinned, heavily thinned delayed first thinning, extra heavily thinned and thinned from the top. Basal area (m²ha^?1) density (stems ha^?1) and diameter sum (m ha^?1) were plotted against CFP. Basal area was the best practical measure of stand in this study. Generally Scots pine stands have higher CFP and the curves are steeper than in Norway spruce stands. Depending on the grade of thinning, heavily and very heavily thinned spruce stands, delayed first thinning included, have CFP values of 10–15% and stands thinned from the top, 20–40%, compared with 30–80% and 30–60% respectively in pine stands. Extra heavily thinned stands have the highest CFP, 20–80% in spruce and 50–90% in pine stands. The CFP levels after thinning are too high in pine stands for avoidance of sucker and sprout production of aspen and birch. In dense Norway spruce stands thinned from the top or heavily and very heavily thinned, the CFP values are low enough (≤30%) to diminish the production of suckers.     

Individual-based measurement and analysis of root system development: case studies for <Emphasis Type="Italic">Larix gmelinii</Emphasis> trees growing on the permafrost region in Siberia

We present results of individual-based root system measurement and analysis applied for Larix gmelinii trees growing on the continuous permafrost region of central Siberia. The data of root excavation taken from the three stands were used for the analyses; young (26 years old), mature (105 years old), and uneven-aged over-mature stand (220 years old). In this article, we highlight two topics: (1) factors affecting spatio-temporal pattern of root system development, and (2) interactions between aboveground (i.e., crown) and belowground (i.e., root) competition. For the first topic, the detailed observation of lateral roots was applied to one sample tree of the overmature stand. The tree constructed a superficial (<30 cm in depth) and rather asymmetric root system, and each lateral root expanded mainly into elevated mounds rather than depressed troughs. This indicated that spatial development of an individual root system was largely affected by microtopography (i.e., earth hummocks). For these lateral roots, elongation growth curves were reconstructed using annual-ring data, and annual growth rates and patterns were compared among them. The comparison suggested that temporal root system development is associated with differences in carbon allocation among the lateral roots. For the second topic, we examined relationships between individual crown projection area (CA) and horizontal rooting area (RA) for the sample trees of each stand. RA was almost equal to CA in the young stand, while RA was much larger (three or four times) than CA in the mature and overmature stands. Two measures of stand-level space occupation, crown area index (aboveground: CAI; sum of CAs per unit land area) and rooting area index (belowground: RAI; sum of RAs), were estimated in each stand. The estimates of RAI (1.3–1.8 m² m₋₂) exceeded unity in all stands. In contrast, CAI exceeded unity (1.3 m² m₋₂) only in the young stand, and was much smaller (<0.3 m² m₋₂) in the two older stands. These between-stand differences in RAI–CAI relationships suggest that intertree competition for both aboveground and belowground spaces occurred in the young stand, but only belowground competition still occurred in the two older stands. Based on this finding, we hypothesized that competition below the ground may become predominant as a stand ages in L. gmelinii forests. Methodological limitations of our analysis are also discussed, especially for the analysis using the two indices of space occupation (CAI, RAI).     

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.     

Timing and intensity of precommercial thinning in Pinus sylvestris stands

The effects of timing and intensity of precommercial thinning were studied in three Scots pine artificially regenerated stands on Vaccinium forest sites in southern Finland. A two-level factorial design (3×3) was used in each stand: thinning at dominant height of 3, 6 and 9 m to 1000, 1600 and 2200 stems ha^?1. The effects of the treatments were analysed after a period of 23–25 yrs when the dominant height was 14–15 m. Early thinning resulted in the highest standing volume and amount of merchantable wood, and also in slightly accelerated height development. Thinning to 1000 stems ha^?1 caused a considerable production loss, but there were no differences between the densities of 1600 and 2200 stems ha^?1. Branches became thicker after early thinning, but the differences between the treatments were negligible for crop trees. Crown ratio was lowest as the result of early or moderate thinning (2200 stems ha^?1).     

Tree morphology responds to neighbourhood competition and slope in species-rich forests of subtropical China

Trees are able to respond to their local biotic and abiotic environment with morphological adjustments which improve resource acquisition and, thus, growth. In forests, light is broadly recognised as one of the major factors determining growth, and morphological responses comprise changes in crown architecture and stem stature. On sloping terrain, the interplay of phototropism and gravitropism may further affect morphological growth characteristics. However, different tree species are expected to show species-specific responses. In this study, we analysed three growth characteristics of tree individuals belonging to four species of two functional groups (evergreen: Schima superba, Castanopsis eyrei, deciduous: Quercus serrata var. brevipetiolata, Castanea henryi) in a species-rich Chinese subtropical forest. Crown projection area, relative crown displacement and stem inclination were related to biotic (local species richness, functional richness, competition, stand age) and abiotic (slope aspect and inclination, soil depth) variables in the local neighbourhood of the tree individuals. We hypothesised that (i) there are species-specific differences in the morphological response of crown architecture and stem stature and (ii) that crown size and asymmetry as well as stem inclination are influenced by both, biotic and abiotic factors. In contrast to our expectations we were unable to reveal any species-specific differences in any of the three growth characteristics. The results of mixed effects models showed that crown area was mainly affected by the target tree's dbh and biotic variables related to neighbours (competition, functional diversity), whereas stem inclination was mainly influenced by slope. Relative crown displacement was influenced by both, biotic and abiotic variables. We conclude that growth responses resulting in crown displacement and stem inclination seem to be an important mechanism to ameliorate foraging for light in our study area, but that these responses appear to be species-independent. The interplay of stem inclination and crown displacement allows for a plastic response of tree individuals in biotically and abiotically heterogeneous environments. Our results indicate that forest management in this region should focus on functionally diverse stands which are promoting crown area positively resulting in increased growth rates of individual trees.     

A comparison of annual transpiration and productivity in monoculture and mixed-species Douglas-fir and red alder stands

Although much is known about drivers of productivity in Douglas-fir and red alder stands, less is known about how productivity may relate to stand transpiration and water use efficiency. We took advantage of a 15-year-old experiment involving Douglas-fir (Pseudotsuga menziesii) and red alder (Alnus rubra) in the western Cascade Range of western Oregon to test the following hypotheses: (a) more productive stands transpire more water, (b) the relationship between productivity and transpiration differs between species, and (c) the relationship between productivity and transpiration differs between sites varying in soil moisture and fertility. Furthermore, the experimental design included alder, a facultative nitrogen-fixing species, which could also affect fertility. Fixed area plots (20 × 20 m) were planted as monocultures of each species or in mixtures at a common density (1100 trees ha⁻¹) in a randomized-block design. Transpiration of Douglas-fir and red alder was measured using heat dissipation sensors installed in eight trees per plot and scaled to the plot level based on sapwood basal area for each species. Although up to 53% of the variability in tree transpiration was explained by basal area, irrespective of species or site conditions, the two stands with the highest biomass and sapwood basal area did not transpire the most. Instead of more productive stands transpiring more water, the greatest variability in both productivity and transpiration was determined by site conditions and to a lesser degree, species composition. For example, 70% of the variation in tree biomass increment (TBI) was determined by leaf area index, which was much higher at the site with higher fertility and soil moisture (p < 0.05). Despite marked phenological and physiological differences, Douglas-fir and red alder performed similarly. Only 19% of annual water use of Douglas-fir occurred between October and March when alder was leafless. Also, there was no evidence of a fertilization effect of the nitrogen-fixing red alder on the Douglas-fir: the nitrogen concentration and N-isotopic ratio of Douglas-fir needles did not differ whether trees were grown in monoculture or in mixtures with red alder. We conclude that lower soil fertility and contrasting microclimate at one site relative to the other suppressed NPP while maintaining higher transpiration, thus reducing water use efficiency.     

Twenty-six-year response of ponderosa pine and Douglas-fir plantations to woody competitor density in treated stands of madrone and whiteleaf manzanita

Ponderosa pine (Pinus ponderosa Dougl.) grown in mixture with whiteleaf manzanita (Arctostaphylos viscida Parry) and Douglas-fir (Pseudotsuga menziesii (Mirb.) Franco var. menziesii) grown in mixture with Pacific madrone (Arbutus menziesii Pursh) in southwestern Oregon showed an increase in growth with removal of competing woody cover. Both conifer species had roughly one-third the volume at plantation ages 26–27 when grown with uncontrolled competition compared to where woody competition was completely controlled at age 2. Intermediate levels of competitors usually led to intermediate levels of growth, but this was more evident with Douglas-fir than pine. When competition was reduced or removed, height/age relationships for Douglas-fir at plantation ages 23 and 27 reflected medium site quality rather than low quality as estimated from adjacent stands, indicating that these sites are potentially more productive than perceived with uncontrolled dense woody cover. These studies support the concept that competition management may allow some poor sites of ponderosa pine or Douglas-fir to be managed on the basis of a higher site potential.     

Linking morphological and ecophysiological leaf traits to canopy dieback in Persian oak trees from central Zagros

Intraspecific variability in morphological and ecophysiological leaf traits might be theorized to be present in declining populations,since they seem to be exposed to stress and plasticity could be advantageous.Here we focused on declining Persian oaks(Quercus brantii Lindl.var.persica(Jaub and Spach)Zohary)in the Zagros Mountains of western Iran,representing the most important tree species of this region.We selected trees with contrasting crown dieback,from healthy to severely defoliated,to investigate the relationships between canopy dieback and leaf morphology,water content and pigments.We also measured esterase and peroxidase,as enzymatic antioxidants and indicators of contrasting genotypes.Trees showing moderate to severe defoliation showed higher leaf mass area(LMA),reduced relative water content(RWC),and lower stomatal density(SD).Increasing LMA indicates a more sclerophyllic structure,according to drier conditions.We did not find significant differences in leaf pigments(chlorophyll a and b,and carotenoids)among crown dieback classes,suggesting that Persian oak trees are able to maintain accurate photochemical efficiency,while reduced RWC and SD suggest hydraulic limitations.Our results do not provide a consistent pattern as regards enzymatic antioxidant defense in Persian oak.Morphological leaf traits would be important drivers of future adaptive evolution in Persian oak,leading to smaller and thicker leaves,which have fitness benefits in dry environments.Nonetheless,drought responses may be critically affecting carbon uptake,as photosynthetic compounds are less effectively used in leaves with higher sclerophylly.     

Determination of mean diameter by basal area in stands of picea abies and pinus sylvestris in southeastern Norway by means of aerial photographs

Mean diameter by basal area (d_g) is an important stand variable for long‐term economic forecasts of forest holdings. In order to use stand‐by‐stand surveys based on aerial photo interpretation as the data basis for forecasts, d_g has to be determined. The objective was to develop and test a regression function for d_g in mature stands of Norway spruce (Picea abies (L.) Karst.) and Scots pine (Pinus sylvestris L.) applicable in southeastern Norway. A study of 700 plots was used to estimate a function for d_g . An additive model was found to be most suitable. The independent variables were potential yield capacity, Lorey's mean tree height, crown closure determined by ocular estimation by means of aerial photographs, and the product of potential yield capacity and crown closure. The R² value was 0.604 and the coefficient of variation was 10.8%. The regression fitted most parts of the calibration data quite well, but it may overestimate the mean diameter in pure spruce stands by 1–2%, and underestimate the diameter in pure pine stands by 3%. For mixed coniferous stands the regression seems satisfactory. Testing by means of an independent data set showed systematic errors of 3–23%. The systematic errors were due partly to calibration problems in connection with the ocular crown closure estimation.     

Production and quality of senesced and green litterfall in a pine–oak forest in central-northwest Mexico

Litterfall is an important ecological process in forest ecosystems, influencing the transfer of organic matter, carbon (C), nitrogen (N), phosphorous (P) and other nutrients from vegetation to the soil. We examined the production of different litterfall fractions as well as nutrient content and nutrient inputs by senesced and green leaf-litter in a semiarid forest from central Mexico. From September 2006 to August 2007, monthly litter sampling was carried out in monospecific and mixed stands of Quercus potosina and Pinus cembroides. Litterfall displayed a marked bimodal pattern with the largest annual amount (5993 ± 655 kg ha⁻¹ yr⁻¹) recorded in mixed stands, followed by Q. potosina (4869 ± 510 kg ha⁻¹ yr⁻¹), and P. cembroides (3023 ± 337 kg ha⁻¹ yr⁻¹). Leaves constituted the largest fraction of total litterfall reaching almost 60%, while small branches contributed with 20–30%. Overall, N content in leaf-litter was higher while lignin content was significantly lower for Q. potosina than for P. cembroides. Thus, greater litter quality together with higher litter production caused the largest C, N and P inputs to forest soils to occur in monospecific Q. potosina stands. Green leaf fall displayed significantly lower lignin:N and C:N ratios in Q. potosina than P. cembroides suggesting faster decomposition and nutrient return rates by the former. Although we recorded only two green leaf fall events, they accounted for 18% and 11% of the total N and P input, respectively, from leaf-litter during the study period. Apart, from the large spatiotemporal heterogeneity introduced by differences in litter quantity and quality of evergreen, deciduous and mixed stands, green litterfall appears to represent a much more important mechanism of nutrient input to semiarid forest ecosystems than previously considered.     

Contribution of a liana species, <Emphasis Type="Italic">Mucuna macrocarpa</Emphasis> Wall., to litterfall production and nitrogen input in a subtropical evergreen broad-leaved forest

Annual amounts of litterfall and nitrogen input by litterfall were measured in a subtropical evergreen broad-leaved forest to examine the contribution of a liana species, Mucuna macrocarpa Wall., to the spatial heterogeneity of litterfall production and nitrogen input. The total litterfall in the study plot was 7.1 t ha⁻¹ year⁻¹. The amount of litterfall varied with topography and was greatest at the valley bottom and decreased toward the ridges. Macuna macrocarpa litterfall was absent on the ridges although it accounted for the largest percentage, 32%, of total leaf litter production in the valley. Nitrogen input by litterfall was 69 kg ha⁻¹ year⁻¹ in the plot. Nitrogen input by litterfall was also largest at the valley bottom and decreased toward the ridges. Leaf litter of M. macrocarpa had approximately twice the nitrogen concentration of litterfall of other species. Macuna macrocarpa accounted for 42% of nitrogen input by leaf litter in the valley. The abundance and the high nitrogen concentration of M. macrocarpa intensified differences in the amount of litterfall and nitrogen input by litterfall between valleys and ridges. It was concluded that a liana species, M. macrocarpa, can contribute to the spatial heterogeneity of litterfall and may subsequently affect nutrient cycling in a subtropical evergreen broad-leaved forest on Okinawa Island.     

Spatial distribution,architecture, and development of the root system of Pinus banksiana Lamb. in natural and planted stands

In the province of Québec, Canada, the majority of planted jack pine (Pinus banksiana Lamb.) seedlings are produced in rigid wall containers. More than 95% of them exhibit deformations of the root system which may induce stem instability. Studies of the root architecture of planted jack pine have been limited to a 30 cm radius from the stem, as barely any studies have been devoted to naturally regenerated stands. Moreover, only a few researches have focused on temporal evolution of root systems. The aim of the present study was to characterize the architectural, spatial, and temporal development of jack pine roots in natural and planted stands. Study sites were located in the continuous boreal forest of Quebec. The plantation was done in 1987, so that the trees were 15 years old at the time of sampling. Trees from natural stand had regenerated after a fire in 1983 and were 13–16 years old. The root systems of 14 jack pine trees per site were manually excavated up to a <5 mm diameter, without regard to their distance from the stem. The number, length, diameter, and the spatial and temporal development of roots were analyzed according to three scales of root architecture: the root system, axes, and segments. Overall, the numbers and lengths of roots were higher with planted pines. However, naturally regenerated trees displayed a better distribution of their roots around the stem and at depth, combined with more rapid length growth during the first years. In natural stands, all the trees had a taproot and 30% of the main roots originated at a depth of more than 20 cm, and they are regularly distributed around the stems. Planted trees did not present a taproot and 97% of the main roots originated in the first 20 cm beneath the soil surface. Moreover, 50% of root length was located in one-third of the area surrounding the stems, an area that corresponded to the furrow. Finally, the annual development of lateral roots in planted stand displayed a 5-year delay when compared with natural stand, which also affected maximum growth length and development of the branching pattern. Root distribution and temporal development are known to play a major role in the stability of aerial parts. Seedling production methods, container type, site preparation and planting techniques need to be examined in greater detail in order to assess their effect throughout the development of the root system. It is necessary to compare different sylvicultural practices and with natural/planted stands to gain a clearer understanding of this problem.     

Development and variation of wood density, kraft pulp yield and fibre dimensions in young Norway spruce (Picea abies)

Important wood, pulp and fibre properties were investigated on small wood samples from two Rumanian, one German and one Norwegian provenance of Norway spruce (Picea abies) grown in Sørkedalen, Norway. Several samples were collected from inside each single tree, both in radial and transversal direction in the stem. Data were collected from a total of 59 trees, each 28 years of age. All investigated properties showed close relationship to ring number (RN) (cambium age). For basic density and fibre wall thickness (FWT), a fast decline was first observed when moving from pith to bark and a minimum value was found around RN 5–8. The declining trend was then followed by an increase. Kraft pulp yield, fibre length and fibre width (FW) also increased with RN, but the increase was most pronounced close to the pith. The pulp yield (PY) more or less stabilised outside of RN 5–6. FW showed a decrease outside of RN 10–12, but this was probably due to the remarkable simultaneous drop in ring width for the investigated trees. FW decreased, while basic density and fibre length increased with increasing height in the tree. PY and FWT were not affected by height in tree. Diameter at breast height (DBH) was the most important variable indicating differences between trees in the investigated material. Basic density, fibre length and FWT decreased, while FW increased with increasing DBH. Height to crown had a positive effect on basic density, but had no influence on any of the remaining properties. Differences between provenances were found for basic density, fibre length and FW. The analyses showed that it is possible to describe the variation inside and between trees satisfactorily for a range of important wood and fibre properties.