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.     

Critical night length for bud set and its variation in two photoperiodic ecotypes of Betula pendula

We studied the variation in critical night length for bud set in two photoperiodic ecotypes (two latitudinally distant stands) of silver birch (Betula pendula Roth) in three phytotron experiments. Seeds from 21 open-pollinated mother trees in a southern (Tuusula, 60 degrees N) and a northern (Kittil?, 67 degrees N) Finnish stand were germinated and grown for 4 weeks in a 24-h photoperiod in a greenhouse and then moved to different night length treatments at 18 degrees C for 4 to 6 weeks. Night lengths from 5 to 8.5 h were used for southern origin seedlings and from 1 to 4.5 h for northern origin seedlings. At the end of the treatments, apical bud set was observed and the percentage of seedlings with bud set calculated for each treatment and tree progeny. The critical night lengths (CNL) for 50% bud set were determined separately for seedlings from each mother tree by regression analysis. In both ecotypes, the mean percentage of seedlings with bud set was lowest for the shortest night lengths and increased rapidly as night lengths increased. Mean CNL with its 95% confidence interval for the southern and northern ecotypes was 6.3 +/- 0.2 and 3.1 +/- 0.3 h, respectively. The CNL of the two ecotypes differed significantly in three experiments. Within-ecotype variance of the CNL was significantly higher in the northern ecotype (0.484) than in the southern ecotype (0.150). Significant differences in CNL were detected between individual mother trees of the southern ecotype, but not between mother trees of the northern ecotype. The ranking of individual mother trees, based on CNL, differed in the three experiments.     

Effect of CCC and daminozide on growth of silver birch container seedlings during three years after spraying

Applications of growth retardants, daminozide and CCC (chlormequatchloride), were studied in order to determine their effects on morphology and post-planting growth of silver birch (Betula pendula Roth) container seedlings. Daminozide was sprayed once (on 28 June at concentrations of 1.0 – 6.0 g l^–1) and CCC was sprayed twice (on 29 June and 27 July at concentrations of 0.5–3.0 gl^–1). Height growth, morphology of seedlings at the end of the growing season, the shoot and root growth potential the following spring and field performance during the following seasons were measured. During the first summer in the nursery, both daminozide and CCC retarded height growth,but daminozide was more effective. The effects of compounds on stem diameter during the summer of application were small. Neither of the compounds affected the field performance of seedlings. The most suitable applications for retardation of height growth, without negative effects on other morphological variables, were 4 g l^–1 (32 mg per seedling) for daminozide and 2 g l^–1 (16 mg per seedling), sprayed in two applications, for CCC.     

Performance and yield of 14 eucalypt species on Imperata cylindrica (L.) Beauv. grassland 3 years after planting

A species trial with 14 eucalypt species with randomized complete blocks design was planted on Imperata cylindrica dominated grassland in South Kalimantan, Indonesia in January 1987. The site was prepared mechanically. No fertilizers were used. At the age of three years E. camaldulensis had clearly the highest survival rate (95%) and mean annual increment (5.9 m³ha^–1a^–1) compared with the other species tried. It was followed by E. urophylla, E. deglupta, E. pellita, E. tereticornis, E. alba, E. torelliana, E. intermedia, E. grandis and E. pilularis. Four other species, E. phaeotricha, E. resinifera, E. cloeziana and E. microcorys died either in the nursery or immediately after planting. All the eucalypts tried had narrow crowns and open canopies. It is suggested that eucalypts are not recommendable for grassland reforestation under minimum plantation tending. Effects of intensive weeding, spacing, fertilizing and performance in second rotation or mixed stands should be tested to elaborate the suitability of eucalypts on grasslands.     

Adaptation of the LIGNUM model for simulations of growth and light response in Jack pine

LIGNUM is a whole tree model, developed for Pinus sylvestris in Finland, that combines tree metabolism with a realistic spatial distribution of morphological parts. We hypothesize that its general concepts, which include the pipe model, functional balance, yearly carbon budget, and a set of architectural growth rules, are applicable to all trees. Adaptation of the model to Pinus banksiana, a widespread species of economic importance in North America, is demonstrated.

Conversion of the model to Jack pine entailed finding new values for 16 physiological and morphological parameters, and three growth functions. Calibration of the LIGNUM Jack pine model for open grown trees up to 15 years of age was achieved by matching crown appearance and structural parameters (height, foliage biomass, aboveground biomass) with those of real trees. A sensitivity study indicated that uncertainty in the photosynthesis and respiration parameters will primarily cause changes to the net annual carbon gain, which can be corrected through calibration of the growth rate. The effect of a decrease in light level on height, biomass, total tree branch length, and productivity were simulated and compared with field data. Additional studies yielded insight into branch pruning, carbon allocation patterns, crown structure, and carbon stress. We discuss the value of the LIGNUM model as a tool for understanding tree growth and survival dynamics in natural and managed forests.     

Predicting spring phenology and frost damage risk of Betula spp. under climatic warming: a comparison of two models

Timing of bud burst and frost damage risk for leaves of Betula spp. in response to climatic warming in Finland was examined with two models. In the first model, ontogenetic development in spring was triggered by an accumulation of chilling temperatures. The second model assumed an additional signal from the light climate. The two models gave radically different estimates of frost damage risk in response to climate warming. The chilling-triggered model forecast a significant and increasing risk with increased warming, whereas the light-climate-triggered model predicted little or no risk. The chilling-triggered model is widely applied in phenological research; however, there is increasing experimental evidence that light conditions play a role in the timing of spring phenology. Although it is not clear if the light response mechanisms are appropriately represented in our model, the results imply that reliance on a light signal for spring development would afford a degree of protection against possible frost damage under climate warming that would not be present if chilling were the sole determinant. Further experimental tests are required to ascertain the light-related mechanisms controlling phenological timing, so that credible model extrapolations can be undertaken.     

Spacing of silver birch seedlings grown in containers of equal size affects their morphology and its variability

Silver birch (Betula pendula Roth) seedlings were grown individually in containers arranged in rows radiating from a central point (Nelder plot) at densities spanning the range from 207 to 891 plants m(-2). Height of one set of seedlings was measured at weekly intervals and additional seedlings were harvested each week for dry mass and leaf area measurements. Height and shoot:root dry mass ratio increased with plant density. Seedling-to-seedling variability in dry mass, but not height, increased with increasing plant density. The red to far-red (R:FR) photon ratio in horizontally propagated radiation decreased with increasing density, even when plant densities and leaf area index values were low. In a separate experiment, elongating internodes of seedlings were irradiated locally by red and far-red light emitting diodes and stem elongation measured. Far-red light markedly increased stem elongation, suggesting that changes in light quality sensed by growing internodes are involved in the observed responses to growth density.     

Effect of tree-level airborne laser-scanning measurement accuracy on the timing and expected value of harvest decisions

The objective was to compare tree-level airborne laser-scanning (ALS) data accuracy with standwise estimation data accuracy as input data for forest planning, using tree- and stand-level simulators. The influence of the input data accuracy was studied with respect to (1) timing of the next thinning or clear-cutting and (2) the relative variation in the predicted income of the next logging expressed as the net present value (NPV). The timing and predicted NPV of thinning and clear-cutting operations were considered separately. The research was based on Monte Carlo simulations carried out with the tree- and stand-level simulators using a simulation and optimisation (SIMO) framework. The simulations used treewise measurements taken on 270 circular plots measured at the Evo Field Station, Finland, as input data. Deviations in the tree data measured were generated according to the mean standard errors found in standwise field estimation and tree-level ALS. The accuracy factors of ALS individual tree detection were based on the EUROSDR/ISPRS Tree Extraction Project. The results show that input data accuracy significantly affects both the timing and relative NPV of loggings. Tree-level ALS produces more accurate simulation results than standwise estimation with the error levels assumed. Diameter-based characteristics are the most important input data in all simulations. Accurate tree height estimates cannot be fully utilised in current simulators.