Understanding changes in black (Picea mariana) and white spruce (Picea glauca) foliage biomass and leaf area characteristics

Key message

Growth conditions related to inter-tree competition greatly influence black and white spruce foliage biomass and projected leaf area characteristics.

Abstract

Foliage characteristics such as biomass and area are important among other reasons because they can be related to tree growth. Despite their economic and ecologic importance, equations to characterize foliage biomass and projected area of black (Picea mariana (Miller) BSP) and white (Picea glauca (Moench) Voss) spruces are sparse. Total foliage biomass and projected leaf area, foliage biomass and leaf area density, and relative vertical distribution of black and white spruces foliage biomass and leaf area were modelled with linear and nonlinear mixed effect models. A total of 65 white spruces and 57 black spruces were destructively sampled at four different locations in Alberta, Québec, and Ontario, Canada. Our results show that for each species, total tree foliage biomass and projected leaf area is proportional to stem diameter, total height, and crown length. The addition of crown length in the equations improved the precision of the predictions of total foliage biomass for both species and diminishes greatly the site level random effect. An increase in DBH for black spruce and in the DBH to total height ratio for white spruce skewed the relative vertical foliage biomass distribution toward the base of the living crown. According to our results, growth conditions or tree development stage influence both foliage biomass and leaf area characteristics of black and white spruces. Our results emphasize the importance of inter-tree competition on foliage biomass characteristics.     

Crown responses to neighbor density and species identity in a young mixed deciduous stand

Key message

Density was more important in shaping crown structure than neighbor species identity. Both species showed high crown plasticity at alternative levels, which may explain species coexistence in mixed broadleaved forests with functionally similar species.

Abstract

Understanding crown response to local competition is essential to predicting stand development in mixed stands. We analyzed data from an 8-year-old field experimental plantation mixing two species according to a crossed gradient of density and species proportion to quantify the effect of a broad range of local neighborhood conditions on the development of young trees at multiple crown levels. We used Fagus sylvatica and Acer pseudoplatanus, as two model deciduous species. They are considered functionally equivalent at the young stages, but with contrasting architectural patterns. For both species: (1) changes in density explained more of the variation on crown development than species proportion (2) much of the effect of competition was accounted for by variables at the stem level, while branch and leaf development within crowns were not directly altered by competition. Both species were able to modify their crowns at the stem level to compete with intra- and inter-specific neighbors: Acer and Fagus were taller with a highest proportion of Fagus as neighbors; Fagus displayed a lower crown base when the proportion of Fagus decreased, while Acer had a lower crown base when the proportion of Fagus around it increased. Both species showed common shapes in allometric relationships but contrasting responses at alternative crown levels. Acer exhibited broader intra-specific variation in its height–diameter relationship and in its crown length, while Fagus displayed higher individual variation of branch development and leaf area than Acer. This study demonstrates that differences in crown development strategy of each species in response to changes in local neighborhood conditions are an important factor in maintaining species coexistence in broadleaved forests and designing mixtures that persist over time.     

Changes in the chemical composition of alder,poplar and willow leaves during decomposition in a river

1. As there was a paucity of information on the aquatic decomposition of leaves of willow (Salix alba), poplar (Populus gr. nigra) and alder (Alnus glutinosa), the net-bag technique was used to study chemical changes in the leaves over a period of six months in the Garonne (France).
2. The disappearance rates of foliar organic matter were about similar for the three species (k = 0.0065 day^?1 for alder, k = 0.0054 day^?1 for poplar and k = 0.0050 day^?1 for willow). Changes in carbon amount were comparable to those of organic matter. Nitrogen and organic phosphorus accumulated during the first months of decomposition. The C/N ratios of foliar matter changed in the same way in the three species, falling steeply for the first month, then levelling off. Amounts of sugar disappeared very rapidly, particularly for alder and willow. Cellulose concentrations were constant throughout the whole decomposition process whilst an increase in the amounts of lignin was observed during the first months; however the latter could be due to an interference by microbial compounds.
3. The changes of some constituents, such as carbon, sugars and cellulose, were described with exponential or multiexponential models. Despite the very different initial ratios of C/N and lignin/N, these changes associated with leaf decomposition were similar for the three species.

     

Plant biomass and net production ofanogeissus latifolia Wall. in forests of semiarid zone of rajasthan (india)

A. latifolia grown in the Borimalan forest block in Prasad range (24°11′N and 73°42′ E) exerts clear positive correlations between CBH (circumference breast height)and number of growth rings of bole and branches, tree height, total biomass and leaf area. The net above-ground biomass is 3.95 × 10⁴ kg ha^-1. The average increment in non-photosynthetic (trunk + branch) biomass shows two peaks, the lower peak at 11–16 growth ring period, and the higher one at 34–36 growth ring period. The ratio of leaf dry weight/leaf area is16.3 to 34.8 mg cm^-2, the ratios between shoot net production: leaf weight and leaf area are1.5 g per g and 212 g m^-2 respectively.     

Stable carbon and nitrogen isotope ratios of Eucalyptus and Acacia species along a seasonal rainfall gradient in Western Australia

Key message

Eucalyptus and Acacia species were surprisingly similar with respect to variations in δ ¹³ C, δ ¹⁵ N. Both genera respond with speciation and associated changes in leaf structure to drought.

Abstract

Stable carbon and nitrogen isotope ratios (δ¹³C and δ¹⁵N) in leaves of eucalypts (Corymbia and Eucalyptus) and Acacia (and some additional Fabaceae) species were investigated together with specific leaf area (SLA), leaf nitrogen (N) and leaf phosphorous (P) concentration along a north–south transect through Western Australia covering winter- and summer-dominated rainfall between 100 and 1,200 mm annually. We investigated 62 eucalypts and 78 woody Fabaceae species, mainly of the genus Acacia. Leaf δ¹³C values of Eucalyptus and Acacia species generally increased linearly with latitude from ?29.5 ± 1.3 ‰ in the summer-dominated rainfall zone (15°S–18°S) to about ?25.7 ± 1.1 ‰ in the winter-dominated rainfall zone (29°S–31°S). δ¹⁵N increased initially with southern latitudes (0.5 ± 1.6 ‰ at 15°S; 5.8 ± 3.3 ‰ at 24–29°S) but decreased again further South (4.6 ± 3.5 ‰ at 31°S). The variation in δ¹³C and δ¹⁵N was probably due to speciation of Eucalyptus and Acacia into very local populations. There were no species that were distributed over the whole sampling area. The variation in leaf traits was larger between species than within species. Average nitrogen concentrations were 11.9 ± 1.05 mg g^?1 in Eucalyptus, and were 18.7 ± 4.1 mg g^?1 in Acacia. Even though the average nitrogen concentration was higher in Acacia than Eucalyptus, δ¹⁵N gave no clear indication for N₂ fixation in Acacia. In a multiple regression, latitude (as a surrogate for rainfall seasonality), mean rainfall, leaf nitrogen concentration, specific leaf area and nitrogen fixation were significant and explained 69 % of the variation of δ¹³C, but only 36 % of the variation of δ¹⁵N. Higher nitrogen and phosphorus concentration could give Acacia an advantage over Eucalyptus in arid regions of undefined rainfall seasonality.     

Specific leaf area of European Larch (<Emphasis Type="Italic">Larix decidua</Emphasis> M<Emphasis Type="SmallCaps">ill</Emphasis>.)

Key message

The specific leaf area of European larch depends on branch height and canopy depth, indicating that both, the effect of hydraulic limitations and low water potentials in greater branch heights, and light availability affect specific leaf area.

Abstract

Specific leaf area (SLA) is defined as the ratio between projected leaf area and needle dry mass. It often serves as parameter in ecosystem modelling as well as indicator for potential growth rate. We explore the SLA of European larch (Larix decidua) and the most important factors which have an influence on it. Data were collected from eight stands in Styria, Austria. The stands varied in age, elevation and species mixture. Four stands were pure larch stands with only minor proportions of Norway spruce (Picea abies), whereas the other four were mixed stands of larch and spruce. In each stand 15 representative sample trees were felled. The crown of each sample tree was divided into three sections of equal length and in each section a random sample of needles was taken for determining projected leaf area and dry mass of 50 needles. The mean SLA of larch was established to be 117 cm² g^?1 with a standard deviation of ±27.9 cm² g^?1. SLA varies within the crown, but neither between different mixtures nor years of observation nor social position of the trees. A mixed-effects model, with the plots as random effect, revealed that SLA of larch decreased with increasing branch height (p = 0.0012) and increased with increasing canopy depth (p = 0.029). We conclude that both the hydraulic limitations due to low water potentials in greater branch heights and light availability affect specific leaf area.

     

Relationships between leaf morphological traits, nutrient concentrations and isotopic signatures for Mediterranean woody plant species and communities

Background and aims

Soil factors are driving forces that influence spatial distribution and functional traits of plant species. We test whether two anchor morphological traits—leaf mass per area (LMA) and leaf dry matter content (LDMC)—are significantly related to a broad range of leaf nutrient concentrations in Mediterranean woody plant species. We also explore the main environmental filters (light availability, soil moisture and soil nutrients) that determine the patterns of these functional traits in a forest stand.

Methods

Four morphological and 19 chemical leaf traits (macronutrients and trace elements and δ¹³C and δ¹⁵N signatures) were analysed in 17 woody plant species. Community-weighted leaf traits were calculated for 57 plots within the forest. Links between LMA, LDMC and other leaf traits were analysed at the species and the community level using standardised major axis (SMA) regressions

Results

LMA and LDMC were significantly related to many leaf nutrient concentrations, but only when using abundance-weighted values at community level. Among-traits links were much weaker for the cross-species analysis. Nitrogen isotopic signatures were useful to understand different resource-use strategies. Community-weighted LMA and LDMC were negatively related to light availability, contrary to what was expected.

Conclusion

Community leaf traits have parallel shifts along the environmental factors that determine the community assembly, even though they are weakly related across individual taxa. Light availability is the main environmental factor determining this convergence of the community leaf traits.     

Xylem sap calcium concentrations do not explain liming-induced inhibition of legume gas exchange

Background and aims

Liming is considered normal agricultural practise for remediating soil acidity and improving crop productivity; however recommended lime applications can reduce yield. We tested the hypothesis that elevated xylem sap Ca²⁺ limited gas exchange of Phaseolus vulgaris L. and Pisum sativum L. plants that exhibited reduced shoot biomass and leaf area when limed.

Methods

We used Scholander and whole-plant pressure chamber techniques to collect root and leaf xylem sap, a calcium-specific ion-selective electrode to measure xylem sap Ca²⁺, infra-red gas analysis to measure gas exchange of limed and unlimed (control) plants, and a detached leaf transpiration bioassay to determine stomatal sensitivity to Ca²⁺.

Results

Liming reduced shoot biomass, leaf area and leaf gas exchange in both species. Root xylem sap Ca²⁺ concentration was only increased in P. vulgaris and not in P. sativum. Detached leaves of both species required 5 mM Ca²⁺ supplied to via the transpiration stream to induce stomatal closure, however, maximum in vivo xylem sap Ca²⁺ concentrations of limed plants was only 1.7 mM and thus not high enough to influence stomata.

Conclusion

We conclude that an alternative xylem-borne antitranspirant other than Ca²⁺ decreases gas exchange of limed plants.     

Differences in responses of moderately salt-tolerant and salt-sensitive tree species to heterogeneous salinity

Growth responses of the moderately salt-tolerant velvet ash (Fraxinus velutina) and salt-sensitive poplar (Populus × euramericana) were investigated under heterogeneous root zone salinity. The salinity treatments imposed on the two root zones (lower-higher) were 137-137 (uniform), 103-171, 68-205, 34-239, and 0-273 mM NaCl for velvet ash, and 51-51 (uniform), 34-68, 17-85, and 0-103 mM NaCl for poplar. The leaf gas exchange of the plants was measured one month after these treatments were implemented, and the plants were sampled 75 d after treatment to measure other physiological parameters. Net photosynthetic rate, transpiration rate, total biomass, and fine root compensatory growth increased as the difference in salinity between the two root zones (i.e., salinity heterogeneity) increased in velvet ash. These parameters showed no significant difference among the treatments in poplar. The leaf Na⁺ content was lower under heterogeneous salinity than under uniform salinity in both tested species. The leaf proline content in velvet ash decreased under heterogeneous salinity compared to that under uniform salinity, whereas that of poplar increased. The soluble sugar content of velvet ash leaves increased under heterogeneous salinity, whereas no changes were observed in poplar. The increased fine root biomass in the lower salinity zone promoted velvet ash growth by decreasing the leaf Na⁺ and Cl^- content under heterogeneous salinity. The poplar’s undifferentiated root distribution and gas exchange in response to the heterogeneous salinity were attributed to its salt sensitivity.     

Within-canopy variation in photosynthetic capacity,SLA and foliar N in temperate broad-leaved trees with contrasting shade tolerance

Key message

The relative shade tolerance of T. cordata , F. sylvatica , and C. betulus in mature stands is based on different species-specific carbon and nitrogen allocation patterns.

Abstract

The leaf morphology and photosynthetic capacity of trees are remarkably plastic in response to intra-canopy light gradients. While most studies examined seedlings, it is not well understood how plasticity differs in mature trees among species with contrasting shade tolerance. We studied light-saturated net photosynthesis (A _max), maximum carboxylation rate (V _cmax), electron transport capacity (J _max) and leaf dark respiration (R _d) along natural light gradients in the canopies of 26 adult trees of five broad-leaved tree species in a mixed temperate old-growth forest (Fraxinus excelsior, Acer pseudoplatanus, Carpinus betulus, Tilia cordata and Fagus sylvatica), representing a sequence from moderately light-demanding to highly shade-tolerant species. We searched for species differences in the dependence of photosynthetic capacity on relative irradiance (RI), specific leaf area (SLA) and nitrogen per leaf area (N _a ). The three shade-tolerant species (C. betulus, T. cordata, F. sylvatica) differed from the two more light-demanding species by the formation of shade leaves with particularly high SLA but relatively low N _a and consequently lower area-based A _max, and a generally higher leaf morphological and functional plasticity across the canopy. Sun leaf morphology and physiology were more similar among the two groups. The three shade-tolerant species differed in their shade acclimation strategies which are primarily determined by the species’ plasticity in SLA. Under low light, T. cordata and F. sylvatica increased SLA, mass-based foliar N and leaf size, while C. betulus increased solely SLA exhibiting only low intra-crown plasticity in leaf morphology and N allocation patterns. This study with mature trees adds to our understanding of tree species differences in shade acclimation strategies under the natural conditions of a mixed old-growth forest.     

Crown structure and vertical foliage distribution in 4-year-old plantation-grown Eucalyptus pilularis and Eucalyptus cloeziana

Tree growth and form are both influenced by crown architecture and how it effects leaf distribution and light interception. This study examined the vertical distribution of foliage in 4-year-old plantation-grown Eucalyptus pilularis Sm. and E. cloeziana F. Muell. trees. Leaf area (LA) distribution was determined at two different sites using allometric approaches to determine LA in crown sections and for whole trees. Leaf area was distributed more towards the upper crowns when canopies had been closed for longer. Leaf area was also skewed more towards the upper crowns for Eucalyptus pilularis than E. cloeziana. These species differences were consistent with differences in vertical light availability gradients as determined by point quantum sensors. Leaf area of individual branches was highly correlated with branch cross-sectional area (CSA) and whole-tree LA was closely related to stem CSA. Branch-level allometric relationships were influenced by site and crown position. However, the general allometric equations between stem size and whole-tree leaf area could be applied across sites. Results from this study suggest that pruning of live branches in these species should follow species-specific guides for the timing and height of pruning to optimise the effects on stem growth and form.     

Ontogenetic changes in crown architecture and leaf arrangement: effects on light capture efficiency in three tree species differing in leaf longevity

Pronounced strategy shifts along ontogeny have been observed in several tree species, mainly because of the trend to maximize growth during the seedling stage, which constitutes the most vulnerable part of the tree’s life cycle. Our aim here was to analyze the ontogenetic changes in crown characteristics and light capture patterns in three Quercus species: the evergreens Quercus ilex and Quercus suber and the deciduous Quercus faginea co-occurring in a Mediterranean open woodland. The seedlings were distributed in the large clearings among the adults and received full sunlight. We constructed three-dimensional models of the aerial parts of seedlings and mature trees of the three species, using the YplantQMC program. Large differences between growth stages were observed for all variables. The seedlings exhibited smaller branch sizes and crown densities than those observed in the adult trees. Leaf angles to horizontal also tended to increase during ontogeny, whereas leaf dispersion and the observed distances between leaves tended to decrease. The amount of photosynthetic radiation absorbed per unit leaf area throughout the growing season was lower in adult specimens than in young specimens. Changes in absorption efficiency during ontogeny were more intense for the species with longer leaf life span at maturity. We conclude that more intense ontogenetic shifts in species with longer leaf life span reflect the priority change from the maximization of short-term productivity at the seedling stage to maximizing leaf longevity during the adult stage.     

Allometric and growth data of an evergreen oak, <Emphasis Type="Italic">Quercus glauca</Emphasis>, in a secondary broadleaved forest

The evergreen oak Quercus glauca often dominates secondary broadleaved forests in Western Japan. It is regarded as a mid-successional species, whose diameter and height growth fall between those of early- and late-successional species. Despite the ecological importance of this evergreen oak in the secondary succession of the evergreen broadleaved forest zone in Japan, tree-felling data that allow estimations of tree mass and leaf area from non-destructive measurements are lacking. This paper provides stem growth data, read from tree rings on disks sampled from 13 Q. glauca stems, and their allometric data. The samples were collected in 1994 from the Ginkakuji-san National Forest, Kyoto City, Japan. Allometric data comprised data on stem age, diameter at breast height, diameter at 10% height, tree height, height of the lowest living branch, height of the lowest living leaf in the crown, volume of the main stem, squared stem diameter just below the lowest living branch, total leaf area of the stem, dry weight of the total leaves, dry weight of all branches, dry weight of the main stem, total aboveground dry weight, mean relative photosynthetic photon flux density (PPFD) above the crown, mean relative PPFD below the crown, crown projection area, and specific leaf area. These data can be helpful for estimating the biomass and leaf area index of a Q. glauca stand by enabling the derivation of allometric relationships between non-destructive measurements (such as stem diameter at breast height, and tree height) and tree mass or leaf area. Diameters (including bark thickness) at ground height and above (at 0.5- or 1-m intervals) for each stem are also provided. Stem growth data were based on tree-ring reads from disks taken from heights of 0.0 and 0.3 m, and at 0.5-m (stem height <7 m) or 1.0-m (stem height ≥7 m) intervals above that. Stem volume growth derived from these tree-ring data can be converted into stem mass growth if combined with an analysis of the allometric data, which may serve as a useful resource for the estimation of carbon fixation by evergreen oaks in relation to global climate change.     

Carbon assimilation,nitrogen, and photochemical efficiency of different Himalayan tree species along an altitudinal gradient

In the area of Jumla region in Western Nepal, measurements of saturated leaf net photosynthetic rate (P_sat), nitrogen content, leaf fluorescence, carbon isotopic composition, and water status were performed on woody coniferous (Pinus wallichiana, Picea smithiana, Abies spectabilis, Juniperus wallichiana, Taxus baccata), evergreen (Quercus semecarpifolia, Rhododendron campanulatum), and deciduous broadleaved species (Betula utilis, Populus ciliata, Sorbus cuspidata) spreading from 2 400 m up to the treeline at 4 200 m a.s.l. With the exception of J. wallichiana, P_sat values were lower in coniferous than broadleaved species. Q. semecarpifolia, that in this area grows above the coniferous belt between 3 000 and 4 000 m, showed the highest P_sat at saturating irradiance and the highest leaf N content. This N content was higher and P_sat lower than those of evergreen oak species of tempe forests at middle and low altitudes. For all species, P_sat and N content were linearly correlated, but instantaneous nitrogen use efficiency was lower than values measured in lowland and temperate plant communities. The values of carbon isotopic composition, estimated by δ₁₃C, showed the same range reported for temperate tree species. The ranking of δ₁₃C values for the different tree types was conifers < evergreen broadleaved<deciduous, suggesting tighter stomatal closure and higher water use efficiency for the evergreen types, confirming trends found elsewhere. No relevant differences of δ₁₃C were found along the altitudinal gradient. Quantum yield of photochemistry at saturating irradiance, measured by leaf fluorescence (δF/F_m’), was highest in J. wallichiana and lowest in T. baccata. Overall, photochemical efficiency was more strongly related to species than to altitude. Interestingly, changes of .δF/F_m’ along the altitudinal gradient correlated well with the reported altitudinal distribution of the species.     

Root distribution of different mature tree species growing on contrasting textured soils in temperate windbreaks

Aim

We studied the vertical and lateral root distribution of tree species from three genera (Populus spp. - poplar, Picea spp. - spruce, Salix spp. - willow) that were planted in temperate windbreaks and assessed the effects of soil texture on root density.

Methods

Root distribution to depths of up to 1 m was assessed using the trench-profile method at different distances from the tree rows (2, 6 and 9 m) in 18 mature (average age, 25 years-old) windbreak-sites that were located on light- or heavy-textured agricultural soils in southeastern Québec, Canada. Roots were classified into three diameter classes: fine (<1 mm), medium-size (1–5 mm), and coarse (>5 mm).

Results

Tree fine-root density in poplar and willow windbreaks was higher than in spruce windbreaks at 2 m from the tree row. Root densities were higher in light compared to heavy soils, but these differences were specific to poplar and spruce. Across species groups and soil types, 67 % of the roots occurred in the uppermost 30 cm. In this soil zone, different soil fertility variables (pH, clay content, CEC) were negatively correlated with root density. Densities of spruce and willow roots at 6 m from the tree row were much lower (and often unobserved) than that of poplar. At 9 m, low root densities were observed at only two sites.

Conclusions

We conclude that tree identity and soil type are important drivers of root distribution in temperate agroforestry systems. These results may have important implications for the management of tree competition in agroforestry systems and several ecosystem services that are provided by roots, including C-sequestration, erosion control and water infiltration.     

Oxidation and reduction of sulfite contribute to susceptibility and detoxification of SO2 in Populus × canescens leaves

Key Message

The critical level for SO ₂ susceptibility of Populus × canescens is approximately 1.2 μL L ^?1 SO ₂ . Both sulfite oxidation and sulfite reduction and assimilation contribute to SO ₂ detoxification.

Abstract

In the present study, uptake, susceptibility and metabolism of SO₂ were analyzed in the deciduous tree species poplar (Populus × canescens). A particular focus was on the significance of sulfite oxidase (SO) for sulfite detoxification, as SO has been characterized as a safety valve for SO₂ detoxification in herbaceous plants. For this purpose, poplar plants were exposed to different levels of SO₂ (0.65, 0.8, 1.0, 1.2 μL L^?1) and were characterized by visible injuries and at the physiological level. Gas exchange parameters (stomatal conductance for water vapor, CO₂ assimilation, SO₂ uptake) of the shoots were compared with metabolite levels (sulfate, thiols) and enzyme activities [SO, adenosine 5′-phosphosulfate reductase (APR)] in expanding leaves (80–90 % expanded). The critical dosage of SO₂ that confers injury to the leaves was 1.2 μL L^?1 SO₂. The observed increase in sulfur containing compounds (sulfate and thiols) in the expanding leaves strongly correlated with total SO₂ uptake of the plant shoot, whereas SO₂ uptake rate was strongly correlated with stomatal conductance for water vapor. Furthermore, exposure to high concentration of SO₂ revealed channeling of sulfite through assimilatory sulfate reduction that contributes in addition to SO-mediated sulfite oxidation to sulfite detoxification in expanding leaves of this woody plant species.     

Plant species effect on the decomposition and chemical changes of leaf litter in grassland and pine and oak forest soils

Aims

The aim of this study was to examine the effect of plant species differing in functional and phylogenetic traits on the decomposition processes of leaf litter in a grassland of Japanese pampas grass (Miscanthus sinensis) and adjacent forests of Japanese red pine (Pinus densiflora) and Japanese oak (Quercus crispula), representing sequential stages of secondary succession.

Methods

The litterbag experiments were carried out for 3 years in a temperate region of central Japan.

Results

The decomposition constant (Olson’s k) was 0.49, 0.39, and 0.56/year for grass, pine, and oak, respectively. Nitrogen mass decreased in grass leaf litter during decomposition, whereas the absolute amount of nitrogen increased in leaf litter of pine and oak during the first year. Holocellulose in grass leaf litter decomposed selectively over acid-unhydrolyzable residues more markedly than in leaf litter of pine and oak. ¹³C nuclear magnetic resonance analysis also revealed a decrease in the relative area of O-alkyl-C in grass.

Conclusions

The different decomposition among the three litter species implied that the secondary succession from grassland to pine forest and from pine to oak forests could decrease and increase, respectively, the rate of accumulation and turnover of organic materials and N in soils.     

Annual and spatial variation in shoot demography associated with masting in Betula grossa: comparison between mature trees and saplings

Backgrounds and Aims

Shoot demography affects the growth of the tree crown and the number of leaves on a tree. Masting may cause inter-annual and spatial variation in shoot demography of mature trees, which may in turn affect the resource budget of the tree. The aim of this study was to evaluate the effect of masting on the temporal and spatial variations in shoot demography of mature Betula grossa.

Methods

The shoot demography was analysed in the upper and lower parts of the tree crown in mature trees and saplings over 7 years. Mature trees and saplings were compared to differentiate the effect of masting from the effect of exogenous environment on shoot demography. The fate of different shoot types (reproductive, vegetative, short, long), shoot length and leaf area were investigated by monitoring and by retrospective survey using morphological markers on branches. The effects of year and branch position on demographic parameters were evaluated.

Key Results

Shoot increase rate, production of long shoots, bud mortality, length of long shoots and leaf area of a branch fluctuated periodically from year to year in mature trees over 7 years, in which two masting events occurred. Branches within a crown showed synchronized annual variation, and the extent of fluctuation was larger in the upper branches than the lower branches. Vegetative shoots varied in their bud differentiation each year and contributed to the dynamic shoot demography as much as did reproductive shoots, suggesting physiological integration in shoot demography through hormonal regulation and resource allocation.

Conclusions

Masting caused periodic annual variation in shoot demography of the mature trees and the effect was spatially variable within a tree crown. Since masting is a common phenomenon among tree species, annual variation in shoot demography and leaf area should be incorporated into resource allocation models of mature masting trees.     

Nonlinear mixed-effects branch diameter and length models for natural Dahurian larch (<Emphasis Type="Italic">Larix gmelini</Emphasis>) forest in northeast China

Key message

We developed the generalized branch diameter and length models using the multi-level nonlinear mixed-effects techniques for the natural Dahurian larch ( Larix gmelini ) forest in northeast China.

Abstract

Dahurian larch (Larix gmelini) is the most commercially cultivated timber species in northeastern China due to its ecological prevalence and its superior wood attribute. However, its timber quality was largely driven by the crown architecture, i.e., the number, size and distribution of branches. The majority of branch-level models in the literature are focused on planted forests, which have substantially different crown architecture than that grown in natural mixed forests. Therefore, the goal of this investigation was to develop branch diameter and length models for Dahurian larch that are grown in natural mixed forests. A multi-level nonlinear mixed-effects model technique, including the fixed-effects, random-effects, variance functions and correlation structures, was employed to develop the branch growth models. The results suggested that the cumulative branch diameter and length were both increased with the increases of branch depth into the crown. Diameter at breast height (DBH) had significant positive influences on the branch size; however, tree height (HT) produced negative influences on the branch size, i.e., larger DBH and smaller HT could lead to larger branch size. Model fitting and validation results confirmed that we should avoid developing over-complex models from the perspective of application. As for the branch diameter and length models in our study, addressing the stand and tree level effects as random component were quite reliable and accurate for predicting the branch growth process of Dahurian larch in northeastern China.

     

Spatial variation in sapwood area to leaf area ratio and specific leaf area within a crown of silver birch

Spatial variation in sapwood area to leaf area ratio (Huber value, HV) and specific leaf area (SLA) was examined in branches of closed-canopy trees of silver birch (Betula pendula Roth). HV increased basipetally within a crown and decreased with increasing branch order, but exhibited no significant radial trend along a primary branch. HV was primarily determined by branch position in a crown and branch diameter at the sampling point, being independent of the size of the tree and branch. Greater HV in the lower-crown branches is considered a means to mitigate differences in hydraulic transport capacity between the branches located in different canopy layers. Beside branch position and sampling location on a branch, SLA depended significantly on several other variables characterising tree and branch size. SLA increased basipetally within a crown and along a primary branch, but exhibited no significant trend with branch orders. Because height caused leaf area (A_L) to diminish more rapidly than leaf dry weight, A_L primarily determined the vertical variation in SLA.