Variations in dark respiration and mitochondrial numbers within needles of Pinus radiata grown in ambient or elevated CO2 partial pressure

Within-leaf variations in cell size, mitochondrial numbers and dark respiration rates were compared in the most recently expanded tip, the mid-section and base of needles of Pinus radiata D. Don trees grown for 4 years in open-top chambers at ambient (36 Pa) or elevated (65 Pa) carbon dioxide partial pressure (p(CO2)a). Mitochondrial numbers and respiratory activity varied along the length of the needle, with the highest number of mitochondria per unit cytoplasm and the highest rate of respiration per unit leaf area at the base of the needle. Regardless of the location of the cells (tip, middle or basal sections), needles collected from trees grown in elevated p(CO2)a had nearly twice the number of mitochondria per unit cytoplasm as those grown in ambient p(CO2)a. This stimulation of mitochondrial density by growth at elevated p(CO2)a was greater at the tip of the needle (2.7 times more mitochondria than in needles grown in ambient CO2) than at the base of the needle (1.7 times). The mean size of individual mitochondria was unaffected either by growth at elevated p(CO2)a or by position along the needle. Tree growth at elevated p(CO2)a had a variable effect on respiration per unit leaf area, significantly increasing respiration in the tip of the needles (+25%) and decreasing respiration at the mid-section and base of the needles (-14% and -25%, respectively). Although a simple relationship between respiration per unit leaf area and mitochondrial number per unit cytoplasm was found within each CO2 treatment, the variable effect of growth at elevated p(CO2)a on respiration along the length of the needles indicates that a more complex relationship must determine the association between structure and function in these needles.     

Leaf respiration at different canopy positions in sweetgum (Liquidambar styraciflua) grown in ambient and elevated concentrations of carbon dioxide in the field

Trees exposed to elevated CO2 partial pressure ([CO2]) generally show increased rates of photosynthesis and growth, but effects on leaf respiration are more variable. The causes of this variable response are unresolved. We grew 12-year-old sweetgum trees (Liquidambar styraciflua L.) in a Free-Air CO2 Enrichment (FACE) facility in ambient [CO2] (37/44 Pa daytime/nighttime) and elevated [CO2] (57/65 Pa daytime/nighttime) in native soil at Oak Ridge National Environmental Research Park. Nighttime respiration (R(N)) was measured on leaves in the upper and lower canopy in the second (1999) and third (2000) growing seasons of CO2 fumigation. Leaf respiration in the light (R(L)) was estimated by the technique of Brooks and Farquhar (1985) in the upper canopy during the third growing season. There were no significant short-term effects of elevated [CO2] on R(N) or long-term effects on R(N) or R(L), when expressed on an area, mass or nitrogen (N) basis. Upper-canopy leaves had 54% higher R(N) (area basis) than lower-canopy leaves, but this relationship was unaffected by CO2 growth treatment. In August 2000, R(L) was about 40% of R(N) in the upper canopy. Elevated [CO(2)] significantly increased the number of leaf mitochondria (62%), leaf mass per unit area (LMA; 9%), and leaf starch (31%) compared with leaves in ambient [CO(2)]. Upper-canopy leaves had a significantly higher number of mitochondria (73%), N (53%), LMA (38%), sugar (117%) and starch (23%) than lower-canopy leaves. Growth in elevated [CO2] did not affect the relationships (i.e., intercept and slope) between R(N) and the measured leaf characteristics. Although no factor explained more than 45% of the variation in R(N), leaf N and LMA were the best predictors for R(N). Therefore, the response of RN to CO2 treatment and canopy position was largely dependent on the magnitude of the effect of elevated [CO2] or canopy position on these characteristics. Because elevated [CO2] had little or no effect on N or LMA, there was no effect on R(N). Canopy position had large effects on these leaf characteristics, however, such that upper-canopy leaves exhibited higher R(N) than lower-canopy leaves. We conclude that elevated [CO2] does not directly impact leaf respiration in sweetgum and that barring changes in leaf nitrogen or leaf chemical composition, long-term effects of elevated [CO2] on respiration in this species will be minimal.     

Seasonal and interannual variability of canopy transpiration of a hedgerow in southern England

Transpiration from a hawthorn (Crataegus monogyna L.) dominated hedgerow in southern England was measured continuously over two growing seasons by the sap flow technique. Accompanying measurements of structural parameters, microclimate and leaf stomatal and boundary layer conductances were used to establish the driving factors of hedgerow transpiration. Observed transpiration rates, reaching peak values of around 8 mm day(-1) and a seasonal mean of about 3.5 mm day(-1), were higher than those reported for most other temperate deciduous woodlands, except short-rotation coppice and wet woodlands. The high rates were caused by the structural and physiological characteristics of hawthorn leaves, which exhibited much higher stomatal and boundary-layer conductances than those of the second-most abundant woody species in the hedgerow, field maple (Acer campestre L.). Only in the hot summer of 2003 did stomatal conductance, and thus transpiration, decrease substantially. The hedgerow canopy was always closely coupled to the atmosphere. Hedgerow transpiration equaled potential evaporation (calculated by the Priestley-Taylor formula) in 2003 and exceeded it in 2004, which meant that a substantial fraction of the energy (21% in 2003 and more than 37% in 2004) came from advection. Hedgerow canopy conductance (g(c)), as inferred from the sap flow data by inverting the Penman-Monteith equation, responded to solar radiation (R(G)) and vapor pressure deficit (D). Although the response to R(G) showed no systematic temporal variation, the response to D, described as g(c)(D) = g(cref) - mln(D), changed seasonally. The reference g(c) depended on leaf area index and the ratio of -m/g(cref) on long-term mean daytime D. A model is proposed based on these observations that predicts canopy conductance for the hawthorn hedge from standard weather data.     

Branch and foliage morphological plasticity in old-growth Thuja plicata

At the Wind River Canopy Crane Facility in southeastern Washington State, USA, we examined phenotypic variation between upper- and lower-canopy branches of old-growth Thuja plicata J. Donn ex D. Don (western red cedar). Lower-canopy branches were longer, sprouted fewer daughter branches per unit stem length and were more horizontal than upper-canopy branches. Thuja plicata holds its foliage in fronds, and these had less projected area per unit mass, measured by specific frond area, and less overlap, measured by silhouette to projected area ratio (SPARmax), in the lower canopy than in the upper canopy. The value of SPARmax, used as an indicator of sun and shade foliage in needle-bearing species, did not differ greatly between upper- and lower-canopy branches. We suggest that branching patterns, as well as frond structure, are important components of morphological plasticity in T. plicata. Our results imply that branches of old-growth T. plicata trees have a guerilla growth pattern, responding to changes in solar irradiance in a localized manner.     

Seasonal and temperature dependence of photosynthesis and respiration for two co-occurring broad-leaved tree species with contrasting leaf phenology

We measured the seasonal and temperature responses of leaf photosynthesis and respiration of two co-occurring native New Zealand tree species with contrasting leaf phenology: winter-deciduous fuchsia (Fuchsia excorticata J. R. Forst & G. Forst) and annual evergreen wineberry (Aristotelia serrata J. R. Forst & G. Forst). There was no difference in the amount of nitrogen per unit leaf area (Narea, range 40-160 mmol m-2, P = 0.18) or specific leaf area (S, range 8-27 m2 kg-1, P = 0.87) in summer leaves of wineberry or fuchsia. The amount of nitrogen per unit leaf area and S varied significantly with height of leaves in the canopy for both species (r2 range 0.61-0.87). Parameters describing the maximum rates of rubisco carboxylation (Vcmax) and electron transport (Jmax) were related significantly to Narea, and were 60% higher on average in spring and summer leaves than in autumn and winter leaves for both species. The seasonal effect remained significant (P < 0.001) when Narea was included in a regression model, indicating that seasonal changes were not only due to changes in Narea. Values for Vcmax and Jmax were 30% lower in wineberry leaves than in fuchsia leaves on average, although the difference ranged from 15% in summer leaves to 39% in autumn leaves. Activation energies describing the temperature dependence of Vcmax and Jmax in wineberry were 111 and 114% of corresponding values for fuchsia (Ea (Vcmax) = 39.1 kJ mol-1, Ea (Jmax) = 32.9 kJ mol-1). Respiration at night was the same (P = 0.34) at 10 degrees C for both species (R10 = 0.7 micromol m-2 s-1), although activation energies (E0) were higher in wineberry than in fuchsia (47.4 and 32.9 kJ mol-1 K-1, respectively). These results show that rates of photosynthesis are higher in winter-deciduous fuchsia than in annual evergreen wineberry.     

Application of real options theory to forestry investment analysis

This paper demonstrates the utility of the real options approach to forestry investment analysis. The main objectives are to discuss the real option theory and show how it can be adopted to model uncertainty and managerial flexibility in forest management and investment. Secondly, we show how to calculate the option values of selected options that may be available to managers of forest industry firms. The paper provides an empirical application, which compares a forestry investment using the static Faustmann model and the real options approach. Four management options are used for the real options approach: an option to delay reforestation, an option to expand the size of the wood processing plant, an option to abandon the processing plant if timber prices fall below a certain level or due to corporate take-over, and multiple options that evaluated all three options together. All options were evaluated using the binomial option-pricing model, where timber values are assumed to follow a multiplicative binomial process. The results show that the Faustmann analysis rejected the investments as unprofitable, while the option analysis showed that all three options were highly valuable if exercised. When real options are considered, the traditional Faustmann model for assessing the profitability of a forestry investment may fail to provide an adequate decision-making framework because it does not properly value management's ability to adjust to shocks in the economy, as well as risks and uncertainty.     

Assessing nitrogen fixation in mixed- and single-species plantations of Eucalyptus globulus and Acacia mearnsii

Mixtures of Eucalyptus globulus Labill. and Acacia mearnsii de Wildeman are twice as productive as E. globulus monocultures growing on the same site in East Gippsland, Victoria, Australia, possibly because of increased nitrogen (N) availability owing to N(2) fixation by A. mearnsii. To investigate whether N(2) fixation by A. mearnsii could account for the mixed-species growth responses, we assessed N(2) fixation by the accretion method and the (15)N natural abundance method. Nitrogen gained by E. globulus and A. mearnsii mixtures and monocultures was calculated by the accretion method with plant and soil samples collected 10 years after plantation establishment. Nitrogen in biomass and soil confirmed that A. mearnsii influenced N dynamics. Assuming that the differences in soil, forest floor litter and biomass N of plots containing A. mearnsii compared with E. globulus monocultures were due to N(2) fixation, the 10-year annual mean rates of N(2) fixation were 38 and 86 kg ha(-1) year(-1) in 1:1 mixtures and A. mearnsii monocultures, respectively. Nitrogen fixation by A. mearnsii could not be quantified on the basis of the natural abundance of (15)N because such factors as mycorrhization type and fractionation of N isotopes during N cycling within the plant confounded the effect of the N source on the N isotopic signature of plants. This study shows that A. mearnsii fixed significant quantities of N(2) when mixed with E. globulus. A decline in delta(15)N values of E. globulus and A. mearnsii with time, from 2 to 10 years, is further evidence that N(2) was fixed and cycled through the stands. The increased aboveground biomass production of E. globulus trees in mixtures when compared with monocultures can be attributed to increases in N availability.     

Large-scale structural variation and long-term growth dynamics of Juniperus thurifera trees in a managed woodland in Soria, central Spain

? Juniper woodlands have a great conservational and productive importance in Spain. However, basic traits of their structural variation and growth dynamics are unknown.

? To characterize the structural variation and assess both height and radial growth patterns, we sampled and measured stem cross-sections every 0.5 m in height from 107 Spanish juniper (Juniperus thurifera L.) trees, which were selected based on a 350 m regular grid over a whole woodland of 3 300 ha in area. We used dendroecological techniques and spatial analysis to study tree structure and growth.

? Structural traits such as bole diameter and height were mainly determined by tree life-related parameters such as tree age and growth rate variation, while abiotic factors such as altitude played a minor role in determining structural variation. Over the last 300 years, juniper establishment has been continuous in time but discontinuous in space. Large-scale spatial heterogeneity of tree establishment, and the presence of an early growth suppression and abrupt growth changes in junipers > 100 years old are consequences of an intense management that almost ceased in the late 19th century.

? Abrupt growth releases and suppressions were synchronic among different age classes, and coincided with documentary records on livestock decrease and key historical changes in land use. Growth patterns suggest that juniper is a brow sing-tolerant species that is able to survive large periods of intense browsing pressure.