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Thinning, fertilization, and crown position interact to control physiological responses of loblolly pine.

To examine physiological responses to thinning, fertilization, and crown position, we measured net photosynthesis (P(n)), transpiration (E), vapor pressure difference (VPD), stomatal conductance (g(s)), and xylem pressure potential (Psi(1)) between 0930 and 1130 h under ambient conditions in the upper and lower crowns of a 13-year-old loblolly pine (Pinus taeda L.) plantation six years (1994) after the treatments were applied. Photosynthetic photon flux density (PPFD) and air temperature (T(a)) within the canopy were also recorded. Needle P(n) of thinned trees was significantly enhanced by 22-54% in the lower crown, because canopy PPFD increased by 28-52%. Lower crown foliage of thinned plots also had higher E and g(s) than foliage of unthinned plots, but thinning had no effect on needle Psi(1) and predawn xylem pressure potential (0430-0530 h; Psi(pd)). Tree water status did not limit P(n), E and g(s) during the late-morning measurements. Fertilization significantly decreased within-canopy PPFD and T(a). Needle Psi(1) was increased in fertilized stands, whereas P(n), E and g(s) were not significantly altered. Upper crown foliage had significantly greater PPFD, P(n), VPD, g(s), E, and more negative Psi(1) than lower crown foliage. In both crown positions, needle P(n) was closely related to g(s), PPFD and T(a) (R(2) = 0.77 for the upper crown and 0.82 for the lower crown). We conclude that (1) silvicultural manipulation causes microclimate changes within the crowns of large trees, and (2) needle physiology adjusts to the within-crown environmental conditions.

Journal Article↗

Transpiration and forest structure in relation to soil waterlogging in a Hawaiian montane cloud forest.

Transpiration, leaf characteristics and forest structure in Metrosideros polymorpha Gaud. stands growing in East Maui, Hawaii were investigated to assess physiological limitations associated with flooding as a mechanism of reduced canopy leaf area in waterlogged sites. Whole-tree sap flow, stomatal conductance, microclimate, soil oxidation-reduction potential, stand basal area and leaf area index (LAI) were measured on moderately sloped, drained sites with closed canopies (90%) and on level, waterlogged sites with open canopies (50-60%). The LAI was measured with a new technique based on enlarged photographs of individual tree crowns and allometric relationships. Sap flow was scaled to the stand level by multiplying basal area-normalized sap flow by stand basal area. Level sites had lower soil redox potentials, lower mean stand basal area, lower LAI, and a higher degree of soil avoidance by roots than sloped sites. Foliar nutrients and leaf mass per area (LMA) in M. polymorpha were similar between level and sloped sites. Stomatal conductance was similar for M. polymorpha saplings on both sites, but decreased with increasing tree height (r(2) = 0.72; P < 0.001). Stand transpiration estimates ranged from 79 to 89% of potential evapotranspiration (PET) for sloped sites and from 28 to 51% of PET for level sites. Stand transpiration estimates were strongly correlated with LAI (r(2) = 0.96; P < 0.001). Whole-tree transpiration was lower at level sites with waterlogged soils, but was similar or higher for trees on level sites when normalized by leaf area. Trees on level sites had a smaller leaf area per stem diameter than trees on sloped sites, suggesting that soil oxygen deficiency may reduce leaf area. However, transpiration per unit leaf area did not vary substantially, so leaf-level physiological behavior was conserved, regardless of differences in tree leaf area.

Journal Article↗

Decreased needle longevity of fertilized Douglas-fir and grand fir in the northern Rockies.

Changes in nutrient availability significantly affect canopy dynamics in conifers. To elucidate these effects, we experimentally fertilized mixed conifer stands at several sites across the northern Rocky Mountains. We measured needle longevity, total branch length and foliated length along the main branch axis, and determined mean retained cohort length on mid-canopy branches of shade-intolerant Douglas-fir (Pseudotsuga menziesii var. glauca (Beissn.) Franco) and shade-tolerant grand fir (Abies grandis Lindl.). Needle longevity ranged from 6 to 8 years in Douglas-fir and from 7 to 8 years in grand fir on unfertilized plots. Fertilization significantly decreased needle longevity by 26 and 27% in Douglas-fir and grand fir, respectively. However, the foliated branch length remained unchanged following fertilization and was similar for both species, indicating a 33% increase in mean branch length per needle cohort in Douglas-fir and a 27% increase in grand fir. These data are consistent with the theory that foliated branch length and needle longevity are a result of the ecological light compensation point (ELCP), which links the inherent physiology of the leaf with the availability of resources in the leaf environment. Mid-canopy ELCP was approximately 74 and 71 cm from the branch terminus in Douglas-fir and grand fir, respectively, regardless of fertilization. We hypothesize that fertilization-enhanced needle production and annual shoot growth resulted in a higher rate of shading of older needles. The shaded needles were unable to maintain a positive carbon balance and abscised. The results demonstrate that foliated branch length of Douglas-fir and grand fir in the northern Rocky Mountains can be treated as a homeostatic response to fertilization, whereas foliar turnover is plastic.

Journal Article↗

A method for estimating light interception by a conifer shoot.

We present an operational method for estimating the amount of PAR intercepted by a coniferous shoot. Interception of PAR by a shoot is divided into three components: the amount of radiation coming from the sky, the transmission of radiation through the surrounding vegetation, and the shoot' s silhouette area facing the direction of the incoming radiation. All three components usually vary with direction. Radiation incident from the sky consists of direct and diffuse radiation. The well-known equation of motion for the sun and Beer' s Law for atmospheric transmittance are used to simulate the directional distribution of direct sunlight for any given period of time. The diffuse component is assumed to be uniform. Meteorological field measurements are used to calibrate the absolute amounts of the direct and diffuse components. The gap fraction (proportion of visible sky) in different directions around a shoot is measured by analyzing a hemispherical fish-eye photograph, taken at the location of the shoot, with an image processing program. Similarly, the shoot silhouette area (SSA) is measured by photographing the shoot from many different directions. The measurements of SSA are interpolated by a method called trigonometric interpolation to obtain the directional distribution of SSA over the entire hemisphere. This distribution is then rotated according to the shoot' s position in the canopy. Multiplying incoming PAR, canopy gap fraction and SSA in different directions, and summing over all directions, gives an estimate of PAR intercepted by the shoot during the chosen period of time. The method is described step by step, and applied, as an example, to a shoot from a Scots pine (Pinus sylvestris L.) stand in central Finland. Differences in radiation interception properties between sun and shade shoots and their relevance to canopy-scale models are discussed.

Cycadopsida↗

Site fertility and the morphological and photosynthetic acclimation of Pinus sylvestris needles to light.

Morphological and photosynthetic acclimation of current-year needles to canopy gradients in light availability (seasonal mean integrated quantum flux density, Q(int)) was studied in the temperate conifer, Pinus sylvestris L., at two sites of contrasting nutrient availability. The nutrient-rich site supported a monospecific P. sylvestris stand on an old-field. The trees were approximately 30 years old and 19-21 m tall. Mean foliar N and P contents (+/- SD) were 1.53 +/- 0.11% and 0.196 +/- 0.017%, respectively. The nutrient-poor site was located on a raised bog supporting a sparse stand of 50- to 100-year-old trees, with a height of 1-2 m, and mean needle N and P contents of 0.86 +/- 0.12% and 0.074 +/- 0.010%, respectively. At both sites, needle thickness (T) and width (W) increased with increasing Qint, and leaf dry mass per unit leaf area (MA) was also greater at higher irradiance. The light effects on MA-the product of needle density (D) and volume to total area ratio (V/AT)-resulted primarily from large increases in V/AT with Qint rather than from modifications of D, which was relatively insensitive to light. Although needle morphology versus light relationships were qualitatively similar at both sites, needles were shorter, and the slopes of W, T, MA and V/AT versus light relationships were lower, at the nutrient-poor than at the nutrient-rich site, indicating that the plasticity of foliar morphological characteristics was affected by nutrient availability. As a result of lower plasticity, needles at the nutrient-poor site were narrower, thinner, and had lower MA at high irradiance than needles at the nutrient-rich site. The maximum carboxylase activity of ribulose-1,5-bisphosphate carboxylase/oxygenase (Vcmax) and the maximum photosynthetic electron transport rate (Jmax) scaled positively with foliar N and P contents. The correlations were generally stronger with P than with N, suggesting that needle photosynthetic capacity was more heavily limited by the availability of P than of N. The Jmax/Vcmax ratio was positively related to the foliar P/N ratio, indicating that Jmax was more strongly suppressed than Vcmax under conditions of low P availability. Phosphorus and N deficiency also limited the plasticity of foliar photosynthetic characteristics. There was a moderate increase in needle photosynthetic capacity of up to 1.6-fold from the bottom to the top of the canopy at the nutrient-rich site, but net assimilation rates were essentially independent of canopy position at the nutrient-poor site. Stomatal constraints on photosynthesis were similar between the sites, indicating that photosynthetic acclimation was curtailed at the biochemical level. We conclude that the foliar capacity for morphological and physiological acclimation to high light significantly decreases with decreasing nutrient availability in P. sylvestris, and that both N and P availability are potentially important determinants of foliar carbon gain capacities.

Chlorophyll↗

Leaf area dynamics in a closed poplar plantation under free-air carbon dioxide enrichment.

Three Populus genotypes (P. alba L. (Clone 2AS-11), P. nigra L. (Clone Jean Pourtet) and P. x euramericana (Clone I-214)) growing in a managed, high-density forest plantation were exposed to free-air CO(2) enrichment (FACE) at CO(2) concentrations expected to occur in the future (550 ppm). Leaf area index (optical LAI), measured with a fish-eye-type plant canopy analyzer, was not significantly affected by FACE after canopy closure in the second growing season. However, when stands of similar size were compared, optical LAI and number of main stem leaves were reduced by FACE. Allometric relationships, which were established to scale-up leaf area to the stand level (allometric LAI), did not differ between the FACE and control plots. Allometric LAI increased in response to FACE, as a result of increased tree dimensions and increased individual leaf size. We postulate that, although FACE increased allometric LAI, FACE had no effect on optical LAI after canopy closure, because FACE caused increased shading and competition resulting in enhanced leaf fall or leaf turnover. Specific leaf area (SLA) was unaffected by FACE. Allometric relationships and relative responses to atmospheric CO(2) enrichment were genotype-dependent.

Carbon Dioxide↗

Enhanced transpiration in response to wind effects at the edge of a blue gum (Eucalyptus globulus) plantation.

In Australia, tree planting has been widely promoted to alleviate dryland salinity and one proposed planting configuration is that of strategically placed interception belts. We conducted an experiment to determine the effect of tree position in a belt on transpiration rate. We also assessed how much the effect of tree position can be explained by advection and environmental conditions. Daily transpiration rates were determined by the heat pulse velocity technique for four edge and 12 inner trees in a 7-year-old Tasmanian blue gum (Eucalyptus globulus) plantation in South Australia. Various climatic variables were logged automatically at one edge of the plantation. The relationship between daily sap flow and sapwood area was strongly linear for the edge trees (r2 = 0.97), but only moderately correlated for the inner trees (r2 = 0.46), suggesting an edge effect. For all trees, sap flow normalized to sapwood area (Qs) increased with potential evaporation (PE) initially and then became independent as PE increased further. There was a fairly close correlation between transpiration of the edge and inner trees, implying that water availability was partially responsible for the difference between inner and edge trees. However, the ratio of edge tree to inner tree transpiration differed from unity, indicating differences in canopy conductance, which were estimated by an inverse form of the Penman-Monteith equation. When canopy conductances were less than a critical value, there was a strong linear relationship between Qs of the edge and inner trees. When canopy conductances of the edge trees were greater than the critical value, the slope of the linear relationship was steeper, indicating greater transpiration of the edge trees compared with the inner trees. This was interpreted as evidence for enhancement of transpiration of the edge trees by advection of wind energy.

Eucalyptus↗

Age-related variations in delta(13)C of ecosystem respiration across a coniferous forest chronosequence in the Pacific Northwest.

We tested the hypothesis that forest age influences the carbon isotope ratio (delta13C) of carbon reservoirs and CO2 at local and regional levels. Carbon isotope ratios of ecosystem respiration (delta13C(R)), soil respiration (delta13C(R-soil)), bulk needle tissue (delta13C(P)) and soil organic carbon (delta(13)C(SOC)) were measured in > 450-, 40- and 20-year-old temperate, mixed coniferous forests in southern Washington, USA. Values of delta13C(R), delta13C(R-soil), delta13C(P) and delta13C(SOC) showed consistent enrichment with increasing stand age. Between the youngest and oldest forests there was an approximately 1 per thousand enrichment in delta13C(P) (at similar canopy levels), delta13C(SOC) (throughout the soil column), delta13C(R-soil) (during the wet season) and delta13C(R) (during the dry season). Mean values of delta13C(R) were -25.9, -26.5 and -27.0 per thousand for the 450-, 40- and 20-year-old forests, respectively. Both delta13C(R-soil) and the difference between delta13C(R) and delta13C(R-soil) were more 13C enriched in older forests than in young forest: delta13C(R) - delta13C(R-soil) = 2.3, 1.1 and 0.5 per thousand for the 450-, 40- and 20-year-old forests, respectively. Values of delta(13)C(P) were proportionally more depleted relative to delta13C(R): delta13C(R) - delta13C(P) = 0.5, 2.2 and 2.5 per thousand for the 450-, 40- and 20-year-old forests, respectively. Values of delta13C(P) were most 13C-enriched at the top of the canopy and in the oldest forest regardless of season (overall values were -26.9, -28.7 and -29.4 per thousand for the 450-, 40- and 20-year-old forests, respectively). Values of delta13C(SOC) from shallow soil depths were similar to delta13C(P) values of upper- and mid-canopy needles. All delta13C data are consistent with the hypothesis that a decrease in stomatal conductance associated with decreased hydraulic conductance leads to increased CO2 diffusional limitations in older coniferous trees. The strong associations between delta13C(P) in needles with delta13C(R) and delta13C(R-soil) at the forest level suggest that 13C observations scale between leaf and ecosystem levels.

Carbon Dioxide↗

Tracing carbon uptake from a natural CO2 spring into tree rings: an isotope approach.

We analyzed 14C, 13C and 18O isotope variations over a 50-year period in tree rings of Quercus ilex L. trees growing at a natural CO2 spring in a Mediterranean ecosystem. We compared trees from two sites, one with high and one with low exposure to CO2 from the spring. The spring CO2 is free of 14C. Thus, this carbon can be traced in the wood, and the amount originating from the spring calculated. The amount decreased over time, from about 40% in 1950 to 15% at present for the site near the spring, indicating a potential difficulty in the use of natural CO2 springs for elevated CO2 research. The reason for the decrease may be decreasing emission from the spring or changes in stand structure, e.g., growth of the canopy into regions with lower concentrations. We used the 14C-calculated CO2 concentration in the canopy to determine the 13C discrimination of the plants growing under elevated CO2 by calculating the effective canopy air 13C/12C isotopic composition. The trees near the spring showed a 2.5 per thousand larger 13C discrimination than the more distant trees at the beginning of the investigated period, i.e., for the young trees, but this difference gradually disappeared. Higher discrimination under elevated CO2 indicated reduced photosynthetic capacity or increased stomatal conductance. The latter assumption is unlikely as inferred from the 18O data, which were insensitive to CO2 concentration. In conclusion, we found evidence for a downward adjustment of photosynthesis under elevated CO2 in Q. ilex in this dry, nutrient-poor environment.

Carbon Dioxide↗

A method for 3D reconstruction of tree crown volume from photographs: assessment with 3D-digitized plants.

We developed a method for reconstructing tree crown volume from a set of eight photographs taken from the N, S, E, W, NE, NW, SE and SW. This photographic method of reconstruction includes three steps. First, canopy height and diameter are estimated from each image from the location of the topmost, rightmost and leftmost vegetated pixel; second, a rectangular bounding box around the tree is constructed from canopy dimensions derived in Step 1, and the bounding box is divided into an array of voxels; and third, each tree image is divided into a set of picture zones. The gap fraction of each picture zone is calculated from image processing. A vegetated picture zone corresponds to a gap fraction of less than 1. Each picture zone corresponds to a beam direction from the camera to the target tree, the equation of which is computed from the zone location on the picture and the camera parameters. For each vegetated picture zone, the ray-box intersection algorithm (Glassner 1989) is used to compute the sequence of voxels intersected by the beam. After processing all vegetated zones, voxels that have not been intersected by any beam are presumed to be empty and are removed from the bounding box. The estimation of crown volume can be refined by combining several photographs from different view angles. The method has been implemented in a software package called Tree Analyzer written in C++. The photographic method was tested with three-dimensional (3D) digitized plants of walnut, peach, mango and olive. The 3D-digitized plants were used to estimate crown volume directly and generate virtual perspective photographs with POV-Ray Version 3.5 (Persistence of Vision Development Team). The locations and view angles of the camera were manually controlled by input parameters. Good agreement between measured data and values inferred from the photographic method were found for canopy height, diameter and volume. The effects of voxel size, size of picture zoning, location of camera and number of pictures were also examined.

Imaging, Three-Dimensional↗

Stomatal conductance, transpiration and sap flow of tropical montane rain forest trees in the southern Ecuadorian Andes.

We investigated tree water relations in a lower tropical montane rain forest at 1950-1975 m a.s.l. in southern Ecuador. During two field campaigns, sap flow measurements (Granier-type) were carried out on 16 trees (14 species) differing in size and position within the forest stand. Stomatal conductance (g(s)) and leaf transpiration (E(l)) were measured on five canopy trees and 10 understory plants. Atmospheric coupling of stomatal transpiration was good (decoupling coefficient Omega = 0.25-0.43), but the response of g(s) and E(l) to the atmospheric environment appeared to be weak as a result of the offsetting effects of vapor pressure deficit (VPD) and photosynthetic photon flux (PPF) on g(s). In contrast, sap flow (F) followed these atmospheric parameters more precisely. Daily F depended chiefly on PPF sums, whereas on short time scales, VPD impeded transpiration when it exceeded a value of 1-1.2 kPa. This indicates an upper limit to transpiration in the investigated trees, even when soil water supply was not limiting. Mean g(s) was 165 mmol m(-2) s(-1) for the canopy trees and about 90 mmol m(-2) s(-1) for the understory species, but leaf-to-leaf as well as tree-to-tree variation was large. Considering whole-plant water use, variation in the daily course of F was more pronounced among trees differing in size and crown status than among species. Daily F increased sharply with stem diameter and tree height, and ranged between 80 and 120 kg day(-1) for dominant canopy trees, but was typically well below 10 kg day(-1) for intermediate and suppressed trees of the forest interior.

Ecuador↗

Calibration and assessment of seasonal changes in leaf area index of a tropical dry forest in different stages of succession.

A simple measure of the amount of foliage present in a forest is leaf area index (LAI; the amount of foliage per unit ground surface area), which can be determined by optical estimation (gap fraction method) with an instrument such as the Li-Cor LAI-2000 Plant Canopy Analyzer. However, optical instruments such as the LAI-2000 cannot directly differentiate between foliage and woody components of the canopy. Studies investigating LAI and its calibration (extracting foliar LAI from optical estimates) in tropical forests are rare. We calibrated optical estimates of LAI from the LAI-2000 with leaf litter data for a tropical dry forest. We also developed a robust method for determining LAI from leaf litter data in a tropical dry forest environment. We found that, depending on the successional stage of the canopy and the season, the LAI-2000 may underestimate LAI by 17% to over 40%. In the dry season, the instrument overestimated LAI by the contribution of the woody area index. Examination of the seasonal variation in LAI for three successional stages in a tropical dry forest indicated differences in timing of leaf fall according to successional stage and functional group (i.e., lianas and trees). We conclude that when calculating LAI from optical estimates, it is necessary to account for the differences between values obtained from optical and semi-direct techniques. In addition, to calculate LAI from litter collected in traps, specific leaf area must be calculated for each species rather than from a mean value for multiple species.

Calibration↗

Variability in net ecosystem exchange from hourly to inter-annual time scales at adjacent pine and hardwood forests: a wavelet analysis.

Orthonormal wavelet transformation (OWT) is a computationally efficient technique for quantifying underlying frequencies in nonstationary and gap-infested time series, such as eddy-covariance-measured net ecosystem exchange of CO2 (NEE). We employed OWT to analyze the frequency characteristics of synchronously measured and modeled NEE at adjacent pine (PP) and hardwood (HW) ecosystems. Wavelet cospectral analysis showed that NEE at PP was more correlated to light and vapor pressure deficit at the daily time scale, and NEE at HW was more correlated to leaf area index (LAI) and temperature, especially soil temperature, at seasonal time scales. Models were required to disentangle the impacts of environmental drivers on the components of NEE, ecosystem carbon assimilation (Ac) and ecosystem respiration (RE). Sensitivity analyses revealed that using air temperature rather than soil temperature in RE models improved the modeled wavelet spectral frequency response on time scales longer than 1 day at both ecosystems. Including LAI improved RE model fit on seasonal time scales at HW, and incorporating parameter variability improved the RE model response at annual time scales at both ecosystems. Resolving variability in canopy conductance, rather than leaf-internal CO2, was more important for modeling Ac at both ecosystems. The PP ecosystem was more sensitive to hydrologic variables that regulate canopy conductance: vapor pressure deficit on weekly time scales and soil moisture on seasonal to interannual time scales. The HW ecosystem was sensitive to water limitation on weekly time scales. A combination of intrinsic drought sensitivity and non-conservative water use at PP was the basis for this response. At both ecosystems, incorporating variability in LAI was required for an accurate spectral representation of modeled NEE. However, nonlinearities imposed by canopy light attenuation were of little importance to spectral fit. The OWT revealed similarities and differences in the scale-wise control of NEE by vegetation with implications for model simplification and improvement.

Carbon Dioxide↗

A photographic gap fraction method for estimating leaf area of isolated trees: Assessment with 3D digitized plants.

A method for computing leaf area of isolated trees from perspective photographs was developed. The method is based on gap fraction inversion. Photographs are discretized into picture zones where gap fraction is computed from image processing. Canopy volume and leaf area density associated with each picture zone are computed from geometrical considerations and inversion of gap fraction equations. Total leaf area and the vertical profile of leaf area are computed from the product of associated volume and its density. The method has been implemented in software called Tree Analyser, written in C++. The method has been tested by comparison with direct estimation of leaf area of three-dimensional (3D) digitized trees of walnut, peach, mango, olive and rubber. Estimated leaf area was sensitive to picture discretization, individual leaf size and leaf inclination distribution. Optimal size of picture discretization was 17 times projected leaf size. Total leaf area was estimated by using a set of eight photographs taken around the tree in the main horizontal directions: deviation ranged from -11% in peach tree to +5% in rubber tree. The method allows fast and nondestructive monitoring of leaf area of individual tree canopies. The next version of the method will include the estimation of 3D leaf area distribution within the canopy.

Models, Biological↗

Turbulent transfer in a deciduous forest.

Carbon dioxide, water vapor and other passive scalars are physically transferred between a plant canopy and the atmosphere by turbulence. Intense and intermittent sweep and ejection events transfer most of the mass. Although the capacity for turbulence to transfer material is high, mass transfer is coupled to the diffusive source or sink strength of the foliage and soil and is ultimately limited to a minimum level set by the supply of material, or the demand for it. The diffusive source/sink strength of material leaving or entering leaves and the soil is a function of many physical, biological and chemical attributes and processes. These attributes and processes include the amount and distribution of foliage, the leaf boundary layer and surface resistances, the turbulence and radiative regimes in the canopy, biochemical and photochemical reactions and the scalar concentration field within and above the canopy and inside leaves and the soil. Here we discuss how these factors contribute to turbulent transfer in a deciduous forest.

Journal Article↗

Plant area index in Eucalyptus globulus plantations determined indirectly by a light interception method.

Plant area index (PAI) was estimated in 3-year-old plantations of Eucalyptus globulus Labill. from (i) allometric relationships obtained from biomass harvesting, and (ii) canopy transmittance measurements. Diffuse light canopy transmittance was measured with a multi-cone sensor consisting of five inverted cones to scan a hemisphere. Plant area index was calculated from transmittance measurements by the inversion method assuming an ellipsoidal angle distribution of uniformly distributed canopy elements, or by application of the Beer-Lambert Law using an empirically determined extinction coefficient (k) for diffuse light of 0.5. Plant area index values of the experimental plantations varied as a consequence of diffferent water and mineral nutrient application rates. The PAI estimated from biomass harvesting ranged from 1.61 to 4.04. The estimates of PAI obtained by the ellipsoidal angle distribution method were significantly different from those based on the allometric relationships or from transmittance measurements assuming a value of k for diffuse light of 0.5.

Journal Article↗

Models of water flux through forest stands: critical leaf and stand parameters.

The basis for estimating water flux through forest stands is described using models. By means of the Penman-Monteith equation, transpiration is described in relation to the degree of coupling between the canopy and the surrounding air. Models of canopy and aerodynamic conductance are available, but more mechanistic models of stomatal conductance, and further development of turbulence theory, are needed along with improved estimates of leaf area index, leaf area distribution, and seasonal dynamics. Three models are presented to show current capabilities in estimating water uptake and flux through tree components, including the effects of capacitance. Defining conductance to water movement through tree components in terms of the properties of the pathway (sapwood area, sapwood relative conductivity, leaf area) is a useful functional approach that can be tested for a range of species at different sites. Further research is required to relate water conducting properties of tree components to architectural arrangement, especially for roots, and improved methods for measuring water potentials and partial flows at different points within the system are necessary. The role of water potential and the significance of its variability within a canopy are discussed. Relating growth processes to an integral of water potential (an accumulated product of water potential and time) is recommended. The need for scaling and integrating information about processes from one level to higher levels is recognized. The importance of understanding the roles that temporal, spatial, and developmental levels have on the ability to scale or integrate individual leaf measurements of, for example, leaf conductance to the stand level is emphasized.

Journal Article↗

Acclimation of rice photosynthesis to irradiance under field conditions.

Acclimation to irradiance was measured in terms of light-saturated photosynthetic carbon assimilation rates (P(max)), Rubisco, and pigment content in mature field-grown rice (Oryza sativa) plants in tropical conditions. Measurements were made at different positions within the canopy alongside irradiance and daylight spectra. These data were compared with a second experiment in which acclimation to irradiance was assessed in uppermost leaves within whole-plant shading regimes (10% low light [LL], 40% medium light [ML], and 100% high light [HL] of full natural sunlight). Two varieties, japonica (tropical; new plant type [NPT]) and indica (IR72) were compared. Values for Rubisco amount, chlorophyll a/b, and P(max) all declined from the top to the base of the canopy. In the artificial shading experiment, acclimation of P(max) (measured at 350 microL L(-1) CO(2)) occurred between LL and ML for IR72 with no difference observed between ML and HL. The Rubisco amount increased between ML and HL in IR72. A different pattern was seen for NPT with higher P(max) (measured at 350 microL L(-1) CO(2)) at LL than IR72 and some acclimation of this parameter between ML and HL. Rubisco levels were higher in NPT than IR72 contrasting with P(max). Comparison of data from both experiments suggests a leaf aging effect between the uppermost two leaf positions, which was not a result of irradiance acclimation. Results are discussed in terms of: (a) acclimation of photosynthesis and radiation use efficiency at high irradiance in rice, and (b) factors controlling photosynthetic rates of leaves within the canopy.

Acclimatization↗