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Seasonal changes in canopy photosynthesis and respiration, and partitioning of photosynthate, in rice (Oryza sativa L.) grown under free-air CO2 enrichment.

An increase in atmospheric CO(2) concentration ( [CO(2)]) is generally expected to enhance photosynthesis and biomass. Rice plants (Oryza sativa L.) were grown in ambient CO(2) (AMB) or free-air CO(2)-enrichment (FACE), in which the target [CO(2)] was 200 micromol mol(-1) above AMB. (13)CO(2) was fed to the plants at different stages so we could examine the partitioning of photosynthates. Furthermore, canopy photosynthesis and respiration were measured at those stages. The ratio of (13)C content in the whole plant to the amount of fixed (13)C under FACE was similar to that under AMB at the vegetative stage. However, the ratio under FACE was greater than the ratio under AMB at the grain-filling stage. At the vegetative stage, plants grown under FACE had a larger biomass than those grown under AMB owing to enhancement of canopy photosynthesis by the increased [CO(2)]. On the other hand, at the grain-filling stage, CO(2) enrichment promoted the partitioning of photosynthate to ears, and plants grown under FACE had a greater weight of ears. However, enhancement of ear weight by CO(2) enrichment was not as great as that of biomass at the vegetative stage. Plants grown under FACE did not necessarily show higher canopy photosynthetic rates at the grain-filling stage. Therefore, we concluded that the ear weight did not increase as much as biomass at the vegetative stage owing to a loss of the advantage in CO(2) gain during the grain-filling period.

Air↗

Diurnal patterns of leaf photosynthesis, conductance and water potential at the top of a lowland rain forest canopy in Cameroon: measurements from the Radeau des Cimes.

Diurnal patterns of leaf conductance, net photosynthesis and water potential of five tree species were measured at the top of the canopy in a tropical lowland rain forest in southwestern Cameroon. Access to the 40 m canopy was by a large canopy-supported raft, the Radeau des Cimes. The measurements were made under ambient conditions, but the raft altered the local energy balance at times, resulting in elevated leaf temperatures. Leaf water potential was equal to or greater than the gravitational potential at 40 m in the early morning, falling to values as low as -3.0 MPa near midday. Net photosynthesis and conductance were typically highest during midmorning, with values of about 10-12 micro mol CO(2) m(-2) s(-1) and 0.2-0.3 mol H(2)O m(-2) s(-1), respectively. Leaf conductance and net photosynthesis commonly declined through midday with occasional recovery late in the day. Photosynthesis was negatively related to leaf temperature above midday air temperature maxima. These patterns were similar to those observed in other seasonally droughted evergreen communities, such as Mediterranean-climate shrubs, and indicate that environmental factors may cause stomatal closure and limit photosynthesis in tropical rain forests during the midday period.

Journal Article↗

Effects of light, temperature and canopy position on net photosynthesis and isoprene emission from sweetgum (Liquidambar styraciflua) leaves.

In June 1993, net photosynthetic rates, stomatal conductance and isoprene emission rates of sweetgum leaves (Liquidambar styraciflua L.) were measured at the top of the forest canopy (sun leaves) and within the canopy at a height of 8-10 m above ground level (shade leaves). Large differences in net photosynthetic rates and stomatal conductance were found between sun and shade leaves. Mean rates of isoprene emission, expressed on a leaf area basis, were significantly lower in shade leaves than in sun leaves (4.1 versus 17.1 nmol m(-2) s(-1)); however, because specific leaf area of sun leaves was lower than that of shade leaves (0.0121 versus 0.0334 m(2) g(-1)), the difference between sun and shade leaves was less, though still significant, when isoprene emissions were expressed on a dry mass basis (45.5 versus 29.0 micro g C g(-1) h(-1)). Saturation of both net photosynthesis and isoprene emission occurred at lower PPFDs in shade leaves than in sun leaves. The effect of leaf temperature on isoprene emissions also differed between sun and shade leaves. Sun leaves lost a significantly greater percentage of fixed carbon as isoprene than shade leaves. The leaf-level physiological measurements were used to derive parameters for a canopy-level isoprene flux model. The importance of incorporating differences between sun- and shade-leaf properties into existing models is discussed.

Journal Article↗

Diurnal changes in leaf gas exchange characteristics in the uppermost canopy of a rain forest tree, Dryobalanops aromatica Gaertn. f.

Dryobalanops aromatica Gaertn. f. is a major tropical canopy species in lowland tropical rain forests in Peninsular Malaysia. Diurnal changes in net photosynthetic rate (A) and stomatal conductance to water vapor (g(s)) were measured in fully expanded young and old leaves in the uppermost canopy (35 m above ground). Maximum A was 12 and 10 micro mol m(-2) s(-1) in young and old leaves, respectively; however, because of large variation in A among leaves, mean maximum A in young and old leaves was only 6.6 and 5.5 micro mol m(-2) s(-1), respectively. Both g(s) and A declined in young leaves when T(leaf) exceeded 34 degrees C and leaf-to-air vapor pressure deficit (DeltaW) exceeded 0.025, whereas in old leaves, g(s) and A did not start to decline until T(leaf) and DeltaW exceeded 36 degrees C and 0.035, respectively. Under saturating light conditions, A was linearly related to g(s). The coefficient of variation (CV) for the difference between the CO(2) concentrations of ambient air and the leaf intercellular air space (C(a) - C(i)) was smaller than the CV for A or g(s), suggesting that maximum g(s) was mainly controlled by mesophyll assimilation (A/C(i)). Minimum C(i)/C(a) ratios were relatively high (0.72-0.73), indicating a small drought-induced stomatal limitation to A and non-conservative water use in the uppermost canopy leaves.

Journal Article↗

Crown architecture of Abies balsamea from four canopy positions.

Data collected from four distinct canopy positions from each of 39 Abies balsamea (L.) Miller trees were used to construct models to describe the cumulative leaf area distribution within the crown and to predict the needle mass of individual branches, the average branch angle, branch diameter, branch length, and crown radius per whorl, and the average number of living branches per whorl. We tested the hypotheses that regression models are equal among canopy positions and that a model to predict branch needle mass is valid at the northern and southern extremes of the central climatic zone of Maine. Canopy position had an effect on the models constructed to predict needle mass, branch angle, branch diameter, branch length, crown radius, and the number of living branches per whorl. However, compared with an expanded model that incorporated parameters calculated for each crown class, there was only a small loss in model precision when a general model constructed from data pooled from all crown classes was used to predict needle mass, branch angle, and branch diameter. Regression equations unique to each crown class were needed to predict crown shape and leaf area distribution in the crown satisfactorily. Our branch needle mass model, which was constructed from data collected at the southern extreme of the central climatic zone of Maine, consistently underestimated needle branch mass when applied to the northern extreme of the central climatic zone.

Journal Article↗

Transpiration rates and canopy conductance of Pinus radiata growing with different pasture understories in agroforestry systems.

We measured tree transpiration and canopy conductance in Pinus radiata D. Don at two low rainfall sites of differing soil fertility in Canterbury, New Zealand. At the more fertile Lincoln site, we also assessed the effects of two common pasture grasses on tree transpiration and canopy conductance. At the less fertile Eyrewell Forest site, the effect of no understory, and the effects of irrigation in combination with mixtures of grass or legume species were determined. Tree xylem sap flux (F(d)') was measured by the heat pulse method. Total canopy conductance to diffusion of water vapor (G(t)) was calculated by inverting a simplified Penman-Monteith model. The different treatment effects were modeled by the simple decaying exponential relationship G(t) = G(tmax)e((-bD)), where D = air saturation deficit. At the Lincoln site, trees with an understory of cocksfoot had lower F(d)' and G(tmax) than trees with an understory of ryegrass, although the sensitivity of G(t) to increasing D (i.e., the value of b) did not differ between treatments. At the Eyrewell site, irrigation only increased F(d)' in the absence of an understory, whereas the presence of understory vegetation, or lack of irrigation, or both, significantly reduced G(tmax) and increased b. We conclude that the selection of understory species is critical in designing successful agroforestry systems for low rainfall areas.

Journal Article↗

Morphological and physiological adjustment to N and P fertilization in nutrient-limited Metrosideros polymorpha canopy trees in Hawaii.

Leaf-level studies of Metrosideros polymorpha Gaud. (Myrtaceae) canopy trees at both ends of a substrate age gradient in the Hawaiian Islands pointed to differential patterns of adjustment to both nutrient limitation and removal of this limitation by long-term (8-14 years) nitrogen (N), phosphorus (P) and N + P fertilizations. The two study sites were located at the same elevation, had similar annual precipitation, and supported forests dominated by M. polymorpha, but differed in the age of the underlying volcanic substrate, and in soil nutrient availability, with relatively low N at the young site (300 years, Thurston, Hawaii) and relatively low P at the oldest site (4,100,000 years, Kokee, Kauai). Within each site, responses to N and P fertilization were similar, regardless of the difference in soil N and P availability between sites. At the young substrate site, nutrient addition led to a larger mean leaf size (about 7.4 versus 4.8 cm2), resulting in a larger canopy leaf surface area. Differences in foliar N and P content, chlorophyll concentrations and carboxylation capacity between the fertilized and control plots were small. At the old substrate site, nutrient addition led to an increase in photosynthetic rate per unit leaf surface area from 4.5 to 7.6 micromol m(-2) s(-1), without a concomitant change in leaf size. At this site, leaves had substantially greater nutrient concentrations, chlorophyll content and carboxylation capacity in the fertilized plots than in the control plots. These contrasting acclimation responses to fertilization at the young and old sites led to significant increases in total carbon gain of M. polymorpha canopy trees at both sites. At the young substrate site, acclimation to fertilization was morphological, resulting in larger leaves, whereas at the old substrate site, physiological acclimation resulted in higher leaf carboxylation capacity and chlorophyll content.

Hawaii↗

Sensitivity of photosynthetic electron transport to photoinhibition in a temperate deciduous forest canopy: Photosystem II center openness, non-radiative energy dissipation and excess irradiance under field conditions.

We used chlorophyll fluorescence techniques to investigate responses of Photosystem II (PSII) quantum yield to light availability in the short term (quantum flux density integrated over the measurement day, Qd) and in the long term (Qd averaged over the season, Qs) in a mixed deciduous forest comprising shade-tolerant and water-stress-sensitive Tilia cordata Mill. in the lower canopy and shade-intolerant and water-stress-resistant Populus tremula L. in the upper canopy. In both species, intrinsic efficiency of PSII in the dark-adapted state (Fv/Fm) was lower during the day than during the night, and the difference in Fv/Fm between day and night increased with increasing Qs. Although the capacity for photosynthetic electron transport increased with increasing Qs in both species, maximum quantum efficiency of PSII in the light-adapted state (alpha) decreased with increasing Qs. At a common Qs, alpha was lower in T. cordata than in P. tremula primarily because of a higher fraction of closed PSII centers, and to a smaller extent because of limited, non-radiative, excitation energy dissipation in the pigment bed in T. cordata. Across both species, photochemical quenching (qP), which measures the openness of PSII centers, varied more than fivefold, but the efficiency of excitation energy capture by open PSII centers (Fv'/Fm'), which is an estimate of non-radiative excitation energy dissipation in PSII antennae, varied by only 50%. Chlorophyll turnover rates increased with increasing irradiance, especially in T. cordata, possibly because of increased photodestruction. Diurnal measurements of PSII quantum yields (PhiPSII) indicated that, under similar environmental conditions, PhiPSII was always lower in the afternoon than in the morning, and the fraction of daily integrated photosynthetic electron transport lost because of diurnal declines in PhiPSII (Delta) increased with increasing Qd. At a common Qd, mean daily PSII center reduction state, the fraction of light in excess (1 - fractions of light used in photochemistry and dissipated as heat) and Delta were higher in T. cordata than in P. tremula. This was attributed to greater stomatal closure during the day, which led to a greater reduction in the requirement for assimilative electron flow in T. cordata. Across both species, Delta scaled negatively with the fraction of light utilized photochemically, demonstrating the leading role of PSII center openness in maintaining high PSII efficiency. Because photosynthesis (A) at current ambient carbon dioxide concentration is limited by CO2 availability in high light and mainly by photosynthetic electron transport rates in low light, overall daily down-regulation of PhiPSII primarily influences A in low light. Given that foliar water stress scales positively with Qs in both species, we conclude that the inverse patterns of variation in water and light availabilities in the canopy result in a greater decline in A than is predicted by decreases in stomatal conductance alone.

Chlorophyll↗

Leaf morphology and photosynthetic adjustments among deciduous broad-leaved trees within the vertical canopy profile.

Photosynthetic acclimation of deciduous broad-leaved tree species was studied along a vertical gradient within the canopy of a multi-species deciduous forest in northern Japan. We investigated variations in (1) local light regime and CO2 concentration ([CO2]), and (2) morphological (area, thickness and area per mass), biochemical (nitrogen and chlorophyll concentrations) and physiological (light-saturated photosynthetic rate) attributes of leaves of seven major species on three occasions (June, August and October). We studied early successional species, alder (Alnus hirsuta (Spach) Rupr.) and birch (Betula platyphylla var. japonica (Miq.) Hara); gap phase species, walnut (Juglans ailanthifolia Carrière) and ash (Fraxinus mandshurica var. japonica Rupr.); mid-successional species, basswood (Tilia japonica (Miq.) Simonk.) and elm (Ulmus davidiana var. japonica (Rehd.) Nakai); and the late-successional species, maple (Acer mono Bunge). All but maple initiated leaf unfolding from the lower part of the crown. The [CO2] within the vertical profile ranged from 320-350 ppm in the upper canopy to 405-560 ppm near the ground. The lowest and highest ambient [CO2] occurred during the day and during the night, respectively. This trend was observed consistently during the summer, but not when trees were leafless. Chlorophyll concentration was positively related to maximum photosynthetic rate within, but not among, species. Leaf senescence started from the inner part of the crown in alder and birch, but started either in the outer or top portion of the canopy of ash, basswood and maple. Chlorophyll (Chl) to nitrogen ratio in leaves increased with decreasing photon flux density. However, Chl b concentration in all species remained stable until the beginning of leaf senescence. Maximum photosynthetic rates observed in sun leaves of early successional species, gap phase or mid-successional species, and late successional species were 12.5-14.8 micromol m(-2) s(-1), 4.1-7.8 micromol m(-2) s(-1) and 3.1 micromol m(-2) s(-1), respectively.

Carbon Dioxide↗

Gas exchange characteristics of a Canarian laurel forest tree species (Laurus azorica) in relation to environmental conditions and leaf canopy position.

Diurnal courses of gas exchange were measured over a 1-year period in fully expanded current-year leaves in the upper (sun-exposed, 18 m above ground) and the lower (shaded, 12 m above ground) canopy of Laurus azorica (Seub.) Franco, a major canopy species of the Canarian laurel forest in Tenerife, Canary Islands, Spain. Laurus azorica exhibited high leaf plasticity in gas exchange characteristics, with a maximum carbon assimilation rate (Amax) of shade leaves about 50% that of sun leaves. This difference reflects the high leaf area index (LAI) of the stand and the correspondingly sharp light attenuation with increasing canopy depth. In sun leaves, Amax peaked at about 11 micromol m-2 s-1 and maximum transpiration (E) was about 8 mmol m-2 s-1, which corresponded with a maximum stomatal conductance (gs) of about 650 mmol m-2 s-1. Mean maximum instantaneous water-use efficiency (WUE) was 1.5 mmol mol-1 and the mean maximum A/gs was 20-35 micromol mol-1. Mean minimum internal CO2 concentration (Ci) was 225 micromol mol-1. Although high air vapor pressure deficit (VPD) caused a small decrease in gs, it remained high enough to maintain relatively high A and E. These gas exchange characteristics indicate a non-conservative use of water, which is appropriate for a species subject to droughts that are mild or of short duration. In this respect, Laurus azorica differs from its congener, L. nobilis L., of the Mediterranean region and other shrubs growing in Mediterranean-type climates in California and Chile that have to withstand more severe or more prolonged droughts.

Atlantic Islands↗

Interannual consistency in canopy stomatal conductance control of leaf water potential across seven tree species.

We investigated interannual variability of canopy transpiration per unit ground area (E (C)) and per unit leaf area (E (L)) across seven tree species in northern Wisconsin over two years. These species have previously been shown to be sufficient to upscale stand-level transpiration to the landscape level during one growing season. Our objective was to test whether a simple plant hydraulic model could capture interannual variation in transpiration. Three species, wetland balsam fir (Abies balsamea (L.) Mill), basswood (Tilia Americana L.) and speckled alder (Alnus rugosa (DuRoi) Spreng), had no change in E (C) or E (L) between 2000 and 2001. Red pine (Pinus resinosa Ait) had a 57 and 19% increase in E (C) and E (L), respectively, and sugar maple (Acer saccharum Marsh) had an 83 and 41% increase in E (C) and E (L), respectively, from 2000 to 2001. Quaking aspen (Populus tremuloides Michx) had a 50 and 21% decrease in E (C) and E (L), respectively, from 2000 to 2001 in response to complete defoliation by forest tent caterpillar (Malascoma distria Hüber) and subsequent lower total leaf area index of the reflushed foliage. White cedar (Thuja occidentalis L.) had a 20% decrease in both E (C) and E (L) caused by lowered surface water in wetlands in 2001 because of lower precipitation and wetland flow management. Upland A. balsamea increased E (L) and E (C) by 55 and 53%, respectively, as a result of release from light competition of the defoliated, overstory P. tremuloides. We hypothesized that regardless of different drivers of interannual variability in E (C) and E (L), minimum leaf water potential would be regulated at the same value. Minimum midday water potentials were consistent over the two years within each of the seven species despite large changes in transpiration between years. This regulation was independently verified by the exponential saturation between daily E (C) and vapor pressure deficit (D) and the tradeoff between a reference canopy stomatal conductance (G (S)) and the sensitivity of G (S) to D, indicating that trees with high G (S) must decrease G (S) in response to atmospheric drought faster than trees with low G (S). Our results show that models of forest canopy transpiration can be simplified by incorporating G (S) regulation of minimum leaf water potential for isohydric species.

Plant Leaves↗

Performance of a canopy light interception model for conifer shoots, trees and stands.

A hierarchical approach was used to evaluate the behavior of a canopy light interception model using submodels for interception at levels of the shoot, crown and stand. Mean silhouette areas of individual shoots of lodgepole pine (Pinus contorta var. latifolia Engelm.) calculated assuming spherical shoot orientation agreed well with silhouette areas measured with a video projection system. A theoretical mean silhouette to total leaf area ratio (STAR) calculated using measured shoot characteristics and a model of shoot geometry agreed reasonably well with STAR values from measured shoot silhouette and leaf areas, although the range of calculated STAR values was slightly lower than that of the measured values. The shadow areas of tree crowns were underestimated for lodgepole pine; however, a model using regular shoot dispersion in the crown gave better agreement than a model based on random shoot dispersion. The underestimates may have been caused by underestimation either of leaf or branch area for the crowns or of the extinction coefficient. Two equations used to calculate leaf area from sapwood area measurements both underestimated leaf area. Accordingly, there are too many uncertainties to determine whether crown interception is best represented by a random or regular distribution of shoots. Mean canopy transmission in the Engelmann spruce (Picea engelmannii Parry ex Engelm.) stand was estimated very well by a random (Poisson) model of the spatial distribution of trees, whereas, in the lodgepole pine stand, a regular (hexagonal) model gave the best results. The Engelmann spruce stand had less regular tree spacing than the lodgepole pine stand. Taken as a whole, the weakest submodel for calculating canopy interception was the model for individual crown transmission, which underestimated crown shadow areas. However, the problem may be in leaf area estimates rather than in model performance.

Journal Article↗

Techniques of ozone monitoring in a mountain forest region: passive and continuous sampling, vertical and canopy profiles.

Ozone is the most harmful air pollutant for plant ecosystems in the Mediterranean and Alpine areas due to its biological and economic damage to crops and forests. In order to evaluate the relation between ozone exposure and vegetation injury under on-field conditions, suitable ozone monitoring techniques were investigated. In the framework of a 5-year research project aimed at ozone risk assessment on forests, both continuous analysers and passive samplers were employed during the summer seasons (1994-1998) in different sites of a wide mountain region (80 x 40 km2) on the southern slope of the European Alps. Continuous analysers allowed the recording of ozone hourly concentration means necessary both to calculate specific exposure indexes (such as AOT, SUM, W126) and to record daily time-courses. Passive samplers, even though supplied only weekly mean concentration values, made it possible to estimate the altitude concentration gradient useful to correct the altitude dependence of ozone concentrations to be inserted into exposure indexes. In-canopy ozone profiles were also determined by placing passive samplers at different heights inside the forest canopy. Vertical ozone soundings by means of tethered balloons (kytoons) allowed the measurement of the vertical concentration gradient above the forest canopy. They also revealed ozone reservoirs aloft and were useful to explain the ozone advection dynamic in mountain slopes where ground measurement proved to be inadequate. An intercomparison between passive (PASSAM, CH) and continuous measurements highlighted the necessity to accurately standardize all the exposure operations, particularly the pre- and postexposure conservation at cold temperature to avoid dye (DPE) activity. Advantages and disadvantages from each mentioned technique are discussed.

Air Pollutants↗

Phytochrome A Mediates the Promotion of Seed Germination by Very Low Fluences of Light and Canopy Shade Light in Arabidopsis.

Seeds of the wild type (WT) and of the phyA and phyB mutants of Arabidopsis thaliana were exposed to single red light (R)/far-red light (FR) pulses predicted to establish a series of calculated phytochrome photoequilibria (Pfr/P). WT and phyB seeds showed biphasic responses to Pfr/P. The first phase, i.e. the very-low-fluence response (VLFR), occurred below Pfr/P = 10-1%. The second phase, i.e. the low-fluence response, occurred above Pfr/P = 3%. The VLFR was similarly induced by either a FR pulse saturating photoconversion or a subsaturating R pulse predicted to establish the same Pfr/P. The VLFR was absent in phyA seeds, which showed a strong low-fluence response. In the field, even brief exposures to the very low fluences of canopy shade light (R/FR ratio < 0.05) promoted germination above dark controls in WT and phyB seeds but not in the phyA mutant. Seeds of the phyA mutant germinated normally under canopies providing higher R/FR ratios or under deep canopy shade light supplemented with R from light-emitting diodes. We propose that phytochrome A mediates VLFR of A. thaliana seeds.

Journal Article↗

Photosynthetic induction state of leaves in a soybean canopy in relation to light regulation of ribulose-1-5-bisphosphate carboxylase and stomatal conductance.

Photosynthetic induction state, stomatal conductance and light regulation of ribulose-1,5-bisphosphate carboxylase (rubisco) were examined for leaves in a mature, closed soybean (Glycine max) canopy (leaf area index approximately 5) with the objective to determine the extent to which these factors may be limiting the capacity to respond to light transients during sunflecks. When sampled along a vertical gradient, leaves near the bottom of the canopy had lower rubisco contents and chlorophyll a/b ratios as compared with upper leaves. Leaves sampled at midcanopy showed a wide variation in photosynthetic induction state (ratio of the photosynthetic rate achieved after 1 minute exposure to high light to the steady-state assimilation rate achieved after 20 minutes exposure). Both photosynthetic induction state and the initial rubisco activity varied in parallel with stomatal conductance. By contrast there was no correlation between total rubisco activity and stomatal conductance. The results indicate that induction state, as determined by the light regulation of both rubisco activity and stomatal conductance, is an important limitation to the ability of leaves in a soybean canopy to respond to light transients that occur during sunflecks.

Journal Article↗

Laser scanning reveals bryophyte canopy structure.

We evaluated laser scanning as a method to provide depth measurements for bryophyte canopies at fine spatial scales to derive surface roughness (Lr), a structural parameter. Depths to the first vertical canopy contact were measured on 5 x 5 cm2 areas of 27 bryophyte canopies using a contact probe, a commercial laser scanner and a scanner employing a laser diode striper (LED scanner). Laser scanning adequately distinguished structural types, but scanner configuration led to differences in the magnitude of Lr. LED scanning did not damage photosystem II function in three bryophyte species, Bazzania trilobata, Sphagnum girgensohnii and Pleurozium schreberi, as evidenced by no change in the chlorophyll fluorescence parameter FV/FM following LED scanning, but a decrease when subjected to high light. A previously published boundary layer conductance model was parameterized with surface roughness values determined using a laser scanner and compared with the results obtained with contact probe measures. The resulting parameters of the functional models did not differ significantly from each other.

Bryophyta↗

Canopy and environmental control of root dynamics in a long-term study of Concord grape.

Below-ground carbon allocation represents a substantial fraction of net photosynthesis in plants, yet timing of below-ground allocation and endogenous and exogenous factors controlling it are poorly understood. Minirhizotron techniques were used to examine root populations of Vitis labruscana Bailey cv. Concord under two levels of dormant-season canopy removal and irrigation. Root production, pigmentation, death and disappearance to a depth of 110 cm were determined over two wet and two dry years (1997-2000). There was continual root production and senescence, with peak root production rates occurring by midseason. Later in the season, when reproductive demands for carbon were highest and physical conditions limiting, few roots were produced, especially in dry years in nonirrigated vines. Root production under minimal canopy pruning was generally greater and occurred several weeks earlier than root production under heavy pruning, corresponding to earlier canopy development. Initial root production occurred in shallow soils, likely due to temperatures at shallow depths being warmer early in the season. Our study showed intricate relationships between internal carbon demands and environmental conditions regulating root allocation.

Agriculture↗

Large daily variation in 13C-enrichment of leaf-respired CO2 in two Quercus forest canopies.

The use of the 13C : 12C isotopic ratio (delta13C) of leaf-respired CO2 to trace carbon fluxes in plants and ecosystems is limited by little information on temporal variations in delta13C of leaf dark-respired CO2 (delta13Cr) under field conditions. Here, we explored variability in delta13Cr and its relationship to key respiratory substrates from collections of leaf dark-respired CO2, carbohydrate extractions and gas exchange measurements over 24-h periods in two Quercus canopies. Throughout both canopies, delta13Cr became progressively 13C-enriched during the photoperiod, by up to 7%, then 13C-depleted at night relative to the photoperiod. This cycle could not be reconciled with delta13C of soluble sugars (delta13Css), starch (delta13Cst), lipids (delta13Cl), cellulose (delta13Cc) or with calculated photosynthetic discrimination (Delta). However, photoperiod progressive enrichment in delta13Cr was correlated with cumulative carbon assimilation (r2 = 0.91). We concluded that there is considerable short-term variation in delta13Cr in forest canopies, that it is consistent with current hypotheses for 13C fractionation during leaf respiration, that leaf carbohydrates cannot be used as surrogates for delta13Cr, and that diel changes in leaf carbohydrate status could be used to predict changes in delta13Cr empirically.

Carbon Dioxide↗