Relationships between hydraulic architecture and leaf photosynthetic capacity in nitrogen-fertilized Eucalyptus grandis trees

Size: px
Start display at page:

Download "Relationships between hydraulic architecture and leaf photosynthetic capacity in nitrogen-fertilized Eucalyptus grandis trees"

Transcription

1 Tree Physiology 21, Heron Publishing Victoria, Canada Relationships between hydraulic architecture and leaf photosynthetic capacity in nitrogen-fertilized Eucalyptus grandis trees MICHAEL J. CLEARWATER 1,2 and FREDERICK C. MEINZER 1,3 1 Hawaii Agriculture Research Center, Aiea Heights Drive, Aiea, HI 96701, USA 2 Present address: Horticulture and Food Research Institute of New Zealand, Te Puke Research Center, RD2 Te Puke, New Zealand 3 Present address: USDA Forest Service, Forestry Sciences Laboratory, 3200 SW Jefferson Way, Corvallis, OR 97331, USA Received October 28, 2000 Summary We compared the effects of nitrogen fertilization on shoot hydraulic architecture and leaf photosynthetic properties of Eucalyptus grandis Hill ex Maiden trees in Hawaii. It was hypothesized that water transport capacity would adjust to nutrient availability, with leaf specific hydraulic conductivity (k l ) increasing in fertilized trees in coordination with higher photosynthetic capacity per unit leaf area. Trees were grown from seedlings in the field for 10 months at four rates of nitrogen (N) fertilization between 0 and 336 kg ha 1. Leaf water potentials, photosynthetic capacity and k l were measured before whole shoots were harvested to determine total growth, leaf area and sapwood density. Mean tree height increased from 4 to 5.3 m, stem basal area increased from 27 to 67 cm 2 and total leaf area increased from 15 to 40 m 2 between the lowest and highest rates of fertilizer addition. When trees were compared on the basis of leaf nitrogen per unit area (N area ), light-saturated rates of photosynthesis on an area and mass basis and the maximum rate of electron transport all increased from 50% to more than 100% as N area increased from 0.8 to 2.1 g m 2. Branch specific hydraulic conductivity (k s ) and k l increased with height in the crown. However, there was no change in branch k l or the ratio of leaf area to sapwood area of the whole shoot in response to fertilization, and k s and density of the sapwood were unrelated to leaf N area. In contrast to photosynthesis, stomatal conductance did not respond to fertilization, leading to decreased internal carbon dioxide partial pressure (p i /p a ) in fertilized plants and similar leaf water potentials in all plants. Consistent with the behavior of p i /p a, carbon isotope discrimination decreased by 2 with increasing leaf N area, supporting the conclusion that intrinsic water-use efficiency was enhanced by fertilization. Increased growth in response to fertilization involved adjustment at the leaf level rather than a change in the balance between water transport capacity and leaf area. It is proposed that, when there are changes in leaf properties without any external change in water availability or evaporative demand, leaf photosynthesis and stomatal conductance are partially constrained by the hydraulic architecture of the tree. Keywords: carbon isotope discrimination, hydraulic conductance, nitrogen, photosynthesis, tree water transport. Introduction Resistance to water flow through the xylem from the roots to the tree crown lowers leaf water potential and may limit water supply to the leaf. The structure of the water transport system, or hydraulic architecture, of the tree is therefore a major determinant of the leaf water balance (Zimmermann 1978, Tyree and Ewers 1991, Tyree and Ewers 1996), and must be considered when interpreting leaf physiological behavior. It has been hypothesized that stomata respond to hydraulic signals transmitted through the xylem (Whitehead 1998), and that the coordination of stomatal functioning with apparent hydraulic capacity acts to maintain water potential above a threshold minimum (Andrade et al. 1998, Bond and Kavanagh 1999, Salleo et al. 2000). Stomata also control the supply of CO 2 to the leaf interior, and photosynthesis and stomatal conductance are normally tightly coupled (e.g., Leuning 1995). The hydraulic properties of the tree thus also influence photosynthesis and growth in addition to stomatal behavior and leaf water balance, but the relationship between leaf physiology and the development of hydraulic architecture is poorly understood (Gartner 1995). Can hydraulic architecture vary in a coordinated manner with changes in leaf physiology, or is leaf physiology constrained by hydraulic architecture? In this study, we compared leaf physiology and stem hydraulic architecture in Eucalyptus grandis Hill ex Maiden trees receiving different rates of nitrogen (N) fertilization. Allocation of resources to hydraulic capacity is not fixed for a particular species, but varies with genotype, in response to the environment and with tree age. For example, Franks et al. (1995) compared provenances of Eucalyptus camaldulensis Dehnh. from contrasting climatic zones and found that seedlings from the more arid habitat had more permeable sapwood and lower vulnerability to xylem cavitation compared with seedlings from the wetter site. The hydraulic architecture of Pinus halepensis Mill. seedlings from a range of provenances was similar, but provenances from arid environments had lower amounts of xylem embolism and higher soil-to-leaf hydraulic conductance during a drought period (Tognetti et al. 1997). The opposite trend occurs when different species are compared, with drought-adapted species generally having

2 684 CLEARWATER AND MEINZER lower hydraulic conductivities than mesic species (Nardini and Tyree 1999). The light environment may also influence hydraulic architecture, with leaf specific hydraulic conductivity (k l ), a measure of hydraulic conductance per unit leaf area, normally being reduced in plants developing in the shade (Schultz and Matthews 1993, Shumway et al. 1993). Within coniferous species, the ratio of sapwood area to leaf area, a major determinant of k l, is known to vary with climate (Mencuccini and Grace 1995), stand density (Pothier and Margolis 1991) and site fertility (Espinosa-Bancalari et al. 1987). Heath et al. (1997) found that elevated CO 2 reduced whole-shoot leaf specific conductance and stomatal conductance in Quercus robur L. seedlings, but not in Fagus sylvatica L. The preceding examples highlight the plasticity of tree hydraulic architecture and the uncertainty involved in predicting its response to environmental variables. In the present study, we measured branch hydraulic conductivity and trunk sapwood area and compared them with leaf area, photosynthetic capacity, stomatal conductance (g s ), leaf water potential (Ψ L ) and carbon isotope discrimination in trees subjected to different N fertilization regimes in the field. We expected increased photosynthesis and g s in response to N fertilization. Our hypothesis was that N fertilization would result in increased leaf specific hydraulic conductivity so that higher photosynthetic rates, g s and transpiration per unit leaf area could be accommodated without a concomitant decline in Ψ L. We predicted that fertilized trees would grow faster and produce sapwood with lower density and higher porosity than unfertilized trees. Alternative hypotheses included no change in either photosynthesis or hydraulic architecture or an increase in photosynthesis and g s without change in hydraulic capacity, and hence a decrease in Ψ L. Materials and methods Site description, plant material and treatments The selected trees were grown as part of a separate study of fertilizer regimes for the establishment of commercial Eucalyptus plantations in Hawaii (Santo 2000). The study site was in an area previously used for sugarcane cultivation on the Hamakua Coast of the Island of Hawaii, 1 km east of the township of Pa auilo ( N W) at an altitude of 250 m, and exposed to the prevailing northeasterly trade winds. The soil is Kukaiau silty clay loam. Rainfall recorded in the township of Pa auilo (Station 221) during October 1997 to September 1998 was 2620 mm, 11% more than the historical average for the same period, although the period from January to March 1998 was drier than average. Before planting, the site was cleared of weeds by spraying glyphosate herbicide, burning, disc harrowing and in-row subsoiling. Seedlings of E. grandis were grown from seed (Mondi Forest Orchards, South Africa) in a nursery for 12 weeks and hand planted in October 1997 at 2-m intervals in rows 3 m apart. Approximately 500 ml of water was applied per seedling at planting, but there was no further irrigation. Fertilizer treatments were applied to plots of trees three rows wide and seven trees deep (21 trees in total). The study trees received 0, 112, 224 or 336 kg N ha 1 year 1, applied as CO(NH 2 ) 2 at 2-month intervals over the first year. Potassium was applied at the same time as N, with all treatments receiving 224 kg K 2 Oha 1 year 1 as muriate of potash, and all treatments received 224 kg P 2 O 5 ha 1 as triple superphosphate and 224 kg Ca ha 1 as CaCO 3 at the time of planting. Starting 2 days after transplanting, fertilizers were applied as a surface band around each tree with a radius of about 0.5 m. Treatments were replicated in three blocks (12 plots in total). Growth, hydraulic conductance, and bulk foliage nitrogen and carbon isotope measurements were made on two randomly selected trees from the outer rows of each plot (six trees in total per treatment), the inner row being reserved for the fertilizer study. The more timeconsuming ambient photosynthesis and water potential measurements were made only on trees in the 0 and 336 kg N ha 1 year 1 plots within each block. Photosynthesis, stomatal conductance and leaf water potentials During June 1998, a closed photosynthesis system (LI-6200, Li-Cor, Inc., Lincoln, NE) was used to record the photosynthetic response to CO 2 of mature, fully expanded single leaves from 13 trees (two to four leaves per N treatment). A source of air with a CO 2 concentration of at least 1000 µmol mol 1 was produced by blowing into a large plastic bag initially filled with ambient air. The leaf was enclosed in a 1-liter chamber where constant photosynthetically active radiation (PAR) of 1000 µmol m 2 s 1 was provided by an LED light source (Q-Beam 2001-A, Quantum Devices Inc., Barneveld, WI). The CO 2 -enriched air was supplied by means of a bypass switch on the photosynthesis system. Once photosynthesis and g s had reached constant values in CO 2 -enriched air, the bypass switch was closed and the rate of CO 2 assimilation recorded every minute as the CO 2 concentration decreased to 50 µmol mol 1 through the action of photosynthesis over a period of at least 60 min. Photosynthetic measurements were corrected for leaks after measuring the leak rate of the closed system before each leaf was enclosed in the chamber. Plots of photosynthesis as a function of intercellular CO 2 concentration were fitted with the mechanistic model of von Caemmerer and Farquhar (1981) and the parameters J max and V cmax were estimated by nonlinear regression (Photosyn Assistant, Dundee Scientific, Dundee, U.K.). On 5 days in July 1998, measurements of CO 2 assimilation and g s under ambient conditions were made with an open photosynthesis system (Ciras, PP Systems, Hitchin, U.K.). Three mature, fully expanded leaves were selected from a range of heights and azimuths within the crown of each tree and marked with flagging tape. Most leaves received at least some direct sunlight. Senescent older leaves located in the shaded lower portions of the crown were avoided. Measurements began at about 0800 h and were repeated every 1 to 2 hours until about 1700 h on two trees in one 0 and one 336 kg Nha 1 year 1 plot from the same replicate. On subsequent days, measurements were repeated on different trees in differ- TREE PHYSIOLOGY VOLUME 21, 2001

3 EUCALYPTUS HYDRAULIC ARCHITECTURE AND PHOTOSYNTHESIS 685 ent replicates of the fertilizer trial. For each measurement, the leaf was enclosed in the chamber, held at its natural orientation and photosynthesis recorded after readings had stabilized (usually within 1 min). Average chamber CO 2 concentration and leaf temperature for each leaf varied between 340 and 370 µmol mol 1 and 24 and 28 C, respectively. During measurements, PAR measured by the external quantum sensor attached to the cuvette varied between 3 and 1960 µmol m 2 s 1. No consistent difference was found between leaves from different positions within the canopy. Results for each tree were therefore pooled and fitted with a non-rectangular hyperbola describing the response of CO 2 assimilation to PAR (Ogren and Evans 1993), with the light-saturated rate of photosynthesis estimated by nonlinear regression (Photosyn Assistant, Dundee Scientific). Leaf water potential was measured with a pressure chamber on the same days as ambient photosynthesis. Measurements began at dawn and were alternated with photosynthesis measurements on the same tree. At least two leaves were enclosed in a plastic bag and removed from the tree for each measurement. Measurements of xylem water potential were also made around midday on adjacent, non-transpiring leaves that had been covered with a plastic bag and aluminum foil on the previous evening. All leaves used for photosynthesis measurements were retained, their area measured and dry mass recorded after drying at 75 C for 24 h. Total N content of finely ground samples was determined by the micro-kjeldahl technique at the Agricultural Diagnostic Service Center, University of Hawaii, Manoa. Branch hydraulic conductivity The hydraulic conductance of proximal pieces of stem from three branches per tree was measured with a high-pressure flow meter (HPFM) (Tyree et al. 1995) during August To account for any vertical gradients in branch conductance, the height of the tree was first measured with a measuring pole, divided into three layers of equal height, and one branch cut from each layer. Branches were chosen to represent a range of azimuths and branch sizes, and the height of insertion of each branch was recorded. A piece of stem approximately 15 cm long was immediately cut from the proximal end of the branch and stored in water for measurement of conductance in the laboratory. The remainder of the branch was retained separately for measurement of leaf area. In the laboratory, the stem segment was recut and smoothed with a razor blade, a pressure of at least 70 kpa was then applied to the proximal end and the rate of flow through the segment recorded with the HPFM. A 10 mm solution of oxalic acid was used as the conductivity solution to inhibit microbial growth, and indigo carmine dye was added to the solution for some stem segments to check that the entire cross section was conducting. Measured conductance was independent of the applied pressure and constant for as long as pressure was applied. Stem temperatures, measured with a thermocouple during conductance measurements, varied between 22 and 26 C. Flow estimates were corrected for differences between HPFM measurement temperature and calibration temperature (Tyree et al. 1995). Specific conductivity (k s ) and leaf specific conductivity (k l ) were calculated according to Tyree and Ewers (1991). Growth On August 20, 1998, two trees from each plot were cut at ground level and tree height and stem diameter at ground level recorded. A slice at least 3-cm thick was cut from the base of the main stem, wet volume under bark determined by displacement and the sample dried at 75 C to a constant weight for calculation of wood density. The crown of each tree was separated into stems >1 cm in diameter and stems <1 cm in diameter (including leaves) and the two fractions weighed with a hanging scale. A subsample of six branches was randomly selected from the stems <1 cm in diameter, separated into leaves and stems and the wet mass of the two fractions recorded. The ratio of leaf mass to total (stem plus leaf) mass was later used to determine total leaf area for the tree. The area of the leaf sample was measured with a leaf area meter, dry mass was recorded after drying at 75 C for 24 h, and the sample finely ground before N content was measured as described above. Subsamples of leaf tissue were also combusted and the relative abundance of 13 C and 12 C in the CO 2 produced determined with an SIRA series II isotope ratio mass spectrometer (VGIsotech, Middlewich, U.K.) at the Duke University Phytotron. Stable carbon isotope composition was expressed as the 13 C/ 12 C ratio relative to that of the PeeDee belemnite standard with a precision of ± 0.2. The resulting δ 13 C values were used to estimate isotopic discrimination as δ =(δ a δ p )/(1 + δ p ), where δ p is the isotopic composition of the plant material and δ a is the isotopic composition of the air (Farquhar et al. 1982). The δ 13 C value of air was assumed to be 8, the value at Mauna Loa, Hawaii (Anonymous 1984). Results Growth Nitrogen-fertilized trees were taller and had at least twice the leaf area of unfertilized trees (Figures 1a and 1b). Mean stem basal area at ground level ranged from 27±5cm 2 in unfertilized trees to 67±3cm 2 in plants receiving 336 kg N ha 1 (F = 11.93, P < 0.01). Mean leaf N concentration per unit area (N area ) was also higher in fertilized trees than in unfertilized trees when measured on a bulk sample of leaves from throughout the crown (Figure 1c), but variation within treatments was high and the overall treatment effect was not significant. Differences in N area were significant when the fertilized treatments were grouped and contrasted with the zero N treatment (t = 2.18, P = 0.02). Although growth and leaf N area increased in response to N fertilization, differences between the N fertilization treatments were small and variable. Photosynthesis Light-saturated photosynthetic rate on a leaf area basis (A max area ) increased with increasing N area (Figure 2a) as a result TREE PHYSIOLOGY ONLINE at

4 686 CLEARWATER AND MEINZER of increases in leaf N per unit mass (N mass ) and photosynthesis per unit mass (A max mass, Figure 2b). Specific leaf area (SLA) was less important as a determinant of A max area (r 2 = 0.02, P = 0.60). The maximum rate of electron transport (J max ), expressed on an area basis, increased significantly with increasing N area, whereas the maximum rate of carboxylation by Rubisco (V cmax ) was insensitive to N area (Figure 3a). Correlations between N concentration and both J max and V cmax improved when the parameters were expressed per unit mass, again indicating that increases in photosynthetic capacity were more the result of increased N concentration and allocation to photosynthetic components than changes in SLA (Figure 3b). Parameters J max and V cmax were correlated with each other (area basis r 2 = 0.65, P < 0.01; mass basis r 2 = 0.87, P < 0.01), indicating that leaves with high electron transport capacity also had high carboxylation capacity. Figure 1. Tree height (a), total leaf area (b) and nitrogen per unit leaf area (c) for E. grandis trees in four nitrogen fertilization regimes. Values are means ± 1 SE, n = 6 trees per treatment. Results of ANOVA: (a) F = 6.1, P<0.01; (b) F = 11.2, P<0.01 and (c) F = 1.4, P = Hydraulic architecture Hydraulic capacity at the whole-tree and branch level did not vary in response to N treatment. Leaf area per tree and sapwood area at the base increased in constant proportion to each other (Figure 4a), and there was no relationship between the ratio of leaf area to sapwood area (LA/SA) and N concentration (N area or N mass ) of the foliage (r 2 = 0.02, P = 0.5). Wood density at the base of the tree did not vary with N area (r 2 = 0.06, P = 0.24), but was weakly correlated with basal stem diameter (y =15x + 269, r 2 = 0.38, P < 0.01). Overall mean wood density was 382 ± 16 kg m 3. Similarly, branch leaf area was closely related to branch sapwood area (Figure 4b) and branch LA/SA for each tree did not vary with tree N area (r 2 = 0.06, P = 0.23). Specific conductivity (k s ) and leaf specific conductivity (k l ) of proximal stem pieces from branches did not vary significantly with tree N area, regardless of height of branch insertion within the crown (k s versus N area, r 2 = 0.02, P = 0.52; k l versus N area, r 2 < 0.01, P = 0.93). Figure 2. Light-saturated photosynthetic rate as a function of leaf nitrogen concentration, with parameters expressed per unit leaf area (a) and per unit leaf mass (b). Values are derived from light response curves fitted to data from three leaves per tree, measured under ambient conditions; n = 16 trees. Linear regressions: (a) y = 8.6x , P<0.01 and (b) y = 12.9x + 0.1, P<0.01. Figure 3. Maximum rates of electron transport (J max, ) and carboxylation (V cmax, ) as a function of leaf nitrogen concentration, with parameters expressed per unit leaf area (a) and per unit leaf mass (b). Parameters are fitted to data for single leaves from 13 trees. Linear regressions: (a) J max =56x + 83, P = 0.03, V cmax =6x + 67, P = 0.65 and (b) J max = 102x + 0.2, P<0.01, V cmax =37x + 0.3, P = TREE PHYSIOLOGY VOLUME 21, 2001

5 EUCALYPTUS HYDRAULIC ARCHITECTURE AND PHOTOSYNTHESIS 687 branches, most of the xylem was later-formed secondary xylem with narrow vessels and low vessel density (mean vessel diameter ± 1 SE, upper branches = 52 ± 2 µm, and lower branches = 44 ± 3 µm, t = 6.2, P = 0.03; mean vessel density, upper branches = 39 ±5mm 2 and lower branches = 32 ± 8mm 2, t = 1.1, P = 0.18). Figure 4. Leaf area versus sapwood area for whole trees (a) and individual branches at all heights of insertion (b). Inset in (b) shows detail for smaller branches. Linear regressions: (a) y = 0.70x , n = 24, P < 0.01 and (b) y = 0.99x 0.06, n = 69, P < Leaf water relations Stomatal conductance under light-saturated conditions did not vary with leaf nitrogen concentration (Figure 6a), even though photosynthetic rates increased with increasing N concentration (Figure 2a). The ratio of internal to external CO 2 partial pressure (p i /p a ) therefore decreased with increasing leaf N concentration (Figure 6b). Leaf water potential measured at midday on transpiring and non-transpiring leaves did not vary significantly with leaf N concentration (Figure 6c). Overall, mean midday Ψ L of non-transpiring leaves was less than half that of transpiring leaves ( 0.38 versus 0.91 MPa) (Figure 6c). Mean predawn water potential during the same period was 0.1 MPa. Carbon isotope discrimination ( ) measured on the bulk leaf sample from throughout the crown was lower in trees with Although branch hydraulic capacity was not correlated with foliage N concentration, k s and k l increased with branch insertion height (Figure 5). Upper branches were younger and had smaller diameters than lower branches. Comparison of vessel diameter and density in stem transverse sections from three lower and three upper branches indicated that higher specific conductivity in the upper branches was the result of larger vessel diameter and possibly higher vessel density in the firstformed secondary xylem of upper branches. In the lower Figure 5. Specific conductivity (a) and leaf specific conductivity (b) as a function of height of branch insertion normalized by tree height. Values are means ±1SE,n = 19 for each height class. Results of ANOVA for differences between height classes: (a) F = 9.5, P < 0.01 (b) F = 23.6, P<0.01. Figure 6. Light-saturated stomatal conductance (a), ratio of internal to external CO 2 concentration (b) and midday leaf water potential for transpiring ( ) and non-transpiring ( ) leaves (c), as functions of leaf nitrogen concentration. Values for g s and p i /p a represent the mean of all measurements for three leaves per tree under ambient conditions when PAR > 700 µmol m 2 s 1, n = 16 trees. Values for midday Ψ L represent the mean of all measurements made between 1000 and 1400 h, n = 15 trees. Linear regressions: (a) y = 0.0x , P = 0.93; (b) y = 0.08x , P<0.01 and (c) transpiring = 5x 16, P = 0.15 and non-transpiring = 1.5x 6,P = TREE PHYSIOLOGY ONLINE at

6 688 CLEARWATER AND MEINZER high leaf N concentration than in trees with low leaf N concentration (Figure 7). Total variation in discrimination was approximately 2, corresponding to variation in N area from 1.0 to 1.8 g m 2. Discussion Leaf specific hydraulic capacity in the E. grandis trees did not change in response to fertilization, despite increases in leaf N concentration and photosynthetic capacity. Stomatal conductance was also unresponsive to fertilization and, given the exposed windy location of the study site, it is reasonable to assume that water use per unit leaf area was also unchanged. Calculated p i /p a was therefore reduced in trees with high leaf N concentration, an observation strongly supported by independent measurements of reduced discrimination against 13 C with increasing leaf N concentration. Our hypothesis that hydraulic capacity increases in coordination with increased leaf photosynthetic capacity was therefore rejected. Instead, we propose that stem hydraulic architecture was conserved, and that stomatal control of transpiration was coordinated with hydraulic capacity rather than with leaf photosynthetic capacity. Although a causal relationship was not demonstrated, it is possible that aboveground hydraulic architecture places a limitation on crown photosynthetic carbon assimilation. Water-use efficiency increased in N-fertilized trees as leaf photosynthetic capacity increased, but hydraulic and stomatal limitations on photosynthesis remained the same. Stomatal control of transpiration is often coordinated with leaf-area-based water transport capacity (Meinzer and Grantz 1990, Whitehead 1998). Partial removal or covering of tree foliage usually induces a rapid increase in g s in the remaining foliage (Whitehead et al. 1996, Pataki et al. 1998), whereas cavitation or stem wounding induces a decline in g s (Sperry et al. 1993, Sperry and Pockman 1993). Therefore, stomata may respond to some form of hydraulic signal, generated by perturbation of the hydraulic pathway, even though bulk Ψ L may remain relatively constant, or increase during the change (Whitehead 1998). Instead of a simple negative feedback response to bulk Ψ L, stomatal coordination of transport capacity Figure 7. Relationship between carbon isotope discrimination and leaf nitrogen concentration for a bulk sample of leaves from throughout the tree crown. Linear regression: y = 2x + 25, P<0.01, n = 24. may be a threshold response to a critical value of water potential (Bond and Kavanagh 1999). We did not determine the threshold water potential for decreased g s, but because of frequent rainfall and high water potentials it is possible that g s was maximal in all trees irrespective of N status. Partial defoliation or root pressurization could be used to determine whether g s increases in response to an increase in leaf specific hydraulic capacity. Regardless of whether g s was constrained by maximal stomatal aperture and density, or by a mechanism that maintained a threshold minimum Ψ L, g s and stem hydraulic capacity remained constant despite a fertilization-induced enhancement of leaf photosynthetic capacity. Why was there no increase in stem hydraulic capacity in fertilized plants in coordination with increased leaf photosynthetic capacity? Our results suggest that stem hydraulic architecture partially determines leaf and crown physiology, rather than the reverse (Gartner 1995). The xylem in new branches develops before the leaves are fully expanded, and stems serve a structural as well as hydraulic function. A trade-off between mechanical and hydraulic performance may limit phenotypic variation in hydraulic capacity in response to changes in leaf photosynthetic capacity (Gartner 1995). The tendency for hydraulic capacity to be established before leaf maturation is illustrated by the strong increases in k s and k l with height of branch insertion. The concept of hydraulic segmentation suggests that leaf specific conductivity normally decreases toward the shoot apex (Zimmermann 1978). However, increasing k l toward the apex in trees is a known trait associated with strong apical control that is thought to hydraulically favor the faster growing leader and upper branches (Tyree and Ewers 1996). In new branches, a large hydraulic capacity is established first, the first leaves mature later, and k l declines as the branch grows larger. The lack of any change in k l in response to fertilization conflicts with previous reports of plasticity in hydraulic architecture in response to climate, stand density or light environment (Pothier and Margolis 1991, Schultz and Matthews 1993, Mencuccini and Grace 1995). However, in many of these examples, adjustments in hydraulic architecture may have been induced by a change in water supply in relation to evaporative demand, such as a decrease in soil water content in more densely planted stands with an initially higher leaf area index. In contrast, in our study, N fertilization induced changes in the intrinsic leaf photosynthetic capacity of young trees that experienced a relatively constant water supply. Our result reflects the short-term response before fertilization affected conditions at the stand level. Differences in hydraulic architecture might have occurred if this experiment had been continued until canopy closure was complete and N-induced differences in tree size and LAI began to affect soil water content. In a long-term study, Brix and Mitchell (1983) found that LA/SA was increased by fertilization, but there is generally little information on hydraulic effects in forest fertilization experiments. Other studies have shown that water-use efficiency in trees can increase in response to N fertilization (Brix and Mitchell TREE PHYSIOLOGY VOLUME 21, 2001

7 EUCALYPTUS HYDRAULIC ARCHITECTURE AND PHOTOSYNTHESIS , Sands and Mulligan 1990, Liu and Dickmann 1996), but few have considered how changes in hydraulic architecture or sapwood properties might contribute to the fertilizer response (Harvey and van den Driessche 1997). Increases in water-use efficiency with N fertilization are more apparent during periods of drought (Liu and Dickmann 1996). Nitrogen-fertilized trees may be more vulnerable to drought stress than unfertilized trees because of their increased size and transpiration rates, and because of decreased allocation to roots (Harvey and van den Driessche 1999). However, increases in growth in response to fertilization are generally larger than any negative effects on growth during periods of drought (Nilsen 1995). Harvey and van den Driessche (1997, 1999) grew poplars in containers and found that a high N supply increased growth and water-use efficiency through an increase in photosynthesis without any increase in g s, as observed in this study. They also found that added N altered vessel pit pore structure and increased vessel diameter, cavitation, and loss of conductivity, although these responses may have been partially caused by decreased soil water content at high-n supply (Harvey and van den Driessche 1997, 1999). In our study, wood density at the base of the main stem was unrelated to foliage N concentration, suggesting that there was no significant change in sapwood anatomy or porosity with fertilization. Wood density was higher in larger trees, reflecting a normal trend of increasing density with increasing tree size and maturity. Vulnerability to cavitation was not measured, but xylem water potentials were above the threshold of approximately 1.3 MPa required for significant cavitation in another study that included E. grandis (Vander Willigen and Pammenter 1998). Given high rainfall and high predawn Ψ L, it is unlikely that loss of conductivity caused by cavitation was significant during the measurement period in our study. It is well known that increased N supply results in decreased allocation to roots in many species (e.g., Gleeson 1993). In an experiment located near the present study, balanced fertilization of E. saligna trees resulted in increased gross primary production and decreased relative allocation below ground (C.P. Giardina, University of Hawaii, HI and M.G. Ryan, US Forest Service, Fort Collins, CO, unpublished observations). It is possible that N fertilization caused decreased allocation to roots in our study, thereby diminishing total soil-to-leaf hydraulic conductance. However, if leaf-area-specific, root hydraulic conductance was reduced in fertilized trees, the change was not large enough to be detectable as more negative water potentials during peak transpiration on sunny days. During dry periods, root hydraulic conductance may increasingly limit total hydraulic conductance; nevertheless, the overall effect is likely to be a more rapid onset of symptoms of drought stress, such as stomatal closure and further increases in water-use efficiency in fertilized trees. Tree to tree variation was high. There was noticeable genotypic variation in leaf thickness and shape, attributable to the use of a bulk Eucalyptus seed source intended for commercial forestry. Therefore, some of the observed variation in foliage N concentration may have been the result of genetic variation among trees, as well as the fertilizer treatments. Genetic variation helps to explain the variation between N treatments and was the rationale for using leaf N concentration as the independent variable. Despite this added component to the underlying cause of variation, shoot hydraulic properties were conserved even though there were clear differences in leaf photosynthetic properties associated with variation in leaf N concentration. Acknowledgments This study was supported by USDA/ARS cooperative agreement 58-91H with the Hawaii Agriculture Research Center. Lance Santo and Hamakua Timber provided access to the Eucalyptus fertilizer trial, Aileen Yeh, Teresa Restom and Linda Lenz gave assistance in the field and Christian Giardina generously provided laboratory facilities in Hilo. References Andrade, J.L., F.C. Meinzer, G. Goldstein, N.M. Holbrook, J. Cavelier, P. Jackson and K. Silvera Regulation of water flux through trunks, branches and leaves in trees of a lowland tropical forest. Oecologia 115: Anonymous Modeling 13 C. In Geophysical Monitoring for Climatic Change No. 12. Eds. Harris, J.M. and E.C. Nickerson. U.S. Department of Commerce, Boulder, CO, pp Bond, B.J. and K.L. Kavanagh Stomatal behavior of four woody species in relation to leaf-specific hydraulic conductance and threshold water potential. Tree Physiol. 19: Brix, H. and A.K. Mitchell Thinning and nitrogen fertilization effects on sapwood development and the relationships of foliage quantity to sapwood area and basal area in Douglas-fir. Can. J. For. Res. 13: Brix, H. and A.K. Mitchell Thinning and nitrogen fertilization effects on soil and tree water stress in a Douglas-fir stand. Can. J. For. Res. 16: Espinosa-Bancalari, M.A., D.A. Perry and J.D. Marshall Leaf area-sapwood area relationships in adjacent young Douglas-fir stands with different growth rates. Can. J. For. Res.17: Farquhar, G.D., M.C. Ball, S. Von Caemmerer and Z. Roksandic Effect of salinity and humidity on the 13 C value of halophytes evidence for diffusional isotope fractionation determined by the ratio of intercellular/atmospheric partial pressure of CO 2 under different environmental conditions. Oecologia 52: Franks, P.J., A. Gibson and E.P. Bachelard Xylem permeability and embolism susceptibility in seedlings of Eucalyptus camaldulensis Dehnh. from two different climatic zones. Aust. J. Plant Physiol. 22: Gartner, B.L Patterns of xylem variation within a tree and their hydraulic and mechanical consequences. In Plant Stems: Physiological and Functional Morphology. Ed. B.L. Gartner. Academic Press, San Diego, pp Gleeson, S.K Optimization of tissue nitrogen and root shoot allocation. Ann. Bot. 71: Harvey, H.P. and R. van den Driessche Nutrition, xylem cavitation and drought resistance in hybrid poplar. Tree Physiol. 17: Harvey, H.P. and R. van den Driessche Nitrogen and potassium effects on xylem cavitation and water-use efficiency in poplars. Tree Physiol. 19: TREE PHYSIOLOGY ONLINE at

8 690 CLEARWATER AND MEINZER Heath, J., G. Kerstiens and M.T. Tyree Stem hydraulic conductance of European beech (Fagus sylvatica L.) and pedunculate oak (Quercus robur L.) grown in elevated CO 2. J. Exp. Bot. 48: Leuning, R A critical-appraisal of a combined stomatal-photosynthesis model for C-3 plants. Plant Cell Environ. 18: Liu, Z.J. and D.I. Dickmann Effects of water and nitrogen interaction on net photosynthesis, stomatal conductance, and water-use efficiency in two hybrid poplar clones. Physiol. Plant. 97: Meinzer, F.C. and D.A. Grantz Stomatal and hydraulic conductance in growing sugarcane stomatal adjustment to water transport capacity. Plant Cell Environ. 13: Mencuccini, M. and J. Grace Climate influences the leaf-area sapwood area ratio in scots pine. Tree Physiol. 15:1 10. Nardini, A. and M.T. Tyree Root and shoot hydraulic conductance of seven Quercus species. Ann. For. Sci. 56: Nilsen, P Effect of nitrogen on drought strain and nutrient-uptake in norway spruce (Picea abies (L.) Karst) trees. Plant Soil 172: Ogren, E. and J.R. Evans Photosynthetic light-response curves. 1. The influence of CO 2 partial-pressure and leaf inversion. Planta 189: Pataki, D.E., R. Oren and N. Phillips Responses of sap flux and stomatal conductance of Pinus taeda L. trees to stepwise reductions in leaf area. J. Exp. Bot. 49: Pothier, D. and A. Margolis Analysis of growth and light interception of balsam fir and white birch saplings following precommercial thinning. Ann. Sci. For. 48: Salleo, S., A. Nardini, F. Pitt and M.A. Lo Gullo Xylem cavitation and hydraulic control of stomatal conductance in laurel (Laurus nobilis L.). Plant Cell Environ. 23: Sands, R. and D.R. Mulligan Water and nutrient dynamics and tree growth. For. Ecol. Manage. 30: Santo, L.T Fertilization of Eucalyptus for rapid canopy closure on the Hamakua coast at Pa auilo. Technical supplement to forestry report. Hawaii Agriculture Research Center, Aiea, Hawaii, 31 p. Schultz, H.R. and M.A. Matthews Xylem development and hydraulic conductance in sun and shade shoots of grapevine (Vitis vinifera L.): evidence that low light uncouples water transport capacity from leaf area. Planta 190: Shumway, D.L., K.C. Steiner and T.E. Kolb Variation in seedling hydraulic architecture as a function of species and environment. Tree Physiol. 12: Sperry, J.S., N.N. Alder and S.E. Eastlack The effect of reduced hydraulic conductance on stomatal conductance and xylem cavitation. J. Exp. Bot. 44: Sperry, J.S. and W.T. Pockman Limitation of transpiration by hydraulic conductance and xylem cavitation in Betula occidentalis. Plant Cell Environ. 16: Tognetti, R., M. Michelozzi and A. Giovannelli Geographical variation in water relations, hydraulic architecture and terpene composition of aleppo pine seedlings from Italian provenances. Tree Physiol. 17: Tyree, M.T. and F.W. Ewers The hydraulic architecture of trees and other woody-plants. New Phytol. 119: Tyree, M.T. and F.W. Ewers Hydraulic architecture of woody tropical plants. In Tropical Forest Plant Ecophysiology. Eds. S.S. Mulkey, R.L. Chazdon and A.P. Smith. Chapman and Hall, New York, pp Tyree, M.T., S. Patino, J. Bennink and J. Alexander Dynamic measurements of root hydraulic conductance using a high-pressure flowmeter in the laboratory and field. J. Exp. Bot. 46: Vander Willigen, C. and N.W. Pammenter Relationship between growth and xylem hydraulic characteristics of clones of Eucalyptus spp. at contrasting sites. Tree Physiol. 18: von Caemmerer, S. and G.D. Farquhar Some relationships between the biochemistry of photosynthesis and the gas-exchange of leaves. Planta 153: Whitehead, D Regulation of stomatal conductance and transpiration in forest canopies. Tree Physiol. 18: Whitehead, D., N.J. Livingston, F.M. Kelliher, K.P. Hogan, S. Pepin, T.M. McSeveny and J.N. Byers Response of transpiration and photosynthesis to a transient change in illuminated foliage area for a Pinus radiata D. Don. tree. Plant Cell Environ. 19: Zimmermann, M.H Hydraulic architecture of some diffuse-porous trees. Can. J. Bot. 56: TREE PHYSIOLOGY VOLUME 21, 2001

Whole-plant hydraulic resistance and vulnerability segmentation in Acer saccharinum

Whole-plant hydraulic resistance and vulnerability segmentation in Acer saccharinum Tree Physiology 17, 351--357 1997 Heron Publishing----Victoria, Canada Whole-plant hydraulic resistance and vulnerability segmentation in Acer saccharinum MAKOTO TSUDA 1,3 and MELVIN T. TYREE 1,2 1 Department

More information

Reduced photosynthesis in old oak (Quercus robur): the impact of crown and hydraulic architecture

Reduced photosynthesis in old oak (Quercus robur): the impact of crown and hydraulic architecture Tree Physiology 22, 597 601 2002 Heron Publishing Victoria, Canada Reduced photosynthesis in old oak (Quercus robur): the impact of crown and hydraulic architecture STEFFEN RUST 1,2 and ANDREAS ROLOFF

More information

Xylem cavitation and loss of hydraulic conductance in western hemlock following planting

Xylem cavitation and loss of hydraulic conductance in western hemlock following planting Tree Physiology 17, 59--63 1997 Heron Publishing----Victoria, Canada Xylem cavitation and loss of hydraulic conductance in western hemlock following planting K. L. KAVANAGH 1 and J. B. ZAERR 2 1 Department

More information

! P = -2T/r. Example: calculate! P for r = 1 x 10-6 m and 1 x 10-7 m. About -0.15MPa for 1!m, and -1.5 MPa for 0.1!m.

! P = -2T/r. Example: calculate! P for r = 1 x 10-6 m and 1 x 10-7 m. About -0.15MPa for 1!m, and -1.5 MPa for 0.1!m. ! P = -2T/r Example: calculate! P for r = 1 x 10-6 m and 1 x 10-7 m. About -0.15MPa for 1!m, and -1.5 MPa for 0.1!m. Getting water from the soil into the plant.! root

More information

Breeding for Drought Resistance in Cacao Paul Hadley

Breeding for Drought Resistance in Cacao Paul Hadley Breeding for Drought Resistance in Cacao Paul Hadley University of Reading Second American Cocoa Breeders Meeting, El Salvador, 9-11 September 215 9 September 215 University of Reading 26 www.reading.ac.uk

More information

Ecological relevance of minimum seasonal water potentials

Ecological relevance of minimum seasonal water potentials Physiologia Plantarum 127: 353 359. 2006 Copyright ß Physiologia Plantarum 2006, ISSN 0031-9317 Ecological relevance of minimum seasonal water potentials R. Bhaskar a,1, * and D.D. Ackerly b a Department

More information

Gas exchange and water relations of evergreen and deciduous tropical savanna trees

Gas exchange and water relations of evergreen and deciduous tropical savanna trees Gas exchange and water relations of evergreen and deciduous tropical savanna trees G. Goldstein, F. Rada, P. Rundel, A. Azocar, A. Orozco To cite this version: G. Goldstein, F. Rada, P. Rundel, A. Azocar,

More information

Department of Dendrology, University of Forestry, 10 Kl. Ohridski blvd., Sofia 1756, Bulgaria, tel.: *441

Department of Dendrology, University of Forestry, 10 Kl. Ohridski blvd., Sofia 1756, Bulgaria, tel.: *441 General and Applied Plant Physiology 2009, Volume 35 (3 4), pp. 122 126 2009 ISSN 1312-8183 Published by the Institute of Plant Physiology Bulgarian Academy of Sciences Available online at http://www.bio21.bas.bg/ipp/

More information

TREES. Functions, structure, physiology

TREES. Functions, structure, physiology TREES Functions, structure, physiology Trees in Agroecosystems - 1 Microclimate effects lower soil temperature alter soil moisture reduce temperature fluctuations Maintain or increase soil fertility biological

More information

Carbon Input to Ecosystems

Carbon Input to Ecosystems Objectives Carbon Input Leaves Photosynthetic pathways Canopies (i.e., ecosystems) Controls over carbon input Leaves Canopies (i.e., ecosystems) Terminology Photosynthesis vs. net photosynthesis vs. gross

More information

Trees are: woody complex, large, long-lived self-feeding shedding generating systems compartmented, self optimizing

Trees are: woody complex, large, long-lived self-feeding shedding generating systems compartmented, self optimizing BASIC TREE BIOLOGY Trees are: woody complex, large, long-lived self-feeding shedding generating systems compartmented, self optimizing Roots: absorb water and minerals store energy support and anchor

More information

POTASSIUM IN PLANT GROWTH AND YIELD. by Ismail Cakmak Sabanci University Istanbul, Turkey

POTASSIUM IN PLANT GROWTH AND YIELD. by Ismail Cakmak Sabanci University Istanbul, Turkey POTASSIUM IN PLANT GROWTH AND YIELD by Ismail Cakmak Sabanci University Istanbul, Turkey Low K High K High K Low K Low K High K Low K High K Control K Deficiency Cakmak et al., 1994, J. Experimental Bot.

More information

Understanding how vines deal with heat and water deficit

Understanding how vines deal with heat and water deficit Understanding how vines deal with heat and water deficit Everard Edwards CSIRO AGRICULTURE & FOOD How hot is too hot? Cell death will occur in any vine tissue beyond a threshold (lethal) temperature cell

More information

Diurnal variation in xylem hydraulic conductivity in white ash (Fraxinus americana L.), red maple (Acer rubrum L.) and red spruce (Picea rubens Sarg.

Diurnal variation in xylem hydraulic conductivity in white ash (Fraxinus americana L.), red maple (Acer rubrum L.) and red spruce (Picea rubens Sarg. Plant, Cell and Environment (1998) 21, 1173 1180 ORIGINAL ARTICLE OA 220 EN Diurnal variation in xylem hydraulic conductivity in white ash (Fraxinus americana L.), red maple (Acer rubrum L.) and red spruce

More information

Effect of 1-MCP on Water Relations Parameters of Well-Watered and Water-Stressed Cotton Plants

Effect of 1-MCP on Water Relations Parameters of Well-Watered and Water-Stressed Cotton Plants Effect of 1-MCP on Water Relations Parameters of Well-Watered and Water-Stressed Cotton Plants Eduardo M. Kawakami, Derrick M. Oosterhuis, and John L. Snider 1 RESEARCH PROBLEM The cotton crop in the U.S.

More information

Stomatal conductance has a strong dependence upon humidity deficits

Stomatal conductance has a strong dependence upon humidity deficits Stomatal conductance has a strong dependence upon humidity deficits 1 There is no universal function between stomatal conductance and humidity deficits. Some plants are more sensitive than others Hall

More information

Acoustic Emission Technique for the Detection of Abnormal Cavitation in Pine Trees Infected with Pine Wilt Disease

Acoustic Emission Technique for the Detection of Abnormal Cavitation in Pine Trees Infected with Pine Wilt Disease Acoustic Emission Technique for the Detection of Abnormal Cavitation in Pine Trees Infected with Pine Wilt Disease Keiko Kuroda (Kansai Research Center, Forestry and Forest Products Research Institute,

More information

Impact of genetic variation in stomatal conductance on water use efficiency in Quercus robur. Oliver Brendel. INRA Nancy France

Impact of genetic variation in stomatal conductance on water use efficiency in Quercus robur. Oliver Brendel. INRA Nancy France Impact of genetic variation in stomatal conductance on water use efficiency in Quercus robur Oliver Brendel INRA Nancy France Unit of Forest Ecology and Ecophysiology In collaboration with INRA Pierroton

More information

LEAF WATER POTENTIAL AND STOMATAL CONDUCTANCE OF RUBBER (Hevea brasiliensis) AS INFLUENCED BY SOIL MOISTURE AND LEAF AGE LALANI SAMARAPPULI ABSTRACT

LEAF WATER POTENTIAL AND STOMATAL CONDUCTANCE OF RUBBER (Hevea brasiliensis) AS INFLUENCED BY SOIL MOISTURE AND LEAF AGE LALANI SAMARAPPULI ABSTRACT LEAF WATER POTENTIAL AND STOMATAL CONDUCTANCE OF RUBBER (Hevea brasiliensis) AS INFLUENCED BY SOIL MOISTURE AND LEAF AGE J BY LALANI SAMARAPPULI ABSTRACT Stomatal conductance and transpiration and leaf

More information

% FOREST LEAF AREA. Figure I. Structure of the forest in proximity of the Proctor Maple Research Center -~--~ ~

% FOREST LEAF AREA. Figure I. Structure of the forest in proximity of the Proctor Maple Research Center -~--~ ~ NTRODUCTON There is a critical need to develop methods to address issues of forest canopy productivity and the role of environmental conditions in regulating forest productivity. Recent observations of

More information

Stomata and water fluxes through plants

Stomata and water fluxes through plants Stomata and water fluxes through plants Bill Davies The Lancaster Environment Centre, UK Summary Stomata and responses to the environment Conductance, a function of frequency and aperture Measuring/estimating

More information

Xylem cavitation, leaf growth and leaf water potential in Eucalyptus globulus clones under well-watered and drought conditions

Xylem cavitation, leaf growth and leaf water potential in Eucalyptus globulus clones under well-watered and drought conditions CSIRO PUBLISHING www.publish.csiro.au/journals/fpb Functional Plant Biology, 23, 3, 891 899 Xylem cavitation, leaf growth and leaf water potential in Eucalyptus globulus clones under well-watered and drought

More information

Water use efficiency in agriculture

Water use efficiency in agriculture Water use efficiency in agriculture Bill Davies The Lancaster Environment Centre, UK Summary Introduction and definitions Impacts of stomata, environment and leaf metabolism on WUE Estimating WUE and modifications

More information

fr>uafcji *> \E % jw r"'''f^,""'i;- ~^H^^

fr>uafcji *> \E % jw r'''f^,'i;- ~^H^^ NAME DATE Carolina Transpiration Kit for AP Biology Imagine that your family has received a bouquet of cut flowers as a gift. You place the flowers in a vase with a small volume of water, and return the

More information

Xylem Hydraulics - Theory. I. Hydraulic Architecture Definition: i) The structure of water transport system in plants. (Tyree and Ewers 1991) V t

Xylem Hydraulics - Theory. I. Hydraulic Architecture Definition: i) The structure of water transport system in plants. (Tyree and Ewers 1991) V t Plant Ecophysiological Measurement Techniques - BOT 6935 January 27, 2014 - Theory. I. Hydraulic Architecture Definition: i) The structure of water transport system in plants. (Tyree and Ewers 1991) ii)

More information

Terrestrial land surfacesa pot pourri

Terrestrial land surfacesa pot pourri CALTECH JPL Center for Climate Sciences March 26, 2018 Terrestrial land surfacesa pot pourri Graham Farquhar Australian National University What do we want from our models? Timescale is a key issue What

More information

References. 1 Introduction

References. 1 Introduction 1 Introduction 3 tion, conservation of soil water may result in greater soil evaporation, especially if the top soil layers remain wetter, and the full benefit of sustained plant physiological activity

More information

Xylem embolism and stomatal regulation in two rubber clones (Hevea brasiliensis Muell. Arg.)

Xylem embolism and stomatal regulation in two rubber clones (Hevea brasiliensis Muell. Arg.) Page 1 of 11 Trees Structure and Function Springer-Verlag 2003 Original Article Xylem embolism and stomatal regulation in two rubber clones (Hevea brasiliensis Muell. Arg.) Krissada Sangsing 1, 2, Poonpipope

More information

Hormonal and other chemical effects on plant growth and functioning. Bill Davies Lancaster Environment Centre, UK

Hormonal and other chemical effects on plant growth and functioning. Bill Davies Lancaster Environment Centre, UK Hormonal and other chemical effects on plant growth and functioning Bill Davies Lancaster Environment Centre, UK Integrating the impacts of soil drought and atmospheric stress High radiant load Reduced

More information

" " " " " " " " " " " " " " " " " " " " " The Effect of Sunlight on the Rate of Photosynthesis in Trachelospermum Jasmenoides" Nika Vafadari "

                     The Effect of Sunlight on the Rate of Photosynthesis in Trachelospermum Jasmenoides Nika Vafadari The Effect of Sunlight on the Rate of Photosynthesis in Trachelospermum Jasmenoides Nika Vafadari Biology Lab 111 Section 05 1 2 Introduction Different types of plant species, such as CAM plants, have

More information

Carbon Cycle, part 2 Ecophysiology of Leaves. ESPM 111 Ecosystem Ecology. Outline

Carbon Cycle, part 2 Ecophysiology of Leaves. ESPM 111 Ecosystem Ecology. Outline Carbon Cycle, part 2 Ecophysiology of Leaves Dennis Baldocchi ESPM UC Berkeley Courtesy of Rob Jackson, Duke 3/13/2013 Outline Photosynthetic Pathways and Cycles Environmental Physiology of Photosynthesis

More information

Carbon isotope variation in Douglas-fir foliage: improving the δ 13 C--climate relationship

Carbon isotope variation in Douglas-fir foliage: improving the δ 13 C--climate relationship Tree Physiology 15, 657--663 1995 Heron Publishing----Victoria, Canada Carbon isotope variation in Douglas-fir foliage: improving the δ 13 C--climate relationship JEANNE A. PANEK and RICHARD H. WARING

More information

Downward shoot positioning affects water transport in field-grown grapevines

Downward shoot positioning affects water transport in field-grown grapevines Vitis 39 (2), 49 53 (2000) Downward shoot positioning affects water transport in field-grown grapevines C. LOVISOLO 1) and A. SCHUBERT 2) 1) Dipartimento di Colture Arboree dell Università di Torino, Italia

More information

Stomatal conductance and leaf water potential responses to hydraulic conductance variation in Pinus pinaster seedlings

Stomatal conductance and leaf water potential responses to hydraulic conductance variation in Pinus pinaster seedlings Trees (7) 21:371 378 DOI 1.17/s468-7-13-6 ORIGINAL PAPER Stomatal conductance and leaf water potential responses to hydraulic conductance variation in Pinus pinaster seedlings Francesco Ripullone Æ Maria

More information

Water deficits and hydraulic limits to leaf water supply

Water deficits and hydraulic limits to leaf water supply Blackwell Science, LtdOxford, UK PCEPlant, Cell and Environment16-825Blackwell Science Ltd 21 25 799 Water deficits and hydraulic limits to leaf water supply J. S. Sperry et al. 1.146/j.16-825.21.799.x

More information

Water relations in tree physiology: where to from here? *Corresponding author: Joe Landsberg

Water relations in tree physiology: where to from here? *Corresponding author: Joe Landsberg 1 1 2 Water relations in tree physiology: where to from here? 3 4 Joe Landsberg 1* and Richard Waring 2 5 6 7 8 9 1 Withycombe, Church Lane, Mt Wilson, NSW 2786, Australia 2 College of Forestry, Oregon

More information

Basic stoichiometric equation on photosynthesis and the production of sugar and oxygen via the consumption of CO2, water, and light

Basic stoichiometric equation on photosynthesis and the production of sugar and oxygen via the consumption of CO2, water, and light 1 2 Basic stoichiometric equation on photosynthesis and the production of sugar and oxygen via the consumption of CO2, water, and light 3 Several pathways exist for fixing CO2 into sugar 4 Photosynthesis

More information

Water Acquisition and Transport - Whole Plants. 3 possible pathways for water movement across the soil-plant-atmosphere continuum

Water Acquisition and Transport - Whole Plants. 3 possible pathways for water movement across the soil-plant-atmosphere continuum Water transport across the entire soil-plant-atmosphere continuum Water Acquisition and Transport - Whole Plants 3 possible pathways for water movement across the soil-plant-atmosphere continuum Apoplast

More information

Effects of rising temperatures and [CO 2 ] on physiology of tropical forests

Effects of rising temperatures and [CO 2 ] on physiology of tropical forests Effects of rising temperatures and [CO 2 ] on physiology of tropical forests We are happy to advise that reports of our impending demise may have been very much exaggerated Jon Lloyd and Graham Farquhar

More information

Sap flow technique as a tool for irrigation schedule in grapevines: control of the plant physiological status

Sap flow technique as a tool for irrigation schedule in grapevines: control of the plant physiological status Sap flow technique as a tool for irrigation schedule in grapevines: control of the plant physiological status Pons P.J., Truyols M., Flexas J., Cifre J., Medrano H., Ribas-Carbó M. in López-Francos A.

More information

Dry season conditions determine wet season water use in the wet dry tropical savannas of northern Australia

Dry season conditions determine wet season water use in the wet dry tropical savannas of northern Australia Tree Physiology 20, 1219 1226 2000 Heron Publishing Victoria, Canada Dry season conditions determine wet season water use in the wet dry tropical savannas of northern Australia D. EAMUS, 1,2 A. P. O GRADY

More information

The Effect of Night Temperature on Cotton Reproductive Development

The Effect of Night Temperature on Cotton Reproductive Development The Effect of Night Temperature on Cotton Reproductive Development Item Type text; Article Authors Zeiher, Carolyn A.; Brown, Paul W.; Silvertooth, Jeffrey C.; Matumba, Nkonko; Mitton, Nancy Publisher

More information

Variation in hydraulic architecture and gas-exchange in two desert sub-shrubs, Hymenoclea salsola (T. & G.) and Ambrosia dumosa (Payne)

Variation in hydraulic architecture and gas-exchange in two desert sub-shrubs, Hymenoclea salsola (T. & G.) and Ambrosia dumosa (Payne) Oecologia (2000) 125:1 10 Springer-Verlag 2000 Jonathan P. Comstock Variation in hydraulic architecture and gas-exchange in two desert sub-shrubs, Hymenoclea salsola (T. & G.) and Ambrosia dumosa (Payne)

More information

A test of gas exchange measurements on excised canopy branches of ten tropical tree species

A test of gas exchange measurements on excised canopy branches of ten tropical tree species PHOTOSYNTHETICA 41 (3): 343-347, 2003 A test of gas exchange measurements on excised canopy branches of ten tropical tree species L.S. SANTIAGO and S.S. MULKEY Department of Botany, University of Florida,

More information

Temperature effects on hydraulic conductance and water relations of Quercus robur L.

Temperature effects on hydraulic conductance and water relations of Quercus robur L. Journal of Experimental Botany, Vol. 51, No. 348, pp. 1255 1259, July 2000 Temperature effects on hydraulic conductance and water relations of Quercus robur L. Hervé Cochard1,3, Rodolphe Martin2, Patrick

More information

ASSESSMENT OF WATER STATUS IN TREES FROM MEASUREMENTS OF STOMATAL CONDUCTANCE AND WATER POTENTIAL

ASSESSMENT OF WATER STATUS IN TREES FROM MEASUREMENTS OF STOMATAL CONDUCTANCE AND WATER POTENTIAL No. 1 159 ASSESSMENT OF WATER STATUS IN TREES FROM MEASUREMENTS OF STOMATAL CONDUCTANCE AND WATER POTENTIAL DAVID WHITEHEAD Forest Research Institute, Rotorua, New Zealand (Received for publication 24

More information

Temperature and light as ecological factors for plants

Temperature and light as ecological factors for plants PLB/EVE 117 Plant Ecology Fall 2005 1 Temperature and light as ecological factors for plants I. Temperature as an environmental factor A. The influence of temperature as an environmental factor is pervasive

More information

Description of 3-PG. Peter Sands. CSIRO Forestry and Forest Products and CRC for Sustainable Production Forestry

Description of 3-PG. Peter Sands. CSIRO Forestry and Forest Products and CRC for Sustainable Production Forestry Description of 3-PG Peter Sands CSIRO Forestry and Forest Products and CRC for Sustainable Production Forestry 1 What is 3-PG? Simple, process-based model to predict growth and development of even-aged

More information

Water Relations in Viticulture BRIANNA HOGE AND JIM KAMAS

Water Relations in Viticulture BRIANNA HOGE AND JIM KAMAS Water Relations in Viticulture BRIANNA HOGE AND JIM KAMAS Overview Introduction Important Concepts for Understanding water Movement through Vines Osmosis Water Potential Cell Expansion and the Acid Growth

More information

The three principal organs of seed plants are roots, stems, and leaves.

The three principal organs of seed plants are roots, stems, and leaves. 23 1 Specialized Tissues in Plants Seed Plant Structure The three principal organs of seed plants are roots, stems, and leaves. 1 of 34 23 1 Specialized Tissues in Plants Seed Plant Structure Roots: absorb

More information

Stomatal regulation in Douglas fir following a fungal-mediated chronic reduction in leaf area

Stomatal regulation in Douglas fir following a fungal-mediated chronic reduction in leaf area Trees (2003) 17:485 491 DOI 10.1007/s00468-003-0262-2 ORIGINAL ARTICLE Daniel K. Manter Kathleen L. Kavanagh Stomatal regulation in Douglas fir following a fungal-mediated chronic reduction in leaf area

More information

Using Pressure-Volume Analysis to Determine the Effect of the Hydrostatic Gradient on Cell Turgidity

Using Pressure-Volume Analysis to Determine the Effect of the Hydrostatic Gradient on Cell Turgidity Using Pressure-Volume Analysis to Determine the Effect of the Hydrostatic Gradient on Cell Turgidity Sarah Elizabeth Reed Global Change Education Program 2001 Mentor: Barbara J. Bond Abstract. The physiological

More information

Comparative Plant Ecophysiology

Comparative Plant Ecophysiology Comparative Plant Ecophysiology 2. Plant traits and climate factors that form bases for eco- physiological comparison 3. Life form comparisons of: Stomatal conductance Photosynthesis Xylem Anatomy Leaf

More information

Forestry and Forest Products

Forestry and Forest Products Forestry and Forest Products TECHNICAL REPORT No. 120 Carbon Isotope Discrimination Of Irrigated And Fertilised Eucalyptus grandis Grown In Different Climates. Ronni L. Korol, Miko U.F. Kirschbaum and

More information

Optical measurement of Leaf Area Index at Falkenberg site. G. Vogel, U. Rummel and J.-P.Schulz

Optical measurement of Leaf Area Index at Falkenberg site. G. Vogel, U. Rummel and J.-P.Schulz Optical measurement of Leaf Area Index at Falkenberg site G. Vogel, U. Rummel and J.-P.Schulz Three variants Plant type/land use type LAI_Wikipedia LAImax- GLC2009 farmland (winter) 0,2 intensively used

More information

New type of vulnerability curve gives insight in the hydraulic capacitance and conductivity of the xylem

New type of vulnerability curve gives insight in the hydraulic capacitance and conductivity of the xylem New type of vulnerability curve gives insight in the hydraulic capacitance and conductivity of the xylem Lidewei L Vergeynst, Jan Bogaerts, Annelies Baert, Lies Kips and Kathy Steppe Laboratory of Plantecology,

More information

in angiosperms 10/29/08 Roots take up water via roots Large surface area is needed Roots branch and have root hairs Cortex structure also helps uptake

in angiosperms 10/29/08 Roots take up water via roots Large surface area is needed Roots branch and have root hairs Cortex structure also helps uptake in angiosperms A. Root System Roots take up water via roots Large surface area is needed Roots branch and have root hairs Cortex structure also helps uptake 1 B. Minerals Nitrogen (NO 3-,NH 4+ ) Potassium

More information

Tree Physiology. Sara Rose

Tree Physiology. Sara Rose Tree Physiology Sara Rose What is a Tree? U.S. Forest Service Woody plants that have well-developed stems and that usually are more than 12 feet tall at maturity. Merriam-Webster A woody perennial plant

More information

Pinaceae in the Pacific Northern USA

Pinaceae in the Pacific Northern USA Functional Ecology 2000 Ecological implications of xylem cavitation for several Blackwell Science, Ltd Pinaceae in the Pacific Northern USA J. PIÑOL* and A. SALA *Centre de Recerca Ecològica i Aplicacions

More information

Relationship between Leaf Water Potential and Photosynthesis in Rice Plants

Relationship between Leaf Water Potential and Photosynthesis in Rice Plants Relationship between Leaf Water Potential and Photosynthesis in Rice Plants By KUNI ISHIHARA and HIDEO SAITO Faculty of Agriculture, Tokyo University of Agriculture and Technology (Saiwaicho,Fuchu, Tokyo,

More information

Mlchio KANECHI, Naotsugu UCHIDA, Takeshl YASUDA and Tadashi YAMAGUCHI Graduate School of Science and Technology, Kobe University, Rokko, Kobe 657

Mlchio KANECHI, Naotsugu UCHIDA, Takeshl YASUDA and Tadashi YAMAGUCHI Graduate School of Science and Technology, Kobe University, Rokko, Kobe 657 Japan. J. Trop. Agr. 32 (1) : 16-21, 1988 Relationships between Leaf Water Potential and Photosynthesis of Coffea arabica L. Grown under Various Environmental Conditions as Affected by Withholding Irrigation

More information

Introduction. Populus trichocarpa TORR. and GRAY. By M. G. R. CANNELL and S. C. WILLETT

Introduction. Populus trichocarpa TORR. and GRAY. By M. G. R. CANNELL and S. C. WILLETT Shoot Growth Phenology, Dry Matter Distribution and Root: Shoot Ratios of Provenances of Populus trichocarpa, Picea sitchensis and Pinus contorta growing in Scotland By M. G. R. CANNELL and S. C. WILLETT

More information

CAMBIUM, meristem, heartwood, and lenticel are

CAMBIUM, meristem, heartwood, and lenticel are Examining the Structures of a Tree CAMBIUM, meristem, heartwood, and lenticel are some terms that may be new to you. These terms are used to describe various tree structures. Not surprisingly, many terms

More information

WATER RELATIONS OF WHEAT ALTERNATED BETWEEN TWO ROOT TEMPERATURES

WATER RELATIONS OF WHEAT ALTERNATED BETWEEN TWO ROOT TEMPERATURES Nezo Phytol. (1979) 82, 89-96 89 WATER RELATIONS OF WHEAT ALTERNATED BETWEEN TWO ROOT TEMPERATURES BY M. B. KIRKHAM Department of Agrofiomy, Oklahoma State University^ StiHwater, Oklahoma 74074, U,S.A.

More information

Drought Tolerant Criterion of Wheat Genotypes Using Carbon Isotopes Discrimination Technique

Drought Tolerant Criterion of Wheat Genotypes Using Carbon Isotopes Discrimination Technique Journal of Earth Science and Engineering 5 (2015) 256-261 doi: 10.17265/2159-581X/2015. 01. 005 D DAVID PUBLISHING Drought Tolerant Criterion of Wheat Genotypes Using Carbon Isotopes Discrimination Technique

More information

Spatial scales resolved. Four hierarchical scales: Leaf. Plant. Canopy. Landscape

Spatial scales resolved. Four hierarchical scales: Leaf. Plant. Canopy. Landscape Spatial scales resolved Four hierarchical scales: Leaf Plant Canopy Landscape Canopy embedded in a turbulent air stream - extracting momentum and releasing/uptaking scalars at the solid-fluid interface.

More information

Analysis and modeling of gas exchange processes in Scaevola aemula

Analysis and modeling of gas exchange processes in Scaevola aemula Scientia Horticulturae 114 (2007) 170 176 www.elsevier.com/locate/scihorti Analysis and modeling of gas exchange processes in Scaevola aemula Soo-Hyung Kim a, *, Paul R. Fisher b, J. Heinrich Lieth c a

More information

6 Heat Ratio Method Theory

6 Heat Ratio Method Theory 6 Heat Ratio Method Theory The Heat Ratio Method (HRM) can measure both sap velocity (Vs) and volumetric water flow in xylem tissue using a short pulse of heat as a tracer. It is a modification of the

More information

Avocado Tree Physiology Understanding the Basis of Productivity

Avocado Tree Physiology Understanding the Basis of Productivity Avocado Tree Physiology Understanding the Basis of Productivity R. L. Heath, M. L. Arpaia UC, Riverside M. V. Mickelbart Purdue University Raw Materials Labor Product Light Carbon Dioxide Temperature Water

More information

AP Biology Transpiration and Stomata

AP Biology Transpiration and Stomata AP Biology Transpiration and Stomata Living things must exchange matter with the environment to survive, Example: Gas Exchange in Plants photosynthesis cellular respiration 1. During which hours does a

More information

Evaluating shrub architectural performance in sun and shade environments with the 3-D model Y-plant: are there optimal strategies?

Evaluating shrub architectural performance in sun and shade environments with the 3-D model Y-plant: are there optimal strategies? Evaluating shrub architectural performance in sun and shade environments with the 3-D model Y-plant: are there optimal strategies? Robert W. Pearcy 1, Hiroyuki Muraoka 2 and Fernando Valladares 3 1 Section

More information

Resource acquisition and transport in vascular plants

Resource acquisition and transport in vascular plants Resource acquisition and transport in vascular plants Overview of what a plant does Chapter 36 CO 2 O 2 O 2 and and CO 2 CO 2 O 2 Sugar Light Shoots are optimized to capture light and reduce water loss

More information

Growth indices and stomatal control of transpiration in Acacia koa stands planted at different densities

Growth indices and stomatal control of transpiration in Acacia koa stands planted at different densities Tree Physiology 16, 607--615 1996 Heron Publishing----Victoria, Canada Growth indices and stomatal control of transpiration in Acacia koa stands planted at different densities FREDERICK C. MEINZER, 1 JAMES

More information

Contents. 1. Evaporation

Contents. 1. Evaporation Contents 1 Evaporation 1 1a Evaporation from Wet Surfaces................... 1 1b Evaporation from Wet Surfaces in the absence of Advection... 4 1c Bowen Ratio Method........................ 4 1d Potential

More information

Interactions between ozone and drought stress in plants: mechanisms and implications. Sally Wilkinson and William J. Davies, Lancaster University

Interactions between ozone and drought stress in plants: mechanisms and implications. Sally Wilkinson and William J. Davies, Lancaster University Interactions between ozone and drought stress in plants: mechanisms and implications Sally Wilkinson and William J. Davies, Lancaster University STOMATA: At the leaf surface water is lost to the atmosphere

More information

Research Proposal: Tara Gupta (CSE Style)

Research Proposal: Tara Gupta (CSE Style) Research Proposal: Tara Gupta (CSE Style) Specific and informative title, name, and other relevant information centered on title page Field Measurements of Photosynthesis and Transpiration Rates in Dwarf

More information

Objective: To teach students the basic anatomy of trees and how different cells function to promote tree survival.

Objective: To teach students the basic anatomy of trees and how different cells function to promote tree survival. Objective: To teach students the basic anatomy of trees and how different cells function to promote tree survival. Materials: Paper Markers/Crayons Tree Anatomy Labels Tree Cookie examples White Paper

More information

Lecture notes on stomatal conductance. Agron 516: Crop physiology. Dr. Mark Westgate.

Lecture notes on stomatal conductance. Agron 516: Crop physiology. Dr. Mark Westgate. Lecture notes on stomatal conductance. Agron 516: Crop physiology. Dr. Mark Westgate. Diurnal variation of stomatal conductance has direct consequences for leaf and canopy gas exchange Measure diurnal

More information

Published Research in Journal: Plant and Soil - PLANT SOIL, vol. 332, no. 1, pp , 2010

Published Research in Journal: Plant and Soil - PLANT SOIL, vol. 332, no. 1, pp , 2010 Published Research in Journal: Plant and Soil - PLANT SOIL, vol. 332, no. 1, pp. 147-162, 2010 Enhancement of morphological, anatomical and physiological characteristics of seedlings of the mangrove Avicennia

More information

Effects of Water Stress during Growth on Xylem Anatomy, Xylem Functioning and Vase Life in Three Zinnia elegans Cultivars

Effects of Water Stress during Growth on Xylem Anatomy, Xylem Functioning and Vase Life in Three Zinnia elegans Cultivars Effects of Water Stress during Growth on Xylem Anatomy, Xylem Functioning and Vase Life in Three Zinnia elegans Cultivars P. Twumasi 1, 2, W. van Ieperen 1*, E.J. Woltering 3, A.M.C. Emons 2, J.H.N. Schel

More information

Tree Anatomy. Arlington and Alexandria Tree Stewards Jim McGlone Urban Forest Conservationist Virginia Department of Forestry

Tree Anatomy. Arlington and Alexandria Tree Stewards Jim McGlone Urban Forest Conservationist Virginia Department of Forestry Tree Anatomy Arlington and Alexandria Tree Stewards Jim McGlone Urban Forest Conservationist Virginia Department of Forestry Tree Structure https://thelandscapebelowground.com/wp-content/uploads/2018/03/poster-tree-100dpi-black-and-white.jpg

More information

Nutrient Cycling in Land Vegetation and Soils

Nutrient Cycling in Land Vegetation and Soils Nutrient Cycling in Land Vegetation and Soils OCN 401 - Biogeochemical Systems 13 September 2012 Reading: Schlesinger, Chapter 6 Outline 1. The annual Intrasystem Nutrient Cycle 2. Mass balance of the

More information

Comparison of physiological responses of pearl millet and sorghum to water stress

Comparison of physiological responses of pearl millet and sorghum to water stress Proc. Indian Acad. Sci. (Plant Sci.), Vol. 99, No. 6, December 1989, pp. 517-522. (~ Printed in India. Comparison of physiological responses of pearl millet and sorghum to water stress V BALA SUBRAMANIAN

More information

Name Date Block. Plant Structures

Name Date Block. Plant Structures Name Date Block What are the Functions of Roots, Stems, and Leaves? Plant Structures Each part of a plant plays an important role in its structure and function. Roots, stems, and leaves are just three

More information

The spatial pattern of air seeding thresholds in mature sugar maple trees

The spatial pattern of air seeding thresholds in mature sugar maple trees Blackwell Science, LtdOxford, UKPCEPlant, Cell and Environment0016-8025Blackwell Science Ltd 2005? 2005 28?10821089 Original Article Plant, Cell and Environment (2005) 28, 1082 1089 Air seeding threshold

More information

To Understand How Trees Decline and Die, We Must: What is Stress? Tree Physiology. Understand stress and how it affects trees. Why Do Trees Die?

To Understand How Trees Decline and Die, We Must: What is Stress? Tree Physiology. Understand stress and how it affects trees. Why Do Trees Die? To Understand How Trees Decline and Die, We Must: Why Do Trees Die? Rex Bastian, Ph.D. The Davey Tree Expert Co./The Care of Trees Wheeling, IL Understand stress and how it affects trees» To do this, we

More information

What is a TREE? Woody plant (stems, branches, roots) Long lived (typically 100 s of yrs. sometimes 10

What is a TREE? Woody plant (stems, branches, roots) Long lived (typically 100 s of yrs. sometimes 10 What is a TREE? Woody plant (stems, branches, roots) Long lived (typically 100 s of yrs. sometimes 10 s or 1000 s) Single main axis (stem, trunk, bole) Typically 20-30 tall at maturity (note variability)

More information

LEAF PHOTOSYNTHETIC CAPACITY, CHLOROPHYLL CONTENT, AND SPECIFIC LEAF NITROGEN RELATIONSHIPS IN SUNFLOWER CROPS PRIOR TO ANTHESIS

LEAF PHOTOSYNTHETIC CAPACITY, CHLOROPHYLL CONTENT, AND SPECIFIC LEAF NITROGEN RELATIONSHIPS IN SUNFLOWER CROPS PRIOR TO ANTHESIS LEAF PHOTOSYNTHETIC CAPACITY, CHLOROPHYLL CONTENT, AND SPECIFIC LEAF NITROGEN RELATIONSHIPS IN SUNFLOWER CROPS PRIOR TO ANTHESIS M. Cecilia Rousseaux, IFEVA Facultad de Agronomía, Universidad de Buenos

More information

BioWash as an Adjuvant, Translocation Promoter, and Cationic Exchange Stimulator Overview of Processes within the Plant

BioWash as an Adjuvant, Translocation Promoter, and Cationic Exchange Stimulator Overview of Processes within the Plant BioWash as an Adjuvant, Translocation Promoter, and Cationic Exchange Stimulator Overview of Processes within the Plant Photosynthesis is the primary driver of the plant. Through a series of complex steps,

More information

TRANSPIRATION COEFFICIENT AND TRANSPIRATION RATE OF THREE GRAIN SPECIES IN GROWTH CHAMBERS

TRANSPIRATION COEFFICIENT AND TRANSPIRATION RATE OF THREE GRAIN SPECIES IN GROWTH CHAMBERS REPRINTED FROM: JAARB. I.B.S. 96, 73-8 TRANSPIRATION COEFFICIENT AND TRANSPIRATION RATE OF THREE GRAIN SPECIES IN GROWTH CHAMBERS C. T. DE WIT and TH. ALBERDA lnstituut voor Biologisch en Scheikundig Onderzoek

More information

Radiation transfer in vegetation canopies Part I plants architecture

Radiation transfer in vegetation canopies Part I plants architecture Radiation Transfer in Environmental Science with emphasis on aquatic and vegetation canopy medias Radiation transfer in vegetation canopies Part I plants architecture Autumn 2008 Prof. Emmanuel Boss, Dr.

More information

How drought stress and CO2 concentration influence stomatal conductance and photosynthesis? Abstract. Introduction

How drought stress and CO2 concentration influence stomatal conductance and photosynthesis? Abstract. Introduction How drought stress and CO2 concentration influence stomatal conductance and photosynthesis? Simon Keck 1, Julian Müller 1, Dominik Guttschick 1, Kaisa Pajusalu 2, Elodie Quer 3, Maria Majekova 4 1 University

More information

LI-COR LI-6400XT Training Course

LI-COR LI-6400XT Training Course LI-COR LI-6400XT Training Course Lingling Yuan and Shannon Loriaux, LI-COR Science and Support September 15-16, 2015 Beijing, China Tentative Agenda Day 1 Theory and hardware overview Preparation Checklists

More information

Finite element tree crown hydrodynamics model (FETCH) using porous media flow within branching elements: A new representation of tree hydrodynamics

Finite element tree crown hydrodynamics model (FETCH) using porous media flow within branching elements: A new representation of tree hydrodynamics WATER RESOURCES RESEARCH, VOL. 41,, doi:10.1029/2005wr004181, 2005 Finite element tree crown hydrodynamics model (FETCH) using porous media flow within branching elements: A new representation of tree

More information

Coordinating leaf functional traits with branch hydraulic conductivity: resource substitution and implications for carbon gain

Coordinating leaf functional traits with branch hydraulic conductivity: resource substitution and implications for carbon gain Tree Physiology 28, 1169 1177 2008 Heron Publishing Victoria, Canada Coordinating leaf functional traits with branch hydraulic conductivity: resource substitution and implications for carbon gain DANIEL

More information

GENETIC ANALYSES OF ROOT SYSTEM DEVELOPMENT IN THE TOMATO CROP MODEL

GENETIC ANALYSES OF ROOT SYSTEM DEVELOPMENT IN THE TOMATO CROP MODEL GENETIC ANALYSES OF ROOT SYSTEM DEVELOPMENT IN THE TOMATO CROP MODEL Kelsey Hoth 1 Dr. Maria Ivanchenko 2 Bioresourse Research 1, Department of Botany and Plant Physiology 2, Oregon State University, Corvallis,

More information

K. A. McCULLOH,* J. S. SPERRY* and F. R. ADLER*

K. A. McCULLOH,* J. S. SPERRY* and F. R. ADLER* Functional Ecology 2004 Murray s law and the hydraulic vs mechanical functioning Blackwell Publishing, Ltd. of wood K. A. McCULLOH,* J. S. SPERRY* and F. R. ADLER* *Department of Biology, and Department

More information

Earth Has a Rich Diversity of Plants. Plant Structure, Nutrition, and Transport. Angiosperms: Monocots and Dicots. Angiosperms: Dicots

Earth Has a Rich Diversity of Plants. Plant Structure, Nutrition, and Transport. Angiosperms: Monocots and Dicots. Angiosperms: Dicots Plant Structure, Nutrition, and Transport Earth Has a Rich Diversity of Plants There are over 280,000 different plant species organized into four major groups: bryophytes (mosses), seedless vascular plants,

More information

Shoot growth patterns in saplings of Cleyera japonica in relation to light and architectural position

Shoot growth patterns in saplings of Cleyera japonica in relation to light and architectural position Tree Physiology 23, 67 71 2003 Heron Publishing Victoria, Canada Shoot growth patterns in saplings of Cleyera japonica in relation to light and architectural position ARATA ANTONIO SUZUKI 1,2 1 Laboratory

More information

Oxygen and Hydrogen in Plants

Oxygen and Hydrogen in Plants Oxygen and Hydrogen in Plants Outline: Environmental factors Fractionation associated with uptake of water Metabolic Fractionation C3, CAM and C4 plants Environmental factors Regional Precipitation d 18

More information