A comparison of three approaches to modeling leaf gas exchange in annually drought-stressed ponderosa pine forests

We tested, compared and modified three models of stomatal conductance at the leaf level in a forest ecosystem where drought stress is a major factor controlling forest productivity. The models were tested against 2 years (1999 and 2000) of leaf-level measurements on ponderosa pine (Pinus ponderosa Dougl. ex Laws.) growing in the Mediterranean climate of California, USA. The Ball, Woodrow and Berry (1987) (BWB) model was modified to account for soil water stress. Among the models, results of the modified BWB model were in the closest agreement with observations (r2 = 0.71). The Jarvis (1976) model showed systematic simulation errors related to vapor pressure deficit (r2 = 0.65). Results of the Williams, Rastetter, Fernandes et al. (1996) (SPA) model showed the poorest correlation with empirical data, but this model has only one calibration parameter (r2 = 0.60). Sensitivity analyses showed that, in all three models, predictions of stomatal conductance were most responsive to photosynthetically active radiation and soil water content. Stomatal conductance showed little sensitivity to vapor pressure deficit in the Jarvis model, whereas in both the BWB and SPA models, vapor pressure deficit (or relative humidity) was the third most important variable. Parameterization of the SPA model was in accordance with the parameterization of the modified BWB model, although the two models differ greatly. Measured and modeled results indicate that stomatal behavior is not water conservative during spring; however, during summer, when soil water content is low and vapor pressure deficit is high, stomatal conductance decreases and, according to the models, intrinsic water- use efficiency increases.     

Nocturnal transpiration in riparian Tamarix thickets authenticated by sap flux, eddy covariance and leaf gas exchange measurements

Tamarix chinensis Lour., which is common throughout the southwestern USA, is a phreatophytic riparian tree capable of high water use. We investigated temporal congruence between daily total evapotranspiration (E) estimated from stem sap flux (J(s)) measurements (E(sf)) and eddy covariance (E(cv)), both seasonally and immediately following rain events, and used measurements of leaf-level gas exchange, stem water content and diurnal changes in leaf water potential to track drivers of transpiration. In one study, conducted near the end of the growing season in a pure T. chinensis stand adjacent to the Rio Grande River in central New Mexico, nighttime E(sf) as a proportion of daily E(sf) increased with water availability to a peak of 36.6%. High nighttime E(sf) was associated with underestimates of nighttime E(cv). A second study, conducted in west Texas, beside the Pecos River, investigated the relationships between nighttime J(s) and stem tissue rehydration, on the one hand, and nighttime E, on the other hand. Leaf gas exchange measurements and stomatal impressions suggested that nighttime J(s) was primarily attributed to concurrent transpiration, although there were small overnight changes in stem water content. Both vapor pressure deficit and soil water availability were positively related to nighttime J(s), especially following rainfall events. Thus, both studies indicate that T. chinensis can transpire large amounts at night, a fact that must be considered when attempting to quantify E either by eddy covariance or sap flux methods.     

Accentuation of gas exchange gradients in flushes of ponderosa pine exposed to ozone

Two-year-old ponderosa pine seedlings (Pinus ponderosa Laws.) were exposed to episodic O(3) concentrations in open-top chambers for two consecutive growing seasons (June through September of 1990 and 1991). Near the end of the second season of O(3) exposure, gas exchange was measured on needles of surviving flushes at saturating CO(2) and photosynthetic photon flux density (PPFD). Both photosynthetic capacity (A(sat)) and stomatal conductance to water vapor (g(wv)) declined linearly with needle age but differences within a flush were also found. Gas exchange rates of needles from the base of the current-year flush were significantly lower than those of needles from the top of the flush, even though age differences between needles were negligible. Although most measurements were conducted at saturating CO(2), similar patterns of gas exchange were also found at 350 micro mol mol(-1) CO(2), indicating that photosynthesis of needles at the bottom of the flush was more strongly affected by O(3) than that of needles at the top of the flush, even though the potential for O(3) uptake was probably less in needles at the bottom of the flush because of reduced stomatal conductance. Carboxylation efficiency (deltaA/deltaC(i)), the linear slope of the A/C(i) response, was highly correlated with A(sat), varying with needle age, needle position in the flush and O(3) exposure, but the magnitude of the reductions was greater than for A(sat). We conclude that susceptibility to O(3) damage among needles of an individual seedling varies not only with needle age but also with needle position, and that reductions in photosynthetic capacity may not be directly attributable to increased uptake of the pollutant. The data also indicate the need to consider within-flush variation when estimating whole-plant carbon gain and O(3) uptake.     

Carbon isotopic composition, gas exchange, and growth of three populations of ponderosa pine differing in drought tolerance

Effects of water supply on gas exchange, carbon isotopic composition, and relative growth rate were compared among seedlings from three populations of ponderosa pine (Pinus ponderosa Dougl. ex Laws.) grown in a controlled environment chamber. The three populations were chosen to represent high, moderate and low drought tolerance. There was no indication that drought tolerance was related to high water-use efficiency. Populations differed (P < 0.05) in relative growth rate (RGR), but did not differ (P > 0.10) in gas exchange variables or carbon isotope ratio (delta(13)C). Well-watered seedlings had significantly higher RGR, xylem pressure potential (Psi(xpp)), net photosynthesis (A), stomatal conductance to water vapor (g), and lower delta(13)C and instantaneous water-use efficiency than water-stressed seedlings. With decreasing Psi(xpp), A decreased linearly, whereas g decreased exponentially. Seedlings of the highly drought-tolerant population were more sensitive to water availability than seedlings from the other populations; they used water quickly when water was available, but closed their stomata in response to water stress. We conclude that, in ponderosa pine, the drought avoidance mechanism is more important for survival and growth in arid and semiarid environments than the efficient use of water.     

Fall lifting and long-term freezer storage of ponderosa pine seedlings: effects on post-storage leaf water potential, stomatal conductance, and root growth potential

Post-storage water relations, stomatal conductance, and root growth potential were examined in ponderosa pine (Pinus ponderosa Dougl. ex Laws.) seedlings from high- and low-elevation seed sources that had been lifted either in October or November and freezer stored, or in March, and then grown hydroponically in a greenhouse for 31 days. Seedlings lifted in October had poor root initiation (< 17 new roots per seedling), low predawn leaf water potentials (< -1.5 MPa), and low stomatal conductance (7.10 mmol m(-2) s(-1)) compared with seedlings lifted in November or March. There was little difference in post-storage water relations and stomatal conductance between seedlings lifted in November and those lifted in March. Throughout the 31-day test, seedlings from the high-elevation seed source produced 3-9 times more new roots, had higher predawn leaf water potentials (-0.6 to -0.7 MPa versus -1.1 to -1.6 MPa), and 1.3-5 times greater stomatal conductance than seedlings from the low-elevation seed source. For all seedlings on Day 31, the number of new roots was significantly related to predawn leaf water potential (r(2) = 0.65) and stomatal conductance (r(2) = 0.82). Similarly, the dry weight of new roots per seedling on Day 31 accounted for a significant amount of the variation in predawn leaf water potential (r(2) = 0.81) and stomatal conductance (r(2) = 0.49).     

Assessing and modeling total height and diameter increment of ponderosa pine planted in Minnesota,USA

New Forests - Forest managers are increasingly planting non-native tree species that are adapted to anticipated future conditions such as increased droughts. This work quantified individual tree...     

Water limitations to carbon exchange in old-growth and young ponderosa pine stands

We investigated the impact of seasonal soil water deficit on the processes driving net ecosystem exchange of carbon (NEE) in old-growth and recently regenerating ponderosa pine (Pinus ponderosa Doug. ex Laws.) stands in Oregon. We measured seasonal patterns of transpiration, canopy conductance and NEE, as well as soil water, soil temperature and soil respiration. The old-growth stand (O) included two primary age classes (50 and 250 years), had a leaf area index (LAI) of 2.1 and had never been logged. The recently regenerating stand (Y) consisted predominantly of 14-year-old ponderosa pine with an LAI of 1.0. Both stands experienced similar meteorological conditions with moderately cold wet winters and hot dry summers. By August, soil volumetric water content within the upper 30 cm had declined to a seasonal minimum of 0.07 at both sites. Between April and June, both stands showed similar rates of transpiration peaking at 0.96 mm day(-1); thereafter, trees at the Y site showed increasing drought stress with canopy stomatal resistance increasing 6-fold by mid-August relative to values for trees at the O site. Over the same period, predawn water potential (psi(pd)) of trees at the Y site declined from -0.54 to -1.24 MPa, whereas psi(pd) of trees at the O site remained greater than -0.8 MPa throughout the season. Soil respiration at the O site showed a strong seasonal correlation with soil temperature with no discernible constraints imposed by declining soil water. In contrast, soil respiration at the Y site peaked before seasonal maximal soil temperatures and declined thereafter with declining soil water. No pronounced seasonal pattern in daytime NEE was observed at either site between April and September. At the Y site this behavior was driven by concurrent soil water limitations on soil respiration and assimilation, whereas there was no evidence of seasonal soil water limitations on either process at the O site.     

Bole girdling affects metabolic properties and root, trunk and branch hydraulics of young ponderosa pine trees

Effects of trunk girdling on seasonal patterns of xylem water status, water transport and woody tissue metabolic properties were investigated in ponderosa pine (Pinus ponderosa Dougl. ex P. Laws.) trees. At the onset of summer, there was a sharp decrease in stomatal conductance (g(s)) in girdled trees followed by a full recovery after the first major rainfall in September. Eliminating the root as a carbohydrate sink by girdling induced a rapid reversible reduction in g(s). Respiratory potential (a laboratory measure of tissue-level respiration) increased above the girdle (branches and upper trunk) and decreased below the girdle (lower trunk and roots) relative to control trees during the growing season, but the effect was reversed after the first major rainfall. The increase in branch respiratory potential induced by girdling suggests that the decrease in g(s) was caused by the accumulation of carbohydrates above the girdle, which is consistent with an observed increase in leaf mass per area in the girdled trees. Trunk girdling did not affect native xylem embolism or xylem conductivity. Both treated and control trunks experienced loss of xylem conductivity ranging from 10% in spring to 30% in summer. Girdling reduced xylem growth and sapwood to leaf area ratio, which in turn reduced branch leaf specific conductivity (LSC). The girdling-induced reductions in g(s) and transpiration were associated with a decrease in leaf hydraulic conductance. Two years after girdling, when root-to-shoot phloem continuity had been restored, girdled trees had a reduced density of new wood, which increased xylem conductivity and whole-tree LSC, but also vulnerability to embolism.     

Response of stomatal conductance to drought in ponderosa pine: implications for carbon and ozone uptake

To gain insight into the limitations imposed by a typical Mediterranean-climate summer drought on the uptake of carbon and ozone in the ponderosa pine (Pinus ponderosa Dougl. ex Laws.) ecosystem, we compared diurnal trends in leaf physiology of young trees in a watered and a control plot located in the Sierra Nevada Mountains, CA, USA (Blodgett Forest, 38 degrees 53' N, 120 degrees 37' W, 1315 m elevation). Predawn water potential of trees in the watered plot remained above -0.3 MPa throughout the growing season, whereas it dropped in the control plot from -0.24 to -0.52 MPa between late May and mid-August. Photosynthesis and stomatal conductance of trees in the watered plot were relatively insensitive to atmospheric vapor pressure deficit (VPD), whereas gas exchange of trees in the control plot varied with changes in soil water, VPD and temperature. Although the 1998 growing season was abnormally wet, we saw a pronounced drought effect at the control site. Over the 2 months following the onset of watering, carbon and ozone uptake were measured on three days at widely spaced intervals. Carbon uptake per unit leaf area by 1-year-old foliage of trees in the control plot was 39, 35 and 30% less, respectively, than in the watered plot, and estimated ozone deposition per unit leaf area (ozone concentration times stomatal conductance) was 36, 46 and 41% less.     

Co-optimal distribution of leaf nitrogen and hydraulic conductance in plant canopies

Leaf properties vary significantly within plant canopies, due to the strong gradient in light availability through the canopy, and the need for plants to use resources efficiently. At high light, photosynthesis is maximized when leaves have a high nitrogen content and water supply, whereas at low light leaves have a lower requirement for both nitrogen and water. Studies of the distribution of leaf nitrogen (N) within canopies have shown that, if water supply is ignored, the optimal distribution is that where N is proportional to light, but that the gradient of N in real canopies is shallower than the optimal distribution. We extend this work by considering the optimal co-allocation of nitrogen and water supply within plant canopies. We developed a simple 'toy' two-leaf canopy model and optimized the distribution of N and hydraulic conductance (K) between the two leaves. We asked whether hydraulic constraints to water supply can explain shallow N gradients in canopies. We found that the optimal N distribution within plant canopies is proportional to the light distribution only if hydraulic conductance, K, is also optimally distributed. The optimal distribution of K is that where K and N are both proportional to incident light, such that optimal K is highest to the upper canopy. If the plant is constrained in its ability to construct higher K to sun-exposed leaves, the optimal N distribution does not follow the gradient in light within canopies, but instead follows a shallower gradient. We therefore hypothesize that measured deviations from the predicted optimal distribution of N could be explained by constraints on the distribution of K within canopies. Further empirical research is required on the extent to which plants can construct optimal K distributions, and whether shallow within-canopy N distributions can be explained by sub-optimal K distributions.     

Age-related changes in ecosystem structure and function and effects on water and carbon exchange in ponderosa pine

As forests age, their structure and productivity change, yet in some cases, annual rates of water loss remain unchanged. To identify mechanisms that might explain such observations, and to determine if widely different age classes of forests differ functionally, we examined young (Y, approximately 25 years), mature (M, approximately 90 years) and old (O, approximately 250 years) ponderosa pine (Pinus ponderosa Dougl. ex P. Laws.) stands growing in a drought-prone region of central Oregon. Although the stands differed in tree leaf area index (LAIT) (Y = 0.9, M = 2.8, O = 2.1), cumulative tree transpiration measured by sap flow did not differ substantially during the growing season (100-112 mm). Yet when water was readily available, transpiration per unit leaf area of the youngest trees was about three times that of M trees and five times that of O trees. These patterns resulted from a nearly sixfold difference in leaf specific conductance (KL) between the youngest and oldest trees. At the time of maximum transpiration in the Y stand in May-June, gross carbon uptake (gross ecosystem production, GEP) was similar for Y and O stands despite an almost twofold difference in stand leaf area index (LAIS). However, the higher rate of water use by Y trees was not sustainable in the drought-prone environment, and between spring and late summer, KL of Y trees declined fivefold compared with a nearly twofold decline for M trees and a < 30% reduction in O trees. Because the Y stand contained a significant shrub understory and more exposed soil, there was no appreciable difference in mean daily latent energy fluxes between the Y stand and the older stands as measured by the eddy-covariance technique. These patterns resulted in 60 to 85% higher seasonal GEP and 55 to 65% higher water-use efficiency at the M and O stands compared with the Y stand.     

Analyses of assumptions and errors in the calculation of stomatal conductance from sap flux measurements

We analyzed assumptions and measurement errors in estimating canopy transpiration (E(L)) from sap flux (J(S)) measured with Granier-type sensors, and in calculating canopy stomatal conductance (G(S)) from E(L) and vapor pressure deficit (D). The study was performed in 12-year-old Pinus taeda L. stands with a wide range in leaf area index (L) and growth rate. No systematic differences in J(S) were found between the north and south sides of trees. However, J(S) in xylem between 20 and 40 mm from the cambium was 50 and 39% of J(S) in the outer 20-mm band of xylem in slow- and fast-growing trees, respectively. Sap flux measured in stems did not lag J(S) measured in branches, and time and frequency domain analyses of time series indicated that variability in J(S) in stems and branches is mostly explained by variation in D. Therefore, J(S) was used to estimate transpiration, after accounting for radial patterns. There was no difference between D and leaf-to-air vapor pressure gradient, and D did not have a vertical profile in stands of either low or high L suggesting a strong canopy-atmosphere coupling. Therefore, D estimated at one point in the canopy can be used to calculate G(S) in such stands. Given the uncertainties in J(S), relative humidity, and temperature measurements, to keep errors in G(S) estimates to less than 10%, estimates of G(S) should be limited to conditions in which D >/= 0.6 kPa.     

Use of a simulation model and ecosystem flux data to examine carbon-water interactions in ponderosa pine

Drought stress plays an important role in determining both the structure and function of forest ecosystems, because of the close association between the carbon (C) and hydrological cycles. We used a detailed model of the soil-plant-atmosphere continuum to investigate the links between carbon uptake and the hydrological cycle in a mature, open stand of ponderosa pine (Pinus ponderosa Dougl. ex Laws.) at the Metolius river in eastern Oregon over a 2-year period (1996-1997). The model was parameterized from local measurements of vegetation structure, soil properties and meteorology, and tested against independent measurements of ecosystem latent energy (LE) and carbon fluxes and soil water content. Although the 2 years had very different precipitation regimes, annual uptake of C and total transpiration were similar in both years, according to both direct observation and simulations. There were important differences in ratios of evaporation to transpiration, and in the patterns of water abstraction from the soil profile, depending on the frequency of summer storms. Simulations showed that, during periods of maximum water limitation in late summer, plants maintained a remarkably constant evapotranspirative flux because of deep rooting, whereas changes in rates of C accumulation were determined by interactions between atmospheric vapor pressure deficit and stomatal conductance. Sensitivity analyses with the model suggest a highly conservative allocation strategy in the vegetation, focused belowground on accessing a soil volume large enough to buffer summer droughts, and optimized to account for interannual variability in precipitation. The model suggests that increased allocation to leaf area would greatly increase productivity, but with the associated risk of greater soil water depletion and drought stress in some years. By constructing sparse canopies and deep rooting systems, these stands balance reduced productivity in the short term with risk avoidance over the long term.     

Influence of moisture stress and induced resistance in ponderosa pine,Pinus ponderosa Dougl. ex. Laws,on the pine sawfly,Neodiprion autumnalis Smith

Plant moisture stress is artificially maintained for ponderosa pine, Pinus ponderosa Dougl. ex Laws, seedlings using polyethylene glycol. Stressed seedlings produce significantly lower amounts of needle nitrogen, phenols, and tannins than non-stressed seedlings. Field tests reveal that previously damaged foliage is a significantly poorer food source for the pine sawfly, Neodiprion autumnalis Smith (Hymenoptera: Diprionidae), than non-damaged foliage. Field studies further indicate that the effect of induction by defoliation is more important than the effect of stress level. These results are discussed in terms of their potential impact on future studies on preferential selection by herbivores of stressed host plants.     

Responses of foliar delta13C, gas exchange and leaf morphology to reduced hydraulic conductivity in Pinus monticola branches

We tested the hypothesis that branch hydraulic conductivity partly controls foliar stable carbon isotope ratio (delta13C) by its influence on stomatal conductance in Pinus monticola Dougl. Notching and phloem-girdling treatments were applied to reduce branch conductivity over the course of a growing season. Notching and phloem girdling reduced leaf-specific conductivity (LSC) by about 30 and 90%, respectively. The 90% reduction in LSC increased foliar delta13C by about 1 per thousand (P < 0.0001, n = 65), whereas the 30% reduction in LSC had no effect on foliar delta13C (P = 0.90, n = 65). Variation in the delta13C of dark respiration was similar to that of whole-tissues when compared among treatments. These isotopic measurements, in addition to instantaneous gas exchange measurements, suggested only minor adjustments in the ratio of intercellular to atmospheric CO2 partial pressures (ci/ca) in response to experimentally reduced hydraulic conductivity. A strong correlation was observed between stomatal conductance (gs) and photosynthetic demand over a tenfold range in gs. Although ci/ca and delta13C appeared to be relatively homeostatic, current-year leaf area varied linearly as a function of branch hydraulic conductivity (r2 = 0.69, P < 0.0001, n = 18). These results suggest that, for Pinus monticola, adjustment of leaf area is a more important response to reduced branch conductivity than adjustment of ci/ca.     

NaCl胁迫对青檀种子发芽率、叶气体交换和苗木生长的影响

在水培条件下,本文探讨了NaCl胁迫对青檀种子发芽率、叶气体交换和苗木生长的影响。研究结果表明,NaCl胁迫延长了青檀种子的发芽时间,特别是当NaCl浓度超过17mM(1.0g/L)时,发芽率明显降低;NaCl胁迫降低了青檀叶的呼吸速率、气孔导度和净光合速率,并随着NaCl胁迫浓度的提高,下降的幅度更大,但NaCl胁迫苗木胞间隙CO2浓度明显高于对照;与对照相比,在NaCl胁迫50天后,青檀苗木的成活率、苗高和地径生长以及根系、茎和叶生物量显著降低,但茎和叶生物量下降更为明显。本文的研究结果认为,青檀是一种对盐胁迫较敏感的植物,一年生幼苗的耐盐阈值在34mM(2.0g/L)左右。图2表3参27。     

Effects of NaCl stress on seed germination, leaf gas exchange and seedling growth of Pteroceltis tatarinowii

Effects of NaCl on seed germination, leaf gas exchange and seedling growth were investigated. The germination of P. tatarinowii seeds was delayed in the presence of NaCl, and was significantly inhibited when NaCl concentration exceeded 17 mM (1.0 g·L⁻¹). NaCl treatments induced reductions of transpiration rate, stomatal conductance and net photosynthetic rate and their reductions were significant with the increasing of NaCl concentration. However, intercellular CO₂ concentration in seedling leaves under salt stress was significantly higher than that of controls. There was a significant reduction in survival, basal diameter increment and height increment with increasing NaCl concentration 50 days after NaCl treatment, and a significant reduction in the dry weight increment of roots, stems, and leaves was also observed in the NaCl treatments. The deleterious effect on the biomass increment was more evident in the case of stems and leaves. Results from this study suggest that P. tatarinowii is sensitive to salinity stress and the salinity threshold for seedling growth might be less 34 mM (2.0 g·L⁻¹). Biography: FANG Sheng-zuo (1963–), male, Professor, College of Forest Resources and Environment, Nanjing Forestry University, Nanjing 210037, P. R. China.     

叶形在树木叶片气体交换应对风胁迫中的作用

作为环境因子,风与树木的关系较其他因子复杂,树木对风胁迫的响应与适应是植物逆境生理生态学研究的热点和难点。本文综述了不同叶形树种叶片气体交换对风胁迫的响应差异,总结发现风对不同叶形树种叶片气体交换影响的研究结论差异较大,有的甚至截然相反,表明了不同叶形树种叶片气体交换对风胁迫响应的复杂性。同时分析了由叶形引起的微环境、边界层导度和理化特征对叶片气孔交换的影响。     

Diurnal and seasonal variations in leaf hydraulic conductance in evergreen and deciduous trees

We studied changes in the hydraulic conductance of leaves (K(leaf)) between dawn and dusk during the growth period (July) and at midday at the beginning of autumn in four tree species. The main objectives of the study were to check the extent of diurnal and seasonal changes in K(leaf) and the relationships between K(leaf), irradiance and leaf gas exchange. Two evergreen (Aleurites moluccana and Persea americana) and two deciduous trees (Platanus orientalis and Quercus rubra) were studied. Leaf hydraulic conductance was measured every 2 h between 0700 and 1900 h in July and compared with values measured between 0900 and 1300 h in October. Other variables measured were photosynthetically active radiation (PAR), leaf conductance to water vapor (gL) and water potential (psiL). In July, K(leaf) varied by up to 75% in Pe. americana on a diurnal basis and by at least 44% in Q. rubra. The diurnal time course of K(leaf) showed a distinct increase between dawn and late morning (1100 h) and a subsequent decrease in the evening in A. moluccana and Pl. orientalis, whereas in the other two species, K(leaf) was highest just after dawn and lowest in the evening. In October, K(leaf) of all the species studied was lower than in July, with differences of 20 to 28% for A. moluccana and Pl. orientalis and of 66 to over 70% in Pe. americana and Q. rubra, respectively. Significant correlations were found between PAR and K(leaf) (in all species) as well as between gL and K(leaf) (in three out of four species). Leaf habit (evergreen or deciduous) did not influence absolute values of K(leaf) or its diurnal variation.