Leaf- and shoot-level plasticity in response to different nutrient and water availabilities

Phenotypic plasticity in response to environmental variation occurs at all levels of organization and across temporal scales within plants. However, the magnitude and functional significance of plasticity is largely unexplored in perennial species. We measured the plasticity of leaf- and shoot-level physiological, morphological and developmental traits in nursery-grown Populus deltoides Bartr. ex Marsh. individuals subjected to different nutrient and water availabilities. We also examined the extent to which nutrient and water availability influenced the relationships between these traits and productivity. Populus deltoides responded to changes in resource availability with high plasticity in shoot-level traits and moderate plasticity in leaf-level traits. Although shoot-level traits generally correlated strongly with productivity across fertilization and irrigation treatments, few leaf-level traits correlated with productivity, and the relationships depended on the resource examined. In fertilized plants, leaf nitrogen concentration was negatively correlated with productivity, suggesting that growth, rather than enhanced leaf quality, is an important response to fertilization in this species. With the exception of photosynthetic nitrogen-use efficiency, traits associated with resource conservation (leaf senescence rate, water-use efficiency and leaf mass per area) were uncorrelated with short-term productivity in nutrient- and water-stressed plants. Our results suggest that plasticity in shoot-level growth traits has a greater impact on plant productivity than does plasticity in leaf-level traits and that the relationships between traits and productivity are highly resource dependent.     

Transpiration by two poplar varieties grown as coppice for biomass production

Fast-growing tree clones selected for biomass plantations are highly productive and therefore likely to use more water than the agricultural crops they replace. We report field measurements of transpiration through the summer of 1994 from two poplar clones, Beaupré (Populus trichocarpa Torr. & A. Gray x P. deltoides Bartr. ex Marsh.) and Dorschkamp (P. deltoides x P. nigra L.), grown as unirrigated short-rotation coppice in southern England. Stand transpiration was quantified by scaling up from sap flow measurements made with the heat balance method in a sample of stems. Leaf conductances, leaf area development, meteorological variables and soil water deficit were also measured to investigate the response of the trees to the environment. High rates of transpiration were found for Beaupré. In June, when soil water was plentiful, the mean (+/- SD) transpiration rate over an 18-day period was 5.0 +/- 1.8 mm day(-1), reaching a maximum of 7.9 mm day(-1). Transpiration rates from Dorschkamp were lower, as a result of its lower leaf area index. High total leaf conductances were measured for both Beaupré (0.34 +/- 0.17 mol m(-2) s(-1)) and Dorschkamp (0.39 +/- 0.16 mol m(-2) s(-1)). Leaf conductance declined slightly with increasing atmospheric vapor pressure deficit in both clones, but only in Beaupré did leaf conductance decrease as soil water deficit increased.     

Regulation of transpirational water loss in Quercus suber trees in a Mediterranean-type ecosystem

Sap flux density in branches, leaf transpiration, stomatal conductance and leaf water potentials were measured in 16-year-old Quercus suber L. trees growing in a plantation in southern Portugal to understand how evergreen Mediterranean trees regulate water loss during summer drought. Leaf specific hydraulic conductance and leaf gas exchange were monitored during the progressive summer drought to establish how changes along the hydraulic pathway influence shoot responses. As soil water became limiting, leaf water potential, stomatal conductance and leaf transpiration declined significantly. Predawn leaf water potential reflected soil water potential measured at 1-m depth in the rhizospheres of most trees. The lowest predawn leaf water potential recorded during this period was -1.8 MPa. Mean maximum stomatal conductance declined from 300 to 50 mmol m(-2) s(-1), reducing transpiration from 6 to 2 mmol m(-2) s(-1). Changes in leaf gas exchange were attributed to reduced soil water availability, increased resistances along the hydraulic pathway and, hence, reduced leaf water supply. There was a strong coupling between changes in soil water content and stomatal conductance as well as between stomatal conductance and leaf specific hydraulic conductance. Despite significant seasonal differences among trees in predawn leaf water potential, stomatal conductance, leaf transpiration and leaf specific hydraulic conductance, there were no differences in midday leaf water potentials. The strong regulation of changes in leaf water potential in Q. suber both diurnally and seasonally is achieved through stomatal closure, which is sensitive to changes in both liquid and vapor phase conductance. This sensitivity allows for optimization of carbon and water resource use without compromising the root-shoot hydraulic link.     

Age-related effects on leaf area/sapwood area relationships,canopy transpiration and carbon gain of Norway spruce stands (Picea abies) in the Fichtelgebirge,Germany

Stand age is an important structural determinant of canopy transpiration (E(c)) and carbon gain. Another more functional parameter of forest structure is the leaf area/sapwood area relationship, A(L)/A(S), which changes with site conditions and has been used to estimate leaf area index of forest canopies. The interpretation of age-related changes in A(L)/A(S) and the question of how A(L)/A(S) is related to forest functions are of current interest because they may help to explain forest canopy fluxes and growth. We conducted studies in mature stands of Picea abies (L.) Karst. varying in age from 40 to 140 years, in tree density from 1680 to 320 trees ha(-1), and in tree height from 15 to 30 m. Structural parameters were measured by biomass harvests of individual trees and stand biometry. We estimated E(c) from scaled-up xylem sap flux of trees, and canopy-level fluxes were predicted by a three-dimensional microclimate and gas exchange model (STANDFLUX). In contrast to pine species, A(L)/A(S) of P. abies increased with stand age from 0.26 to 0.48 m(2) cm(-2). Agreement between E(c) derived from scaled-up sap flux and modeled canopy transpiration was obtained with the same parameterization of needle physiology independent of stand age. Reduced light interception per leaf area and, as a consequence, reductions in net canopy photosynthesis (A(c)), canopy conductance (g(c)) and E(c) were predicted by the model in the older stands. Seasonal water-use efficiency (WUE = A(c)/E(c)), derived from scaled-up sap flux and stem growth as well as from model simulation, declined with increasing A(L)/A(S) and stand age. Based on the different behavior of age-related A(L)/A(S) in Norway spruce stands compared with other tree species, we conclude that WUE rather than A(L)/A(S) could represent a common age-related property of all species. We also conclude that, in addition to hydraulic limitations reducing carbon gain in old stands, a functional change in A(L)/A(S) that is related to reduced light interception per leaf area provides another potential explanation for reduced carbon gain in old stands of P. abies, even when hydraulic constraints increase in response to changes in canopy architecture and aging.     

Effects of branch height on leaf gas exchange,branch hydraulic conductance and branch sap flux in open-grown ponderosa pine

Recent studies have shown that stomata respond to changes in hydraulic conductance of the flow path from soil to leaf. In open-grown tall trees, branches of different heights may have different hydraulic conductances because of differences in path length and growth. We determined if leaf gas exchange, branch sap flux, leaf specific hydraulic conductance, foliar carbon isotope composition (delta13C) and ratios of leaf area to sapwood area within branches were dependent on branch height (10 and 25 m) within the crowns of four open-grown ponderosa pine (Pinus ponderosa Laws.) trees. We found no difference in leaf gas exchange or leaf specific hydraulic conductance from soil to leaf between the upper and lower canopy of our study trees. Branch sap flux per unit leaf area and per unit sapwood area did not differ between the 10- and 25-m canopy positions; however, branch sap flux per unit sapwood area at the 25-m position had consistently lower values. Branches at the 25-m canopy position had lower leaf to sapwood area ratios (0.17 m2 cm-2) compared with branches at the 10-m position (0.27 m2 cm-2) (P = 0.03). Leaf specific conductance of branches in the upper crown did not differ from that in the lower crown. Other studies at our site indicate lower hydraulic conductance, sap flux, whole-tree canopy conductance and photosynthesis in old trees compared with young trees. This study suggests that height alone may not explain these differences.     

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

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

Does initial spacing influence crown and hydraulic architecture of Eucalyptus marginata?

Long-term declines in rainfall in south-western Australia have resulted in increased interest in the hydraulic characteristics of jarrah (Eucalyptus marginata Donn ex Smith) forest established in the region's drinking water catchments on rehabilitated bauxite mining sites. We hypothesized that in jarrah forest established on rehabilitated mine sites: (1) leaf area index (L) is independent of initial tree spacing; and (2) more densely planted trees have less leaf area for the same leaf mass, or the same sapwood area, and have denser sapwood. Initial stand densities ranged from about 600 to 9000 stems ha(-1), and trees were 18 years old at the time of sampling. Leaf area index was unaffected by initial stand density, except in the most sparsely stocked stands where L was 1.2 compared with 2.0-2.5 in stands at other spacings. The ratio of leaf area to sapwood area (A(l):A(s)) was unaffected by tree spacing or tree size and was 0.2 at 1.3 m height and 0.25 at the crown base. There were small increases in sapwood density and decreases in leaf specific area with increased spacing. Tree diameter or basal area was a better predictor of leaf area than sapwood area. At the stand scale, basal area was a good predictor of L (r(2) = 0.98, n = 15) except in the densest stands. We conclude that the hydraulic attributes of this forest type are largely independent of initial tree spacing, thus simplifying parameterization of stand and catchment water balance models.     

Transpiration and water relations of poplar trees growing close to the water table

Sap flow was measured on five branches of two poplar (Populus trichocarpa Torr. & A. Gray x P. tacamahaca L.) trees from June to September 1994 in the south of England with stem-surface, heat balance gauges, and was scaled up to estimate transpiration from single trees on the basis of leaf area. On six days, stomatal conductance and plant water potential were measured simultaneously with a porometer and pressure chamber, respectively. The effects of solar radiation (S), vapor pressure deficit (D) and stomatal conductance on transpiration were evaluated. Sap flow per unit leaf area (F(a)) was closely related to the time course of demand attributable to S and D throughout the season, and only slightly affected by the water content of the top 120 cm of soil. Although F(a) increased linearly at low values of D, it showed a plateau with increases in D above 1.2 kPa. The canopy coupling coefficient (1 - Omega) ranged from 0.48 to 0.78 with a mean of 0.65 +/- 0.01, indicating that transpiration was controlled more by stomatal conductance than by incident radiation. The seasonal pattern of tree water loss followed potential evaporation with a peak in late June or early July. On bright days, daily transpiration over the projected crown area was 3.6 mm early in the season, 3.8 mm in mid-season, and 2.7 mm late in the season. The water balance of the system indicated that poplar trees took 15-60% of water transpired from groundwater, with the proportion increasing as the soil in the unsaturated zone dried out. Access to the water table resulted in high predawn water potentials throughout the season. Estimated hydraulic resistance to water flow in the soil-tree system was in the range of 1.5 to 1.93 x 10(6) MPa s m(-3).     

Restoration thinning and influence of tree size and leaf area to sapwood area ratio on water relations of Pinus ponderosa

Ponderosa pine (Pinus ponderosa Dougl. ex P. Laws) forest stand density has increased significantly over the last century (Covington et al. 1997). To understand the effect of increased intraspecific competition, tree size (height and diameter at breast height (DBH)) and leaf area to sapwood area ratio (A(L):A(S)) on water relations, we compared hydraulic conductance from soil to leaf (kl) and transpiration per unit leaf area (Q(L)) of ponderosa pine trees in an unthinned plot to trees in a thinned plot in the first and second years after thinning in a dense Arizona forest. We calculated kl and Q(L) based on whole- tree sap flux measured with heat dissipation sensors. Thinning increased tree predawn water potential within two weeks of treatment. Effects of thinning on kl and Q(L) depended on DBH, A(L):A(S) and drought severity. During severe drought in the first growing season after thinning, kl and Q(L) of trees with low A(L):A(S) (160-250 mm DBH; 9-11 m height) were lower in the thinned plot than the unthinned plot, suggesting a reduction in stomatal conductance (g(s)) or reduced sapwood specific conductivity (K(S)), or both, in response to thinning. In contrast kl and Q(L) were similar in the thinned plot and unthinned plot for trees with high A(L):A(S) (260-360 mm DBH; 13-16 m height). During non-drought periods, kl and Q(L) were greater in the thinned plot than in the unthinned plot for all but the largest trees. Contrary to previous studies of ponderosa pine, A(L):A(S) was positively correlated with tree height and DBH. Furthermore, kl and Q(L) showed a weak negative correlation with tree height and a strong negative correlation with A(S) and thus A(L):A(S) in both the thinned and unthinned plots, suggesting that trees with high A(L):A(S) had lower g(s). Our results highlight the important influence of stand competitive environment on tree-size-related variation in A(L):A(S) and the roles of A(L):A(S) and drought on whole-tree water relations in response to thinning.     

Influence of soil water on the physiological and morphological components of plant water balance in Populus trichocarpa, Populus deltoides and their F(1) hybrids

Patterns of leaf growth, transpiration and whole-plant water balance in Populus trichocarpa, P. deltoides and their F(1) hybrids were studied during a soil drying cycle. Plant responses were analyzed during three distinct stages of dehydration. In stage I, the transpiration rate of drought-stressed plants remained constant and equal to that of well-watered plants even though soil water content declined by more than 40%. Stage II began as soil and plant water deficits induced stomatal closure. When soil water was expressed as a fraction of transpirable soil water, the transition from stage I to stage II occurred at soil water fractions of 0.35, 0.45 and 0.60 for P. trichocarpa, P. deltoides and their F(1) hybrids, respectively. Reductions in leaf growth coincided with the shift from stage I to stage II. As soil water declined further, decreases in relative transpiration and whole-plant leaf area were significantly greater in parental species than in F(1) hybrids. Inherent feedbacks controlling stomatal water loss and the maintenance and growth of leaf tissue appeared to differ between F(1) and parental genotypes in a pattern characteristic of an overdominant mode of inheritance.Stage III began once the ability of stomata to compensate for water loss had been exhausted. Substantial differences were found in plant survival during stage III, with F(1) hybrids surviving longer than parental species. Survival was more strongly correlated with the hydraulic conductivity of xylem tissues than with the dehydration tolerance of leaf tissues. Collectively, these responses suggest that F(1) hybrids were more drought resistant than either parental species and highlight the importance of whole-plant studies of functional relationships between plant growth, water balance and hydraulic conductivity.     

宝天曼三桠乌药对降雨减少后的生理生态响应

[目的]在宝天曼森林原位建立降水减少(截雨)样地后,样地内的三桠乌药出现了顶端枯死,本研究从水和碳的角度,探讨三桠乌药顶端枯死的原因。[方法]2013年4月在宝天曼锐齿栎林原位建立了3块截雨样地,在2014年生长季干旱时,采用压力室、低压液流系统和蒽酮硫酸法测定了三桠乌药的水力系统特征和非结构性碳等指标。[结果]显示:(1)三桠乌药在截雨处理后出现了顶端枯死;三桠乌药最大导管长度约为60 cm,栓塞50%时的水势(P50)为-1.43 MPa,其木质部栓塞脆弱性较大。在截雨处理1年后生长季较干旱时,净光合速率、凌晨水势、中午水势显著低于对照,中午栓塞显著高于对照,且水力安全边际为负值。(2)三桠乌药气孔导度、蒸腾速率、叶面积、导管直径、边材比导率、叶片比导率显著下降,Huber值、导管密度显著升高;而叶片、韧皮部、木质部3个器官的可溶性糖、淀粉、总非结构性碳与对照差异不显著。[结论]水力失衡是三桠乌药顶端枯死的主要原因,而蒸腾面积、水分输导系统等的变化表明三桠乌药在缺水环境下进行了一定的适应性调节,但这些调节不足以使三桠乌药在缺水情况下避免水力失衡而导致的顶端枯死。     

Actual and potential transpiration and carbon assimilation in an irrigated poplar plantation

We examined the tradeoffs between stand-level water use and carbon uptake that result when biomass production of trees in plantations is maximized by removing nutrient and water limitations. A Populus trichocarpa Torr. x P. deltoides Bartr. & Marsh. plantation was irrigated and received frequent additions of nutrients to optimize biomass production. Sap flux density was measured continuously over four of the six growing-season months, supplemented with periodic measurements of leaf gas exchange and water potential. Measurements of tree diameter and height were used to estimate leaf area and biomass production based on allometric relationships. Sap flux was converted to canopy conductance and analyzed with an empirical model to isolate the effects of water limitation. Actual and soil-water-unlimited potential CO(2) uptakes were estimated with a canopy conductance constrained carbon assimilation (4C-A) scheme, which couples actual or potential canopy conductance with vertical gradients of light distribution, leaf-level conductance, maximum Rubisco capacity and maximum electron transport. Net primary production (NPP) was about 43% of gross primary production (GPP); when estimated for individual trees, this ratio was independent of tree size. Based on the NPP/GPP ratio, we found that current irrigation reduced growth by about 18% compared with growth with no water limitation. To achieve maximum growth, however, would require 70% more water for transpiration, and would reduce water-use efficiency by 27%, from 1.57 to 1.15 g stem wood C kg(-1) water. Given the economic and social values of water, plantation managers appear to have optimized water use.     

Water utilization,plant hydraulic properties and xylem vulnerability in three contrasting coffee (Coffea arabica) cultivars

Water use, hydraulic properties and xylem vulnerability to cavitation were studied in the coffee (Coffea arabica L.) cultivars San Ramon, Yellow Caturra and Typica growing in the field under similar environmental conditions. The cultivars differed in growth habit, crown morphology and total leaf surface area. Sap flow, stomatal conductance (g(s)), crown conductance (g(c)), apparent hydraulic conductance of the soil-leaf pathway (G(t)), leaf water potential (Psi(L)) and xylem vulnerability to loss of hydraulic conductivity were assessed under well-watered conditions and during a 21-day period when irrigation was withheld. Sap flow, g(c), and G(t) were greatest in Typica both with and without irrigation, lowest in San Ramon, which was relatively unresponsive to the withholding of irrigation, and intermediate in Yellow Caturra. The cultivars had similar g(s) when well watered, but withholding water decreased g(s) more in Typica and Yellow Caturra than in San Ramon. Typica had substantially lower Psi(L) near the end of the unirrigated period than the other cultivars (-2.5 versus -1.8 MPa), consistent with the relatively high sap flow in this cultivar. Xylem vulnerability curves indicated that Typica was less susceptible to loss of hydraulic conductivity than the other cultivars, consistent with the more negative Psi(L) values of Typica in the field during the period of low soil water availability. During soil drying, water use declined linearly with relative conductivity loss predicted from vulnerability curves. However, cultivar-specific relationships between water use and predicted conductivity loss were not observed because of pronounced hysteresis during recovery of water use following soil water recharge. All cultivars shared the same functional relationship between integrated daily sap flow and G(t), but they had different operating ranges. The three cultivars also shared common functional relationships between hydraulic architecture and water use despite consistent differences in water use under irrigated and dry soil conditions. We conclude that hydraulic architectural traits, rate of water use per plant and crown architecture are important determinants of short- and long-term variations in the water balance of Coffea arabica.     

Stable annual pattern of water use by Acacia tortilis in Sahelian Africa

Water use by mature trees of Acacia tortilis (Forsk.) Hayne ssp. raddiana (Savi) Brenan var. raddiana growing in the northern Sahel was continuously recorded over 4 years. Water use was estimated from xylem sap flow measured by transient heat dissipation. Concurrently, cambial growth, canopy phenology, leaf water potential, climatic conditions and soil water availability (SWA) were monitored. In addition to the variation attributable to interannual variation in rainfall, SWA was increased by irrigation during one wet season. The wet season lasted from July to September, and annual rainfall ranged between 146 and 367 mm. The annual amount and pattern of tree water use were stable from year-to-year despite interannual and seasonal variations in SWA in the upper soil layers. Acacia tortilis transpired readily throughout the year, except for one month during the dry season when defoliation was at a maximum. Maximum water use of about 23 l (dm sapwood area)(-2) day(-1) was recorded at the end of the wet season. While trees retained foliage in the dry season, the decline in water use was modest at around 30%. Variation in predawn leaf water potential indicated that the trees were subject to soil water constraint. The rapid depletion of water in the uppermost soil layers after the wet season implies that there was extensive use of water from deep soil layers. The deep soil profile revealed (1) the existence of living roots at 25 m and (2) that the availability of soil water was low (-1.6 MPa) down to the water table at a depth of 31 m. However, transpiration was recorded at a predawn leaf water potential of -2.0 MPa, indicating that the trees used water from both intermediary soil layers and the water table. During the full canopy stage, mean values of whole-tree hydraulic conductance were similar in the wet and dry seasons. We propose that the stability of water use at the seasonal and annual scales resulted from a combination of features, including an extensive rooting habit related to deep water availability and an effective regulation of canopy conductance. Despite a limited effect on tree water use, irrigation during the wet season sharply increased predawn leaf water potential and cambial growth of trunks and branches.     

Effects of irrigation on water use and water use efficiency in two fast growing Eucalyptus plantations

Eucalyptus plantations occupy almost 20 million ha worldwide and exceed 3.7 million ha in Brazil alone. Improved genetics and silviculture have led to as much as a three-fold increase in productivity in Eucalyptus plantations in Brazil and the large land area occupied by these highly productive ecosystems raises concern over their effect on local water supplies. As part of the Brazil Potential Productivity Project, we measured water use of Eucalyptus grandis × urophylla clones in rainfed and irrigated stands in two plantations differing in productivity. The Aracruz (lower productivity) site is located in the state of Espirito Santo and the Veracel (higher productivity) site in Bahia state. At each plantation, we measured stand water use using homemade sap flow sensors and a calibration curve using the clones and probes we utilized in the study. We also quantified changes in growth, leaf area and water use efficiency (the amount of wood produced per unit of water transpired). Measurements were conducted for 1 year during 2005 at Aracruz and from August through December 2005 at Veracel. Transpiration at both sites was high compared to other studies but annual estimates at Aracruz for the rainfed treatment compared well with a process model calibrated for the Aracruz site (within 10%). Annual water use at Aracruz was 1394 mm in rainfed treatments versus 1779 mm in irrigated treatments and accounted for approximately 67% and 58% of annual precipitation and irrigation inputs respectively. Increased water use in the irrigated stands at Aracruz was associated with higher sapwood area, leaf area index and transpiration per unit leaf area but there was no difference in the response of canopy conductance with air saturation deficit between treatments. Water use efficiency at the Aracruz site was also not influenced by irrigation and was similar to the rainfed treatment. During the period of overlapping measurements, the response to irrigation treatments at the more productive Veracel site was similar to Aracruz. Stand water use at the Veracel site totaled 975 mm and 1102 mm in rainfed and irrigated treatments during the 5-month measurement period respectively. Irrigated stands at Veracel also had higher leaf area with no difference in the response of canopy conductance with air saturation deficit between treatments. Water use efficiency was also unaffected by irrigation at Veracel. Results from this and other studies suggest that improved resource availability does not negatively impact water use efficiency but increased productivity of these plantations is associated with higher water use and should be given consideration during plantation management decision making processes aimed at increasing productivity.     

Size-mediated tree transpiration along soil drainage gradients in a boreal black spruce forest wildfire chronosequence

Boreal forests are crucial to climate change predictions because of their large land area and ability to sequester and store carbon, which is controlled by water availability. Heterogeneity of these forests is predicted to increase with climate change through more frequent wildfires, warmer, longer growing seasons and potential drainage of forested wetlands. This study aims at quantifying controls over tree transpiration with drainage condition, stand age and species in a central Canadian black spruce boreal forest. Heat dissipation sensors were installed in 2007 and data were collected through 2008 on 118 trees (69 Picea mariana (Mill.) Britton, Sterns & Poggenb. (black spruce), 25 Populus tremuloides Michx. (trembling aspen), 19 Pinus banksiana Lamb. (jack pine), 3 Larix laricina (Du Roi) K. Koch (tamarack) and 2 Salix spp. (willow)) at four stand ages (18, 43, 77 and 157 years old) each containing a well- and poorly-drained stand. Transpiration estimates from sap flux were expressed per unit xylem area, J(S), per unit ground area, E(C) and per unit leaf area, E(L), using sapwood (A(S)) and leaf (A(L)) area calculated from stand- and species-specific allometry. Soil drainage differences in transpiration were variable; only the 43- and 157-year-old poorly-drained stands had?～?50% higher total stand E(C) than well-drained locations. Total stand E(C) tended to decrease with stand age after an initial increase between the 18- and 43-year-old stands. Soil drainage differences in transpiration were controlled primarily by short-term physiological drivers such as vapor pressure deficit and soil moisture whereas stand age differences were controlled by successional species shifts and changes in tree size (i.e., A(S)). Future predictions of boreal climate change must include stand age, species and soil drainage heterogeneity to avoid biased estimates of forest water loss and latent energy exchanges.     

Hydraulic conductance, light interception and needle nutrient concentration in Scots pine stands and their relations with net primary productivity

Aboveground xylem hydraulic conductance was determined in Scots pine (Pinus sylvestris L.) trees and stands from 7 to about 60 years of age. At the stand scale, leaf area index and net primary productivity (NPP, above- plus belowground) increased and reached a plateau at about 25-30 and 15-20 years, respectively; both parameters declined in mature stands. Stand hydraulic conductance followed a similar trend to NPP, with a maximum at about 15-20 years and a pronounced reduction in old stands. At the tree scale, annual biomass growth per unit of leaf area (growth efficiency) declined with tree age, whereas aboveground sapwood volume per unit leaf area, which is linearly related to maintenance respiration costs, steadily increased. Radiation interception per unit leaf area increased significantly with reduced leaf area index of mature stands, despite increased foliage clumping in the canopies of mature trees. Needle nutrient concentration did not change in the chronosequence. Tree hydraulic conductance per unit leaf area was strongly and positively correlated with growth efficiency. We discuss our findings in the context of growth reductions in mature and old trees, and suggest that increased hydraulic resistance and maintenance respiration costs may be the main causes of reduced carbon gain in mature and old trees.     

Water loss regulation in mature Hevea brasiliensis: effects of intermittent drought in the rainy season and hydraulic regulation

Effects of soil and atmospheric drought on whole-tree transpiration (E(T)), leaf water potential (Ψ(L)) and whole-tree hydraulic conductance (K(T)) were investigated in mature rubber trees (Hevea brasiliensis, clone RRIM 600) during the full canopy stage in the rainy season in a drought-prone area of northeast Thailand. Under well-watered soil conditions, transpiration was tightly regulated in response to high evaporative demand, i.e., above reference evapotranspiration (ET(0)) ~2.2 mm day(-1) or maximum vapor pressure deficit ~1.8 kPa. When the trees experienced intermittent soil drought E(T) decreased sharply when relative extractable water in the top soil was?相似文献   

Mean canopy stomatal conductance responses to water and nutrient availabilities in Picea abies and Pinus taeda

We compared sap-flux-scaled, mean, canopy stomatal conductance (GS) between Picea abies (L.) Karst. in Sweden and Pinus taeda (L.) in North Carolina, both growing on nutritionally poor soils. Stomatal conductance of Picea abies was approximately half that of Pinus taeda and the sensitivity of GS in Picea abies to vapor pressure deficit (D) was lower than in Pinus taeda. Optimal fertilization increased leaf area index (L) two- and threefold in Pinus taeda and Picea abies, respectively, regardless of whether irrigation was increased. Although it increased L, fertilization did not increase GS in Picea abies unless irrigation was also provided. In Pinus taeda growing on coarse, sandy soils, the doubling of L in response to fertilization reduced GS sharply unless irrigation was also provided. The reduction in GS with fertilization in the absence of irrigation resulted from the production of fine roots with low saturated hydraulic conductivity. When Pinus taeda received both fertilization and irrigation, the increase in L was accompanied by a large increase in GS. In Pinus taeda, a reference GS (defined as GS at D = 1 kPa; GSR) decreased in all treatments with decreasing volumetric soil water content (theta). In Picea abies, theta varied little within a treatment, but overall, GSR declined with theta, reaching lowest values when drought was imposed by the interception of precipitation. Despite the large difference in GS both between Picea abies and Pinus taeda and among treatments, stem growth was related to absorbed radiation, and stem growth response to treatment reflected mostly the changes in L.     

Correlations between stable carbon-isotope abundance and hydraulic conductivity in Douglas-fir across a climate gradient in Oregon, USA

Stomatal conductance in trees is related to both foliar carbon-isotope abundance and stem hydraulic properties. By combining these relationships, I hypothesized that carbon-isotope abundance in foliage should vary with limitations to water movement through supporting branches. I sampled Douglas-fir branches (Pseudotsuga menziesii (Mirb.) Franco) from six sites across a climate gradient in Oregon, USA for foliar carbon-isotope abundance and stem hydraulic properties. I used a forest growth model to quantify climate-induced stomatal limitations, expressed as reduced potential transpiration, across the gradient. Foliar stable carbon-isotope abundance showed a strong inverse relationship with branch specific conductivity (hydraulic conductivity per unit functional sapwood area) and leaf-specific conductivity (hydraulic conductivity per unit leaf area). Foliar stable carbon-isotope abundance was correlated with modeled reductions in potential transpiration; however, the inclusion of leaf-specific conductivity improved the correlation by more than 30%. Combined, leaf-specific conductivity and climate-induced stomatal constraints explained 84% of the variation in foliar isotope abundance in 1994 foliage. This model was confirmed on foliage classes 1990-1993.