Detection of xylem cavitation in field-grown pine trees using the acoustic emission technique

The relationship between the ultrasonic acoustic emission (AE) and the needle water potential in field-grown Japanese black pine,Pinus thunbergii, and excised-pine shoots was investigated in summer. The AE technique was employed as a reliable indicator of cavitation events in the water columns of xylem tracheids even under field conditions. In excised shoots. AE events detected when the needle water potential fell below about-0.9 MPa. In the case of field-grown pine trees, however, relatively few AE event were detected when needle water potential was lower than-0.9 MPa. This suggests that the occurrence of cavitation events in Japanese black pine growing under field conditions is comparatively rare, even in summer.     

Sap Ascent in Vascular Plants: Challengers to the Cohesion Theory Ignore the Significance of Immature Xylem and the Recycling of Munch Water

In recent years the cohesion theory has been attacked on thegrounds that direct measurements made with the pressure probeindicate that sap tensions are much less (maximum tension approx.0.7 MPa) than indicated by parallel measurements made with themore conventional methods: osmotic methods, pressure bomb, orpsychrometer. It has also been claimed that other direct methodsdo not support the cohesion theory. Thus a re-examination usingthe Renner technique indicated sap tensions of approx. 2.5 MPa.Also an independent method based on mercury penetrometry providesevidence that sap tensions of at least 2.0 MPa can be demonstrateddirectly implying, that serious limitations arise from the pressureprobe method itself. Without tensions exceeding 2.0 MPa mangroveswould be unable to extract fresh water for transpiration fromseawater. It is suggested that the pressure probe is susceptibleto bias because it investigates the least mature xylem conduitswhile they are still under varying degrees of turgor pressureand only partially interconnected with the main xylem system.This supposition is supported by claims that the xylem sap sampledby the probe contains significant concentrations of solutes.Additionally water, supplied by reverse osmosis from the sievetubes (‘Münch water’), is continually beingliberated in the vicinity of the outermost xylem vessels hydratingthem to an atypical degree which can explain several of thediscrepancies claimed. These results, which are supported bythe work of others, demonstrate that the challenges to the cohesiontheory for the ascent of sap are ill-founded. The release ofwater from the phloem can explain not only some discrepanciesclaimed by the cohesion challengers, but also explain the refillingof cavitated xylem conduits: a hitherto unsuspected role forthe phloem transport system. Cohesion theory; sap ascent; cavitation; pressure probe; xylem transport; vessel development; recycled water; reverse osmosis     

The canopy water relations of old-growth Douglas-fir trees

 We investigated whole tree water relations in 56–65 m tall, old-growth Pseudotsuga menziesii trees within the Wind River Canopy Crane site, Carson, Washington, USA. We measured at predawn and solar noon the vertical gradients in xylem pressure potential using a pressure chamber. On an Abies amabilis sapling located in the understory at the base of one of the study trees, predawn and solar noon xylem pressure potentials were also measured. Xylem pressure potential data were measured from late June through early September 1996 on foliage sampled from 1 to 64.5 m. Over this height gradient, predawn water potentials ranged from –0.23 to –1.10 MPa. Solar noon values showed an even greater range (from –0.44 to –2.51 MPa). At predawn, the water potential gradient approached the theoretical hydrostatic gradient (–0.0105 vs –0.010 MPa m^–1). The gradient at solar noon was steeper (–0.0331 MPa m^–1). Instantaneous stomatal conductances were not greatly different between young, sapling-sized and old-growth trees [0.094±0.033 (SD) vs 0.086±0.045 cm s^–1, respectively]. Stomata of both size classes of trees appeared very sensitive to increasing vapor pressure deficits. A comparison of stable carbon isotope values from the old-growth and sapling-sized trees indicated lower stomatal conductances in the old-growth. This study provides sound documentation regarding the utility of the cohesion theory in the interpretation of water potential gradients. This study also emphasizes inherent differences between sapling-sized and tall, old-growth trees. Received: 10 January 1998 / Accepted: 12 October 1998     

Native root xylem embolism and stomatal closure in stands of Douglas-fir and ponderosa pine: mitigation by hydraulic redistribution

Hydraulic redistribution (HR), the passive movement of water via roots from moist to drier portions of the soil, occurs in many ecosystems, influencing both plant and ecosystem-water use. We examined the effects of HR on root hydraulic functioning during drought in young and old-growth Douglas-fir [Pseudotsuga menziesii (Mirb.) Franco] and ponderosa pine (Pinus ponderosa Dougl. Ex Laws) trees growing in four sites. During the 2002 growing season, in situ xylem embolism, water deficit and xylem vulnerability to embolism were measured on medium roots (2–4-mm diameter) collected at 20–30 cm depth. Soil water content and water potentials were monitored concurrently to determine the extent of HR. Additionally, the water potential and stomatal conductance (g_s) of upper canopy leaves were measured throughout the growing season. In the site with young Douglas-fir trees, root embolism increased from 20 to 55 percent loss of conductivity (PLC) as the dry season progressed. In young ponderosa pine, root embolism increased from 45 to 75 PLC. In contrast, roots of old-growth Douglas-fir and ponderosa pine trees never experienced more than 30 and 40 PLC, respectively. HR kept soil water potential at 20–30 cm depth above –0.5 MPa in the old-growth Douglas-fir site and –1.8 MPa in the old-growth ponderosa pine site, which significantly reduced loss of shallow root function. In the young ponderosa pine stand, where little HR occurred, the water potential in the upper soil layers fell to about –2.8 MPa, which severely impaired root functioning and limited recovery when the fall rains returned. In both species, daily maximum g_s decreased linearly with increasing root PLC, suggesting that root xylem embolism acted in concert with stomata to limit water loss, thereby maintaining minimum leaf water potential above critical values. HR appears to be an important mechanism for maintaining shallow root function during drought and preventing total stomatal closure.     

Xylem sap as a pathway for total mercury and methylmercury transport from soils to tree canopy in the boreal forest

Conifer needles are an important link in the cycling of Total Mercury (THg) and Methylmercury (MeHg) in the boreal ecosystem due to the high THg and MeHg concentrations in litterfall. Translocation within the tree of Hg from soils to the crown canopy has been assumed to be a minor source of the Hg in litterfall. This paper, however, is the first to present direct observations of THg/MeHg transport from the soil via xylem sap. Xylem sap concentrations of THg and MeHg were measured in sap drained from different levels along the boles of freshly cut 100 year old Norway spruce (Picea abies) and Scots pine (Pinus sylvestris). The trees came from a mixed stand growing on podzolized till soils at the Svartberget Forest Research Station in N. Sweden. Soil solution concentrations of THg and MeHg at different levels in the soil profile were measured for comparison.Concentrations of THg in xylem sap ranged from 10–15 ng L^-1 in both the Scots pine and Norway spruce. Concentrations of MeHg varied from 0.03 ng L^-1to 0.16 ng L^-1, with higher values in Scots pine than Norway spruce. If these concentrations are representative of the transport from soils to needles in xylem sap at this site, then only 3% of the MeHg in litterfall (0.12 mg ha^-1 yr^-1) and 11% of the THg (26 mg ha^-1 yr^-1) can originate via this pathway. The upward transport via xylem sap is larger relative to the open field inputs (84% of THg and 17% of MeHg). Comparison of soil solution and xylem sap THg/MeHg suggested some degree of THg exclusion during water uptake in Scots pine and Norway spruce, but MeHg exclusion only in Norway spruce.     

Long-term implications of industrial pollution stress on lipids composition in Scots pine (Pinus sylvestris L.) roots

The influence of pollution stress (SO₂, Cu²⁺, Pb²⁺, Zn²⁺ and fluoride) on composition changes in cellular membranes of roots of three European of Scots pine (Pinus sylvestris L.) populations were examined. Plant material growing in three experimental areas: Kórnik relatively free of air pollution (control), Luboń: SO₂ and HF and Głogów: SO₂ and heavy metals. Analysis of total phospholipids and their composition indicates that the phosphatidylcholine (PC) and phosphatidylethanolamine (PE) were present in lower concentration in the root tissues from both polluted sites (Luboń and Głogów). The difference in PC:PE ratio between control and both polluted sites was greater in root of Scots pine population from Russia than in the population from Slovakia. Under pollution conditions the content of lipid soluble antioxidant α-tocopherol was lower about 220 %, in comparison to the control. The action of pollution stress also lead to lowering of unsaturated:saturated ratio of total fatty acid, and lower content of polyunsaturated fatty acids, linoleic acid (18:2) and eicosatrienoic acid (20:3). We concluded that the long-term pollution stress markedly inhibited lipid biosynthesis in root tissue of Scots pine and it is probably contribute to the reduction of productivity of forests. These results also suggest that lipid composition can be used as an indicator of changes in tissue roots of Scots pine caused by air and/or soil long-term pollution.     

Hydraulic properties of living late metaxylem and interactions between transpiration and xylem pressure in maize

A new approach to study dynamic interactions between transpiration and xylem pressure in intact plants is presented. Pressure probe measurements were preformed in living (immature) late metaxylem of maize roots rather than in adjacent mature xylem. This eliminated technical limitations related to the measurement of negative pressures. Water relations of single cells showed that turgor and volumetric elastic modulus were significantly larger in living metaxylem than in cortical cells; hydraulic conductivity was similar in both types of root cells. Increasing transpiration induced an immediate decrease of xylem pressure, and vice versa. Turgor in the living metaxylem could be continuously recorded for more than 1 h. The relationship between xylem pressure and transpiration yielded a root hydraulic resistance of 1.3 x 10⁹ MPa s m^-3. Control experiments indicated that the response of living xylem in the positive pressure range essentially paralleled that of mature root xylem in the negative range. In mature xylem, pressures as low as -0.55 MPa were recorded for short periods (several minutes). Several tests verified that the pressure probe was in contact with mature xylem during the measurements of tensions. The results demonstrate convincingly that transpiration generates an effective driving force for water uptake in roots, a central feature of the cohesion theory.Key words: Hydraulic conductivity, negative pressure, root development, turgor, water transport, Zea mays.      

Water content, hydraulic conductivity, and ice formation in winter stems of Pinus contorta: a TDR case study

Stem water content, ice fraction, and losses in xylem conductivity were monitored from November 1996 to October 1997 in an even-aged stand of Pinus contorta (lodgepole pine) near Potlatch, Idaho, USA. A time domain reflectometry (TDR) probe was used to continuously monitor stem water contents and ice fractions. Stem sapwood water contents measured with TDR were not different from water contents measured gravimetrically. The liquid water content of stems ranged from 0.70 m³ m^-3 to 0.20 m³ m^-3 associated with freezing and thawing of the wood tissue. Ice fraction of the stem varied from 0-75% during the winter suggesting liquid water was always present even at ambient temperatures below -20°C. Shoot xylem tensions decreased through the winter to a minimum of ca. -1.4 MPa in February then increased to -0.4 MPa in May. Shoot xylem tensions decreased during the growing season reaching -1.7 MPa by September. Annually, low shoot water potentials were not correlated to decreases in stem hydraulic conductivity. Xylem conductivity decreased due to cavitation through the winter and was 70% of summer values by March. Decreases in xylem conductivity were correlated to low shoot water potentials and cumulative freezing and thawing events within the xylem. Xylem conductivity increased to pre-winter values by May and no reductions in xylem conductivity were observed during the growing season.     

The role of below-ground competition during early stages of secondary succession: the case of 3-year-old Scots pine (Pinus sylvestris L.) seedlings in an abandoned grassland

In abandoned or extensively managed grasslands, the mechanisms involved in pioneer tree species success are not fully explained. Resource competition among plants and microclimate modifications have been emphasised as possible mechanisms to explain variation of survivorship and growth. In this study, we evaluated a number of mechanisms that may lead to successful survival and growth of seedlings of a pioneer tree species (Pinus sylvestris) in a grass-dominated grassland. Three-year-old Scots pines were planted in an extensively managed grassland of the French Massif Central and for 2 years were either maintained in bare soil or subjected to aerial and below-ground interactions induced by grass vegetation. Soil temperatures were slightly higher in bare soil than under the grass vegetation, but not to an extent explaining pine growth differences. The tall grass canopy reduced light transmission by 77% at ground level and by 20% in the upper part of Scots pine seedlings. Grass vegetation presence also significantly decreased soil volumetric water content (Hv) and soil nitrate in spring and in summer. In these conditions, the average tree height was reduced by 5% compared to trees grown in bare soil, and plant biomass was reduced by 85%. Scots pine intrinsic water-use efficiency (A/g), measured by leaf gas-exchange, increased when Hv decreased owing to a rapid decline of stomatal conductance (g). This result was also confirmed by δ ¹³C analyses of needles. A summer ¹⁵N labelling of seedlings and grass vegetation confirmed the higher NO₃ capture capacity of grass vegetation in comparison with Scots pine seedlings. Our results provide evidence that the seedlings' success was linked to tolerance of below-ground resource depletion (particularly water) induced by grass vegetation based on morphological and physiological plasticity as well as to resource conservation.     

Indications of Hydraulic Lift by Pinus halepensis and Its Effects on the Water Relations of Neighbour Shrubs

We measured the stable deuterium isotopic composition of xylem sap, the shoot predawn and midday water potentials, and the leaf δ¹³C of Mediterranean shrubs Pistacia lentiscus, Globularia alypum and Rosmarinus officinalis in a south-oriented transect from a large (12 m tall) Aleppo pine tree, Pinus halepensis. We aimed to study the possibility of hydraulic lift from the deep roots of this pine tree to the shallow soil layers and its influence on these neighbour shrubs. These same traits were also studied in several individuals of the shrub Pistacia lentiscus growing with different types of neighbours: just shrubs, a small (3 – 4 m) pine tree, and the above mentioned large pine tree. The greater the distance from P. halepensis the plants grew, the higher xylem water δD, the lower the soil water content, and, the lower the predawn and midday water potentials were found. These results suggest the existence of an hydraulic lift from deep roots to shallow soil in this big tree. Further indication of this existence is provided by the improved water status of P. lentiscus (higher water potentials and δD, and lower δ¹³C and, therefore, lower water use efficiencies) when growing close to the big pine in comparison with the same shrub species growing close to small pines or just surrounded by other shrubs. Moreover, all these trends occurred in the dry summer season, but disappeared in the wet spring season. This revised version was published online in July 2006 with corrections to the Cover Date.     

Natural 15N abundance in two nitrogen saturated forest ecosystems

Natural ¹⁵N abundance values were measured in needles, twigs, wood, soil, bulk precipitation, throughfall and soil water in a Douglas fir (Pseudotsuga menziesii (Mirb.) and a Scots pine (Pinus sylvestris L.) stand receiving high loads of nitrogen in throughfall (>50 kg N ha⁻¹ year⁻¹). In the Douglas fir stand δ¹⁵N values of the vegetation ranged between −5.7 and −4.2‰ with little variation between different compartments. The vegetation of the Scots pine stand was less depleted in ¹⁵N and varied from −3.3 to −1.2‰δ¹⁵N. At both sites δ¹⁵N values increased with soil depth, from −5.7‰ and −1.2‰ in the organic layer to +4.1‰ and +4.7‰ at 70 cm soil depth in the Douglas fir and Scots pine stand, respectively. The δ¹⁵N values of inorganic nitrogen in bulk precipitation showed a seasonal variation with a mean in NH₄ ⁺-N of −0.6‰ at the Douglas fir stand and +10.8‰ at the Scots pine stand. In soil water below the organic layer NH₄ ⁺-N was enriched and NO₃ ⁻-N depleted in ¹⁵N, which was interpreted as being caused by isotope fractionation accompanying high nitrification rates in the organic layers. Mean δ¹⁵N values of NH₄ ⁺ and NO₃ ⁻ were very similar in the drainage water at 90 cm soil depth at both sites (−7.1 to −3.8‰). A dynamic N cycling model was used to test the sensitivity of the natural abundance values for the amount of N deposition, the ¹⁵N ratio of atmospheric N deposited and for the intrinsic isotope discrimination factors associated with N transformation processes. Simulated δ¹⁵N values for the N saturated ecosystems appeared particularly sensitive to the ¹⁵N ratio of atmospheric N inputs and discrimination factors during nitrification and mineralization. The N-saturated coniferous forest ecosystems studied were not characterized by elevated natural ¹⁵N abundance values. The results indicated that the natural ¹⁵N abundance values can only be used as indicators for the stage of nitrogen saturation of an ecosystem if the δ¹⁵N values of the deposited N and isotope fractionation factors are taken into consideration. Combining dynamic isotope models and natural ¹⁵N abundance values seems a promising technique for interpreting natural ¹⁵N abundance values found in these forest ecosystems. Received: 5 May 1996 / Accepted: 10 April 1997     

Characterization of the photosynthetic apparatus in cortical bark chlorenchyma of Scots pine

Winter-induced inhibition of photosynthesis in Scots pine (Pinus sylvestris L.) needles is accompanied by a 65% reduction of the maximum photochemical efficiency of photosystem II (PSII), measured as F _v/F _m, but relatively stable photosystem I (PSI) activity. In contrast, the photochemical efficiency of PSII in bark chlorenchyma of Scots pine twigs was shown to be well preserved, while PSI capacity was severely decreased. Low-temperature (77 K) chlorophyll fluorescence measurements also revealed lower relative fluorescence intensity emitted from PSI in bark chlorenchyma compared to needles regardless of the growing season. Nondenaturating SDS-PAGE analysis of the chlorophyll–protein complexes also revealed much lower abundance of LHCI and the CPI band related to light harvesting and the core complex of PSI, respectively, in bark chlorenchyma. These changes were associated with a 38% reduction in the total amount of chlorophyll in the bark chlorenchyma relative to winter needles, but the Chl a/b ratio and carotenoid composition were similar in the two tissues. As distinct from winter pine needles exhibiting ATP/ADP ratio of 11.3, the total adenylate content in winter bark chlorenchyma was 2.5-fold higher and the estimated ATP/ADP ratio was 20.7. The photochemical efficiency of PSII in needles attached to the twig recovered significantly faster (28–30 h) then in detached needles. Fluorescence quenching analysis revealed a high reduction state of Q _A and the PQ-pool in the green bark tissue. The role of bark chlorenchyma and its photochemical performance during the recovery of photosynthesis from winter stress in Scots pine is discussed.     

Transpiration of Pinus rotundata on a wooded peat bog in central Europe

Transpiration of a central European endemic tree species, Pinus rotundata Link, growing on a wooded peat bog in the Třeboň Basin, Czech Republic, was studied in 1999–2000. Transpiration was measured by sap flow techniques (heat field deformation method) on individual trees and scaled up to stand level. The radial patterns of sap flow density showed narrow peaks in the outer part of the xylem, sapwood accounted for 47–60% of the xylem radius and 72–84% of the xylem basal area. Adult trees tolerated well both short-term flooding during the growing season and drawdown of the water table to a depth of 60 cm below ground level. The maximum and mean daily transpiration rates were 3.0 and 1.8 mm per day, and were thus similar to published data for Scots pine. The seasonal total transpiration (25 April–20 October 2000, 180 days) amounted to 322 mm, or 62% of the potential evapotranspiration over this period. This canopy transpiration was compensated by 319 mm of precipitation. The difference between the accumulated precipitation and the accumulated transpiration (derived from seasonal sap flow measurements) closely mimicked the seasonal course of the water table.     

Ectomycorrhizal root growth in scots pine stands in response to manipulation of litter and humus layers

 The effect on ectomycorrhizal root growth in a nitrogen-enriched planted stand of Scots pine (Pinus sylvestris L.) on podzolic sandy soil to manipulation of litter and humus layers (removal, doubling and control treatments) was examined, and compared to ectomycorrhizal root growth in an untreated naturally established Scots pine stand on nutrient-poor non-podzolic sandy soil. Half a year after manipulation of litter and humus layers in the planted stand, ingrowth-cores to a depth of 60 cm were installed in both stands. Scots pine roots were sampled four times during two growing seasons. Ectomycorrhizal roots were found at all sampled soil depths to 60 cm in all plots. Root growth and ectomycorrhizal development were greater in the naturally established stand than in all plots in the planted stand. Numbers of ectomycorrhizal root tips in the litter and humus removal plots were generally higher than in the control plots in the planted stand until May 1992. Doubling litter and humus did not significantly affect root length or the numbers of ectomycorrhizal root tips. The N _dissolved , NH₄ ⁺ and NO₃ ^– concentrations and the organic matter content in the upper 5 cm of the mineral soil in the planted stand on podzolic sandy soil were generally higher and the pH significantly lower than in the naturally established stand on non-podzolic sandy soil. Root growth and ectomycorrhizal development in the secondary stand may have been negatively affected by the chemical composition of the podzolic sandy soil. Accepted: 19 March 1997     

Short-term effects of manipulated increase in acid deposition on soil, soil solution chemistry and fine roots in Scots pine (Pinus sylvestris) stand on a podzol

A manipulated increase in acid deposition (15 kg S ha⁻¹), carried out for three months in a mature Scots pine (Pinus sylvestris) stand on a podzol, acidified the soil and raised dissolved Al at concentrations above the critical level of 5 mg l⁻¹ previously determined in a controlled experiment with Scots pine seedlings. The induced soil acidification reduced tree fine root density and biomass significantly in the top 15 cm of soil in the field. The results suggested that the reduction in fine root growth was a response not simply to high Al in solution but to the depletion of exchangeable Ca and Mg in the organic layer, K deficiency, the increase in NH₄:NO₃ ratio in solution and the high proton input to the soil by the acid manipulation. The results from this study could not justify the hypothesis of Al-induced root damage under field conditions, at least not in the short term. However, the study suggests that a short exposure to soil acidity may affect the fine root growth of mature Scots pine.     

Short communication. Comparative measurements of xylem pressure in transpiring and non-transpiring leaves by means of the pressure chamber and the xylem pressure probe

Simultaneous measurements were made with the xylem pressure probe on exposed, transpiring leaves and with the Scholander pressure chamber on both transpiring and covered, non-transpiring leaves of sugarcane and maize plants. Xylem tensions inferred from pressure chamber balancing pressures on non-transpiring leaves were similar to those measured directly with the xylem pressure probe in transpiring leaves. However, tensions inferred with the pressure chamber on transpiring leaves that were placed in plastics bags just prior to excision were up to 0.6 MPa greater than those measured concurrently with the xylem pressure probe. These findings suggest that relatively large differences in water potential between the xylem and bulk leaf tissue can exist during periods of rapid transpiration, and they confirm that the balance pressure of an excised, previously transpiring leaf is only a measure of the bulk average equilibrium leaf water potential and not of the true xylem pressure that existed prior to excision.Key words: Cohesion-Tension theory, xylem pressure probe, pressure chamber, xylem tension.      

Predawn disequilibrium between plant and soil water potentials in two cold-desert shrubs

Classical water relations theory predicts that predawn plant water potential should be in equilibrium with soil water potential (soil Ψ_w) around roots, and many interpretations of plant water status in natural populations are based on this expectation. We examined this expectation for two salt-tolerant, cold-desert shrub species in glasshouse experiments where frequent watering assured homogeneity in soil Ψ_w and soil-root hydraulic continuity and where NaCl controlled soil Ψ_w. Plant water potentials were measured with a pressure chamber (xylem Ψ_p) and thermocouple psychrometers (leaf Ψ_w). Soil Ψ_w was measured with in situ thermocouple psychrometers. Predawn leaf Ψ_w and xylem Ψ_p were significantly more negative than soil Ψ_w, for many treatments, indicating large predawn soil-plant Ψ_w disequilibria: up to 1.2 MPa for Chrysothamnus nauseosus (0 and 100 mm NaCl) and 1.8 MPa for Sarcobatus vermiculatus (0, 100, 300, and 600 mm NaCl). Significant nighttime canopy water loss was one mechanism contributing to predawn disequilibrium, assessed by comparison of xylem Ψ_p for bagged (to minimize transpiration) and unbagged canopies, and by gas exchange measurements. However, nighttime transpiration accounted for only part of the predawn disequilibrium. Other mechanisms that could act with nighttime transpiration to generate large predawn disequilibria are described and include a model of how leaf apoplastic solutes could contribute to the phenomenon. This study is among the first to conclusively document such large departures from the expectation of predawn soil-plant equilibrium for C₃ shrubs, and provides a general framework for considering relative contributions of nighttime transpiration and other plant-related mechanisms to predawn disequilibrium. Received: 12 November 1998 / Accepted: 5 May 1999     

Effects of long-term open-air exposure to fluoride,nitrogen compounds and SO2 on visible symptoms,pollutant accumulation and ultrastructure of Scots pine and Norway spruce seedlings

 Effects of SO₂, aqueous fluoride (NaF) and a solution of nitrogen compounds (NH₄NO₃) on the visible symptoms, pollutant accumulation and ultrastructure of Scots pine (Pinus sylvestris L.) and Norway spruce [Picea abies (L.) Karst.] seedlings were studied in an open-air experiment lasting for 3 consecutive years. Visible injury symptoms were most pronounced in combination exposures and whenever F was applied. Visible symptoms correlated well with needle pollutant concentrations. Exposure to NaF increased needle F contents particularly when F was applied with SO₂ or NH₄NO₃. This suggests that a reduction in N or SO₂ emissions, in F polluted areas, could improve the condition of conifers via decreased accumulation of phytotoxic F in the needles. Norway spruce needles accumulated 2 – 10 times as much S and F as those of Scots pine. Microscopic observations showed various changes in the needle mesophyll cell ultrastructure. In both species, exposure to SO₂ increased significantly the amount of cytoplasmic vacuoles, suggesting detoxification of excess sulphate or low pH. F treatments resulted in a significant enlargement of plastoglobuli in Scots pine and a darkening of plastoglobuli in Norway spruce. All exposures enhanced the accumulation of lipid bodies. An increased portion of translucent plastoglobuli was most pronounced in N treatments. Many of the ultrastructural changes and visible symptoms appeared only as number of years exposed increased, indicating that long-term experiments are needed. Both visible symptoms and ultrastructural changes pointed to the more pronounced sensitivity of Norway spruce compared to Scots pine. Ultrastructural results mostly supported earlier qualitative observations of F, N and SO₂ effects on needle mesophyll cell ultrastructure. However, no reduction of thylakoids in SO₂ containing exposure or curling of thylakoids in F exposure could be detected in the present study. Received: 5 December 1994 / Accepted: 28 April 1995     

Time lags for xylem and stem diameter variations in a Scots pine tree

Diameter variations in the xylem and whole stem (i.e. over bark) stem of a Scots pine (Pinus sylvestris L.) tree were measured at four heights over a 23 d period at 5 min intervals. Cross‐correlation analysis was used to calculate time lags between the measurements. Xylem diameter measurements at the different heights had time lags varying from 10 to 50 min, measurements at the lower heights lagging behind the most. This result was in good agreement with the cohesion theory of transpiration. For the whole stem diameter measurements, the treetop lagged behind all other heights and the shortest lags were midway along the stem. Changes in whole stem diameter always lagged behind those of xylem stem diameter (30–110 min), and at all heights. The considerable differences in the behaviour of xylem and whole stem diameter support the Münch hypothesis of phloem flow. Time lags calculated separately for the shrinkage (morning) and swelling (afternoon) periods indicated shorter time lags during the swelling periods. The non‐destructive methods used show promise in the simultaneous study of flow dynamics of xylem and phloem in trees.     

Coordination of leaf and stem water transport properties in tropical forest trees

Stomatal regulation of transpiration constrains leaf water potential (Ψ_L) within species-specific ranges that presumably avoid excessive tension and embolism in the stem xylem upstream. However, the hydraulic resistance of leaves can be highly variable over short time scales, uncoupling tension in the xylem of leaves from that in the stems to which they are attached. We evaluated a suite of leaf and stem functional traits governing water relations in individuals of 11 lowland tropical forest tree species to determine the manner in which the traits were coordinated with stem xylem vulnerability to embolism. Stomatal regulation of Ψ_L was associated with minimum values of water potential in branches (Ψ_br) whose functional significance was similar across species. Minimum values of Ψ_br coincided with the bulk sapwood tissue osmotic potential at zero turgor derived from pressure–volume curves and with the transition from a linear to exponential increase in xylem embolism with increasing sapwood water deficits. Branch xylem pressure corresponding to 50% loss of hydraulic conductivity (P ₅₀) declined linearly with daily minimum Ψ_br in a manner that caused the difference between Ψ_br and P ₅₀ to increase from 0.4 MPa in the species with the least negative Ψ_br to 1.2 MPa in the species with the most negative Ψ_br. Both branch P ₅₀ and minimum Ψ_br increased linearly with sapwood capacitance (C) such that the difference between Ψ_br and P ₅₀, an estimate of the safety margin for avoiding runaway embolism, decreased with increasing sapwood C. The results implied a trade-off between maximizing water transport and minimizing the risk of xylem embolism, suggesting a prominent role for the buffering effect of C in preserving the integrity of xylem water transport. At the whole-tree level, discharge and recharge of internal C appeared to generate variations in apparent leaf-specific conductance to which stomata respond dynamically.