Detection of ultrasonic cavitation based on low-frequency analysis of acoustic signal

The acoustic cavitation phenomenon constitutes a potential hazard in ultrasound diagnostics and therapy so that early and effective detection of cavitation is of great interest. However, cavitation might even bring a higher risk especially when an echocontrast agent based on microbubbles is used. The major goal of the present work was to develop a cavitation detection method based on increased level of cavitation noise in the range of low frequencies (about 1 Hz). This method was applied in vitro using a model of body fluid containing a model echocontrast agent, such as 5% solution of lyophilized egg albumin, which was sonicated by ultrasound disintegrator. Ultrasound signal evokes cavitation in microbubble suspension accompanied by a certain level of cavitation acoustic noise. The level of noise voltage increased in the frequency range of 0.1 to 2 Hz in the presence of cavitation. Hence, this method makes it possible to determine the value of cavitation threshold. In addition, we examined how the cavitation threshold is affected by temperature and viscosity. It was found that the cavitation threshold decreased with growing temperature while it increased with growing viscosity.     

Characterization and propagation of acoustic emission signals in woody plants: towards an improved acoustic emission counter

Abstract. The physics of ultrasonic acoustic emissions (AEs) was investigated for AE transmission through wood and transducers. The physical properties measured were velocity, attenuation and frequency composition of AEs produced by two sources: cavitation events in xylem and pencil lead breaks. The authors also measured the relative sensitivity of various combinations of ultrasound transducers and amplifiers to aid in the selection of a measuring system optimized for cavitation detection in woody plants. Some of the authors' conclusions are: (1) Softwoods ( Thuja, Pinus ) attenuate AEs more rapidly than hardwoods (maple, birch). (2) The velocity of AEs in wood exceeds that measured by others in water so the main medium of AE transmission must be the cellulose. (3) The strongest frequencies of AEs are in the range of 100–300 kHz. (4) Cavitation-induced AEs tend to shift to higher frequency as wood dehydration progresses. (5) One cannot determine the locus of origin of AEs from its frequency composition. (6) The frequency composition of the acoustic emissions probably cannot be determined at all with the sensors used because of their tendency to 'ring'. The data collected in this paper were used to aid in the design of an improved AE counter having a seven-fold increase in signal to noise ratio compared to counters previously used in our laboratory. The improved counter, model 4615 Drought Stress Monitor, is now commercially available from Physical Acoustics Corp., Princeton, NJ, U.S.A.     

Ultrasound acoustic emissions from dehydrating leaves of deciduous and evergreen trees

While drying, detached leaves produced ultrasound acoustic emissions (UAE) comparable to emissions from stem and twig wood. Experiments on Ilex aquifolium L. showed that the main source of these signals was cavitation in the veins, to which conduits and fibres probably both contributed. Regions of the leaf blade with abundant mesophyll and only small veins emitted few signals. More signals were counted on the adaxial side of the midrib than on the abaxial one and on the proximal third than on the distal one, in accordance with the anatomical structure. Sound attenuation was pronounced. Eight species were compared with respect to cavitation behaviour, field water relations and pressure–volume curves, and the data showed differences in cumulative number of events and resistance of leaves to cavitation. Data were generally in good agreement with anatomical structure and habitat preferences. The number of signals per conduit counted on cross-sections was in some leaves much higher than unity, which suggests short xylem elements or an acoustic activity of cells other than conduits. There was no correlation between cavitation threshold or cumulative number of signals and the degree of sclerophylly; unexpectedly, there was a correlation between the cumulative number of signals at a water potential of -1.3 MPa and the bulk modulus of elasticity.     

Leaf size serves as a proxy for xylem vulnerability to cavitation in plantation trees

Hybrid poplars are an important renewable forest resource known for their high productivity. At the same time, they are highly vulnerable to water stress. Identifying traits that can serve as indicators for growth performance remains an important task, particularly under field conditions. Understanding which trait combinations translate to improved productivity is key in order to satisfy the demand for poplar wood in an uncertain future climate. In this study, we compared hydraulic and leaf traits among five hybrid poplar clones at 10 plantations in central Alberta. We also assessed the variation of these traits between 2‐ to 3‐year‐old branches from the lower to mid‐crown and current‐year long shoots from the mid to upper crown. Our results showed that (1) hybrid poplars differed in key hydraulic parameters between branch type, (2) variation of hydraulic traits among clones was relatively large for some clones and less for others, and (3) strong relationships between measured hydraulic traits, such as vessel diameter, cavitation resistance, xylem‐specific and leaf‐specific conductivity and leaf area, were observed. Our results suggest that leaf size could serve as an additional screening tool when selecting for drought‐tolerant genotypes in forest management and tree improvement programmes.     

Hydraulic responses to height growth in maritime pine trees

As trees grow taller, decreased xylem path conductance imposes a major constraint on plant water and carbon balance, and is thus a key factor underlying forest productivity decline with age. The responses of stomatal conductance, leaf area: sapwood area ratio (A_L : A_S) and soil–leaf water potential gradient (ΔΨ_S–L) to height growth were investigated in maritime pine trees. Extensive measurements of in situ sap flow, stomatal conductance and (non‐gravitational) needle water potential (_L = Ψ_L ? ρ_wgh) were made during 2 years in a chronosequence of four even‐aged stands, under both wet and dry soil conditions. Under wet soil conditions, _L was systematically lower in taller trees on account of differences in gravitational potential. In contrast, under dry soil conditions, our measurements clearly showed that _L was maintained above a minimum threshold value of ?2.0 MPa independently of tree height, thus limiting the range of compensatory change in ΔΨ_S–L. Although a decrease in the A_L : A_S ratio occurred with tree height, this compensation was not sufficient to prevent a decline in leaf‐specific hydraulic conductance, K_L (50% lower in 30 m trees than in 10 m trees). An associated decline in stomatal conductance with tree height thus occurred to maintain a balance between water supply and demand. Both the increased investment in non‐productive versus productive tissues (A_S : A_L) and stomatal closure may have contributed to the observed decrease in tree growth efficiency with increasing tree height (by a factor of three from smallest to tallest trees), although other growth‐limiting responses (e.g. soil nutrient sequestration, increased respiratory costs) cannot be excluded.     

Water transport through tall trees: A vertically explicit,analytical model of xylem hydraulic conductance in stems

Trees grow by vertically extending their stems, so accurate stem hydraulic models are fundamental to understanding the hydraulic challenges faced by tall trees. Using a literature survey, we showed that many tree species exhibit continuous vertical variation in hydraulic traits. To examine the effects of this variation on hydraulic function, we developed a spatially explicit, analytical water transport model for stems. Our model allows Huber ratio, stem‐saturated conductivity, pressure at 50% loss of conductivity, leaf area, and transpiration rate to vary continuously along the hydraulic path. Predictions from our model differ from a matric flux potential model parameterized with uniform traits. Analyses show that cavitation is a whole‐stem emergent property resulting from non‐linear pressure‐conductivity feedbacks that, with gravity, cause impaired water transport to accumulate along the path. Because of the compounding effects of vertical trait variation on hydraulic function, growing proportionally more sapwood and building tapered xylem with height, as well as reducing xylem vulnerability only at branch tips while maintaining transport capacity at the stem base, can compensate for these effects. We therefore conclude that the adaptive significance of vertical variation in stem hydraulic traits is to allow trees to grow tall and tolerate operating near their hydraulic limits.     

Ultrasonic acoustic emissions in drought-stressed trees--more than signals from cavitation?

Ultrasonic acoustic emission (UAE) in trees is often related to collapsing water columns in the flow path as a result of tensions that are too strong (cavitation). However, in a decibel (dB) range below that associated with cavitation, a close relationship was found between UAE intensities and stem radius changes. UAE was continuously recorded on the stems of mature field-grown trees of Scots pine (Pinus sylvestris) and pubescent oak (Quercus pubescens) at a dry inner-Alpine site in Switzerland over two seasons. The averaged 20-Hz records were related to microclimatic conditions in air and soil, sap-flow rates and stem-radius fluctuations detrended for growth (Delta W). Within a low-dB range (27 +/- 1 dB), UAE regularly increased and decreased in a diurnal rhythm in parallel with DeltaW on cloudy days and at night. These low-dB emissions were interrupted by UAE abruptly switching between the low-dB range and a high-dB range (36 +/- 1 dB) on clear, sunny days, corresponding to the widely supported interpretation of UAE as sound from cavitations. It is hypothesized that the low-dB signals in drought-stressed trees are caused by respiration and/or cambial growth as these physiological activities are tissue water-content dependent and have been shown to produce courses of CO(2) efflux similar to our courses of Delta W and low-dB UAE.     

Xylem cavitation in two mature Scots pine forests growing in a wet and a dry area of Britain

The xylem cavitation rate, stem water content, stomatal conductance and leaf water potential of mature Scots pine (Pinus sylvestris L.) were compared over a period of 18 months at two sites in Britain where trees were planted from the same seed source. The sites were at Thetford in south-east England, where the climate is relatively warm and dry (average rainfall of 600mm per year), and Aberfoyle in central Scotland, where it is relatively cool and wet (average rainfall of 1500 mm year^?1). In the first year of study (1992) the natural difference between the sites was amplified by a severe drought affecting south-east England. Acoustic emissions (as a result of cavitation) were detected at both sites, but were not an everyday occurrence, and rates depended on short-term meteorological variation. The relative water content (RWC) of the trunk at breast height at Thetford was significantly lower than at Aberfoyle, and declined in parallel with increasing severity of drought. Following the release from drought a gradual increase in RWC was found, but the pre-drought RWC was not attained. The same xylem water potential gradient and stomatal conductance was found at the two sites; but on a diurnal basis, as water potential declined, cavitation rates increased. There was no difference in vulnerability to cavitation or in hydraulic conductance between the sites.     

Vulnerability to xylem cavitation and the distribution of Sonoran Desert vegetation

We studied 15 riparian and upland Sonoran desert species to evaluate how the limitation of xylem pressure (Ψ(x)) by cavitation corresponded with plant distribution along a moisture gradient. Riparian species were obligate riparian trees (Fraxinus velutina, Populus fremontii, and Salix gooddingii), native shrubs (Baccharis spp.), and an exotic shrub (Tamarix ramosissima). Upland species were evergreen (Juniperus monosperma, Larrea tridentata), drought-deciduous (Ambrosia dumosa, Encelia farinosa, Fouquieria splendens, Cercidium microphyllum), and winter-deciduous (Acacia spp., Prosopis velutina) trees and shrubs. For each species, we measured the "vulnerability curve" of stem xylem, which shows the decrease in hydraulic conductance from cavitation as a function of Ψ(x) and the Ψ(crit) representing the pressure at complete loss of transport. We also measured minimum in situ Ψ(x)(Ψ(xmin)) during the summer drought. Species in desert upland sites were uniformly less vulnerable to cavitation and exhibited lower Ψ(xmin) than riparian species. Values of Ψ(crit) were correlated with minimum Ψ(x). Safety margins (Ψ(xmin)-Ψ(crit)) tended to increase with decreasing Ψ(xmin) and were small enough that the relatively vulnerable riparian species could not have conducted water at the Ψ(x) experienced in upland habitats (-4 to -10 MPa). Maintenance of positive safety margins in riparian and upland habitats was associated with minimal to no increase in stem cavitation during the summer drought. The absence of less vulnerable species from the riparian zone may have resulted in part from a weak but significant trade-off between decreasing vulnerability to cavitation and conducting efficiency. These data suggest that cavitation vulnerability limits plant distribution by defining maximum drought tolerance across habitats and influencing competitive ability of drought tolerant species in mesic habitats.     

长白山阔叶红松林树木N2O排放及总量初步估算

大气中主要温室气体N2 O的部分来源尚不清楚。以往国内外对森林生态系统N2 O排放通量的测定中 ,只测土壤通量而把树木排除在外。如果树木在自然状态下能排放N2 O ,那么森林生态系统的N2 O排放可能被低估。本文旨在证明自然状态下森林生态系统中树木也是N2 O的主要排放源。采用封闭罩法 ,在树木生长的主要季节 (7～ 9月 )对长白山阔叶红松林的几个主要树种———水曲柳、红松和椴树的连体枝叶释放N2 O的速率进行了原位测定。并在此基础上 ,初步估算出森林树木N2 O年排放量是土壤N2 O年排放量的 0 8～ 1 0 3倍 ,相当于甚至超过了土壤的N2 O排放量。     

Ecological differentiation in xylem cavitation resistance is associated with stem and leaf structural traits

Cavitation resistance is a critical determinant of drought tolerance in tropical tree species, but little is known of its association with life history strategies, particularly for seasonal dry forests, a system critically driven by variation in water availability. We analysed vulnerability curves for saplings of 13 tropical dry forest tree species differing in life history and leaf phenology. We examined how vulnerability to cavitation (P₅₀) related to dry season leaf water potentials and stem and leaf traits. P₅₀‐values ranged from ?0.8 to ?6.2 MPa, with pioneers on average 38% more vulnerable to cavitation than shade‐tolerants. Vulnerability to cavitation was related to structural traits conferring tissue stress vulnerability, being negatively correlated with wood density, and surprisingly maximum vessel length. Vulnerability to cavitation was negatively related to the Huber‐value and leaf dry matter content, and positively with leaf size. It was not related to SLA. We found a strong trade‐off between cavitation resistance and hydraulic efficiency. Most species in the field were operating at leaf water potentials well above their P₅₀, but pioneers and deciduous species had smaller hydraulic safety margins than shade‐tolerants and evergreens. A trade‐off between hydraulic safety and efficiency underlies ecological differentiation across these tropical dry forest tree species.     

Limitation of transpiration by hydraulic conductance and xylem cavitation in Betula occidentalis

The extent to which stomatal conductance (g_s) was capable of responding to reduced hydraulic conductance (k)and preventing cavitation-inducing xylem pressures was evaluated in the small riparian tree, Betula occidentalis Hook. We decreased k by inducing xylem cavitation in shoots using an air-injection technique. From 1 to 18 d after shoot injection we measured midday transpiration rate (E), g_s, and xylem pressure (Ψ_p-xylem) on individual leaves of the crown. We then harvested the shoot and made direct measurements of k from the trunk (2–3 cm diameter) to the distal tip of the petioles of the same leaves measured for E and g_s. The k measurement was expressed per unit leaf area (k_l, leaf-specific conductance). Leaves measured within 2 d of shoot injection showed reduced g_s and E relative to non-injected controls, and both parameters were strongly correlated with k_l At this time, there was no difference in leaf Ψ_p-xylem between injected shoots and controls, and leaf Ψ_p-xylem was not significantly different from the highest cavitation-inducing pressure (Ψ_p-cav) in the branch xylem (-1.43 ± 0.029 MPa, n=8). Leaves measured 7–18 d after shoots were injected exhibited a partial return of g_s and E values to the control range. This was associated with a decrease in leaf Ψ_p-xylem below Ψ_p-cav and loss of foliage. The results suggest the stomata were incapable of long-term regulation of E below control values and that reversion to higher E caused dieback via cavitation.     

Evaluation of centrifugal methods for measuring xylem cavitation in conifers, diffuse- and ring-porous angiosperms

A centrifugal method is used to measure 'vulnerability curves' which show the loss of hydraulic conductivity in xylem by cavitation. Until recently, conductivity was measured between bouts of centrifugation using a gravity-induced head. Now, conductivity can be measured during centrifugation. This 'spin' method is faster than the 'gravity' technique, but correspondence between the two has not been evaluated. The two methods were compared on the same stem segments for two conifer, four diffuse-porous, and four ring-porous species. Only 17 of 60 conductivity measurements differed, with differences in the order of 10%. When different, the spin method gave higher conductivities at the beginning of the curve and lower at the end. Pressure at 50% loss of conductivity, and mean cavitation pressure, were the same in 14 of 20 comparisons. When different, the spin method averaged 0.32 MPa less negative. Ring-porous species showed a precipitous initial drop in conductivity by both techniques. This striking pattern was confirmed by the air-injection method and native embolism measurements. Close correspondence inspires confidence in both methods, each of which has unique advantages. The observation that ring-porous species operate at only a fraction of their potential conductivity at midday demands further study.     

Relations between stomatal closure, leaf turgor and xylem vulnerability in eight tropical dry forest trees

This study examined the linkage between xylem vulnerability, stomatal response to leaf water potential (Ψ_L), and loss of leaf turgor in eight species of seasonally dry tropical forest trees. In order to maximize the potential variation in these traits species that exhibit a range of leaf habits and phenologies were selected. It was found that in all species stomatal conductance was responsive to Ψ_L over a narrow range of water potentials, and that Ψ_L inducing 50% stomatal closure was correlated with both the Ψ_L inducing a 20% loss of xylem hydraulic conductivity and leaf water potential at turgor loss in all species. In contrast, there was no correlation between the water potential causing a 50% loss of conductivity in the stem xylem, and the water potential at stomatal closure (Ψ_SC) amongst species. It was concluded that although both leaf and xylem characters are correlated with the response of stomata to Ψ_L, there is considerable flexibility in this linkage. The range of responses is discussed in terms of the differing leaf‐loss strategies exhibited by these species.     

红松阔叶林倒木贮量动态的研究

在森林倒木研究的基础上探讨长白山红松阔叶林倒木贮量的动态,涉及红松阔叶林倒木分解及其贮量的动态规律。研究表明,倒木分解,除心腐木外,均由表及里进行;倒木分解速率在其它生态条件相同时因树种、直径和部位而异。红松阔叶林倒木贮量动态包括现有倒木贮量和倒木年输入量两个分解动态过程,现有倒木贮量在头100年其干重迅速减少,其中椴树比红松尤速,前者分解91%,后者为72%.林地倒木贮量动态与倒木年输入量分解动态相似,但前者在分解初期贮量增加较大,因为部分现有倒木未完全分解;100年后趋于一致,并恒定于16～17t·hm^-2,直至群落的顶极阶段结束.     

Diurnal changes in water content of the stems of apple trees, as influenced by irrigation

Abstract Groups of apple trees within an orchard were irrigated by either releasing a fine mist within the canopy or spraying water on the soil. Diurnal changes in the water content of the xylem were inferred from measurements of density made with a gamma probe. Transpiration rates were estimated from leaf temperatures, and water potentials and changes in radial dimension of the stem were recorded. Xylem water content and water potential declined each morning and recovered in the afternoon. Radial dimensions of the xylem followed a similar pattern, but the percentage change was very small in relation to the change in water content of the same tissue. The decline in water content is unlikely to have been caused by cavitation, as it was readily reversed in the afternoon. It is more likely to have been caused by changes in the volume of water occupying already-cavitated fibres or intercellular spaces. Water potential and xylem water content were highest in the mist-irrigated trees and least in the unirrigated controls.     

Water storage in the wood and xylem cavitation in 1-year-old twigs of Populus deltoides Bartr.

The possible role of water expelled from cavitated xylem conduits in the rehydration of water-stressed leaves has been studied in one-year-old twigs of populus deltoides Bartr. Twigs were dehydrated in air. At desired values of leaf water potential (Ψ_l) (between near full turgor and -1.62 MPa), twigs were placed in black plastic bags for 1–2h. Leaf water content was measured every 3–5 min before bagging and every 10 min in the dark. Hydraulic conductivity and xylem cavitation were measured both in the open and in the dark. Cavitation was monitored as ultrasound acoustic emissions (AE). A critical Ψ_l value of -0.96 MPa was found, at which AE increased significantly while the leaf water deficit decreased by gain of water. Since the twigs were no longer attached to roots, it was concluded that water expelled from cavitated xylem conduits was transported to the leaves, thus contributing to their rehydration. Xylem cavitation is discussed in terms of a ‘leaf water deficit buffer mechanism’, under not very severe water stress conditions.     

Xylem cavitation and hydraulic control of stomatal conductance in Laurel (Laurus nobilis L.)

The possible link between stomatal conductance (g_L), leaf water potential ( Ψ _L) and xylem cavitation was studied in leaves and shoots of detached branches as well as of whole plants of Laurus nobilis L. (Laurel). Shoot cavitation induced complete stomatal closure in air‐dehydrated detached branches in less than 10 min. By contrast, a fine regulation of g_L in whole plants was the consequence of Ψ _L reaching the cavitation threshold ( Ψ _CAV) for shoots. A pulse of xylem cavitation in the shoots was paralleled by a decrease in g_L of about 50%, while Ψ _L stabilized at values preventing further xylem cavitation. In these experiments, no root signals were likely to be sent to the leaves from the roots in response to soil dryness because branches were either detached or whole plants were growing in constantly wet soil. The stomatal response to increasing evaporative demand appeared therefore to be the result of hydraulic signals generated during shoot cavitation. A negative feedback link is proposed between g_L and Ψ _CAV rather than with Ψ _L itself.