A model of bubble growth leading to xylem conduit embolism

The dynamics of a gas bubble inside a water conduit after a cavitation event was modeled. A distinction was made between a typical angiosperm conduit with a homogeneous pit membrane and a typical gymnosperm conduit with a torus-margo pit membrane structure. For conduits with torus-margo type pits pit membrane deflection was also modeled and pit aspiration, the displacement of the pit membrane to the low pressure side of the pit chamber, was found to be possible while the emboli was still small. Concurrent with pit aspiration, the high resistance to water flow out of the conduit through the cell walls or aspirated pits will make the embolism process slow. In case of no pit aspiration and always for conduits with homogeneous pit membranes, embolism growth is more rapid but still much slower than bubble growth in bulk water under similar water tension. The time needed for the embolism to fill a whole conduit was found to be dependent on pit and cell wall conductance, conduit radius, xylem water tension, pressure rise in adjacent conduits due to water freed from the embolising conduit, and the rigidity and structure of the pits in the case of margo-torus type pit membrane. The water pressure in the conduit hosting the bubble was found to occur almost immediately after bubble induction inside a conduit, creating a sudden tension release in the conduit, which can be detected by acoustic and ultra-acoustic monitoring of xylem cavitation.     

A molecular dynamics analysis of homogeneous bubble nucleation with a noncondensable gas in cryogenic cavitation inception

In this paper, homogeneous bubble nucleation in liquid oxygen (as one of the cryogenic fluids) with a noncondensable gas of nitrogen or that of helium was investigated using molecular dynamics method employing a fitted Lennard-Jones potential. We evaluated the influence of nitrogen gas and helium gas on the SATuration line (SAT) and the spinodal line as the thermodynamic limit of stability (TLS), and on the kinetic limit of stability (KLS) defined from a bubble nucleation rate. As a result, it was obtained that the influence of the noncondensable gases on the SAT and the TLS was negligible at molar fraction less than 1% although helium gas had several times stronger action to decrease the KLS compared with nitrogen gas. On the other hand, it was also indicated that the actual influence of both noncondensable gases on the cavitation inception in liquid oxygen might be negligible at least at standard conditions where the fluid starts to flow around or less than the atmospheric pressure.     

Mathematical models of diffusion-limited gas bubble dynamics in tissue

Mathematical models of bubble evolution in tissue have recentlybeen incorporated into risk functions for predicting the incidence ofdecompression sickness (DCS) in human subjects after diving and/or flying exposures. Bubble dynamics models suitable forthese applications assume the bubble to be either contained in anunstirred tissue (two-region model) or surrounded by a boundary layerwithin a well-stirred tissue (three-region model). The contrastingpremises regarding the bubble-tissue system lead to differentexpressions for bubble dynamics described in terms of ordinarydifferential equations. However, the expressions are shown to bestructurally similar with differences only in the definitions ofcertain parameters that can be transformed to make the modelsequivalent at large tissue volumes. It is also shown that thetwo-region model is applicable only to bubble evolution in tissues ofinfinite extent and cannot be readily applied to bubble evolution infinite tissue volumes to simulate how such evolution is influenced byinteractions among multiple bubbles in a given tissue. Two-regionmodels that are incorrectly applied in such cases yield results thatmay be reinterpreted in terms of their three-region model equivalents but only if the parameters in the two-region model transform into consistent values in the three-region model. When such transforms yieldinconsistent parameter values for the three-region model, results maybe qualitatively correct but are in substantial quantitative error.Obviation of these errors through appropriate use of the differentmodels may improve performance of probabilistic models of DCSoccurrence that express DCS risk in terms of simulated in vivo gas andbubble dynamics.     

The effectiveness of a bubble curtain for preventing physical damage to antarctic krill in captivity

Krill in captivity endure spatial restrictions, regardless of the size or shape of tank used. Visual inspection of freshly caught and captive animals suggests that their appendages, particularly the antennae and antennules, are often subject to injury. We have developed a novel method for reducing physical damage to captive krill based on their reported reaction to feeding whales. This method involves creating a bubble curtain that keeps the krill in the centre of the tank and away from potentially damaging hard surfaces. There was no difference in the length of the antennae between any of the captive animals. Trends in the data suggest that the antennules of krill maintained in a 200 L bubble curtain tank and a1000 L holding tank, were longer and less variable in length, than for krill in a 200 L tank with no bubble curtain. Freshly caught krill from the field were found to have damage to both antennae and antennules. The implications of these findings for husbandry of krill and for behavioural studies in the laboratory are discussed.     

The effectiveness of a bubble curtain for preventing physical damage to antarctic krill in captivity

Krill in captivity endure spatial restrictions, regardless of the size or shape of tank used. Visual inspection of freshly caught and captive animals suggests that their appendages, particularly the antennae and antennules, are often subject to injury. We have developed a novel method for reducing physical damage to captive krill based on their reported reaction to feeding whales. This method involves creating a bubble curtain that keeps the krill in the centre of the tank and away from potentially damaging hard surfaces. There was no difference in the length of the antennae between any of the captive animals. Trends in the data suggest that the antennules of krill maintained in a 200?L bubble curtain tank and a1000?L holding tank, were longer and less variable in length, than for krill in a 200?L tank with no bubble curtain. Freshly caught krill from the field were found to have damage to both antennae and antennules. The implications of these findings for husbandry of krill and for behavioural studies in the laboratory are discussed.     

A parallel bubble column system for the cultivation of phototrophic microorganisms

An incubator with up to 16 parallel bubble columns was equipped with artificial light sources assuring a light supply with a homogenous light spectrum directly above the bioreactors. Cylindrical light reflecting tubes were positioned around every single bubble column to avoid light scattering effects and to redirect the light from the top onto the cylindrical outer glass surface of each bubble column. The light reflecting tubes were equipped with light intensity filters to control the total light intensity for every single photo-bioreactor. Parallel cultivations of the unicellular obligate phototrophic cyanobacterium, Synechococcus PCC7942, were studied under different constant light intensities ranging from 20 to 102 microE m(-2)s(-1) at a constant humidified air flow rate supplemented with CO(2).     

Thermodynamic analysis of the interaction of the xylem water and phloem sugar solution and its significance for the cohesion theory

The cohesion theory explains water transport in trees by the evaporation of water in the leaves (transpiration), which in turn generates the tension required for sap ascent, i.e. the flow of pure water from the soil through the root system and the non-living cells of the tree (xylem tracheids) up to the leaves. Only a small part of this water flow entering the leaves is used in photosynthesis to produce sugar solution, which is transported from the leaves through the living cells (phloem) to everywhere in the tree where it is needed and used. The phloem sieves are connected to the xylem tracheids by water transparent membranes, which means that the upflow of pure water and downflow of sugar solution interact with each other, causing the osmotic pressure in the sugar solution (Münch model). In this paper we analyse this interaction with a thermodynamic approach and we show that some open questions in the cohesion theory can then perhaps be better understood. For example, why under a quite high tension the water can flow in the xylem mostly without any notable cavitation, and how the suction force itself depends on the cavitation. Minimizing Gibbs energy of the system of xylem and phloem, we derive extended vapor pressure and osmotic pressure equations, which include gas bubbles in the xylem conduits as well as the cellulose-air-water interface term. With the aid of the vapor pressure equation derived here, we estimate the suction force that the cavitation controlled by the phloem sugar solution can generate at high moisture contents. We also estimate the suction force that the transpiration can generate by moisture gradient at low moisture contents. From the general osmotic pressure equation we derive an equation for calculating the degree of cavitation with different sugar solution concentrations and we show the conditions under which the cavitation in the xylem is totally avoided. Using recent field measurement results for a Scotch pine, the theory is demonstrated by showing its predictions for possible amounts of cavitation or embolism from morning hours to late afternoon.     

Impact of long-term drought on xylem embolism and growth in Pinus halepensis Mill.

 The present study was carried out to elucidate the response mechanisms of 50-year-old Pinus halepensis Mill. trees to a long-term and severe drought. The amount of water available to trees was artificially restricted for 12 months by covering the soil with a plastic roof. Over the short term a direct and rapid impact of drought was evident on the water relations and gas exchanges of trees: as the soil dried out in the Spring, there was a concurrent decrease of predawn water potential; transpiration was strongly reduced by stomatal closure. Seasonal changes in the water volume fractions of twig and stem xylem were observed and interpreted as the result of cavitation and refilling in the xylem. When droughted trees recovered to a more favourable water status, refilling of embolized xylem was observed; twig predawn water potentials were still negative in the period when the embolism was reversed in the twig xylem. A few months after the removal of the covering, no differences in whole plant hydraulic resistance were observed between droughted and control trees. Needle and shoot elongation and stem radial growth were considerably reduced in droughted trees; no strategy of trees to allocate carbon preferentially to the stem conducting tissues was apparent throughout the experiment. An after-effect of the drought on growth was observed. Received: 4 August 1997 / Accepted: 1 October 1997     

Plasmodesmatal pores in the torus of bordered pit membranes affect cavitation resistance of conifer xylem

The pit membrane in bordered pits of conifer tracheids is characterized by a porous margo and central thickening (torus), which is traditionally considered to function as an impermeable safety valve against air-seeding. However, electron microscopy based on 33 conifer species, including five families and 19 genera, reveals that pores occur in the torus of 13 of the species studied. The pores have a plasmodesmatal origin with an average diameter of 51 nm and grouped arrangement. Evidence for embolism spreading via pores in tori is supported by the pore sizes, which correspond relatively well with the pressure inducing cavitation. Predictions based on earlier correlations between pit structure and cavitation resistance were only weakly supported for species with punctured tori. Moreover, species with punctured tori are significantly less resistant to cavitation than species with non-punctured tori. Nevertheless, absolute pore diameters must be treated with caution and correlations between theoretical and measured air-seeding pressures are weak. Because most pores appear not to traverse the torus but are limited to one torus pad, only complete pores would trigger air-seeding. Embolism spreading through a leaky torus is not universal across gymnosperms and unlikely to represent the only air-seeding mechanism.     

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.     

A method for inducing xylem emboli in situ: experiments with a field-grown tree

A pressure collar, assembled around 25cm sections of 4-year-old willow twigs, was used to examine cavitation events under field conditions. When the air pressure inside the collar was raised to between 1–8 and 2–8MPa, ultrasound acoustic emission signals were triggered which indicated the breaking of water columns in the xylem. The hydraulic conductivity of the twig portion inside the chamber decreased markedly. As a result, water potentials and conductances in leaves at the end of the twig decreased. Similar changes were induced at comparable pressures in detached twigs. The equipment used is described in detail, and evidence is presented that the mechanism of this artificial production of emboli follows the air-seeding principle hypothesized for natural cavitation events.     

Molecular dynamics simulation of the first stages of the cavitation process in amorphous polymers

The first stages of the cavitation process in amorphous polymers submitted to an hydrostatic deformation in the glassy state are studied with coarse grain molecular dynamics simulations for various intermolecular interactions strengths and flexible and semi-flexible chains. For strong intermolecular interactions, the cavitation process is highly localized and the holes have a marked spherical symmetry. The cavitation regions are more diffuse for weaker intermolecular interactions or when the chain stiffness is increased. The mean Voronoï polyhedra volume and the disorder inside the polymer increase until the stress peak observed below the glass transition. High mobility regions are present before the stress peak that may act as nucleation sites for the cavitation process. The localization of these high mobility zones is enhanced for strong intermolecular interactions or a low chain rigidity. Moreover, the velocity fluctuations are more marked in the vicinity of the holes. For strong intermolecular interactions, the holes are not randomly distributed throughout the system and the nucleation of cavities upon deformation occurs preferentially near the chain ends of the polymer.     

Molecular genetic and physical analysis of gas vesicles in buoyant enterobacteria

Different modes of bacterial taxis play important roles in environmental adaptation, survival, colonization and dissemination of disease. One mode of taxis is flotation due to the production of gas vesicles. Gas vesicles are proteinaceous intracellular organelles, permeable only to gas, that enable flotation in aquatic niches. Gene clusters for gas vesicle biosynthesis are partially conserved in various archaea, cyanobacteria, and some proteobacteria, such as the enterobacterium, Serratia sp. ATCC 39006 ( S39006 ). Here we present the first systematic analysis of the genes required to produce gas vesicles in S39006 , identifying how this differs from the archaeon Halobacterium salinarum. We define 11 proteins essential for gas vesicle production. Mutation of gvpN or gvpV produced small bicone gas vesicles, suggesting that the cognate proteins are involved in the morphogenetic assembly pathway from bicones to mature cylindrical forms. Using volumetric compression, gas vesicles were shown to comprise 17% of S39006 cells, whereas in Escherichia coli heterologously expressing the gas vesicle cluster in a deregulated environment, gas vesicles can occupy around half of cellular volume. Gas vesicle production in S39006 and E. coli was exploited to calculate the instantaneous turgor pressure within cultured bacterial cells; the first time this has been performed in either strain.     

Acoustic emission analysis and experiments with physical model systems reveal a peculiar nature of the xylem tension

Advanced acoustic emission analysis, special microscopic examinations and experiments with physical model systems give reasons for the assumption that the tension in the water conducting system of vascular plants is caused by countless minute gas bubbles strongly adhering to the hydrophobic lignin domains of the xylem vessel walls. We ascertained these bubbles for several species of temperate deciduous trees and conifers. It is our hypothesis that the coherent bubble system of the xylem conduits operates as a force-transmitting medium that is capable of transporting water in traveling peristaltic waves. By virtue of the high elasticity of the gas bubbles, the hydro-pneumatic bubble system is capable of cyclic storing and releasing of energy. We consider the abrupt regrouping of the wall adherent bubble system to be the origin of acoustic emissions from plants. For Ulmus glabra, we recorded violent acoustic activity during both transpiration and re-hydration. The frequency spectrum and the waveforms of the detected acoustic emissions contradict traditional assumptions according to which acoustic emissions are caused by cavitation disruption of the stressed water column. We consider negative pressure in terms of the cohesion theory to be mimicked by the tension of the wall adherent bubble system.     

Comparative analysis of end wall resistivity in xylem conduits

The hydraulic resistivity (R, pressure gradient/flow rate) through end walls of xylem conduits was estimated in seven species of diverse anatomy and affinity including a vessel-bearing fern, a tracheid-bearing gymnosperm, and angiosperms with versus without vessels. Conduit lengths were measured with a silicone injection method which was easier and more accurate than the usual paint injection. The R declined linearly with the removal of end walls as stems were shortened from 10 to 0.3 cm. This relationship gave the minimum R with no end walls present, or the lumen resistivity (R_L). This was indistinguishable from the Hagen–Poiseuille value. The maximum R with all end walls present gave R_C, the resistivity of end wall and lumen in series. Average end-wall resistivity (R_W) was the difference R_C − R_L and the ‘wall fraction’ was R_W/R_C. Wall fraction was approximately constant, averaging 0.54 ± 0.07. This suggests that end wall and lumen resistivities are nearly co-limiting in vascular plants. Average conduit length was proportional to the diameter squared across species (r² = 0.94). Together with a constant wall fraction, this was consistent with the end wall resistance (r_w, pressure difference/flow rate) being inversely proportional to conduit length. Lower r_w in longer conduits is consistent with their having more end wall pits than shorter conduits.     

A model of extravascular bubble evolution: effect of changes in breathing gas composition.

Observations of bubble evolution in rats after decompression from air dives (O. Hyldegaard and J. Madsen. Undersea Biomed. Res. 16: 185-193, 1989; O. Hyldegaard and J. Madsen. Undersea Hyperbaric Med. 21: 413-424, 1994; O. Hyldegaard, M. Moller, and J. Madsen. Undersea Biomed. Res. 18: 361-371, 1991) suggest that bubbles may resolve more safely when the breathing gas is a heliox mixture than when it is pure O(2). This is due to a transient period of bubble growth seen during switches to O(2) breathing. In an attempt to understand these experimental results, we have developed a multigas-multipressure mathematical model of bubble evolution, which consists of a bubble in a well-stirred liquid. The liquid exchanges gas with the bubble via diffusion, and the exchange between liquid and blood is described by a single-exponential time constant for each inert gas. The model indicates that bubbles resolve most rapidly in spinal tissue, in adipose tissue, and in aqueous tissues when the breathing gas is switched to O(2) after surfacing. In addition, the model suggests that switching to heliox breathing may prolong the existence of the bubble relative to breathing air for bubbles in spinal and adipose tissues. Some possible explanations for the discrepancy between model and experiment are discussed.