ABA-based chemical signalling: the co-ordination of responses to stress in plants

There is now strong evidence that the plant hormone abscisic acid (ABA) plays an important role in the regulation of stomatal behaviour and gas exchange of droughted plants. This regulation involves both long-distance transport and modulation of ABA concentration at the guard cells, as well as differential responses of the guard cells to a given dose of the hormone. We will describe how a plant can use the ABA signalling mechanism and other chemical signals to adjust the amount of water that it loses through its stomata in response to changes in both the rhizospheric and the aerial environment. The following components of the signalling process can play an important part in regulation: (a) ABA sequestration in the root; (b) ABA synthesis versus catabolism in the root; (c) the efficiency of ABA transfer across the root and into the xylem; (d) the exchange of ABA between the xylem lumen and the xylem parenchyma in the shoot; (e) the amount of ABA in the leaf symplastic reservoir and the efficiency of ABA sequestration and release from this compartment as regulated by factors such as root and leaf-sourced changes in pH; (f) cleavage of ABA from ABA conjugates in the leaf apoplast; (g) transfer of ABA from the leaf into the phloem; (h) the sensitivity of the guard cells to the [ABA] that finally reaches them; and lastly (i) the possible interaction between nitrate stress and the ABA signal.     

Survey and synthesis of intra- and interspecific variation in stomatal sensitivity to vapour pressure deficit

Responses of stomatal conductance (g_s) to increasing vapour pressure deficit (D) generally follow an exponential decrease described equally well by several empirical functions. However, the magnitude of the decrease – the stomatal sensitivity – varies considerably both within and between species. Here we analysed data from a variety of sources employing both porometric and sap flux estimates of g_s to evaluate the hypothesis that stomatal sensitivity is proportional to the magnitude of g_s at low D ( ≤ 1 kPa). To test this relationship we used the function g_s = g_sref–m· lnD where m is the stomatal sensitivity and g_sref = g_s at D = 1 kPa. Regardless of species or methodology, m was highly correlated with g_sref (average r² = 0·75) with a slope of approximately 0·6. We demonstrate that this empirical slope is consistent with the theoretical slope derived from a simple hydraulic model that assumes stomatal regulation of leaf water potential. The theoretical slope is robust to deviations from underlying assumptions and variation in model parameters. The relationships within and among species are close to theoretical predictions, regardless of whether the analysis is based on porometric measurements of g_s in relation to leaf-surface D (D_s), or on sap flux-based stomatal conductance of whole trees (G_Si), or stand-level stomatal conductance (G_S) in relation to D. Thus, individuals, species, and stands with high stomatal conductance at low D show a greater sensitivity to D, as required by the role of stomata in regulating leaf water potential.     

Effects of humidity on short-term responses of stomatal conductance to an increase in carbon dioxide concentration

The magnitude of the response of stomatal conductance to a change in the concentration of carbon dioxide external to the leaf from 350 to 700 cm³ m^–3 was found to be extremely variable from day to day in the field in Glycine max , Hordeum vulgare and Triticum aestivum . It was found that the leaf-to-air water vapour pressure difference (LAVPD) during the midday measurements of the stomatal response to carbon dioxide affected the magnitude of the response. On days when LAVPD was low, no significant change in conductance occurred with the increase in carbon dioxide concentration. When LAVPD was higher, conductance decreased by 24–52% with the increase in carbon dioxide within a few minutes. The sensitivity of conductance was approximately linearly related to LAVPD in wheat and barley. Experiments with G. max in the field indicated that, on days with low LAVPD, increasing the LAVPD just around the measured portion of a leaflet made stomatal conductance responsive to increased carbon dioxide. This result was also obtained under laboratory conditions with G. max , Helianthus annuus and Amaranthus retroflexus . In G. max , it was determined that leaves in which conductance was not responsive to the increase in carbon dioxide could be made responsive even at low LAVPD by the injection of abscisic acid into their petioles. Because it is known that abscisic acid sensitizes stomata to carbon dioxide, these results are consistent with the idea that abscisic acid may be involved in the response of stomatal conductance to changes in LAVPD.     

Effects of water vapour pressure deficit and stomatal conductance on photosynthesis,internal pressurization and convective flow in three emergent wetland plants

Internal pressurization and convective gas flow in emergent wetland plants is a function of the water vapour pressure deficit (WPD) and stomatal conductance (G _s) separating the external atmosphere from the internal aerenchyma. We have compared the effects of WPD and G _s under a range of light intensities on static pressures and convective flows in Phragmites australis, Typha orientalis and Baumea articulata. The capacity of the three species to generate flows per unit leaf area differed, being greatest in P. australisand lowest in B. articulata. In all three species, decreasing light intensity from full sunlight (2200 mol m^–2 s^–1 photosynthetically active photon flux density (PPFD)) to < 200 and < 10 mol m^–2 s^–1PPFD caused immediate decreases in photosynthetic assimilation, followed by more gradual decreases in transpiration and G _s. However, internal pressures and flows in the two low light intensities remained similar to values recorded in full sunlight. WPD was more significantly related to pressures and flows in P. australis and T. orientalis than G _s. In B. articulata, pressures increased at low G _s values but flow rates were unaffected, as predicted by earlier models describing pore size effects on pressures and flows. The data suggest that emergent macrophytes can maintain significant internal convection even at low light intensities, and this may be beneficial for nocturnal aeration, particularly in arid climates where the atmospheric humidity at night is low.     

Stomatal control by fed or endogenous xylem ABA in sunflower: interpretation of correlations between leaf water potential and stomatal conductance in anisohydric species

The stomatal conductance of several anisohydric plant species, including field-grown sunflower, frequently correlates with leaf water potential (φ₁), suggesting that chemical messages travelling from roots to shoots may not play an important role in stomatal control. We have performed a series of experiments in which evaporative demand, soil water status and ABA origin (endogenous or artificial) were varied in order to analyse stomatal control. Sunflower plants were subjected to a range of soil water potentials under contrasting air vapour pressure deficits (VPD, from 0.5 to 2.5 kPa) in the field, in the glasshouse or in a humid chamber. Sunflower plants were also fed through the xylem with varying concentrations of artificial ABA, in the glasshouse and in the field. Finally, detached leaves were fed directly with varying concentrations of ABA under three contrasting VPDs. A unique relationship between stomatal conductance (g_s) and the concentration of ABA in the xylem sap (xylem [ABA]) was observed in all cases. In contrast, the relationship between φ₁ and g_s varied substantially among experiments. Its slope was positive for droughted plants and negative for ABA-fed whole plants or detached leaves, and also varied appreciably with air VPD. All observed relationships could be modelled on the basis of the assumption that φ₁ had no controlling effect on g_s. We conclude that stomatal control depended only on the concentration of ABA in the xylem sap, and that φ₁ was controlled by water flux through the plant (itself controlled by stomatal conductance). The possibility is also raised that differences in stomatal ‘strategy’ between isohydric plants (such as maize, where daytime φ₁ does not vary appreciably with soil water status) and anisohydric plants (such as sunflower) may be accounted for by the degree of influence of φ₁ on stomatal control, for a given level of xylem [ABA]. We propose that statistical relationships between φ₁ and g_s are only observed when φ₁ has no controlling action on stomatal behaviour.     

Effects of carbon dioxide concentration on the interactive effects of temperature and water vapour on stomatal conductance in soybean

Soybeans were grown at three CO₂ concentrations in outdoor growth chambers and at two concentrations in controlled-environment growth chambers to investigate the interactive effects of CO₂, temperature and leaf-to-air vapour pressure difference (LAVPD) on stomatal conductance. The decline in stomatal conductance with CO₂ was a function of both leaf temperature and LAVPD. In the field measurements, stomatal conductance was more sensitive to LAVPD at low CO₂ at 30 °C but not at 35 °C. There was also a direct increase in conductance with temperature, which was greater at the two elevated carbon dioxide concentrations. Environmental growth chamber results showed that the relative stomatal sensitivity to LAVPD decreased with both leaf temperature and CO₂. Measurements in the environmental growth chamber were also performed at the opposing CO₂, and these experiments indicate that the stomatal sensitivity to LAVPD was determined more by growth CO₂ than by measurement CO₂. Two models that describe stomatal responses to LAVPD were compared with the outdoor data to evaluate whether these models described adequately the interactive effects of CO₂, LAVPD and temperature.     

Integration of hydraulic and chemical signalling in the control of stomatal conductance and water status of droughted plants

We describe here an integration of hydraulic and chemical signals which control stomatal conductance of plants in drying soil, and suggest that such a system is more likely than control based on chemical signals or water relations alone. The determination of xylem [ABA] and the stomatal response to xylem [ABA] are likely to involve the water flux through the plant. (1) If, as seems likely, the production of a chemical message depends on the root water status (Ψ_r), it will not depend solely on the soil water potential (Ψ_s) but also on the flux of water through the soil-plant-atmosphere continuum, to which are linked the difference between Ψ_r and Ψ_s. (2) The water flux will also dilute the concentration of the message in the xylem sap. (3) The stomatal sensitivity to the message is increased as leaf water potential falls. Stomatal conductance, which controls the water flux, therefore would be controlled by a water-flux-dependent message, with a water-flux-dependent sensitivity. In such a system, we have to consider a common regulation for stomatal conductance, leaf and root water potentials, water flux and concentration of ABA in the xylem. In order to test this possibility, we have combined equations which describe the generation and effects of chemical signals and classical equations of water flux. When the simulation was run for a variety of conditions, the solution suggested that such common regulation can operate. Simulations suggest that, as well as providing control of stomatal conductance, integration of chemical and hydraulic signalling may also provide a control of leaf water potential and of xylem [ABA], features which are apparent from our experimental data. We conclude that the root message would provide the plant with a means to sense the conditions of water extraction (soil water status and resisance to water flux) on a daily timescale, while the short-term plant response to this message would depend on the evaporative demand.     

Sequential response of whole plant water relations to prolonged soil drying and the involvement of xylem sap ABA in the regulation of stomatal behaviour of sunflower plants

Sunflower plants ( Helianihus animus cv. Tall Single Yellow} were grown in the greenhouse in drain pipes (100 mm inside diameter and 1 m long) rilled with John Innes No. 2 compost. When the fifth leaf had emerged, half of the plants were left unwatered for 6 days, rewatered for 2 days and then not watered for another 12 days. Measurements of water relations and abaxial stomatal conductance were made at each leaf position at regular intervals during the experimental period. Estimates were also made of soil water potentials along the soil profile and of ABA concentrations in xylem sap and leaves.
Soil drying led to some reduction in stomatal conductance alter only 3 days but leaf turgors were not reduced until day 13 (6 days after rewatering). When the water relations of leaves did change, older leases became substantially dehydrated while high turgors were recorded in younger leaves. Leaf ABA content measured on the third youngest leaf hardly changed over the first 13 days of the experiment, despite substantial soil drying, while xylem ABA concentrations changed very significantly and dynamically as soil water status varied, even when there was no effect of soil drying on leaf water relations. We argue that the highest ABA concentrations in the xylem, found as a result of substantial soil drying, arise from synthesis in both the roots and the older leaves, and act to delay the development of water deficit in younger leases.
In other experiments ABA solutions were watered on to the root systems of sunflower plants to increase ABA concentrations in xylem sap. The stomatal response to applied ABA was quantitatively very similar to that to ABA generated as a result of soil drying. There was a log-linear relationship between the reduction of leaf conductance and the increase of ABA concentration m xylem sap.     

ABA-deficient (aba1) and ABA-insensitive (abi1-1, abi2-1) mutants of Arabidopsis have a wild-type stomatal response to humidity

In most plant species, a decrease in atmospheric humidity at the leaf surface triggers a decrease in stomatal conductance. While guard cells appear to respond to humidity‐induced changes in transpiration rate, as opposed to relative humidity or vapour pressure difference, the underlying cellular mechanisms for this response remain unknown. In the present set of experiments, abscisic acid (ABA)‐deficient (aba1) and ABA‐insensitive (abi1‐1 and abi2‐1) mutants of Arabidopsis thaliana were used to test the hypothesis that the humidity signal is transduced by changes in the flux or concentration of ABA delivered to the stomatal complex in the transpiration stream. In gas exchange experiments, stomatal conductance was as sensitive to changes in vapour pressure difference in aba1, abi1‐1 and abi2‐1 mutant plants as in wild‐type plants. These experiments appear to rule out an obligate role for either the concentration or flux of ABA or ABA conjugates as mediators of the guard cell response to atmospheric water potential. The results stand in contrast to the well‐established role of ABA in mediating guard cell responses to decreases in soil water potential.     

Environmental and stomatal control of photosynthetic enhancement in the canopy of a sweetgum (Liquidambar styraciflua L.) plantation during 3 years of CO2 enrichment

Light‐saturated photosynthetic and stomatal responses to elevated CO₂ were measured in upper and mid‐canopy foliage of a sweetgum (Liquidambar styraciflua L) plantation exposed to free‐air CO₂ enrichment (FACE) for 3 years, to characterize environmental interactions with the sustained CO₂ effects in an intact deciduous forest stand. Responses were evaluated in relation to one another, and to seasonal patterns and natural environmental stresses, including high  temperatures, vapour pressure deficits (VPD), and drought. Photosynthetic CO₂ assimilation (A) averaged 46% higher in the +200 µmol mol^?1 CO₂ treatment, in mid‐ and upper canopy foliage. Stomatal conductance (g_s) averaged 14% (mid‐canopy) and 24% (upper canopy) lower under CO₂ enrichment. Variations in the relative responses of A and g_s were linked, such that greater relative stimulation of A was observed on dates when relative reductions in g_s were slight. Dry soils and high VPD reduced g_s and A in both treatments, and tended to diminish treatment differences. The absolute effects of CO₂ on A and g_s were minimized whenever g_s was low (<0·15 mol m^?2 s^?1), but relative effects, as the ratio of elevated to ambient rates, varied greatly under those conditions. Both stomatal and non‐stomatal limitations of A were involved during late season droughts. Leaf temperature had a limited influence on A and g_s, and there was no detectable relationship between prevailing temperature and CO₂ effects on A or g_s. The responsiveness of A and g_s to elevated CO₂, both absolute and relative, was maintained through time and within the canopy of this forest stand, subject to seasonal constraints and variability associated with limiting air and soil moisture.     

Xylem transport and shoot accumulation of lumichrome, a newly recognized rhizobial signal, alters root respiration, stomatal conductance, leaf transpiration and photosynthetic rates in legumes and cereals

* Root respiration, stomatal conductance, leaf transpiration and photosynthetic rates were measured in phytotron and field-grown plants following the application of 5 or 10 nM lumichrome, 10 nM ABA (abscisic acid) and 10 ml of 0.2 OD600 infective rhizobial cells. * Providing soybean and cowpea roots with their respective homologous rhizobia and/or purified lumichrome increased the concentration of this molecule in xylem sap and leaf extracts. Relative to control, rhizobial inoculation and lumichrome application significantly increased root respiration in maize, decreased it in lupin, but had no effect on the other test species. * Applying either lumichrome (10 nM), infective rhizobial cells or ABA to roots of plants for 44 h in growth chambers altered leaf stomatal conductance and transpiration in cowpea, lupin, soybean, Bambara groundnut and maize, but not in pea or sorghum. Where stomatal conductance was increased by lumichrome application or rhizobial inoculation, it resulted in increased leaf transpiration relative to control plants. Treating roots of field plants of cowpea with this metabolite up to 63 d after planting showed decreased stomatal conductance, which affected CO2 intake and reduction by Rubisco. * The effect of rhizobial inoculation closely mirrored that of lumichrome application to roots, indicating that rhizobial effects on these physiological activities were most likely due to lumichrome released into the rhizosphere.     

Ozone suppresses soil drying- and abscisic acid (ABA)-induced stomatal closure via an ethylene-dependent mechanism

Elevated atmospheric ozone concentrations (70 ppb) reduced the sensitivity of stomatal closure to abscisic acid (ABA) in Leontodon hispidus after at least 24 h exposure (1) when detached leaves were fed ABA, and (2) when intact plants were sprayed or injected with ABA. They also reduced the sensitivity of stomatal closure to soil drying around the roots. Such effects could already be occurring under current northern hemisphere peak ambient ozone concentrations. Leaves detached from plants which had been exposed to elevated ozone concentrations generated higher concentrations of ethylene, although leaf tissue ABA concentrations were unaffected. When intact plants were pretreated with the ethylene receptor binding antagonist 1-methylcyclopropene, the stomatal response to both applied ABA and soil drying was fully restored in the presence of elevated ozone. Implications of ethylene's antagonism of the stomatal response to ABA under oxidative stress are discussed. We suggest that this may be one mechanism whereby elevated ozone induces visible injury in sensitive species. We emphasize that drought linked to climate change and tropospheric ozone pollution, are both escalating problems. Ozone will exacerbate the deleterious effects of drought on the many plant species including valuable crops that respond to this pollutant by emitting more ethylene.     

Chemical signals and their regulations on the plant growth and water use efficiency of cotton seedlings under partial root-zone drying and different nitrogen applications

Partial root-zone drying during irrigation (PRD) has been shown effective in enhancing plant water use efficiency (WUE), however, the roles of chemical signals from root and shoot that are involved and the possible interactions affected by nitrogen nutrition are not clear. Pot-grown cotton (Gossypium spp.) seedlings were treated with three levels of N fertilization and PRD. The concentrations of nitrate (NO₃⁻), abscisic acid (ABA) and the pH value of leaf and root xylem saps, biomass and WUE were measured. Results showed that PRD plants produced larger biomass and higher WUE than non-PRD plants, with significant changes in leaf xylem ABA, leaf and root xylem NO₃⁻ concentrations and pH values, under heterogeneous soil moisture conditions. Simultaneously, high-N treated plants displayed larger changes in leaf xylem ABA and higher root xylem NO₃⁻ concentrations, than in the medium- or low-N treated plants. However, the WUE of plants in the low-N treatment was higher than that of those in the high- and medium-N treatments. PRD and nitrogen levels respectively induced signaling responses of ABA/NO₃⁻ and pH in leaf or root xylem to affect WUE and biomass under different watering levels, although significant interactions of PRD and nitrogen levels were found when these signal molecules responded to soil drying. We conclude that these signaling chemicals are regulated by interaction of PRD and nitrogen status to regulate stomatal behavior, either directly or indirectly, and thus increase PRD plant WUE under less irrigation.     

The boundary layer and the apparent responses of stomatal conductance to wind speed and to the mole fractions of CO2 and water vapour in the air

The experiments and simulations reported in this paper show that, for stomata sensitive to both CO₂ and water vapour concentrations, responses of stomatal conductance (g^w_s) to boundary layer thickness have two components, one resulting from changes in intercellular CO₂ concentration (χ^c_i) and another from changes in leaf surface water vapour saturation deficit (D^w_s). The experiments and simulations also show that the boundary layer conductance (g^w_b) can significantly alter the apparent response of g^w_s to ambient air CO₂ mole fraction (χ^c_a) and water vapour mole fraction (χ^w_a). Because of the feedback loop involved the responses of g^w_s for χ^c_a and χ^w_a each include responses to both χ^c_i and D^w_s. The boundary layer alters the state of the variables sensed by the guard cells—i.e. χ^c_i and D^w_s—and so it is a source of feedback. Thus, when scaling up from responses of stomata to the response of g^w_s for a whole leaf, the effect of the boundary layer must be considered. The results indicate that, for given responses of g^w_s to χ^c_i and D^w_s, the apparent responses of g^w_s to D^w_a and χ^c_a depend on the size of the leaf and wind speed, showing that this effect of the boundary layer should be considered when comparing data measured under different conditions, or with different methods.     

Interaction with ethylene: changing views on the role of abscisic acid in root and shoot growth responses to water stress

Shoot and root growth are differentially sensitive to water stress. Interest in the involvement of hormones in regulating these responses has focused on abscisic acid (ABA) because it accumulates in shoot and root tissues under water-limited conditions, and because it usually inhibits growth when applied to well-watered plants. However, the effects of ABA can differ in stressed and non-stressed plants, and it is therefore advantageous to manipulate endogenous ABA levels under water-stressed conditions. Studies utilizing ABA-deficient mutants and inhibitors of ABA synthesis to decrease endogenous ABA levels, and experimental strategies to circumvent variation in plant water status with ABA deficiency, are changing the view of the role of ABA from the traditional idea that the hormone is generally involved in growth inhibition. In particular, studies of several species indicate that an important role of endogenous ABA is to limit ethylene production, and that as a result of this interaction ABA may often function to maintain rather than inhibit shoot and root growth. Despite early speculation that interaction between these hormones may influence many of the effects of water deficit, this topic has received little attention until recently.     

Stomatal response to vapour pressure deficit and the effect of plant water stress

Abstract The dynamic response of stomata to changes in atmospheric humidity was investigated in Fragaria × ananassa Duch., Picea engelmannii Parry, and Pseudotsuga menziesii (Mirb.) Franco; and the effect of water stress on this response was determined in Pseudotsuga menziesii. The plants were rotated through three regimes of ambient temperature and vapour pressure deficit: 35°C–3. 5kPa, 35°C–0. 5 kPa, and 20°C–1. 5kPa. Branch and leaflet conductance were measured with a steady-state porometer, first at ambient vapour pressure deficit and then at one of four treatment conditions achieved by increasing or decreasing vapour pressure within the porometer cuvette. All three species showed similar stomatal response: enhanced conductance at low vapour pressure deficit and depressed conductance at high vapour pressure deficit. Engelmann spruce was more sensitive than Douglas fir and strawberry. Plant water status significantly altered stomatal response to vapour pressure deficit. The relationship of conductance of xylem water potential was linear under ambient conditions but became curvilinear when conductance was measured above and below ambient vapour pressure deficit. Between ?0. 5 MPa and ?2. 0 MPa xylem water potential, the stomata were sensitive to vapour pressure deficit, but below ? 2. 0 MPa, the sensitivity decreased.     

Growth response of barley and tomato to nitrogen stress and its control by abscisic acid,water relations and photosynthesis

Barley (Hordeum vulgare L.) and tomato Lycopersicon esculentum Mill.) were grown hydroponically and examined 2, 5, and 10 d after being deprived of nitrogen (N) supply. Leaf elongation rate declined in both species in response to N stress before there was any reduction in rate of dryweight accumulation. Changes in water transport to the shoot could not explain reduced leaf elongation in tomato because leaf water content and water potential were unaffected by N stress at the time leaf elongation began to decline. Tomato maintained its shoot water status in N-stressed plants, despite reduced water absorption per gram root, because the decline in root hydraulic conductance with N stress was matched by a decline in stomatal conductance. In barley the decline in leaf elongation coincided with a small (8%) decline in water content per unit area of young leaves; this decline occurred because root hydraulic conductance was reduced more strongly by N stress than was stomatal conductance. Nitrogen stress caused a rapid decline in tissue NO ₃ ^- pools and in NO ₃ ^- flux to the xylem, particularly in tomato which had smaller tissue NO ₃ ^- reserves. Even in barley, tissue NO ₃ ^- reserves were too small and were mobilized too slowly (60% in 2 d) to support maximal growth for more than a few hours. Organic N mobilized from old leaves provided an additional N source to support continued growth of N-stressed plants. Abscisic acid (ABA) levels increased in leaves of both species within 2 d in response to N stress. Addition of ABA to roots caused an increase in volume of xylem exudate but had no effect upon NO ₃ ^- flux to the xylem. After leaf-elongation rate had been reduced by N stress, photosynthesis declined in both barley and tomato. This decline was associated with increased leaf ABA content, reduced stomatal conductance and a decrease in organic N content. We suggest that N stress reduces growth by several mechanisms operating on different time scales: (1) increased leaf ABA content causing reduced cell-wall extensibility and leaf elongation and (2) a more gradual decline in photosynthesis caused by ABA-induced stomatal closure and by a decrease in leaf organic N.Abbreviation and symbols ABA abscisic acid - c_i leaf internal CO₂ concentration - L_p root hydraulic conductance     

Growth at reduced turgor: irreversible and reversible cell-wall extension of maize coleoptiles and its implications for the theory of cell growth

The relationship between steady-state elongation rate (G) and turgor pressure (P; G/P curve) was investigated using isolated segments of maize (Zea mays L.) coleoptiles incubated in osmotic solutions of a water potential range of 0 to -10 bar (polyethylene glycol 6000 as osmoticum). Short-term elongation measurements revealed curvilinear G/P curves with a steep slope at high turgor and a shallow slope at low turgor. Owing to a decrease of osmotic pressure and turgor, there was a tendency for straightening of the G/P curves during long-term elongation. An elongation rate of zero was adjusted by lowering the turgor by 4.5 bar at a constant osmotic pressure of 6.7 bar. Auxin increased — whereas abscisic acid decreased — the slope of the G/P curve but these hormones had no effect on the threshold turgor of growth (Y = 2.2 bar). It is concluded that extensibility of the growing cell walls represented by the yielding coefficient of Lockhart's growth equation is turgor-dependent and therefore decreases to a very low value as the turgor approaches Y. When the turgor was kept at Y, a constant segment length was maintained over at least 6 h. However, separation of reversible (l_rev) and irreversible (l_irr) components of total (in vivo) length (l_tot = l_rev + l_irr) W measuring segment length before and after freezing/thawing revealed that l_irr increased continuously and l_rev decreased continuously at constant l_tot. After a step-down in turgor the segments grew in l_irr although they shrank in l_tot over the whole turgor range of 0irr irreversible length - l_rev reversible length - l_tot total length (= l_irr + l_rev) - _i osmotic pressure of cell sap - _i water potential of tissue - _o water potential of incubation medium - ABA abscisic acid - G growth rate - m yielding coefficient - P turgor pressure - PEG polyethylene glycol 6000 - Y yield threshold Supported by Deutsche Forschungsgemeinschaft (SFB 206). We thank R. Hertel for helpful comments.     

A comparison of sap flux and water relations of leaves of various isolated trees with special reference to foundation movement in clay soil

Diurnal variation in sap flux (S) through stems of six trees, two each of Ulmus procera SALISB., Melaleuca styphelioides SM. and Prunus cerasifera EHRH. ‘Nigra’ (referred to hereafter by their generic names), were estimated from measurements of heat pulse velocities. Leaf water potential (ψ), stomatal conductance (g _s ) and transpiration from leaves (T) of all replicate trees were measured at 1300–1500h, once during the summer. On two separate occasions measurements were made of S, ψ, (g _s ) and T for one each of Ulmus and Melaleuca trees to study diurnal variations in these parameters. A 12×12 m² area around each tree was kept covered to simulate the condition of trees growing on pavements adjacent to residential properties. Sap flux for these tree species was in the order Melaleuca>Ulmus>Prunus. It is suggested that the smaller canopy and sapwood area in Prunus compared to the other two species is responsible for lower water potential and lower transpiration rate than the other species. Detailed analysis of the diurnal variation in sap flux and water relation of leaves of Melaleuca and Ulmus indicated sap flux of Melaleuca to be greater than that of Ulmus at the same transpiration rate per unit leaf area although the sapwood area of the two species was marginally different. This may have been due either to the difference in canopy conductance or in leaf area between the two species. With the assumption that sap flux closely resembles the rate of soil water extraction for both species, results indicate that Melaleuca is likely to extract soil water at a higher rate than Ulmus and hence is capable of causing greater shrinkage and soil movement than Ulmus.