Study of xylem pressure potential daily dynamics by means of autocorrelation analysis

The xylem pressure potential (Ψ_xylem) of the leaves ofQuercus cerris, Acer campestre andCarpinus betulus was measured under anticyclonic weather types. The autocorrelation analysis revealed the daily course of the Ψ_xylem values approaching the stationary random process. A close statistical relation was found between the results obtained in three successive measurements of the Ψ_xylem (interval 2 h). A close statistical relation also between the value of the base potential (Ψ_b) measured at dawn and the actual values of the Ψ_xylem allowed the prediction of the Ψ_xylem values on the base of the known Ψ_b-values by means of a simple linear regression model.     

Xylem cavitation vulnerability influences tree species’ habitat preferences in miombo woodlands

Although precipitation plays a central role in structuring Africa’s miombo woodlands, remarkably little is known about plant-water relations in this seasonally dry tropical forest. Therefore, in this study, we investigated xylem vulnerability to cavitation for nine principal tree species of miombo woodlands, which differ in habitat preference and leaf phenology. We measured cavitation vulnerability (Ψ₅₀), stem-area specific hydraulic conductivity (K _S), leaf specific conductivity (K _L), seasonal variation in predawn water potential (Ψ_PD) and xylem anatomical properties [mean vessel diameter, mean hydraulic diameter, mean hydraulic diameter accounting for 95 % flow, and maximum vessel length (V _L)]. Results show that tree species with a narrow habitat range (mesic specialists) were more vulnerable to cavitation than species with a wide habitat range (generalists). Ψ₅₀ for mesic specialists ranged between ?1.5 and ?2.2 MPa and that for generalists between ?2.5 and ?3.6 MPa. While mesic specialists exhibited the lowest seasonal variation in Ψ_PD, generalists displayed significant seasonal variations in Ψ_PD suggesting that the two miombo habitat groups differ in their rooting depth. We observed a strong trade-off between K _S and Ψ₅₀ suggesting that tree hydraulic architecture is one of the decisive factors setting ecological boundaries for principal miombo species. While vessel diameters correlated weakly (P > 0.05) with Ψ₅₀, V _L was positively and significantly correlated with Ψ₅₀. Ψ_PD was significantly correlated with Ψ₅₀ further reinforcing the conclusion that tree hydraulic architecture plays a significant role in species’ habitat preference in miombo woodlands.     

Homeostasis in leaf water potentials on leeward and windward sides of desert shrub crowns: water loss control vs. high hydraulic efficiency

Phenotypic plasticity in morphophysiological leaf traits in response to wind was studied in two dominant shrub species of the Patagonian steppe, used as model systems for understanding effects of high wind speed on leaf water relations and hydraulic properties of small woody plants. Morpho-anatomical traits, hydraulic conductance and conductivity and water relations in leaves of wind-exposed and protected crown sides were examined during the summer with nearly continuous high winds. Although exposed sides of the crowns were subjected to higher wind speeds and air saturation deficits than the protected sides, leaves throughout the crown had similar minimum leaf water potential (Ψ_L). The two species were able to maintain homeostasis in minimum Ψ_L using different physiological mechanisms. Berberis microphylla avoided a decrease in the minimum Ψ_L in the exposed side of the crown by reducing water loss by stomatal control, loss of cell turgor and low epidermal conductance. Colliguaja integerrima increased leaf water transport efficiency to maintain transpiration rates without increasing the driving force for water loss in the wind-exposed crown side. Leaf physiological changes within the crown help to prevent the decrease of minimum Ψ_L and thus contribute to the maintenance of homeostasis, assuring the hydraulic integrity of the plant under unfavorable conditions. The responses of leaf traits that contribute to mechanical resistance (leaf mass per area and thickness) differed from those of large physiological traits by exhibiting low phenotypic plasticity. The results of this study help us to understand the unique properties of shrubs which have different hydraulic architecture compared to trees.     

Seasonal physiological responses of Argania spinosa tree from Mediterranean to semi-arid climate

Argania spinosa (the argan tree) is a slow-growing tree endemic of Morocco, growing on semi-arid areas where no other tree species can live. With the aim of predicting temporal changes in A. spinosa woodlands under a probable increase in aridity, we set off to investigate these questions: how do A. spinosa physiological attributes respond to variations in climatic conditions and seasonality, and which is the set of attributes that most affects tree response to environmental conditions? In three study sites, Beni Snassen (North), High-Atlas (Mountain) and Admine Forest in Agadir (Coastal), gas exchange measurements, photochemical efficiency, leaf water potential and different leaf attributes were monitored in February, July and November of 2006. The Mountain site presents the most continental climate. Trees in this site were the most stressed in summer, having the lowest midday leaf water potential values, photochemical efficiency and assimilation rates. We found a Ψ_md threshold around -4 MPa, below which stomatal conductance responds linearly to Ψ_md. Plants from the North area never reached this threshold during the study period. Although leaf pigments presented a clear seasonal pattern, leaves from Coastal trees exhibit the highest content for each season. The three study sites were separated by two discriminate functions obtained by canonical discriminant analysis. In summer, the Mountain population is separated from the other sites mainly by assimilation rate and F_v/F_m, while in winter transpiration rates and chlorophyll content are the main discriminant variables. Our study shows that A. spinosa trees adjust their physiological status and leaf attributes to environmental conditions allowing plants to thrive under a dry climate. Under a scenario of global change, the distribution of the argan tree likely shifts to milder areas.     

Allocation to leaf area and sapwood area affects water relations of co-occurring savanna and forest trees

Water availability is a principal factor limiting the distribution of closed-canopy forest in the seasonal tropics, suggesting that forest tree species may not be well adapted to cope with seasonal drought. We studied 11 congeneric species pairs, each containing one forest and one savanna species, to test the hypothesis that forest trees have a lower capacity to maintain seasonal homeostasis in water relations relative to savanna species. To quantify this, we measured sap flow, leaf water potential (Ψ_L), stomatal conductance (g _s), wood density, and Huber value (sapwood area:leaf area) of the 22 study species. We found significant differences in the water relations of these two species types. Leaf area specific hydraulic conductance of the soil/root/leaf pathway (G _t) was greater for savanna species than forest species. The lower G _t of forest trees resulted in significantly lower Ψ_L and g _s in the late dry season relative to savanna trees. The differences in G _t can be explained by differences in biomass allocation of savanna and forest trees. Savanna species had higher Huber values relative to forest species, conferring greater transport capacity on a leaf area basis. Forest trees have a lower capacity to maintain homeostasis in Ψ_L due to greater allocation to leaf area relative to savanna species. Despite significant differences in water relations, relationships between traits such as wood density and minimum Ψ_L were indistinguishable for the two species groups, indicating that forest and savanna share a common axis of water-use strategies involving multiple traits.     

Water relations of Caatinga trees in the dry season

The Caatinga is one of the world's richest dry forests. This forest occurs only in Brazil, but is the least studied and protected Brazilian ecosystem. There are few reports about drought tolerance mechanisms in Caatinga trees. This work evaluates water relations of six adult species in the middle of the dry season to further understand water relations in this ecosystem, which will be important for future reforestation and management projects. Based on results, the trees were classified into four groups: (I), Mimosa caesalpiniifolia had low leaf water potential (Ψ_w) at predawn and no significant decrease at midday. Stomatal conductance (g_s) analyses indicates that plants have reached its lowest Ψ_w; (II), Caesalpinia pyramidalis and Auxemma oncocalyx had low Ψ_w at predawn and significant decrease at midday. For these species the recuperation of water status at night may have been sufficient for maintaining stomata open during the day; (III), Caesalpinia ferrea and Calliandra spinosa had relatively high Ψ_w at predawn and a significant decrease at midday. These species might maintain their water status similar to individuals of group II, but they might also have deeper root systems; and (IV), Tabebuia caraiba with the highest Ψ_w at predawn and no significant decrease at midday, possibly indicating a combination of good stomatal control of water loss and a deeper root system. Moreover, except for individuals of group I, both in species with lower and higher Ψ_w at predawn it was not observed strong inhibition of g_s.     

Water potential — water saturation deficit relationship during dehydration and resaturation of leaves

The relationship between the water potential (Ψ_w) and the water saturation deficit (Δ W _sat) in kale and maize leaf tissue was measured during dehydration and resaturation either of leavesin situ or of cut leaves. The curves relating Ψ_w toΔW _sat were similar in all variants, but at the same values ofΔ W _sat corresponding values of Ψ_w were always lower in leavesin situ than in cut leaves and during dehydration than during resaturation.     

The distribution of four Caragana species is related to their differential responses to drought stress

Caragana species are widespread in northwest China. However, species in semiarid areas of Inner Mongolia experience summer and autumn rainfall, whereas morphologically similar close relatives in arid areas in Xinjiang Province experience wet springs, but hot dry summers. We hypothesize that the differences among species in response to soil drought help to explain their distributions. A comparison of the closely related species C. intermedia and C. microphylla from Inner Mongolia (semiarid species) and C. pruinosa and C. spinosa from Xinjiang (arid species) was conducted to examine whether responses to a water deficit in a pot experiment were associated with the environments of their habitat. In the two semiarid species, the stomatal conductance was relatively insensitive to the decrease in predawn leaf water potential (Ψ_leaf), and leaves did not abscise or die even when the Ψ_leaf decreased below ?6.0 MPa, while in seedlings of the two arid species the stomata closed at a relatively high Ψ_leaf, and the leaves abscised gradually when Ψ_leaf fell below about ?3.0 MPa. Furthermore, major vein density, minor vein density and the cubed ratio of the conduit wall thickness to the conduit lumen breadth [(t/b)³] of minor veins increased by an average of 60, 150 and 220 %, respectively, in the semiarid species compared with the arid species. The results indicated that semiarid species have typical anisohydric properties, while arid species have typical isohydric properties. We suggest that the divergence in water-use strategies in the genus may be associated with variation in vein architecture, and may possibly play an important role in determining the distribution of these species in the different environments of northwest China.     

The ecosystem wilting point defines drought response and recovery of a Quercus-Carya forest

Soil and atmospheric droughts increasingly threaten plant survival and productivity around the world. Yet, conceptual gaps constrain our ability to predict ecosystem-scale drought impacts under climate change. Here, we introduce the ecosystem wilting point (Ψ_EWP), a property that integrates the drought response of an ecosystem's plant community across the soil–plant–atmosphere continuum. Specifically, Ψ_EWP defines a threshold below which the capacity of the root system to extract soil water and the ability of the leaves to maintain stomatal function are strongly diminished. We combined ecosystem flux and leaf water potential measurements to derive the Ψ_EWP of a Quercus-Carya forest from an “ecosystem pressure–volume (PV) curve,” which is analogous to the tissue-level technique. When community predawn leaf water potential (Ψ_pd) was above Ψ_EWP (=−2.0 MPa), the forest was highly responsive to environmental dynamics. When Ψ_pd fell below Ψ_EWP, the forest became insensitive to environmental variation and was a net source of carbon dioxide for nearly 2 months. Thus, Ψ_EWP is a threshold defining marked shifts in ecosystem functional state. Though there was rainfall-induced recovery of ecosystem gas exchange following soaking rains, a legacy of structural and physiological damage inhibited canopy photosynthetic capacity. Although over 16 growing seasons, only 10% of Ψ_pd observations fell below Ψ_EWP, the forest is commonly only 2–4 weeks of intense drought away from reaching Ψ_EWP, and thus highly reliant on frequent rainfall to replenish the soil water supply. We propose, based on a bottom-up analysis of root density profiles and soil moisture characteristic curves, that soil water acquisition capacity is the major determinant of Ψ_EWP, and species in an ecosystem require compatible leaf-level traits such as turgor loss point so that leaf wilting is coordinated with the inability to extract further water from the soil.     

Responses of epidermal diffusive conductance to simultaneous changes in two factors: Determination of interactions

Responses of the epidermal diffusive conductance (gep) to irradiance (I) during ontogeny of primary bean leaves or during their wilting were followed. Effects ofI, leaf age and leaf water potential (Ψw) as well as interactive effects (I × leaf age andI × Ψw) ongep were statistically significant.     

Osmotic and elastic adjustments in cold desert shrubs differing in rooting depth: coping with drought and subzero temperatures

Physiological adjustments to enhance tolerance or avoidance of summer drought and winter freezing were studied in shallow- to deep-rooted Patagonian cold desert shrubs. We measured leaf water potential (Ψ_L), osmotic potential, tissue elasticity, stem hydraulic characteristics, and stomatal conductance (g _S) across species throughout the year, and assessed tissue damage by subzero temperatures during winter. Species behavior was highly dependent on rooting depth. Substantial osmotic adjustment (up to 1.2?MPa) was observed in deep-rooted species exhibiting relatively small seasonal variations in Ψ_L and with access to a more stable water source, but having a large difference between predawn and midday Ψ_L. On the other hand, shallow-rooted species exposed to large seasonal changes in Ψ_L showed limited osmotic adjustment and incomplete stomatal closure, resulting in turgor loss during periods of drought. The bulk leaf tissue elastic modulus (ε) was lower in species with relatively shallow roots. Daily variation in g _S was larger in shallow-rooted species (more than 50?% of its maximum) and was negatively associated with the difference between Ψ_L at the turgor loss point and minimum Ψ_L (safety margin for turgor maintenance). All species increased ε by about 10?MPa during winter. Species with rigid tissue walls exhibited low leaf tissue damage at ?20?°C. Our results suggest that osmotic adjustment was the main water relationship adaptation to cope with drought during summer and spring, particularly in deep-rooted plants, and that adjustments in cell wall rigidity during the winter helped to enhance freezing tolerance.     

Changes in leaf hydraulic conductance correlate with leaf vein embolism in<Emphasis Type="Italic"> Cercis siliquastrum</Emphasis> L.

The impact of xylem cavitation and embolism on leaf (K _leaf) and stem (K _stem) hydraulic conductance was measured in current-year shoots of Cercis siliquastrum L. (Judas tree) using the vacuum chamber technique. K _stem decreased at leaf water potentials (Ψ_L) lower than ?1.0 MPa, while K _leaf started to decrease only at Ψ_L L K _leaf changes. Field measurements of leaf conductance to water vapour (g _L) and Ψ_L showed that stomata closed when Ψ_L decreased below the Ψ_L threshold inducing loss of hydraulic conductance in the leaf. The partitioning of hydraulic resistances within shoots and leaves was measured using the high-pressure flow meter method. The ratio of leaf to shoot hydraulic resistance was about 0.8, suggesting that stem cavitation had a limited impact on whole shoot hydraulic conductance. We suggest that stomatal aperture may be regulated by the cavitation-induced reduction of hydraulic conductance of the soil-to-leaf water pathway which, in turn, strongly depends on the hydraulic architecture of the plant and, in particular, on leaf hydraulics.     

Bounds on the total population for species governed by reaction-diffusion equations in arbitrary two-dimensional regions

Analysis based on the integration of differential inequalities is employed to derive upper and lower bounds on the total populationN(t) = ∫ _R θ(x ₁,x ₂,t) dx ₁ dx ₂ of a biological species with an area-density distribution function θ=θ(x ₁,x ₂,t) (≥0) governed by a reaction-diffusion equation of the form ∂θ/∂t =D∇²θ +fθ −gθ ⁿ⁺¹ whereD (>0),n (>0),f andg are constant parameters, θ=0 at all points on the boundary ∂R of an (arbitrary) two-dimensional regionR, and the initial distribution (θ(_{x 1},x ₂, 0) is such thatN(0) is finite. Forg≥0 withR the entire two-dimensional Euclidean space, a lower bound onN(t) is obtained, showing in particular thatN(∞) is bounded below by a finite positive quantity forf≥0 andn>1. An upper bound onN(t) is obtained for arbitrary bounded or unbounded)R withn=1,f andg negative, and ∫ _R θ(x ₁,x ₂, 0)² dx ₁ dx ₂ sufficiently small in magnitude, implying that the population goes to extinction with increasing values of the time,N(∞)=0. Forg≥0 andR of finite area, the analysis yields upper bounds onN(t), predicting eventual extinction of the population if eitherf≤0 or if the area ofR is less than a certain grouping of the parameters in cases for whichf is positive. These results are directly applicable to biological species with distributions satisfying the Fisher equation in two spatial dimensions and to species governed by certain specialized population models.     

Tree size structure of stands and each species in primary warm-temperate rain forests of Southern Japan

The frequency distribution of trunk diameter was analyzed for forest stands of various developmental stages and for each tree species population in primary warm-temperate evergreen rain forests, dominated byDistylium racemosum, on Yakushima Island of southern Kyushu, Japan. Trunk diameter distribution in stands showed the inverse J shape. A regression model, lnf(x)=b+b ₁x+b₂ lnx, expressed the distribution well, wherex is diameter andf(x) trunk density atx. Three coefficients of the model, calculated from the data in 20 stands, clearly showed linear relations to each other and they had high correlations with the basal area per stands. The result suggests thatb andb ₁ increase andb ₂ decreases with stand age. Fourteen populations of abundant tree species also showed inverse J-shaped diameter distribution, which can be well expressed by the above model. In both open sites (gaps) and closed stands, these species were abundant and their characteristics in diameter distribution were persistent.     

Tracing plant source water dynamics during drought by continuous transpiration measurements: An in-situ stable isotope approach

The isotopic composition of xylem water (δ_X) is of considerable interest for plant source water studies. In-situ monitored isotopic composition of transpired water (δ_T) could provide a nondestructive proxy for δ_X-values. Using flow-through leaf chambers, we monitored 2-hourly δ_T-dynamics in two tropical plant species, one canopy-forming tree and one understory herbaceous species. In an enclosed rainforest (Biosphere 2), we observed δ_T-dynamics in response to an experimental severe drought, followed by a ²H deep-water pulse applied belowground before starting regular rain. We also sampled branches to obtain δ_X-values from cryogenic vacuum extraction (CVE). Daily flux-weighted δ¹⁸O_T-values were a good proxy for δ¹⁸O_X-values under well-watered and drought conditions that matched the rainforest's water source. Transpiration-derived δ¹⁸O_X-values were mostly lower than CVE-derived values. Transpiration-derived δ²H_X-values were relatively high compared to source water and consistently higher than CVE-derived values during drought. Tracing the ²H deep-water pulse in real-time showed distinct water uptake and transport responses: a fast and strong contribution of deep water to canopy tree transpiration contrasting with a slow and limited contribution to understory species transpiration. Thus, the in-situ transpiration method is a promising tool to capture rapid dynamics in plant water uptake and use by both woody and nonwoody species.     

Response of eucalypt species to drought

The response of three eucalypt species (Eucalyptus pulchella, Eucalyptus coccifera and Eucalyptus delegatensis) to a severe drought in the summer of 1982/83 was examined at Snug Plains, south-eastern Tasmania. Few large differences in leaf water potential (Ψ_l) or stomatal conductance (gs) were apparent even at the height of the drought when both Ψ_l and soil water potentials (Ψ_s) reached ca. — 4.5 MPa. However, E. pulchella maintained a higher relative water content (RWC) in its leaves than E. coccifera and E. delegatensis, and showed less severe crown damage. After the first light rains substantial interspecific differences in Ψ_l and gs occurred. Eucalyptus pulchella restored normal Ψ_l, gs and RWC more rapidly than the other two species and, even for severely droughted trees, crown growth commenced via epicormic buds near the ends of its branches while for E. delegatensis and E. coccifera crown regeneration was via epicormic buds arising from stems and larger branches. This resulted in a change in dominance in certain stands and showed that E. pulchella was more drought-resistant than E. coccifera, which was in turn more resistant than E. delegatensis. This conclusion was confirmed during competition experiments using potted seedlings. However, potted seedlings differed from mature field trees by maintaining moderate gs at high vapour pressure deficits and closing stomata at Ψ_l below ca. — 2.0 MPa. Substantial variation in the severity of drought symptoms was observed over short distances. This variation appeared to be determined by the moisture-holding capacity of the soil and the biomass of the stand. Although differences in the rooting patterns of seedlings were evident, field measurements of Ψ_l and Ψ_s suggested that all three species were exploiting the same water resource. In contrast to previous studies, the results suggest that large interspecific differences in tissue hydration and crown damage may be present, even though differences in Ψ_l, gs and characteristics of the root system may appear small.     

Leaf anatomy,water relations and crassulacean acid metabolism in the chlorenchyma and colourless internal water-storage tissue of Carpobrotus edulis and Senecio ?mandraliscae

Both Carpobrotus edulis and Senecio ?mandraliscae possess leaves with a peripheral chlorenchyma and colourless internal water-storage tissue. Water stress in C. edulis growing under semi-natural conditions resulted in the induction of weak Crassulacean acid metabolism (CAM) whereas well-watered plants of S. ?mandraliscae exhibited a similar degree of CAM. Titratable acidity in the separated water-storage tissue was substantially lower than in the chlorenchyma in both species but, nevertheless, increased during the night and decreased during the day either when sampled from the intact plant or from incubated tissue slices. Indeed, the increase in nocturnal titratable acidity produced by the water-storage tissue in situ accounted for approx. 30% of total acidification on a per-leaf basis. It appears that during the night the water-storage tissue in these species is able to fix CO₂ which is subsequently released during the day to enter the photosynthetic carbon-reduction cycle of the chlorenchyma. Diurnal rhythms of water potential (Ψ) and osmotic potential (Ψs) were measured in separated chlorenchyma and water-storage tissue by thermocouple psychrometry. Both parameters increased during the latter part of the daytime and initial nocturnal period and decreased during the rest of the night and into the post-dawn period. The chlorenchyma of water-stressed plants of C. edulis appeared to possess a marked negative turgor pressure (as determined from Ψ-Ψs) but this was caused by a severe underestimation in the measurement of the chlorenchyma Ψ. It is suggested that this artefact arose from release of colloidal polysaccharide mucilage, or possibly tannins, from broken tannin cells producing a lowering of water activity when measured using thermocouple psychrometry.     

Drought acclimation of two deciduous tree species of different layers in a temperate forest canopy

Water-use strategies of Populus tremula and Tilia cordata, and the role of abscisic acid in these strategies, were analysed. P. tremula dominated in the overstorey and T. cordata in the lower layer of the tree canopy of the temperate deciduous forest canopy. Shoot water potential (), bulk-leaf abscisic acid concentration ([ABA]_leaf), abscisic acid concentration in xylem sap ([ABA]_xyl), and rate of stomatal closure following the supply of exogenous ABA (v) decreased acropetally through the whole tree canopy, and foliar water content per area (w), concentration of the leaf osmoticum (c), maximum leaf-specific hydraulic conductance of shoot (L), stomatal conductance (g_s), and the threshold dose per leaf area of the exogenous ABA (d_a) required to reduce stomatal conductance increased acropetally through the tree canopy (from the base of the foliage of T. cordata to the top of the foliage of P. tremula) in non-stressed trees. The threshold dose per leaf dry mass of the exogenous ABA (d_w) required to reduce stomatal conductance, was similar through the tree canopy. After a drought period (3 weeks), the , w, L, g_s, d_a and d_w had decreased, and c and v had increased in both species. Yet, the effect of the drought period was more pronounced on L, g_s, d_a, d_w and v in T. cordata, and on , w and c in P. tremula. It was concluded that the water use of the species of the lower canopy layer—T. cordata, is more conservative than that of the species of the overstorey, P. tremula. [ABA]_leaf had not been significantly changed in these trees, and [ABA]_xyl had increased during the drought period only in P. tremula. The relations between [ABA]_leaf, [ABA]_xyl and the stomatal conductance, the osmotic adjustment and the shoot hydraulic conductance are also discussed.     

Reversible decreases in the bulk elastic modulus of mature leaves of deciduous Quercus species subjected to two drought treatments

Changes in leaf water relations under water stress were examined. In experiment 1, water stress was imposed by withholding irrigation to potted seedlings of deciduous oak, Quercus crispula and Q. serrata. Changes in the pressure–volume (P–V) curve in mature leaves were followed. The leaf water potential at turgor loss (Ψ_l,tlp) significantly decreased after 13 d of drought treatment. The bulk elastic modulus (?) significantly decreased, which contributed to the maintenance of cell turgor together with the decrease in osmotic potential. In experiment 2, water stress was imposed by notching a branch of a Q. serrata tree. After the notching, the daily minimum leaf water potential (Ψ_l) decreased, and a significant decrease in Ψ_l,tlp was observed 15 d after notching. The osmotic potential at water saturation (Ψ_π,sat) did not decrease significantly until 25 d after notching whereas, ? had already decreased significantly within 15 d after notching and increased promptly after substantial precipitation. It was confirmed that ? of mature leaves decreased reversibly in water stress. This response of ? was more rapid than that of the osmotic potential and, thus, effectively maintained cell turgor when water stress was suddenly imposed on the leaves.     

Osmoregulation and membrane-mediated responses to altered water potential in plant cells

Plant cells respond to short-term stress dehydration by modification of internal Ψ_π such that an inward gradient of Ψ_ω is maintained. In response to lowered Ψ_ω, increases in internal Ψ_π are created by alteration of cell inorganic ions and small organic solute content. Passive movement of water follows, changing cell hydration and forcing the plasma membrane against the elastic cell wall. The stretched cell wall presses against the cell contents, creating a hydrostatic pressure, Ψ_π, which tends to force water out of the cell. The resulting hydrostatic pressure eventually comes into equilibrium with forces bringing water into the cell, largely Ψ_π, and the net flow of water ceases.The mechanism for sensing cell Ψ_ω changes is unknown but the initial event must be physical, not biochemical. The method of translation of such physical events into biochemical actions is also unknown but the Zimmermann model provides a means of signal transduction and amplification, through the alteration of membrane parameters, which could account for the observed changes. As for animal cells, cell levels of Ca²⁺ are important for their regulation of membrane P_j in these responses but unlike osmoregulation in higher animals, the involvement of plant hormones in these responses have not been clearly established. However, the important role of plant cell limiting membranes in plant cell osmoregulation responses seems obvious.