Thermal and Water Relations of Roots of Desert Succulents

Two succulent perennials from the Sonoran Desert, Agave desertiEngelm. and Ferocactus acanthodes (Lem.) Britton and Rose, loselittle water through their roots during drought, yet respondrapidly to light rainfall. Their roots tend to be shallow, althoughabsent from the upper 20 mm or so of the soil. During 12–15d after a rainfall, new root production increased total rootlength by 47 per cent to 740 m for A. deserti and by 27 percent to 230 m for F. acanthodes; root dry weight then averagedonly 15 per cent of shoot dry weight. The annual carbon allocatedto dry weight of new roots required 11 per cent of shoot carbondioxide uptake for A. deserti and 19 per cent for F. acanthodes.Elongation of new roots was greatest near a soil temperatureof 30°C, and lethal temperature extremes (causing a 50 percent decrease in root parenchyma cells taking up stain) were56°C and -7°C. Soil temperatures annually exceeded themeasured tolerance to high temperature at depths less than 20mm, probably explaining the lack of roots in this zone. Attached roots immersed in solutions with osmotic potentialsabove -2·6 MPa could produce new lateral roots, with50 per cent of maximum elongation occurring near -1·4MPa for both species. Non-droughted roots lost water when immersedin solutions with osmotic potentials below -0·8 MPa,and root hydraulic conductance decreased markedly below about-1·2 MPa. Pressure-volume curves indicated that, fora given change in water potential, non-droughted roots lostthree to five times more water than droughted roots, non-droughtedleaves, or non-droughted stems. Hence, such roots, which couldbe produced in response to a rainfall, will lose the most tissuewater with the onset of drought, the resulting shrinkage beingaccompanied by reduced root hydraulic conductance, less contactwith drying soil, and less water loss from the plant to thesoil. Agave deserti, Ferocactus acanthodes, roots, soil, temperature, water stress, drought, Crassulacean acid metabolism, succulents     

Matric Water Potential of Leaf Tissue--Measurement and Significance

A thermocouple psychrometer method, previously described foruse in determining tissue and sap water potentials, has beenadapted for determining matric potentials. Matric values ranging from approximately zero to –10 x10⁵ Pa were observed with wilting wheat leaves. Mean valuesof total tissue water potential and of its solute (osmotic),matric, and cell wall pressure components were –16.6,–18.3, –30 and +4.7 Pa x 10⁵, respectively. Matric potentials are often ignored in investigations of plantwater relations. This practice is shown to result in underestimationof wall pressures (in this case by a mean of 64 per cent) andsometimes in spuriously negative values.     

Allometric Root/Shoot Relationships and Predicted Water Uptake for Desert Succulents

Root morphology, shoot morphology, and water uptake for Agavedeserti and Ferocactus acanthodes of various sizes were studiedusing allometric relationships (y = ax^b) and a previously developedwater uptake model. Shoot surface area increased with shootvolume with an exponent b of 0.75 for both species. Root lengthand the ground area explored by the roots increased with shootsurface area with b's of 0.72 for A. deserti and 0.92 for F.acanthodes. Various sized individuals had about the same ratioof root length to explored ground area, with higher values occurringfor A. deserti. Predicted water uptake averaged over the exploredground area was approximately constant over a 10⁴-fold rangein shoot surface area, suggesting that shoot size confers nointraspecific competitive advantage for water uptake. For theroot lengths per explored ground area observed in the field,water uptake was predicted to be 85 per cent of maximal; wateruptake could be increased by the production of more rain roots.When differences in shoot volume were accounted for by allometry,small plants had relatively less shoot surface area and relativelymore root length per shoot volume than did large plants, whichmay be important for the water relations of seedling establishment. Agave deserti, Ferocactus acanthodes, allometry, desert succulents, root distribution, root length, seedling growth, seedling establishment, shoot surface area, shoot volume, water uptake     

Water Uptake and Germination of Leguminous Seeds in Soils of Changing Matric and Osmotic Water Potential

Water uptake and germination rate of chickpea and pea seedswere compared under changing water potentials in sand and soilaggregate columns and osmotic solutions. The final water uptake and germination were the same in allcases for a given water potential, but the rates were lowerfor seeds planted in sand columns, probably due to mechanicalconstraints imposed on the swelling seed by the dense sand,since the capillary conductivity, and the diffusivity to waterof the sand were very high. The area of the seed in contact with soil is not of importanceif soil aggregates are small as compared to the seeds but increasesin importance when the seeds and the soil aggregates are ofthe same size and at low water potentials.     

Diurnal Pattern of Transpiration,Water Uptake and Water Budget of Succulents with Different CO2 Fixation Pathways

The water fluxes and the CO₂ exchange of three leaf succulents, Othonna opima, Cotyledon orbiculata and Senecio medley-woodii, with different leaf anatomy, growth form and CO₂ fixation pathways (C₃, CAM) were monitored with a gas exchange cuvette which was combined with a potometric system to quantify water uptake. Measurements, which are primarily valid for plants with a sufficient water supply, were made during 6 to 10 consecutive days under constant experimental conditions. Water uptake for 24 h exceeded water loss by transpiration only for a S, medley-woodii plant with 10 expanding but only 7 mature leaves. In this case the gained water evidently is put into leaf expansion. All other plants showed balanced transpiration and water uptake rates. O. opima and C. orbiculata have a similar life form, similar water storage volumes and the same natural habitat but their diurnal water uptake patterns differ significantly. In the C₃ plant O. opima water uptake increased when the transpiration increased or transpiration rates were higher than uptake rates and vice versa. On the contrary the CAM plant C. orbiculata transpired during the dark period at constant or decreasing rates but showed steadily increasing uptake rates. Senecio medley-woodii- and C. orbiculata are CAM plants with similar diurnal water uptake patterns with its maximum in uptake during or towards the end of the CO₂ dark fixation period. Water uptake of C. orbiculata was at its minimum at the end of the light period despite transpiration being maximal. The results were discussed considering the different CO₂ fixation pathways. In the investigated CAM succulents, C. orbiculata and S. medley-woodii, the CAM influenced water uptake throughout the whole day and not only during the CO₂ dark fixation period.     

Accumulation of Trehalose and Sucrose in Cyanobacteria Exposed to Matric Water Stress

The drought-resistant cyanobacteria Phormidium autumnale, strain LPP₄, and a Chroococcidiopsis sp. accumulated trehalose, sucrose, and both trehalose and sucrose, respectively, in response to matric water stress. Accumulated sugar concentrations reached values of up to 6.2 μg of trehalose per μg of chlorophyll in P. autumnale, 6.9 μg of sucrose per μg of chlorophyll in LPP₄, and 4.1 μg of sucrose and 3.2 μg of trehalose per μg of chlorophyll in the Chroococcidiopsis sp. The same sugars were accumulated by these cyanobacteria in similar concentrations under osmotic water stress. Cyanobacteria that did not show drought resistance (Plectonema boryanum and Synechococcus strain PCC 7942) did not accumulate significant amounts of sugars when matric water stress was applied.     

A Two-dimensional Model for Water Uptake by Desert Succulents: Implications of Root Distribution

Water uptake by Agave deserti and Ferocatus acanthodes was predictedusing a two-dimensional simulation model in which the soil arounda plant was divided into a series of layers and concentric cylindricalshells. Root lengths in 0.05 m thick soil layers were determinedfor both species in the field, where mean root depths were only0.11 m for A. deserti and 0.10 m for F. acanthodes. For a yearwith average precipitation (159 mm), 42 per cent of the annualprecipitation could be taken up by A. deserti and 25 per centby F. acanthodes. Predicted water uptake by both species wasgreater from the upper soil layers (above 0.15 m) for averageand dry years, but was greater from the deeper layers for awet year. The actual root distribution for both species ledto more water uptake than when all of the roots were in a singlelayer. The large number of days per year when the soil temperaturesexceeded 57 °C (the temperature for 50 per cent inhibitionof uptake of a vital stain by root cells) may exclude rootsfrom the 0.00–0.05 m soil layer, even though water uptakewhen all roots were located there was predicted to be maximal.Therefore, the observed root distribution of A. deserti andF. acanthodes may be limited near the soil surface by high temperaturesand at maximum depths by water availability for all but wetyears. Agave deserti, Ferocactus acanthodes, desert succulents, root system, root distribution, soil temperature, water uptake     

Matric potentials of leaves

A pressure chamber was used to measure matric potentials of frozen and thawed leaves. Significant matric potentials were demonstrated in sunflower (Helianthus annuus L.), yew (Taxus cuspidata Sieb. and Zucc.), and rhododendron (Rhododendron roseum Rehd.). Matric potentials were particularly negative in rhododendron and were correlated with the amount of cell wall present and with the volume of water outside the leaf protoplasts at comparable matric potentials. It was concluded that matric forces in leaves are associated mainly with cell walls, at least within the physiological range of water contents. Calculations indicated that the water potential of the solution in the cell wall could be estimated for living tissue from the sum of matric and osmotic potentials acting on water outside the protoplasts.     

The Meaning of Matric Potential

The commonly used equation, = P - + , which describes thepartitioning of plant water potential, , into components ofhydrostatic pressure, P, osmotic pressure, , and matric potential,, is misleading. The term , which is supposed to show the influenceof a solid phase on , is zero if a consistent definition ofpressure is used in the standard thermodynamic derivation. However,it can be usefully defined by = + _D, where _D is the osmoticpressure of the equilibrium dialysate of the system. The practicaland theoretical significance of this definition is discussed.     

Rectifier-like Activities of Roots of Two Desert Succulents

Axial and radial water flows for roots in response to appliedhydrostatic pressure drops, water loss from roots after variousperiods of drying, and development of new roots after rewettingdroughted plants were examined for two sympatric desert succulents.Agave deserti Engelm. and Ferocactus acanthodes (Lemaire) Brittonand Rose. For a 40 kPa hydrostatic pressure drop applied to20 mm long root pieces, radial water flows from the epidermisto the root xylem were 2- to 5-fold greater at the tip thanat midlength and were much less than axial flows along the xylem.Upon drying detached roots in air at 20 °C and a water vapoursaturation deficit of 1.2 kPa (50% relative humidity), radialwater flow decreased more than 10-fold in 3–6 h, and couldrecover to the original level 6 h after rewetting. The rateof water loss from attached roots of plants dried in air at20 °C and a 1.2 kPa saturation deficit decreased about 200-foldin 72 h, which would greatly limit water loss from the plantto a drying soil. At 96 h after rewetting roots of A. desertithat had been exposed to air at 20 °C and a 1.2 kPa saturationdeficit for 120 h, rehydration of existing roots and developmentof new roots contributed about equally to water uptake by thewhole plant. In summary, roots of these desert succulents canreadily take up water from a wet soil but do not lose much waterto a dry soil, thus effectively acting like rectifiers withrespect to plant-soil water movement. Key words: Agave, Cactus, Drought, Root, Water flow, Xylem     

Bound Water and Cryptobiosis:Thermodynamic Properties of Water at Biopolymer Surfaces

The molecular mechanisms underlying the adaptations to water loss developed in several tardigrade species remain poorly understood. It seems, however, that the binding of the disaccharide trehalose to membranes and other cellular components at low water contents is important for the tolerance to extreme drought. Trehalose is thus thought to replace interfacial- or “bound” water and enhance the conformational stability of labile macromolecules. To gain further insight into this we investigate here thermodynamic properties of water bound to the protein lysozyme at low water content (<100 water molecules pr. protein). It appears that this surface water has a higher enthalpy and higher entropy than the bulk liquid. These observations call for re-evaluation of the term “bound water” since “bound” carries the connotation of a low-energy, ordered (i.e. low-entropy) state.

To rationalize these observations it is suggested that — in addition to the self-evident energetic contribution from biopolymer-water contacts — the properties of interfacial water are dominated by two effects. These are i) the ability of water to facilitate fast movements of individual parts of biopolymers and ii) the high molecular cohesion in the aqueous bulk. Thus, the hydration of a surface leads to enhanced flexibility in the biopolymer and breakage in the network of hydrogen bonding in the liquid bulk, and these effects collectively increase the enthalpy and entropy of the system. As a result, the thermodynamic parameters of hydration of lysozyme carry the thermodynamic hallmarks of an order → disorder process, even for the first hundred (i.e. most strongly associated) water molecules. We discuss these data for protein hydration together with some recent, very similar observations for the hydration of lipid bilayer membranes.     

Changes in Structure and Hydraulic Conductivity for Root Junctions of Desert Succulents as Soil Water Status Varies

Variations in hydraulic conductivity (L_P) and the underlying anatomical and morphological changes were investigated for main root-lateral root junctions of Agave deserti and Ferocactus acanthodes under wet, dry, and rewetted soil conditions. During 21 d of drying, L_P and radial conductivity (L_R) increased threefold to fivefold at junctions of both species. The increase in L_R was accompanied by the formation of an apoplastic pathway for radial water movement from the surface of the junction to the stele for A. deserti and by the rupture of periderm by emerging primordia of secondary lateral roots for F. acanthodes. During 7 d of rewetting, L_R decreased for junctions of A. deserti, as apoplastic water movement was not apparent, but L_R was unchanged for F. acanthodes. Axial conductance (K_h) decreased during drying for both species, largely because of embolism related to the degradation of unlignified cell wall areas in tracheary elements at the root junction. The resulting apertures in the cell walls of such elements would admit air bubbles at pressure differences of only 0.12-0.19 MPa. Rewetting restored K_h for both species, but not completely, due to blockage of xylem elements by tyloses. About 40% of the primary lateral roots of the monocotyledon A. deserti abscised during 21 d of drying. For the dicotyledon F. acanthodes, which can form new conduits in its secondary xylem, only 10% of the primary lateral roots abscised during 21 d of drying, consistent with the much greater frequency of lateral roots that persist during drought in the field compared with the case for the sympatric A. deserti.     

Bound Water in Durum Wheat under Drought Stress

To study drought stress effects on bound water, adsorption isotherms and pressure-volume curves were constructed for two durum wheat (Triticum durum Desf.) cultivars: Capeiti 8 (drought tolerant) and Creso (drought sensitive). Plants were grown under well-watered and water-stressed conditions in a controlled environment. Differential enthalpy (ΔH) was calculated through van't Hoff analysis of adsorption isotherms at 5 and 20°C, which allowed us to determine the strength of water binding. ΔH reached the most negative values at approximately 0.06 gram H₂O/gram dry weight and then increased rapidly for well-watered plants (until 0.10 gram H₂O/gram dry weight) or more slowly for drought-stressed plants (until 0.15-0.20 gram H₂O/gram dry weight). Bound water values from pressure-volume curves were greater for water-stressed (0.17 gram H₂O/gram dry weight) than for well-watered plants (0.09 gram H₂O/gram dry weight). They may be estimates of leaf moisture content where ΔH reaches the less negative values and hence some free water appears. With respect to the well-watered plants, tightly bound water tended to be less bound during drought, and more free water was observed in cv Creso compared to cv Capeiti 8 at moisture contents >0.10 gram H₂O/gram dry weight.     

Study of Bound Water of Poly-Adenine Using High Frequency Dielectric Measurements

The high frequency dielectric constant of poly-adenine (poly-A) was measured between 1 MHz and 1 GHz. The purpose of these experiments was to investigate the state of water molecules that are bound to the charged groups of the poly-A molecule. Analysis of the data using the Maxwell's mixture equation revealed the dielectric constant of bound water higher than we expected. Using Onsager's internal field in Debye's equation, we calculated the dielectric constant of water in the vicinity of a charged ion. The result of this computation demonstrates that the dielectric constant of bound water is much smaller than the normal value only in the immediate proximity of charged ions (within 2 Å). The dielectric constant increases rapidly to the normal value as the distance increases from 2 to 4 Å. This observation indicates that charged sites of polyions have only short range interactions with the surrounding water molecules. However, this conclusion pertains only to rotary diffusion of bound water since dielectric measurement is unable to detect translational diffusion.     

An Investigation of the Bound Water in Tendon by Dielectric Measurement

The dielectric constant of natural tendon in the frequency range of 200 Hz to 100 kHz has been determined as a function of temperature (300-450°K) for water concentrations ranging from about 6 to 16% by weight. The results compare well with the approach taken by Haggis and his coworkers. Based on the assumption that at low concentration of water the probability of water-water bonding is small and hence may be disregarded, a structure for water in the collagen matrix is proposed in which the water is either bonded in one of four possible states to the polar groups of the polypeptide chains, or is unbound. In determining the distribution of the water among these states an approach similar to that of Haggis and his co-workers, in conjunction with the order-disorder theory of Bragg and Williams, is used. The number of water molecules per unit volume is then determined experimentally by relating it to weight loss as a function of temperature, as determined by thermogravimetric analysis. The dispersion which is normally found in dipolar substances has not been found for tendon. A maximum in the value of the dielectric constant is observed to occur between 25 and 80°C, the temperature depending upon the heating rate.     

Matric potential of several plant tissues and biocolloids

The pressure membrane apparatus was used to study the matric potential (imbibition pressure or moisture tension) of plant tissues and of several organic colloidal preparations.     

Influence of Soil Water Matric Potential and Hydraulic Conductivity on the Germination of Rape (Brassica napus L.)

The influence of matric potential and hydraulic conductivityon the water absorption and germination of rape (Brassica napusL.) was investigated by comparative studies in three seed-soilwater systems. In all systems the rate of germination and totalgermination were substantially retarded with decreasing matricpotential. Hydraulic conductivity of the soil system was demonstratedto be a limiting factor in the germination process. The effectsof matric potential and hydraulic conductivity on water absorptionfollowed closely those demonstrated for germination. The influenceof matric potential was shown to be comparable to that of osmoticpotential provided consideration was given to the hydraulicproperties of the soil-seed system.