Water flow and water storage in Agave deserti: osmotic implications of crassulacean acid metabolism

Abstract Water flow and water storage were investigated for Agave deserti, a desert succulent showing crassulacean acid metabolism (CAM). The anatomy and water relations of the peripheral chlorenchyma, where CAM occurs, and the central water-storage parenchyma were investigated for its massive leaves so that these tissues could be incorporated as discrete elements into an electrical-circuit analogue of the whole plant. The daily cycling of osmotic pressure was represented by voltage sources in series with the storage capacitors. With soil water potential and leaf transpiration rate as input variables, axial water flow through the vascular bundles and radial flows into and out of storage during the day/night cycle were determined. The predominantly nocturnal transpiration was coincident with increases in cell osmotic pressure and in titratable acid of the leaf chlorenchyma. In the outer layers of the chlorenchyma, water potential was most negative at the beginning of the night when transpiration was maximum, while the water-storage parenchyma reached its minimal water potential 9 h later. The roots plus stem contributed 7% and the leaves contributed 50% to the total water flow during maximal transpiration; peak water flow from the soil to the roots occurred at dawn and was only 58% of the maximal transpiration rate. Over each 24-h period, 39% of the water lost from the plant was derived from storage, with flow into storage occurring mainly during the daytime. Simulations showed that the acid accumulation rhythm of CAM had little impact on water uptake from the soil under the conditions employed. In the outer chlorenchyma, water potential and water flows were more sensitive to the day/night changes in transpiration than in osmotic pressure. Nevertheless, cell osmotic pressure had a large influence on turgor pressure in this tissue and determined the extent to which storage was recharged during the latter part of the night.     

Responses of a CAM Plant to Drought and Rainfall: Capacitance and Osmotic Pressure Influences on Water Movement

Daily and seasonal patterns in water flow and water potentialwere investigated for the Crassulacean acid metabolism succulentAgave deserti during an extended summer drought and for a periodfollowing rainfall. Field measurements of transpiration andof osmotic pressure changes over selected 24 h periods wereused as input variables for a computer model of water flow thatwas based on an electrical circuit analog of the whole plant.Parameters such as root resistance and tissue capacitance werealso varied to reflect the effects of changing plant or soilwater status. The model predicted internal water flow and waterpotential during the drought cycle and was used to assess therole of tissue osmotic properties in water uptake from the soiland in internal water redistribution. For plants under wet soil conditions, 55% of the night-timetranspiration was derived from water storage, this storage beingrecharged during the day. As drought progressed, transpirationand the nocturnal increase in osmotic pressure declined, althoughthe osmotic pressure itself increased. The difference in osmoticpressure between the water storage tissue and the chlorenchymacaused a net flow of water into the chlorenchyma after 3 weeksof drought, thereby increasing chlorenchyma turgor pressure.Simulations also indicated that a large increase in root resistancemust occur to prevent substantial water loss from the plantto the dry soil. After rainfall, recharge of plant water storagewas complete within one week, although full recovery in theamplitude of daily osmotic pressure variations took longer. Key words: Agave deserti, transpiration, water potential, water storage     

Diel water movement between parenchyma and chlorenchyma of two desert CAM plants under dry and wet conditions

Abstract. Electric-circuit analogue models of the water relations of crassulacean acid metabolism (CAM) succulents such as Agave deserti and Ferocactus acanthodes have predicted diel movement of water between the water-storage parenchyma and the photo-synthetic chlorenchyma. Injection of tritiated water into either tissue in the laboratory confirmed substantial and bidirectional water movements, especially under conditions of wet soil. For A. deserti , water movement from the water-storage parenchyma to the chlorenchyma increased at night as the chlorenchyma osmotic pressure increased. Although nocturnal osmotic pressure increases and transpiration for both species were minimal in the field under dry conditions, diel changes in the deuterium: hydrogen ratio (expressed as ΔD) were similar for the water-storage parenchyma and the chlorenchyma. Such indication of [substantial mixing of water between the tissues over a 24-h cycle was more evident under wet conditions in the field. For A. deserti , ΔD then increased by 32%o from the afternoon to midnight and was essentially identical in the water-storage parenchyma and the chlorenchyma. For F. acanthodes , the diel changes in ΔD were one-third those of A. deserti , and ΔD was always slightly higher for the chlorenchyma than for the water-storage parenchyma, apparently reflecting the lower surface-to-volume ratio of A. deserti. In summary, data obtained using radioactive and stable isotopes strongly supported model predictions concerning diel cycles of internal water distribution for these CAM species.     

Day-Night Variations in Malate Concentration, Osmotic Pressure, and Hydrostatic Pressure in Cereus validus

Malate concentration and stem osmotic pressure concomitantly increase during nighttime CO₂ fixation and then decrease during the daytime in the obligate Crassulacean acid metabolism (CAM) plant, Cereus validus (Cactaceae). Changes in malate osmotic pressure calculated using the Van't Hoff relation match the changes in stem osmotic pressure, indicating that changes in malate level affected the water relations of the succulent stems. In contrast to stem osmotic pressure, stem water potential showed little day-night changes, suggesting that changes in cellular hydrostatic pressure occurred. This was corroborated by direct measurements of hydrostatic pressure using the Jülich pressure probe where a small oil-filled micropipette is inserted directly into chlorenchyma cells, which indicated a 4-fold increase in hydrostatic pressure from dusk to dawn. A transient increase of hydrostatic pressure at the beginning of the dark period was correlated with a short period of stomatal closing between afternoon and nighttime CO₂ fixation, suggesting that the rather complex hydrostatic pressure patterns could be explained by an interplay between the effects of transpiration and malate levels. A second CAM plant, Agave deserti, showed similar day-night changes in hydrostatic pressure in its succulent leaves. It is concluded that, in addition to the inverted stomatal rhythm, the oscillations of malate markedly affect osmotic pressures and hence water relations of CAM plants.     

Diel Patterns of Water Potential Components for the Crassulacean Acid Metabolism Plant Opuntia ficus-indica when Well-Watered or Droughted

Under well-watered conditions, chlorenchyma acidity in cladodes of Opuntia ficus-indica increased substantially at night, fully accounting for the 0.26-megapascal nocturnal increase in osmotic pressure in the outer 2 millimeters. Osmotic pressure in the inner part of the chlorenchyma and in the water-storage parenchyma did not change significantly over 24-hour periods. Three months of drought decreased nocturnal acid accumulation by 73% and essentially abolished transpiration; also, 27% of the chlorenchyma water and 61% of the parenchyma water was lost during such drought, but the average tissue osmotic pressure was little affected. Turgor pressure was maintained in the chlorenchyma after 3 months of drought, although it decreased sevenfold in the water-storage parenchyma compared with the well-watered condition. Moreover, the nocturnal increases in turgor pressure of about 0.08 megapascal in the outer part of the chlorenchyma was also unchanged by such drought. The water potential magnitudes favored water movement from the parenchyma to the chlorenchyma at the end of the night and in the reverse direction during the late afternoon. Experiments with tritiated water support this pattern of water movement, which is also in agreement with predictions based on electric-circuit analog models for Crassulacean acid metabolism plants.     

Water relations and photosynthesis of a barrel cactus,Ferocactus acanthodes,in the Colorado desert

Summary The structural characteristics, water relations, and photosynthesis of Ferocactus acanthodes (Lemaire) Britton and Rose, a barrel cactus exhibiting Crassulacean acid metabolism (CAM), were examined in its native habitat in the western Colorado desert. Water storage in its succulent stem permitted nighttime stomatal opening ot continue for about 40 days after the soil water potential became less than that of the stem, a period whe the plant would be unable to extract water from the soil. After 7 months of drought and consequent unreplenished water loss from a plant, diurnal stomatal activity was not observed and the stem osmotic pressure was 6.4 bars, more than double the value measured during wet periods with nighttime stomatal opening. F. acanthodes had a shallow root system (mean depth of 8 cm) which responded within 24 h to rainfall.When the nocturnal stem surface temperature was raised from 8.0° C to 35.0° C, the stomatal resistance increased 4-fold, indicating that cool nighttime temperatures are advantageous for gas exchange by F. acanthodes. Moreover, the optimal temperature for CO₂ uptake in the dark was only 12.6° C. CO₂ uptake at night became maximal for 3.0 mEinsteins cm^-2 of photosynthetically active radiation incident during the preceding day, and the minimum number of incident quanta absorbed per CO₂ fixed was 68. The transpiration ratio (mass of water transpired/mass of CO₂ fixed) had the relatively low value of 70 for an entire year, consistent with values obtained for other CAM plants. The total amount of water annually diverted to the floral structures was about 6% of the stem wet weight. The annual growth increment estimated from the net CO₂ assimilation corresponded to about 10% of the stem mass for barrel cacti 34 cm tall, in agreement with measured dimension changes, and indicated that such plants were about 26 years old.     

Use of the root contact concept,an empirical leaf conductance model and pressure-volume curves in simulating crop water relations

A simulation model “DanStress” was developed for studying the integrated effects of soil, crop and climatic conditions on water relations and water use of field grown cereal crops. The root zone was separated into 0.1 m deep layers of topsoil and subsoil. For each layer the water potential at the root surface was calculated by a single root model, and the uptake of water across the root was calculated by a root contact model. Crop transpiration was calculated by Monteith's combination equation for vapour flow. Crop conductance to water vapour transfer for use in Monteith's combination equation was scaled up from an empirical stomatal conductance model used on sunlit and shaded crop surfaces of different crop layers. In the model, transpirational water loss originates from root water uptake and changes in crop water storage. Crop water capacitance, used for describing the water storage, was derived from the slope of pressure-volume (PV) curves of the leaves. PV curves were also used for deriving crop water potential, osmotic potential, and turgor pressure. The model could simulate detailed diurnal soil-crop water relations during a 23-day-drying cycle with time steps of one hour. During the grain filling period in spring barley (Hordeum distichum L.), grown in a sandy soil in the field, measured and predicted values of leaf water and osmotic potential, RWC, and leaf stomatal conductance were compared. Good agreement was obtained between measured and predicted values at different soil water deficits and climatic conditions. In the field, measured and predicted volumetric soil water contents (θ) of topsoil and subsoil layers were also compared during a drying cycle. Predicted and measured θ-values as a function of soil water deficits were similar suggesting that the root contact model approach was valid. From the investigation we concluded: (I) a model, which takes the degree of contact between root surface and soil water into account, can be used in sandy soil for calculation of root water uptake, so that the root conductance during soil water depletion only varies by the degree of contact; (II) crop conductance, used for calculation of crop transpiration, can be scaled up from an empirical single leaf stomatal conductance model controlled by the level of leaf water potential and micrometeorological conditions; (III) PV curves are usable for describing crop water status including crop water storage.     

Matric Bound Water of Water Tissue from Succulents

Matric bound water was measured as water retained by frozen and thawed tissue after desorption on a pressure membrane filter under 20 bars nitrogen gas pressure. Central water-storage tissue and peripheral chlorenchyma from leaves or stems of 15 taxonomically diverse non-halophytic succulent species were investigated. Matric bound water as a per cent of the dry weight averaged higher in water storage than in chlorenchyma tissue but lower than values reported for many mesophytic leaves. Matric bound water as a proportion of the total water held, however, was lower in water tissues. Osmotic potentials were generally high (solute contents low). It is concluded that matric or osmotic forces cannot account, in any unique way, for the high water content of water tissues. This appears to depend, instead, on the enormous ability of the thin-walled cells to take up available water and expand.     

Parenchyma-chlorenchyma water movement during drought for the hemiepiphytic cactus Hylocereus undatus

BACKGROUND AND AIMS: Hylocereus undatus, a hemiepiphytic cactus cultivated in 20 countries for its fruit, has fleshy stems whose water storage is crucial for surviving drought. Inter-tissue water transfer during drought was therefore analysed based on cell volumes and water potential components. METHODS: In addition to determining cell dimensions, osmotic pressures and water potentials, a novel but simple procedure leading to an external water potential of zero was devised by which cells in thin sections were perfused with distilled water. The resulting volume changes indicated that the parenchyma-chlorenchyma water movement was related to more flexible cell walls in the water-storage parenchyma with its lower internal turgor pressure (P) than in the chlorenchyma. KEY RESULTS: Under wet conditions, P was 0.45 MPa in the chlorenchyma but only 0.10 MPa in the water-storage parenchyma. During 6 weeks of drought, the stems lost one-third of their water content, becoming flaccid. About 95 % of the water lost came from cells in the water-storage parenchyma, which decreased by 44 % in length and volume, whereas cells in the adjacent chlorenchyma decreased by only 6 %; the osmotic pressure concomitantly increased by only 10 % in the chlorenchyma but by 75 % in the water-storage parenchyma. CONCLUSIONS: The concentrating effect that occurred as cellular volume decreased indicated no change in cellular solute amounts during 6 weeks of drought. The ability to shift water from the parenchyma to the chlorenchyma allowed the latter tissue to maintain a positive net CO2 uptake rate during such a drought.     

Time constant for water transport in loblolly pine trees estimated from time series of evaporative demand and stem sapflow

 The use of stem sap flow data to estimate diurnal whole-tree transpiration and canopy stomatal conductance depends critically upon knowledge of the time lag between transpiration and water flux through the stem. In this study, the time constant for water movement in stems of 12-year-old Pinus taeda L. individuals was estimated from analysis of time series data of stem water flux and canopy transpiration computed from mean daytime canopy conductance, and diurnal vapor pressure deficit and solar radiation measurements. Water uptake through stems was measured using a constant-heat sapflow probe. Canopy transpiration was correlated to stem uptake using a resistance-capacitance equation that incorporates a time constant parameter. A least-squares auto-regression determined the parameters of the resistance-capacitance equation. The time constants for ten loblolly pine trees averaged 48.0 (SE = 2.0) min and the time lag for the diurnal frequency averaged 47.0 (SE = 2.0) min. A direct-cross correlation analysis between canopy transpiration and sap flow time series showed maximum correlation at an approximately 30 min lag. Residuals (model-predicted minus actual stem flow data) increased with increasing soil moisture depletion. While the time constants did not vary significantly within the range of tree sizes studied, hydraulic resistance and capacitance terms were individually dependent on stem cross-sectional area: capacitance increased and resistance decreased with stem volume. This result may indicate an inverse adjustment of resistance and capacitance to maintain a similar time constant over the range of tree sizes studied.     

Effects of drought on water relations and nonstructural carbohydrates in cladodes of Opuntia ficus-indica

Nocturnal acid accumulation, water content, osmotic pressure (π), and nonstructural carbohydrates were determined in the chlorenchyma and the water-storage parenchyma of Opuntia ficus-indica (L.) Miller for well-watered plants and those subjected to drought for 15 weeks. During the 15-week drought, total cladode water content decreased by 57%, the water-storage parenchyma losing a greater fraction of water than the chlorenchyma, which most likely helped maintain nocturnal acid accumulation in the latter tissue. Despite the preferential water loss from the water-storage parenchyma, it had a lower π than the chlorenchyma over the 15 weeks of drought, suggesting a substantial decrease in osmotically active solutes in the water-storage parenchyma. Also, the measured π increases of both tissues were much less than those predicted based on the loss of water during drought and the initial content of osmotically active solutes under well-watered conditions. A decrease in the amount of soluble sugars (glucose. fructose and sucrose) occurred in plants subjected to drought. accounting for 46% and 81% of the difference between the measured and the predicted increases in π of the chlorenchyma and the water-storage parenchyma. respectively. The decrease in soluble sugars was associated with an equivalenl increase in polysaccharides, presumably starch, in the water-storage parenchyma. but not in the chlorenchyma.     

Biophysical properties and functional significance of stem water storage tissues in Neotropical savanna trees

Biophysical characteristics of sapwood and outer parenchyma water storage compartments were studied in stems of eight dominant Brazilian Cerrado tree species to assess the impact of differences in tissue capacitance on whole-plant water relations. The rate of decline in tissue water potential with relative water content (RWC) was greater in the outer parenchyma than in the sapwood for most of the species, resulting in tissue-and species-specific differences in capacitance. Sapwood capacitance on a tissue volume basis ranged from 40 to 160 kg m-3 MPa-1, whereas outer parenchyma capacitance ranged from 25 to only 60 kg m-3 MPa-1. In addition, osmotic potentials at full turgor and at the turgor loss point were more negative for the outer parenchyma compared with the sapwood, and the maximum bulk elastic modulus was higher for the outer parenchyma than for the sapwood. Sapwood capacitance decreased linearly with increasing sapwood density across species, but there was no significant correlation between outer parenchyma capacitance and tissue density. Midday leaf water potential, the total hydraulic conductance of the soil/leaf pathway and stomatal conductance to water vapour (gs) all increased with stem volumetric capacitance, or with the relative contribution of stored water to total daily transpiration. However, the difference between the pre-dawn water potential of non-transpiring leaves and the weighted average soil water potential, a measure of the water potential disequilibrium between the plant and soil, increased asymptotically with total stem capacitance across species, implying that overnight recharge of water storage compartments was incomplete in species with greater capacitance. Overall, stem capacitance contributes to homeostasis in the diurnal and seasonal water balance of Cerrado trees.     

Water relations of stem succulent trees in north-central Baja California

Summary Water relations of several stem succulent trees were measured in north-central Baja California in comparisons to other growth forms in the same habitat. Our research concentrated on three stem succulent species (Idria collumnaris, Pachycormus discolor and Bursera microphylla) each with a different succulent stem morphology. The stem succulent trees had 1 to 4 kg H₂O/m³ of trunk while the other trees and shrubs in the same habitat had 0.6 to 0.8 kg H₂O/m³. The diurnal and seasonal variation in leaf water potential was small for the stem succulent species in comparison to deciduous and evergreen species as a consequence of the stem-water, buffering capacity. In addition, the leaf conductance of the stem succulent species was low (60 mmol m^–2 s^–1) and yet, the leaf conductance decreased through the day similar to adjacent evergreen and deciduous species. The leaves of the stem succulent trees lost turgor at low saturated water deficits (0.06 to 0.14), had comparatively high osmotic potentials, and high values of elastic modulus in comparison to adjacent evergreen and deciduous species. The stem acts as an important buffering mechanism allowing for the maintenance of leaf turgor in these stem succulent trees. The low transpiration rates of the stem succulent trees may be a mechanism to minimize leaf saturated water deficit and extend leaf longevity.     

Water relations and mucopolysaccharide increases for a winter hardy cactus during acclimation to subzero temperatures

Summary Thickness, relative water content (RWC), osmotic pressure, water potential isotherms, and mucopolysaccharide content were measured for the photosynthetic chlorenchyma and the water-storage parenchyma of the winter hardy cactus, Opuntia humifusa, after shifting from day/night air temperatures of 25° C/15° C to 5° C/–5° C. After 14 d at 5° C/–5° C, the average fraction of water contained in the symplast decreased from 0.92 to 0.78, the water potential of saturated (fully hydrated) tissue was essentially unchanged, but the osmotic pressure of saturated tissue decreased (by 0.15 MPa for the chlorenchyma and 0.12 MPa for the water-storage parenchyma). After 7 weeks at 5° C/–5° C, tissue thickness was reduced by 61% for the chlorenchyma and 65% for the water-storage parenchyma, and the RWC decreased by 42% and 68%, respectively; these changes contributed to an osmotic pressure increase of 0.55 MPa for the chlorenchyma and 0.34 MPa for the water-storage parenchyma. During the 7 week acclimation to low temperature, mucopolysaccharide increased by 114% for the chlorenchyma and by 89% for the water-storage parenchyma. The water potential of the extracted mucopolysaccharide was relatively constant for an RWC between 1.00 and 0.30, decreasing abruptly below 0.30. Changes in water relations parameters and in mucopolysaccharide content during low-temperature acclimation may reduce water efflux from the cells, and thus reduce damage due to rapid dehydration during extracellular freezing.     

Root Water Uptake, Leaf Water Storage and Gas Exchange of a Desert Succulent: Implications for Root System Redundancy

A technique used for hydroponics was adapted to measure instantaneousroot water uptake from the soil for a leaf succulent CAM species,Agave deserti. Comparisons were made to previously modelledwater fluxes for A. deserti and to Encelia farinosa, a non-succulentC₃species. Net CO₂uptake and transpiration forA. deserti underwell-watered conditions occurred primarily at night whereasroot water uptake was relatively constant over 24 h. Leaf thicknessdecreased when transpiration commenced and then increased whenrecharge from the stem and soil occurred, consistent with previousmodels. A drought of 90 d eliminated net CO₂uptake and transpirationand reduced the water content of leaves by 62%. Rewetting theentire root system for 7 d led to a full recovery of leaf waterstorage but only 56% of maximal net CO₂uptake. Root water uptakewas maximal immediately after rewetting, which replenished rootwater content, and decreased to a steady rate by 14 d. Whenonly the distal 50% of the root system was rewetted, the timefor net CO₂uptake and leaf water storage to recover increased,but by 30 d gas exchange and leaf water storage were similarto 100% rewetting. Rewetting 10 or 20% of the root system resultedin much less water uptake; these plants did not recover leafwater storage or gas exchange by 30 d after rewetting. A redundancyin the root system of A. deserti apparently exists for dailywater uptake requirements under wet conditions but the entireroot system is required for rapid recovery from drought.Copyright1999 Annals of Botany Company Agave deserti Engelm., desert, drought, gas exchange, rewetting, roots, succulent, water uptake.     

Nocturnal water storage in plants having Crassulacean acid metabolism

Measurements of water uptake and transpiration, during the dark period of plants having Crassulacean acid metabolism (CAM) allow calculation of leaf-volume changes (V). Nocturnal leaf-volume changes of CAM plants have also been reported in the literature on the basis of waterdisplacement measurements. A third way of estimation is from measurements of turgor changes and cellular water-storage capacity using the pressure probe, cytomorphometry and the Scholander pressure chamber. An extension of the interpretation of results reported in the literature shows that for leaf succulent CAM plants the three different approaches give similar values of V ranging between 2.3 and 10.7% (v/v). It is evident that nocturnal malic-acid accumulation osmotically drives significant water storage in CAM leaf tissue.Abbreviations and symbols C_c water-storage capacity - E transporation (evaporational water loss) - P turgor pressure - U water uptake - V cell volume - cell-wall elastic modulus - osmotic pressure - CAM Crassulacean acid metabolism     

Water balance in the arborescent palm, Sabal palmetto. II. Transpiration and stem water storage

The contribution of stem water storage to the water balance of the arborescent palm, Sabal palmetto, was investigated using greenhouse studies, field measurements and a tree-cutting experiment. Water balance studies of greenhouse trees (1.5 to 3 m tall) were conducted in which transpiration was measured by weight loss, and changes in soil and stem water content by time-domain reflectometry. When the greenhouse plants were well-watered (soil moisture near saturation), water was withdrawn from the stem during periods of high transpiration and then replenished during the night so that the net transpirational water loss came primarily from the soil. As water was withheld, however, an increasing percentage of daily net transpirational water loss came from water stored in the stem. However, studies on palms growing in their natural environment indicated that during periods of high transpiration leaf water status was somewhat uncoupled from stem water stores. In a tree-cutting experiment, the maintenance of high relative water content of attached leaves was significantly correlated with stem volume/leaf area. Leaves of a 3-m tree remained green and fully hydrated for approximately 100d after it had been cut down, whereas those of a 1-m-tall plant turned brown within one week. The significance of stem water storage may be in buffering stem xylem potentials during periods of high transpiration and in contributing to leaf survival during extended period of low soil water availability.     

Water relations in silver birch during springtime: How is sap pressurised?

• Positive sap pressures are produced in the xylem of birch trees in boreal conditions during the time between the thawing of the soil and bud break. During this period, xylem embolisms accumulated during wintertime are refilled with water. The mechanism for xylem sap pressurization and its environmental drivers are not well known.
• We measured xylem sap flow, xylem sap pressure, xylem sap osmotic concentration, xylem and whole stem diameter changes, and stem and root non‐structural carbohydrate concentrations, along with meteorological conditions at two sites in Finland during and after the sap pressurisation period.
• The diurnal dynamics of xylem sap pressure and sap flow during the sap pressurisation period varied, but were more often opposite to the diurnal pattern after bud burst, i.e. sap pressure increased and sap flow rate mostly decreased when temperature increased. Net conversion of soluble sugars to starch in the stem and roots occurred during the sap pressurisation period. Xylem sap osmotic pressure was small in comparison to total sap pressure, and it did not follow changes in environmental conditions or tree water relations.
• Based on these findings, we suggest that xylem sap pressurisation and embolism refilling occur gradually over a few weeks through water transfer from parenchyma cells to xylem vessels during daytime, and then the parenchyma are refilled mostly during nighttime by water uptake from soil. Possible drivers for water transfer from parenchyma cells to vessels are discussed. Also the functioning of thermal dissipation probes in conditions of changing stem water content is discussed.

     

Development of Chlorenchyma and Window Tissues in Leaves of Peperomia columella

Window leaves consist primarily of tissues specialized for waterstorage (window tissue) and photosynthesis (chlorenchyma). Theobjective of our research was to determine when these specializationsoccur during leaf development in Peperomia columella, a succulentwindow plant, native to deserts of South America. We measuredabsolute and relative volumes of leaf tissues. Young leavesconsist of approximately 75% chlorenchyma and 12% window tissue,suggesting that they are structurally specialized primarilyfor photosynthesis rather than water storage. In mature leavesthe percentages of chlorenchyma and window tissue are approximately20% and 58%, respectively, indicating that specialization forwater storage occurs during later stages of leaf development.The percent window tissue decreases in mature leaves, but increasesin young leaves with water stress.Copyright 1993, 1999 AcademicPress Chlorenchyma, Peperomia columella, succulent, window plant     

华北落叶松夜间树干液流特征及生长季补水格局

在宁夏六盘山北侧半干旱区的叠叠沟小流域,采用热扩散探针法在2011年生长季监测了华北落叶松(Larix principisrupprechtii)人工林的树干液流速率,分析了夜间树干液流和补水量的变化特征及与气象、土壤水分等环境因子的关系.结果表明:树干液流速率日变化表现为典型的单峰宽峰曲线,且整个生长季均存在微弱的夜间液流,一般表现为逐渐减小,特别是在晴天,且晴天的变幅显著大于雨天.除生长季中期雨天夜间液流平均速率显著高于晴天,生长季初期及末期雨天与晴天的差异并不显著.生长季内,夜间树干补水总量为11.03 mm,占总蒸腾量的7.22％;5月份的树干补水量最大(4.19mm),其他月份的树干补水量明显减小,在0.9-1.7mm的范围波动.但不同月份间的补水贡献率存在明显差异,表现为生长季末期(9、10月)＞初期(5月)＞中期(6-8月).相关分析表明,日补水量与各气象因子关系不大,仅与降水量显著正相关(P＜0.05),与土壤含水率、日间蒸腾量、日蒸腾总量极显著正相关(P＜0.01).夜间补水的月蒸腾贡献率与月均土壤含水率、月均气温、月均日间蒸腾量、月总蒸腾量等显著相关(P＜0.05);而夜间补水的日蒸腾贡献率与日最高气温、日均气温、日间蒸腾量、日均饱和水汽压差、日总蒸腾量、日均太阳辐射强度、日最低气温、日均空气相对湿度、日降水量、土壤含水率等极显著相关(P＜0.01),经逐步回归分析建立了日补水量蒸腾贡献率与环境因子的多元线性模型.