Water-storage Parenchyma Research Articles

Leaf Anatomy and CO<sub>2</sub> Recycling during Crassulacean Acid Metabolism in Twelve Epiphytic Species of Tillandsia (Bromeliaceae)

The relationship between leaf anatomy, specifically the percent of leaf volume occupied by waterstorage parenchyma (hydrenchyma), and the contribution of respiratory CO₂ during Crassulacean acid metabolism (CAM) was investigated in 12 epiphytic species of Tillandsia. It has been postulated that the hydrenchyma, which contributes to CO₂ exchange through respiration only, may be causally related to the recently observed phenomenon of CO₂ recycling during CAM. Among the 12 species of Tillandsia, leaves of T. usneoides and T. bergeri exhibited 0% hydrenchyma, while the hydrenchyma in the other species ranged from 2.9% to 53% of leaf cross-sectional area. Diurnal malate fluctuation and nighttime atmospheric CO₂ uptake were measured in at least four individuals of each species. A significant excess of diurnal malate fluctuation as compared with atmospheric CO₂ absorbed overnight was observed only in T. schiedeana. This species had an intermediate proportion (30%) of hydrenchyma in its leaves. Results of this study do not support the hypothesis that CO₂ recycling during CAM may reflect respiratory contributions of CO₂ from the tissue hydrenchyma.

Changes in Osmotic Pressure and Mucilage during Low-Temperature Acclimation of Opuntia ficus-indica

Opuntia ficus-indica, a Crassulacean acid metabolism plant cultivated for its fruits and cladodes, was used to examine chemical and physiological events accompanying low-temperature acclimation. Changes in osmotic pressure, water content, low molecular weight solutes, and extracellular mucilage were monitored in the photosynthetic chlorenchyma and the water-storage parenchyma when plants maintained at day/night air temperatures of 30/20 degrees C were shifted to 10/0 degrees C. An increase in osmotic pressure of 0.13 megapascal occurred after 13 days at 10/0 degrees C. Synthesis of glucose, fructose, and glycerol accounted for most of the observed increase in osmotic pressure during the low-temperature acclimation. Extracellular mucilage and the relative apoplastic water content increased by 24 and 10%, respectively, during exposure to low temperatures. These increases apparently favor the extracellular nucleation of ice closer to the equilibrium freezing temperature for plants at 10/0 degrees C, which could make the cellular dehydration more gradual and less damaging. Nuclear magnetic resonance studies helped elucidate the cellular processes during ice formation, such as those revealed by changes in the relaxation times of two water fractions in the chlorenchyma. The latter results suggested a restricted mobility of intracellular water and an increased mobility of extracellular water for plants at 10/0 degrees C compared with those at 30/20 degrees C. Increased mobility of extracellular water could facilitate extracellular ice growth and thus delay the potentially lethal intracellular freezing during low-temperature acclimation.

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

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.

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.

Differences in Water Relations Parameters for the Chlorenchyma and the Parenchyma of Opuntia ficus-indica Under Wet Versus Dry Conditions

Water relations of the photosynthetic tissue (chlorenchyma) and of the water-storage parenchyma were studied for well watered and droughted Opuntia ficus-indica, a crassulacean acid metabolism plant cultivated worldwide for its fruits and cladodes. For well watered plants, die1 changes in osmotic pressure were evident in the chlorenchyma. Droughting the plants for 4 months resulted in a massive loss of water from the cladodes, particularly from the water-storage parenchyma, which could lose up to 82% of the water present at full turgor without irreversible tissue damage. Pressure-volume curves indicated a decrease in the osmotic pressure at full turgor of about 0.1 MPa for the water-storage parenchyma cells during drought; such a decrease of osmotically active solutes was consistent with the appearance of large numbers of starch grains. The bulk modulus of elasticity was 0.36 MPa for the water-storage parenchyma cells and 2.5-fold higher for the chlorenchyma cells, which were smaller with thicker cell walls than the former cells. Mucilage, a polysaccharide occurring extracellularly, constituted about 14% of the cladode dry weight; it could hold more than 30% of the total water content of the water-storage parenchyma. Polymerisation of sugars, large elastic cells in the water-storage parenchyma and mucilage with its high water-holding capacity helped maintain a positive turgor in the photosynthetic tissue, even after 4 months of drought.

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 storage and osmotic pressure influences on the water relations of a dicotyledonous desert succulent

Abstract Water storage and nocturnal increases in osmotic pressure affect the water relations of the desert succulent Ferocactus acanthodes, which was studied using an electrical circuit analog based on the anatomy and morphology of a representative individual. Transpiration rates and osmotic pressures over a 24‐h period were used as input variables. The model predicted water potential, turgor pressure and water flow for various tissues. Plant capacitances, storage resistances and nocturnal increases in osmotic pressure were varied to determine their role in the water relations of this dicotyledonous succulent. Water coming from storage tissues contributed about one‐third of the water transpired at night: the majority of this water came from the nonphotosynthetic, water storage parenchyma of the stem. Time lags of 4 h were predicted between maximum transpiration and maximum water uptake from the soil. Varying the capacitance of the plant caused proportional changes in osmotically driven water movement but changes in storage resistance had only minor effects. Turgor pressure in the chlorenchyma depended on osmotic pressure, but was fairly insensitive to doubling or halving of the capacitance or storage resistance of the plant. Water uptake from the soil was only slightly affected by osmotic pressure changes in the chlorenchyma. For this stem succulent, the movement of water from the chlorenchyma to the xylem and the internal redistribution of water among stem tissues were dominated by nocturnal changes in chlorenchyma osmotic pressure, not by transpiration.

Carbon balance during CAM: an assessment of respiratory CO2 recycling in the epiphytic bromeliads Aechmea nudicaulis and Aechmea fendleri

Abstract The regulation of crassulacean acid metabolism (CAM) under controlled environmental conditions has been investigated for two tropical epiphytes, relating plant water and carbon balance to growth form and habitat preference under natural conditions. Aechmea fendleri is restricted to wet, upper montane regions of Trinidad, while A. nudicaulis has a wider distribution extending into more arid regions of the island. Morphological characteristics of these plants are related to habitat preference in terms of leaf succulence (0.44 and 0.94 kg m−2 for the two species respectively) and a distinct layer of water storage parenchyma in A. nudicaulis In contrast, the thinner leaves of A. fendleri contain little water‐storage parenchyma and less chlorenchyma per unit area, but the plants have a more open leaf rosette. The two species differ in expression of CAM, since the proportion of respiratory CO2 recycled as part of CAM had been found to be much lower in A. fendleri This study compared the efficiency of water use and role of respiratory CO2 recycling under two PAR regimes (300 and 120 μnol m−2 s−1) and three night temperatures (12, 18 and 25 °C). Dark CO2 uptake rates for both species were comparable to plants in the field (maximum of 2.3 ± 0.2 μmol m−2s−1± SD, n= 3). Total net CO2 uptake at night increased on leaf area basis with temperature for both species under higher PAR, although under the low PAR regime CO2 uptake was maximal at 18 °C. Water‐use efficiency (WUE) increased at 18 °C and 25 °C during dark CO2 uptake (Phase I) and also during late afternoon photosynthesis (Phase IV) in both species. For A. fendleri, dawn to dusk changes in titrable acidity (ΔH +) were similar under high and low PAR, although ΔH+ was correlated to night temperature and PAR in A. nudicaulis. The proportion of ΔH+ derived from respiratory CO2 also varied with experimental conditions. Thus percentage recycling was lower in A. fendleri under high PAR (0–10%), but was only reduced at 18 °C under low PAR. Recycling by A. nudicaulis ranged from 32–42% under high PAR, but was also reduced to 6% under low PAR at 18 °C; at 12 °C and 25 °C, recycling was 37% and 52% respectively. Previous studies have suggested a relationship between the proportion of recycling and degree of water stress. This study indicated that CAM as a CO2 concentrating mechanism regulates both water‐use efficiency and plant carbon balance in these epiphytes, in response to PAR and night temperature. However, the precise relationship between respiratory processes and the balance between external and internal sources of CO2 is as yet unresolved.

COMPARATIVE LEAF ANATOMY OF SOLIDAGO AND RELATED ASTERACEAE

Leaf anatomy of 63 taxa is investigated to elucidate generic relationships among Brachychaeta, Brintonia, Chrysoma, Euthamia, Gundlachia, Oligoneuron, Oreochrysum, Petradoria, and Solidago. All these genera have been included at one time or another within Solidago. Aster ptarmicoides is also studied because it hybridizes with some species of Solidago (sens. str.). Qualitative and quantitative differences in mesophyll, storage parenchyma, secretory apparatus, bundle sheath extensions, and midvein structure allow rather precise grouping of the taxa. Brachychaeta, Brintonia, Oligoneuron, Oreochrysum, and Aster ptarmicoides should be considered as constituents of Solidago. They all have bundle sheath extensions and little or no water storage parenchyma. In Solidago secretory cavities, when present, are shaped and positioned differently from those in Euthamia. The absence of bundle sheath extensions and various combinations of other anatomical features suggest that Chrysoma, Euthamia, Gundlachia, and Petradoria are generically distinct from one another and from Solidago.