Fluctuations In Titratable Acidity Research Articles

Different CO2 acclimation strategies in juvenile and mature leaves of Ottelia alismoides.

The freshwater macrophyte, Ottelia alismoides, is a bicarbonate user performing C4 photosynthesis in the light, and crassulacean acid metabolism (CAM) when acclimated to low CO2. The regulation of the three mechanisms by CO2 concentration was studied in juvenile and mature leaves. For mature leaves, the ratios of phosphoenolpyruvate carboxylase (PEPC) to ribulose-bisphosphate carboxylase/oxygenase (Rubisco) are in the range of that of C4 plants regardless of CO2 concentration (1.5-2.5 at low CO2, 1.8-3.4 at high CO2). In contrast, results for juvenile leaves suggest that C4 is facultative and only present under low CO2. pH-drift experiments showed that both juvenile and mature leaves can use bicarbonate irrespective of CO2 concentration, but mature leaves have a significantly greater carbon-extracting ability than juvenile leaves at low CO2. At high CO2, neither juvenile nor mature leaves perform CAM as indicated by lack of diurnal acid fluctuation. However, CAM was present at low CO2, though the fluctuation of titratable acidity in juvenile leaves (15-17µequiv g-1 FW) was slightly but significantly lower than in mature leaves (19-25µequivg-1 FW), implying that the capacity to perform CAM increases as leaves mature. The increased CAM activity is associated with elevated PEPC activity and large diel changes in starch content. These results show that in O. alismoides, carbon-dioxide concentrating mechanismsare more effective in mature compared to juvenile leaves, and C4 is facultative in juvenile leaves but constitutive in mature leaves.

Source-to-sink relationship between green leaves and green petals of different ages of the CAM orchid Dendrobium cv. Burana Jade

Photosynthetic capacities of green leaves (GL) and green flower petals (GFP) of different ages of the CAM plant Dendrobium cv. Burana Jade were studied through chlorophyll (Chl) content, Chl fluorescence characteristic Fv/Fm, maximal photosynthetic O2 evolution rates (P max), and CAM acidities [dawn/dusk fluctuations in titratable acidity (TA)]. All these photosynthetic parameters were higher in GL than in GFP. Among the different ages of GFP, the young GFP had significant higher readings of all photosynthetic parameters than the oldest GFP, indicating that reduced photosynthesis occurred in the senesced GFP. The source-to-sink relationship between GL and GFP was also studied by comparing the diurnal changes in contents of total soluble and insoluble sugars and TA between the fully irradiated (FI) control (with both irradiated GL and GFP) and GL-darkened plants (covering all GL with aluminium foils, leaving only the GFP exposed to radiation). CAM acidities were much lower in GL darkened with aluminium foils compared to those of FI-GL while there were no differences in CAM acidities of their GFP. The contents of total soluble and insoluble sugars and the CAM acidities of GL towards the end of the day were lower in GL-darkened plants compared to that of FI-plants. Hence CAM acidities of GL depended on their saccharide contents. However, diurnal changes of TA in GFP were similar in all GFP regardless of their ages, with or without GL photosynthetic sources. Thus CAM acidities of GFP are independent of GL saccharides. However, lower saccharide content in GFP (especially the oldest GFP) of GL-darkened plants implies that GFP function as sinks and depend on saccharides exported from GL for its development and growth.

Crassulacean acid metabolism in the ZZ plant, Zamioculcas zamiifolia (Araceae)

Zamioculcas zamiifolia (Araceae), a terrestrial East African aroid, with two defining attributes of crassulacean acid metabolism (CAM) (net CO(2) uptake in the dark and diel fluctuations of titratable acidity) is the only CAM plant described within the Araceae, a mainly tropical taxon that contains the second largest number of epiphytes of any vascular plant family. Within the Alismatales, the order to which the Araceae belong, Z. zamiifolia is the only documented nonaquatic CAM species. Zamioculcas zamiifolia has weak CAM that is upregulated in response to water stress. In well-watered plants, day-night fluctuations in titratable acidity were 2.5 μmol H(+)·(g fresh mass)(-1), and net CO(2) uptake in the dark contributed less than 1% to daily carbon gain. Following 10 d of water stress, net CO(2) uptake in the light fell 94% and net CO(2) uptake in the dark increased 7.5-fold, such that its contribution increased to 19% of daily carbon gain. Following rewatering, dark CO(2) uptake returned to within 5% of prestressed levels. We postulate that CAM assists survival of Z. zamiifolia by reducing water loss and maintaining carbon gain during seasonal droughts characteristic of its natural habitat.

Susceptibility of green leaves and green flower petals of CAM orchid Dendrobium cv. Burana Jade to high irradiance under natural tropical conditions

Photosynthetic rates of green leaves (GL) and green flower petals (GFP) of the CAM plant Dendrobium cv. Burana Jade and their sensitivities to different growth irradiances were studied in shade-grown plants over a period of 4 weeks. Maximal photosynthetic O2 evolution rates and CAM acidities [dawn/dusk fluctuations in titratable acidity] were higher in leaves exposed to intermediate sunlight [a maximal photosynthetic photon flux density (PPFD) of 500-600 µmol m-2 s-1] than in leaves grown under full sunlight (a maximal PPFD of 1 000-1 200 µmol m-2 s-1) and shade (a maximal PPFD of 200-250 µmol m-2 s-1). However, these two parameters of GFP were highest in plants grown under the shade and lowest in full sun-grown plants. Both GL and GFP of plants exposed to full sunlight had lower predawn Fv/Fm [dark adapted ratio of variable to maximal fluorescence (the maximal photosystem 2 yield without actinic irradiation)] than those of shade-grown plants. When exposed to intermediate sunlight, however, there were no significant changes in predawn Fv/Fm in GL whereas a significant decrease in predawn Fv/Fm was found in GFP of the same plant. GFP exposed to full sunlight exhibited a greater decrease in predawn Fv/Fm compared to those exposed to intermediate sunlight. The patterns of changes in total chlorophyll (Chl) content of GL and GFP were similar to those of Fv/Fm. Although midday Fv/Fm fluctuated with prevailing irradiance, changes of midday Fv/Fm after exposure to different growth irradiances were similar to those of predawn Fv/Fm in both GL and GFP. The decreases in predawn and midday Fv/Fm were much more pronounced in GFP than in GL under full sunlight, indicating greater sensitivity in GFP to high irradiance (HI). In the laboratory, electron transport rate and photochemical and non-photochemical quenching of Chl fluorescence were also determined under different irradiances. All results indicated that GFP are more susceptible to HI than GL. Although the GFP of Dendrobium cv. Burana Jade require a lower amount of radiant energy for photosynthesis and this plant is usually grown in the shade, is not necessarily a shade plant.

A phylogenetic view of low‐level CAM in Pelargonium (Geraniaceae)

Crassulacean acid metabolism (CAM) is common in several plant families and is often associated with succulence. Few studies have examined the occurrence of CAM from a phylogenetic perspective. The genus Pelargonium is promising for such a study because members are characterized by dramatic variation in growth form (including geophytes, shrubs, and stem succulents) and because growth form diversity is expressed to the greatest extent in a monophyletic group comprising 80% of Pelargonium species. This clade, predominantly from the winter rainfall region of southern Africa, likely proliferated in response to Miocene or Pliocene aridification. We present a survey for CAM across Pelargonium, emphasizing the winter rainfall clade. Dawn/dusk fluctuations in titratable acidity were examined in 41 species, with detailed measurements of carbon uptake and stomatal conductance under progressive water stress in four species. No species exhibited obligate CAM. When well-watered, most species exhibited stomatal conductances and acid fluctuations characteristic of C(3) photosynthesis, though some exhibited more pronounced increases in nocturnal acidity, suggesting CAM cycling. In four species examined during dry-down, water stress led to increased nighttime acid levels and decreased daytime stomatal conductance. Ultimately, stomata closed and external carbon uptake ceased, consistent with CAM idling. These results are discussed from the perspective of the evolution of CAM flexibility.

The Influence of Leaf "Windows" on Crassulacean Acid Metabolism in the South African Succulent Senecio Rowleyanus (Asteraceae)

Net CO2 exchange rate (PN) of shoots and diel fluctuations in titratable acidity of leaves of Senecio rowleyanus were measured to determine whether penetration of radiant energy through leaf “windows” (narrow, translucent strips on the leaf epidermis) resulted in increased CAM. Nocturnal PN and nighttime increases in acidity were compared among plants with windows covered with reflective adhesive tape, transparent adhesive tape (to control for potential effects of the adhesive), and no tape. The windows did not significantly enhance the degree of CAM in S. rowleyanus.

Seasonal patterns of acid fluctuations and resource storage in the arborescent cactus Opuntia excelsa in relation to light availability and size.

We investigated relationships between light availability, diel acid fluctuation, and resource storage in the arborescent cactus Opuntia excelsa growing in western Mexico. We compared canopy and understory individuals from a deciduous forest as well as open-grown plants of the same approximate size as those in the understory. During the wet season light availability and daily fluctuations in titratable acidity (an index of carbon uptake) were lower in the understory than in unshaded habitats. In the dry season all plants had reduced levels of acid fluctuation, with the smallest individuals, regardless of habitat, showing the greatest reduction. These data suggest that light availability in the forest understory constrains carbon assimilation during the wet season, but that a factor associated with plant size, possibly water status, limits carbon gain during the dry season. Plants in all habitats remained physiologically active for at least five months into the dry season. We suggest that this was possible due to the maintenance of constant concentrations of water and nitrogen in the photosynthetically active chlorenchyma. Parenchyma in terminal cladodes showed a different seasonal pattern of resource storage; water content and nitrogen concentration were reduced from the wet to the dry season in the parenchyma. Using the parenchyma to supply photosynthetic tissues during times of reduced resource availability allows O. excelsa to assimilate carbon during times of the year when most other trees in the forest are leafless.

The appearance of a new molecular species of phosphoenolpyruvate carboxylase (PEPC) and the rapid induction of CAM in Sedum telephium L.

Abstract. When detached leaves of Sedum telephium are incubated in the absence of water, a rapid switch from C3 photosynthesis to CAM (as indicated by the onset of day‐to‐night fluctuations in titratable acidity. ΔH+) occurs within the first dark period. The C3‐CAM switch in intact plants occurs within 3 5d. Extractable activity of phosphoenolpyruvate carboxylase (PEPC) increases five‐fold in intact plants during CAM induction; however, during rapid CAM induction in detached leaves, there is only a very small increase in PEPC activity. Fractionation by anion exchange chromatography of crude extracts from leaves of intact plants subjected to water deficit shows that CAM induction is associated with the appearance of a molecular species of PEPC termed PEPC I. PEPC I is barely detectable in well‐watered plants which are not performing CAM. The major form in these plants is termed PEPC II. In leaves from intact plants, there is a significant positive correlation between PEPC I activity and ΔH+ during a period of increasing water deficit. PEPC I exhibits day to night fluctuations in malate sensitivity, being less sensitive during the dark period. In contrast, PEPC II is more sensitive to inhibition by malate and has no day to night fluctuation in sensitivity. In detached leaves deprived of water, a small increase in PEPC I capacity is detected at the end of the first dark period (20 h after the start of treatment). The results suggest that PEPC I is required for attainment of maximum nocturnal malic acid synthesis. There is a significant correlation between leaf water status (relative water content), ΔH+, total PEPC and PEPC I activity suggesting that the internal water status of the plant may be a trigger for CAM induction. Abscisic acid applied to detached leaves does not cause nocturnal acidification.

A preliminary survey for crassulacean acid metabolism (CAM) in submerged aquatic macrophytes in New Zealand

Abstract This preliminary survey covered 13 aquatic macrophytes present at two sites in Lake Taupo and two sites in the Auckland region (northern half of the North Island, New Zealand). Three species previously unrecorded as CAM plants, Isoetes kirkii A. Braun, Lilaeopsis lacustris Hill, and Vallisneria spiralis Graeb., exhibited diurnal fluctuations in titratable acidity and malic acid content. The changes in titratable acidity varied seasonally from 97 (Dec. 1985) to 46 (June 1986) microequivalents per gram fresh weight for I. kirkii, and from 40 to 23 and 51 to 26 microequivalents per gram fresh weight for L. lacustris and V. spiralis, respectively. The absence of CAM both in other members of the low mixed community and in other hydrocharitacean species at the same sites suggests that CAM can not be unequivocally regarded as conferring an ecological advantage to a species.

Shifts in the Carbon Metabolism of Xerosicyos danguyi H. Humb. (Cucurbitaceae) Brought About by Water Stress

Xerosicyos danguyi H. Humb. (Cucurbitaceae) is an unusual leaf succulent endemic to Madagascar. Under well-watered conditions the plant exhibited Crassulacean acid metabolism (CAM), as characterized by large diurnal changes in titratable acidity, predominantly nighttime stomatal opening and CO(2) uptake, and high delta(13)C values. When plants were exposed to water stress for a minimum of a month, they shifted to a mode of carbon metabolism previously labeled CAM-idling. Under this mode of metabolism, the plants exhibited reduced stomatal opening, reduced CO(2) uptake, dampened diurnal fluctuations in titratable acidity, and no apparent changes in the delta(13)C values. Additionally, investigations showed that the stress hormones 1-aminocyclopropane-1-carboxylic acid (an ethylene precursor) and abscisic acid increased as much as 6-fold in the water-stressed plants. The results are discussed in relation to physiological significance and evolution of the CAM-idling mode of metabolism.

DISTRIBUTION OF DIURNAL ACID METABOLISM IN THE GENUS ISOETES

Previous studies have revealed a crassulacean acid metabolism‐like pathway in the submerged aquatic Isoetes howellii (Isoetaceae). A survey of 11 other species in this largely aquatic genus showed all had diurnal fluctuations in titratable acidity and malic acid concentration. The magnitude of the diurnal fluctuation in titratable acidity varied from 119 (μeq g–1 fresh weight) for I. bolanderi to 44 for I. engelmanni. Although these species were grown under similar conditions in a greenhouse, it is questionable whether or not any significance can be attached to observed species specific differences in magnitude of diurnal acid fluctuation. This is suggested by a single species comparison of plants in the field with greenhouse grown plants which indicated nearly as much variation in magnitude of acid fluctuations as observed across all species in the greenhouse. Much of this variation may be due to seasonal differences. Substrate appears to have very little effect on magnitude of acid fluctuations. Since this survey covered most of the range of the latitudinal, elevational, habitat, and morphological variation in the genus, it is likely that diurnal acid metabolism will prove to be common in the genus Isoetes.

Seasonal patterns of growth, tissue acid fluctuations, and 14CO2 uptake in the crassulacean acid metabolism epiphyte Tjllandsia usneoides L. (Spanish moss).

Seasonal patterns of growth, 14CO2 uptake, and fluctuations in tissue titratable acidity were studied over the course of a year at a study site in the coastal plain of North Carolina.Elongation rates of Spanish moss strands were maximal in the summer and minimal in the winter. Summer maximal biomass addition rates were calculated to be 3.4 mg·month-1. Mortality of the strands was greatest in the winter months. Rates of 14CO2 uptake and fluctuations in tissue acidity were greatest in the summer over a fairly broad spectrum of environmental conditions (day and night temperatures, irradiance, length of drought). Maximal 14CO2 uptake rates (1.2 mg CO2·mg Chl-1 ·h01) were measured in May 1978. Rates of 14CO2 uptake and fluctuations in titratable acidity were inhibited below 5°C and eliminated at 0°C air temperatures.Isothermal diurnal conditions resulted in low rates of 14CO2 uptake. Tissue water content did not appear to be a major factor controlling 14CO2 uptake rates. However, tissue wetting by rain severely reduced nighttime uptake yet stimulated low rates of daytime 14CO2 uptake. This was the only condition in which daytime 14CO2 uptake occurred, excluding the early morning and late afternoon 14CO2 uptake typical of many Crassulacean Acid Metabolism (CAM) plants.The results suggest that tissue water content is not the major factor controlling CO2 uptake as has been found in many other CAM species; and that low temperatures limit the growth of Spanish moss in North Carolina.

Carbon Metabolism in Two Species of Pereskia (Cactaceae)

The Pereskia are morphologically primitive, leafed members of the Cactaceae. Gas exchange characteristics using a dual isotope porometer to monitor (14)CO(2) and tritiated water uptake, diurnal malic acid fluctuations, phosphoenolpyruvate carboxylase, and malate dehydrogenase activities were examined in two species of the genus Pereskia, Pereskia grandifolia and Pereskia aculeata. Investigations were done on well watered (control) and water-stressed plants. Nonstressed plants showed a CO(2) uptake pattern indicating C(3) carbon metabolism. However, diurnal fluctuations in titratable acidity were observed similar to Crassulacean acid metabolism. Plants exposed to 10 days of water stress exhibited stomatal opening only during an early morning period. Titratable acidity, phosphoenolpyruvate carboxylase activity, and malate dehydrogenase activity fluctuations were magnified in the stressed plants, but showed the same diurnal pattern as controls. Water stress causes these cacti to shift to an internal CO(2) recycling ("idling") that has all attributes of Crassulacean acid metabolism except nocturnal stomata opening and CO(2) uptake. The consequences of this shift, which has been observed in other succulents, are unknown, and some possibilities are suggested.

Responses of Succulents to Plant Water Stress

Experiments were performed to test the hypothesis that succulents "shift" their method of photosynthetic metabolism in response to environmental change. Our data showed that there were at least three different responses of succulents to plant water status. When plant water status of Portulacaria afra (L.) Jacq. was lowered either by withholding water or by irrigating with 2% NaCl, a change from C(3)-photosynthesis to Crassulacean acid metabolism (CAM) occurred. Fluctuation of titratable acidity and nocturnal CO(2) uptake was induced in the stressed plants. Stressed Peperomia obtusifolia A. Dietr. plants showed a change from C(3)-photosynthesis to internal cycling of CO(2). Acid fluctuation commenced in response to stress but exogenous CO(2) uptake did not occur. Zygocactus truncatus Haworth plants showed a pattern of acid fluctuation and nocturnal CO(2) uptake typical of CAM even when well irrigated. The cacti converted from CAM to an internal CO(2) cycle similar to Peperomia when plants were water-stressed. Reverse phase gas exchange in succulents results in low water loss to carbon gain. Water is conserved and low levels of metabolic activity are maintained during drought periods by complete stomatal closure and continual fluctuation of organic acids.