The Effect of Temperature and Light on Gas Exchange and Acid Accumulation in the C3-CAM Plant Clusia minor L

Gas exchange and organic acid accumulation of the C₃-CAM intermediateClusia minor L. were investigated in response to various day/nighttemperatures and two light regimes (low and high PAR). For bothlight levels equal day/night temperatures between 20°C and30°C caused a typical C₃ gas exchange pattern with all CO₂uptake occurring during daylight hours. A day/ night temperatureof 15°C caused a negative CO₂ balance over a 24 h periodfor low-PAR-grown plants while high-PAR-grown plants showeda CAM gas exchange pattern with most CO₂ uptake taking placeduring the dark period. However, there was always a considerablenight-time accumulation of malic acid which increased when thenight-time temperature was lowered and had its maximum (54 mmolm^–2) at day/night temperature of 30/15°C. A significantamount of malic acid accumulation (23 mmol m^–2) in low-PAR-grownplants was observed only at 30/15°C. Recycling of respiratoryCO₂ in terms of malic acid accumulation reached between 2·0and 21·5 mmol m_–2 for high-PAR-grown plants whilethere was no significant recycling for low-PAR-grown plants.Both low and high-PAR-grown plants showed considerable night-timeaccumulation of citric acid. Indeed under several temperatureregimes low-PAR-grown plants showed day/night changes in citricacid levels whereas malic acid levels remained approximatelyconstant or slightly decreased. It is hypothesized that lowand high-PAR-grown plants have different requirements for citrate.In high-PAR-grown plants, the breakdown of citrate preventsphotoinhibition by increasing internal CO₂ levels, whereas inlow-PAR-grown plants the night-time accumulation of citric acidmay function as an energy and carbon saving mechanism. Key words: C. minor, C₃, CAM, citric acid, light intensity     

The Response of the C3-CAM Tree, Clusia rosea, to Light and Water Stress: II. INTERNAL CO2 CONCENTRATION AND WATER USE EFFICIENCY

Gas exchange in Clusia rosea has been measured under variousconditions of water status, light and leaf-air vapour pressuredeficit (_w, mbar bar^–1) which produce daytime (C₃), night-time(CAM) or 24 h uptake of CO₂. At high light levels, at a _w of6.6, well-watered plants utilized C₃ photosynthesis while CAMand 24 h uptake occurred under lower light levels and with lowto normal water availability and differing _w (13.5 and 3.4,respectively). CO₂ uptake was highest, stomatal conductanceto water vapour (gH₂o) lowest, and water use efficiency (WUE)highest in plants using C₃ photosynthesis. This latter factis contrary to the accepted view that CAM is most water useefficient, i.e. it optimizes CO₂ uptake with minimal water loss.It is suggested that the low CO₂ uptake in CAM photosynthesismay be related not only to the higher _w but also to the factthat Clusia species accumulate citrate which may originate fromß-carboxylation of fatty acids (i.e. an internal sourceof CO₂) and does not contribute to night-time external CO₂ assimilation.Curves of assimilation (A) versus internal partial pressureof CO₂ (A/c₁) for the three photosynthetic types, under atmosphericconditions, did not produce a single trend. The trends whichwere produced represent the supply function for the interaction,under differing modes of photosynthesis, of the two major enzymesystems involved in CAM. Key words: Clusia rosea, Crassulacean acid metabolism, C₃ photosynthesis, internal CO₂ concentration, 24 h carbon dioxide uptake, water use efficiency.     

The Response of the C3-CAM Tree, Clusia rosea, to Light and Water Stress: I. GAS EXCHANGE CHARACTERISTICS

Gas exchange measurements were undertaken on 2-year-old plantsof Clusia rosea. The plants were shown to have the ability toswitch from C₃-photosynthesis to CAM and vice versa regardlessof leaf age and, under some conditions, CO₂ was taken up continuously,throughout the day and night. The light response was saturatedby 120 µmol m^–2 s^–1 typical of a shade plant. Gas exchange patterns in response to light, water and VPD wereexamined. All combinations of daytime and night-time CO₂ uptakewere observed, with rates of CO₂ uptake ranging from 2 to 11µmol m^–2 s^–1 depending upon water status andlight. Categorization of this plant asC₃, CAM or an intermediateis impossible. Differing VPD affected the magnitude of changesfrom CAM to C₃-photosynthesis (0 to 0.5 and 0 to 6.0 µmolm^–2 s^–1 CO₂, respectively) when plants were watered.Under well-watered conditions, but not under water stress, highPPFD elicited changes from CAM to C₃ gas exchange. This is unusualnot only for a shade plant but also for a plant with CAM. Itis of ecological importance for C. rosea, which may spend theearly years of its life as an epiphyte or in the forest understorey,to be able to maximize photosynthesis with minimal water loss. Key words: Clusia rosea, CAM, C₃, stress     

Comparison of Photosynthetic Responses of the Sympatric Tropical C3 Species Clusia multiflora H. B. K. and the C3-CAM Intermediate Species Clusia minor L. to Irradiance and Drought Stress in a Phytotron1

Abstract: Clusia multiflora H. B. K., an obligate C₃ species and Clusia minor L. a C₃/CAM intermediate species, are two physio-types of a similar morphotype. They can sympatrically occupy secondary savanna sites exposed to high insolation in the tropics. In C. multiflora severe stress, i.e., switching shade-grown plants to high light plus drought, resulted in leaves browning or yellowing and becoming necrotic. However, in long-term light stress C. multiflora was able to grow new leaves with their photosynthetic apparatus fit for high light conditions. Shade-grown C. minor readily overcame switching to high light conditions and drought, responding by a rapid change from C₃ photosynthesis to CAM. Decreasing _soil led to increased abscisic acid levels in the leaves of C. minor, however CAM induction was not directly related to this and was mainly determined by increased PPFD. Both species were capable of rapid accumulation of zea-xanthin for acute photoprotection following high PPFD exposure. The maximum capacity for zeaxanthin accumulation was larger in C. minor, but under steady high PPFD it only partially made use of this capacity, relying on high internal CO₂ concentrations of Phase Ill of CAM, in addition to zeaxanthin, for acute photo-protection. Thus, by different means the two species perform well under high light conditions. However, C. multiflora needs time for development of adapted leaves under such stress conditions while the more flexible C. minor can readily switch from low light to high light conditions.     

Short-term changes in carbon-isotope discrimination in the C3-CAM intermediate Clusia minor L. growing in Trinidad

On-line instantaneous carbon isotope discrimination was measured in conjunction with net uptake of CO₂ in leaves of exposed and shaded plants of the C₃-CAM intermediate Clusia minor growing under natural conditions in Trinidad. At the end of the rainy season (late January-early February, 1992) C₃ photosynthesis predominated although exposed leaves recaptured a small proportion of respiratory CO₂ at night for the synthesis of malic acid. Citric acid was the major organic acid accumulated by exposed leaves at this time with a citric: malic acid ratio of 11:1. Values of instantaneous discrimination () in exposed leaves during the wet season rose from 17.1 shortly after dawn to 22.7 around mid-day just before stomata closed, suggesting that most CO₂ was fixed by Rubisco at this time. During the late afternoon, instantaneous declined from 22.2 to 17, probably reflecting the limited contribution from PEPc activity and an increase in diffusional resistance to CO₂ in exposed leaves. Shaded leaves showed no CAM activity and CO₂ uptake proceeded throughout the day in the wet season. The decrease in instantaneous from 27 in the morning to 19.2 in the late afternoon was therefore entirely due to diffusional limitation. Leaves sampled in the dry season (mid-March, 1992) had by now induced full CAM activity with both malic and citric acids accumulated overnight and stomata closed for 4–5 h over the middle of the day. Values of instantaneous measured over the first 3 h after dawn (6.4–9.1) indicated that C₄ carboxylation dominated CO₂ uptake for most of the morning when rates of photosynthesis were maximal, implying that under natural conditions, the down regulation of PEPc in phase II occurs much more slowly than laboratory-based studies have suggested. The contribution from C₃ carboxylation to CO₂ uptake during phase II was most marked in leaves which accumulated lower quantities of organic acids overnight. In exposed leaves, measurements of instantaneous during the late afternoon illustrated the transition from C₃ to C₄ carboxylation with stomata remaining open during the transition from dusk into the dark period. Uptake of CO₂ by shaded leaves during the late afternoon however appeared to be predominantly limited by decreased stomatal conductance. The short-term measurements of instantaneous were subsequently integrated over 24 h in order to predict the leaf carbon isotope ratios (_p) and to compare this with the _p measured for leaf organic material. Whilst there was close agreement between predicted and measured _p for plants sampled in the wet season, during the dry season the predicted carbon isotope ratios were 5–9 higher than the measured isotope ratios. During the annual cycle of leaf growth most carbon was fixed via the C₃ pathway although CAM clearly plays an important role in maintaining photochemical integrity in the dry season.     

Ecophysiology of the C3-CAM intermediate Clusia minor L. in Trinidad: seasonal and short-term photosynthetic characteristics of sun and shade leaves

The seasonal changes in crassulacean acid metabolism (CAM) activity in response to daily integrated photon flux density (PFD) and precipitation were compared in sun and shade leaves of the C3-CAM intermediate tree Clusia minor L. Measurements of CAM activity (H⁺) showed that maximum leaf acidity consistently occurred 4 h after dawn, suggesting that new sampling procedures need to be adopted in order to quantify CAM in Clusia species. Whilst exposed leaves responded to intermittent dry conditions, shaded leaves showed a clear induction of CAM activity as conditions became drier. The magnitude of CAM activity correlated well with daily integrated PFD, such that the extent of decarboxylation of organic acids was consistently associated with increased acidification during the subsequent dark period. Over two sampling days, both sun and shade leaves exhibited the four phases of CAM, although PEPc remained active throughout phase II with the result that 50% of the maximum leaf acidity in shade leaves was accumulated during this time. During phase III, internal CO2 supply was augmented by substantial citrate decarboxylation, in addition to malic acid. Chlorophyll fluorescence characteristics were dominated by high rates of PSII electron transport, together with an extremely high potential for thermal dissipation, such that excess light was maintained within safe limits at times of maximum PFD. Photochemical stability was maintained by matching supply and demand for internal CO2: in the morning, C3 and C4 carboxylation processes were regulated by extended PEPc activity, so that decarboxylation was delayed until temperature and light stress were highest at midday.     

The Effect of Leaf Age on Gas Exchange and Malate Accumulation in C3-CAM Plant Marrubium Frivaldszkyanum (Lamiaceae)

For the first time the expression of C₃ and CAM in the leaves of different age of Marrubium frivaldszkyanum Boiss, is reported. With increasing leaf age a typical C₃ photosynthesis pattern and high transpiration rate were found. In older leaves a shift to CAM occurred and the 24-h transpiration water loss decreased. A correlation was established between leaf area and accumulation of malate. Water loss at early stages of leaf expansion may be connected with the shift to CAM and the water economy of the whole plant.     

Light and dark CO2 fixation in Clusia uvitana and the effects of plant water status and CO2 availability

Summary In well-watered plants of Clusia uvitana, a species capable of carbon fixation by crassulacean acid metabolism (CAM), recently expanded leaves gained 5 to 13-fold more carbon during 12 h light than during 12 h dark periods. When water was withheld from the plants, daytime net CO₂ uptake strongly decreased over a period of several days, whereas there was a marked increase in nocturnal carbon gain. Photosynthetic rates in the chloroplasts were hardly affected by the water stress treatment, as demonstrated by measurements of chlorophyll a fluorescence of intact leaves, indicating efficient decarboxylation of organic acids and refixation of carbon in the light. Within a few days after rewatering, plants reverted to the original gas exchange pattern with net CO₂ uptake predominantly occurring during daytime. The reversible increase in dark CO₂ fixation was paralleled by a reversible increase in the content of phosphoenolpyruvate (PEP) carboxylase protein. In wellwatered plants, short-term changes in the degree of dark CO₂ fixation were induced by alterations in CO₂ partial pressure during light periods: a decrease from 350 to 170 bar CO₂ caused nocturnal carbon gain, measured in normal air (350 bar), to increase, whereas an increase to 700 bar CO₂, during the day, caused net dark CO₂ fixation to cease. The increased CAM activity in response to water shortage may, at least to some extent, be directly related to the reduced carbon gain during daytime.     

Properties of Phosphoenolpyruvate Carboxylase and Carbohydrate Accumulation in the C3-CAM Intermediate Sedum telephium L. Grown Under Different Light and Watering Regimes

A comparison of the activity and properties of the enzyme phosphoenolpyruvatecarboxylase (PEPC) was made for plants of Sedum telephium L.grown under low (70 µmol m^–2 s^–1) or high(500µmol m^–2 s^–1) PPFD and subjected to varyingdegrees of water stress. Under well-watered conditions onlyplants grown under high PPFD accumulated titratable acidityovernight and the extractable activity of PEPC was almost 2-foldhigher in these plants than in plants grown under low PPFD.Increasing drought stress resulted in a substantial increasein the activity of PEPC extracted both during the light anddark periods and a decrease in the sensitivity to inhibitionby malic acid. The magnitude of these changes was determinedby the severity and duration of drought and by light intensity.A comparison of the kinetic properties of PEPC from severelydroughted plants revealed that plants droughted under high PPFDhad a lower K_m for PEP than plants under low PPFD. Additionof 2·0 mol m^–3 malate resulted in an increase inthe K_m for PEP, with plants draughted under low PPFD havinga significantly higher K_m in the presence of malic acid comparedto those under high _PPFD. Response to the activator glc-6-P,which lowered the K_m for PEP, also varied between plants grownunder the two light regimes. Under well-watered conditions PEPCextracted from plants under high PPFD was more sensitive toactivation by glc-6-P than those under low PPFD. After the severedrought treatment, however, the K_m for PEP in the presence ofglc-6-P was similar for enzyme extracted from plants grown underboth light regimes. Soluble sugars and starch were depletedovernight and were both possible sources of substrate for PEPC.With increasing drought, however, the depletion of starch relativeto soluble sugars increased under both light regimes. The propertiesof PEPC and the characteristics of carbohydrate accumulation/depletionare discussed in relation to the regulation of CAM in S. telephiumgrown under different light and watering regimes. Key words: PEP carboxylase, CAM, carbohydrates, Sedum telephium     

Water Relations and CO2 Exchange of Tropical Bryophytes in a Lower Montane Rain Forest in Panama

For 6 tropical bryophytes, measurements of the diel courses in water status and net CO₂ exchange were made in a submontane tropical rain forest in Panama. In addition, the response of gas exchange to changes in photon flux density (PPFD) and thallus water content (WC) was studied under controlled conditions. Diel variation of WC was pronounced, and both low and high WC limited carbon gain considerably. Low PPFD, e.g. during rain storms, was less important in limiting CO₂ exchange. More than half of the mean diurnal carbon gain of 2.9 mg C per g thallus carbon was lost during the night as respiration. Assuming that the average 24-h carbon gain was representative for the entire year, we estimated the net annual primary productivity of the mosses and liverworts to be 45% of the initial plant carbon content.     

The Role of Dissolved Organic Carbon, Dissolved Organic Nitrogen, and Dissolved Inorganic Nitrogen in a Tropical Wet Forest Ecosystem

Although tropical wet forests play an important role in the global carbon (C) and nitrogen (N) cycles, little is known about the origin, composition, and fate of dissolved organic C (DOC) and N (DON) in these ecosystems. We quantified and characterized fluxes of DOC, DON, and dissolved inorganic N (DIN) in throughfall, litter leachate, and soil solution of an old-growth tropical wet forest to assess their contribution to C stabilization (DOC) and to N export (DON and DIN) from this ecosystem. We found that the forest canopy was a major source of DOC (232 kg C ha^–1 y^–1). Dissolved organic C fluxes decreased with soil depth from 277 kg C ha^–1 y^–1 below the litter layer to around 50 kg C kg C ha^–1 y^–1 between 0.75 and 3.5m depth. Laboratory experiments to quantify biodegradable DOC and DON and to estimate the DOC sorption capacity of the soil, combined with chemical analyses of DOC, revealed that sorption was the dominant process controlling the observed DOC profiles in the soil. This sorption of DOC by the soil matrix has probably led to large soil organic C stores, especially below the rooting zone. Dissolved N fluxes in all strata were dominated by mineral N (mainly NO₃⁻). The dominance of NO₃^– relative to the total amount nitrate of N leaching from the soil shows that NO₃^– is dominant not only in forest ecosystems receiving large anthropogenic nitrogen inputs but also in this old-growth forest ecosystem, which is not N-limited.     

The Influence of Prior Water Stress and Abscisic Acid Foliar Spraying on Stomatal Responses to CO2, IAA, ABA, and Calcium in Leaves of Solanum melongena

The influence of a water stress or foliar ABA spraying pretreatmenton stomatal responses to water loss, exogenous ABA, IAA, Ca²⁺,and CO₂ were studied using excised leaves of Solanum melongena.Both pretreatments increased stomatal sensitivity of water loss,in the presence and absence of CO₂, but decreased stomatal sensitivityto exogenous ABA. CO₂ greatly reduced the effect of exogenouslyapplied ABA. IAA decreased leaf diffusion resistance for controland ABA sprayed leaves, but did not influence the LDR of previouslywater-stressed leaves. CA²⁺ did not influence LDR of any leavesof any treatments. Key words: Water stress, stomatal response, pretreatments     

Field Measurements of Water Relations and CO2 Exchange of the Tropical,Cyanobacterial Basidiolichen Dictyonema glabratum in a Panamanian Rainforest*

Diel time courses of microclimate, hydration, and CO₂ exchange of the basidiolichen Dictyonema glabratum and its responses to experimentally changed conditions were measured for 14 days in a clearing of a premontane, tropical rainforest (Panama). Net photosynthesis (NP) was adapted to high temperatures and there was no depression of CO₂uptake at the highest thallus hydration. The presence of a CO₂ concentrating mechanism was demonstrated. Decreased NP occurred after desiccation, and apparent photon yield of CO₂ fixation was also sensitive to water loss. Natural NP was controlled by the interplay of thallus hydration and radiation. Regular, daily desiccation and dense cloud cover suppressed production. On average, 72 % of the diurnal photosynthetic gain was lost during the night, and there were days with negative carbon balance. Nevertheless, total carbon gain of the lichen was extremely high. A tentative estimation suggests that annual carbon gain amounts to 228 % of initial thallus carbon content. Because of their strong effect on net production, increased nocturnal temperatures as given at lower elevation, would lead to a negative carbon balance. This might explain the lack of abundance of this and other macrolichens in warmer, lowland rainforests.     

Effects of Powdery Mildew and Water Stress on CO(2) Exchange in Uninfected Leaves of Barley

Net photosynthesis is stimulated in third seedling leaves of barley plants whose lower two leaves are heavily infected by Erysiphe graminis f.sp. hordei Marchal. Stimulation is greater in water-stressed than in well-watered plants. In stressed, but not in well-watered plants, stimulation is associated with the maintenance of high leaf water potential and high leaf conductance. A small part of the changes in net photosynthesis is attributable to changes in respiratory metabolism in the third leaf, and other possible causes are discussed.     

Large Canopy Exchange Fluxes of Inorganic and Organic Nitrogen and Preferential Retention of Nitrogen by Epiphytes in a Tropical Lowland Rainforest

Little is known about how tropical forest canopies interact with atmospheric nitrogen deposition and how this affects the internal nutrient dynamics and the processing of external nutrient inputs. The objectives of this study therefore were (1) to investigate gross and net canopy nitrogen (N) fluxes (retention and leaching) and (2) the effect of canopy components on net canopy N retention. Tracers were applied on detached branches in a tropical wet lowland rainforest, Costa Rica. A novel ¹⁵N pool dilution method showed that gross canopy fluxes (retention and leaching) of NO₃ ^?, NH₄ ⁺, and dissolved organic nitrogen (DON) were remarkably higher than net throughfall fluxes. Gross fluxes of NH₄ ⁺ and NO₃ ^? resulted in a negligible net flux whereas DON showed net uptake by the canopy. The highest quantity of ¹⁵N was recovered in epiphytic bryophytes (16.4%) although the largest biomass fraction was made up of leaves. The study demonstrates that tracer applications allow investigation of the dynamic and complex canopy exchange processes and that epiphytic communities play a major role in solute fluxes in tree canopies and therefore in the nutrient dynamics of tropical rain forests.     

Carbon-Isotope Composition of Biochemical Fractions and the Regulation of Carbon Balance in Leaves of the C3-Crassulacean Acid Metabolism Intermediate Clusia minor L. Growing in Trinidad

Carbon-isotope ratios ([delta]13Cs) were measured for various bio-chemical fractions quantitatively extracted from naturally exposed and shaded leaves of the C3-Crassulacean acid metabolism (CAM) intermediate Clusia minor, sampled at dawn and dusk on days during the wet and dry seasons in Trinidad. As the activity of CAM increased in response to decreased availability of water and higher photon flux density, organic acids and soluble sugars were enriched in 13C by approximately 3.5 to 4%[per mille (thousand) sign] compared to plants sampled during the wet season. The induction of CAM was accompanied by a doubling in size of the reserve carbohydrate pools. Moreover, stoichiometric measurements indicated that degradation of both chloroplastic reserves and soluble sugars were necessary to supply phosphoenolpyruvate for the synthesis of organic acids at night. Results also suggest that two pools of soluble sugars exist in leaves of C. minor that perform CAM, one a vacuolar pool enriched in 13C and the second a transport pool depleted in 13C. Estimates of carbon-isotope discrimination expressed during CAM, derived from the trafficking among inorganic carbon, organic acids, and carbohydrate pools overnight, ranged from 0.9 to 3.1%[per mille (thousand) sign]. The [delta]13C of structural material did not change significantly between wet and dry seasons, indicating that most of the carbon used in growth was derived from C3 carboxylation.     

The Direct Effect of Increased CO2 on Gas Exchange and Growth of Forest Tree Species

CO₂ enrichment of the atmosphere is now well documented andits effect on the growth of world forests is being questionedby the scientific community. The direct effects of increasedCO₂ on tree species are reviewed: the different experimentalapproaches are described, as well as the principal results alreadyobtained. Short-term experiments have shown an increased photosyntheticrate, as predicted by leaf models. In longer experiments thisincrease is reduced after a few weeks or months by mechanismsthat remain to be found. Elevated CO₂ seems to decrease thedark respiration rate, but the results are still controversial.Biomass partitioning in elevated CO₂ is clearly related to themineral supply of the trees: An increase in root investmentin elevated CO₂ is related to a poor mineral status. The mineralcontent of trees grown in elevated CO2 is generally loweredcompared to controls. No general rule has yet been found forthe effect of increased CO₂ on leaf area development. The paper emphasizes large areas of ignorance: the reasons forthe different responses of different species, which may be relatedto their developmental strategies, are largely ignored. Muchexperimental effort is needed to parameterize all the physiologicalprocesses which are susceptible to change with an increase inatmospheric CO₂, leading to a change in forest tree growth. Key words: Elevated CO₂, tree, forest, photosynthesis, respiration, biomass, partitioning, mineral nutrition     

Influence of Nitrogen Supply and Water Stress on Growth and Nitrogen, Phosphorus, Potassium and Calcium Contents in Pearl Millet

Influence of supra-optimal concentrations of N on growth and accumulation of N, K, P and Ca in the shoots and roots in Pennisetum glaucum (L.) R.Br. under water stress was assessed in a pot experiment under glasshouse conditions. Thirty four-day-old plants of two lines, ICMV94133 and WCA-78, were subjected to 224, 336, or 448 mg(N) kg^–1(soil) and soil moisture 100 or 30 % of field capacity for 30 d. Increasing soil N supply decreased growth of both lines under water deficit. Nitrogen content in the shoots of both lines was not affected by supra-optimal levels of N or different watering regimes, but in contrast, the root N content was increased consistently in WCA-78 with increase in soil N content. Shoot P content increased considerably in WCA-78 at the two higher N contents, but it was significantly lower at drought stress than at well-watered treatment. In contrast, shoot or root P content in ICMV94133 did not differ under both watering regimes. Potassium content in the shoots of WCA-78 was considerably increased at the two higher N contents under drought conditions. Root K content was increased in WCA-78 at the highest N content under well-watered conditions, whereas the reverse was true in ICMV94133. Calcium content in the shoots of ICMV94133 was higher under drought stress compared with that at well-watered conditions, but such pattern was not observed in WCA-78. However, root Ca content increased in both lines with increase in N supply.     

Influence of Water Stress on Water Relations, Photosynthetic Parameters and Nitrogen Metabolism of Moth Bean Genotypes

Effects of water stress at pre-flowering stage were studied in three genotypes (RMO-40, Maru moth and CZM-32 E) of moth bean [Vigna aconitifolia (Jacq.) Marechal]. Increasing water stress progressively decreased plant water potential, leaf area, net photosynthetic rate, starch and soluble protein contents and nitrate reductase activity while contents of reducing sugars, total soluble sugar, free amino acids and free proline progressively increased. Significant genotypic differences were observed and genotype CZM-32-E displayed a better drought tolerance than other genotypes.     

The Influence of Nitrate and Chloride Uptake on Expressed Sap pH, Organic Acid Synthesis, and Potassium Accumulation in Higher Plants

The influence of NO₃⁻ uptake and reduction on ionic balance in barley seedlings (Hordeum vulgare, cv. Compana) was studied. KNO₃ and KCl treatment solutions were used for comparison of cation and anion uptake. The rate of Cl⁻ uptake was more rapid than the rate of NO₃⁻ uptake during the first 2 to 4 hours of treatment. There was an acceleration in rate of NO₃⁻ uptake after 4 hours resulting in a sustained rate of NO₃⁻ uptake which exceeded the rate of Cl⁻ uptake. The initial (2 to 4 hours) rate of K⁺ uptake appeared to be independent of the rate of anion uptake. After 4 hours the rate of K⁺ uptake was greater with the KNO₃ treatment than with the KCl treatment, and the solution pH, cell sap pH, and organic acid levels with KNO₃ increased, relative to those with the KCl treatment. When absorption experiments were conducted in darkness, K⁺ uptake from KNO₃ did not exceed K⁺ uptake from KCl. We suggest that the greater uptake and accumulation of K⁺ in NO₃⁻-treated plants resulted from (a) a more rapid, sustained uptake and transport of NO₃⁻ providing a mobile counteranion for K⁺ transport, and (b) the synthesis of organic acids in response to NO₃⁻ reduction increasing the capacity for K⁺ accumulation by providing a source of nondiffusible organic anions.