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Diurnal Changes of the Titratable Acidity in a New CAM Plant, Sedum Caucasıcum Leaves

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Diurnal Changes of the Titratable Acidity in a New CAM Plant, Sedum Caucasıcum Leaves Proceedings of ANAS (Biological and Medical Sciences), vol. 72, No 3, p. 26-31 (2017) Diurnal Changes of the Titratable Acidity in A New CAM Plant, Sedum caucasıcum Leaves S.M. Bayramov*, T.Y. Orujova, U.A. Gurbanova, N.M. Guliyev Institute of Molecular Biology & Biotechnologies, Azerbaijan National Academy of Sciences, 2A Matbuat Avenue, Baku AZ1073, Azerbaijan;*E-mail: [email protected] Determination of the photosynthetic pathways of the plants in the studied areas would be very useful for evaluation of the modern vegetation and prediction of changes. Crassulacean acid metabolism (CAM) is a specialized mode of photosynthesis that features nocturnal CO2 uptake, facilitates increased water-use efficiency. A large variation in CAM has been found within the genus Sedum. C3 species, CAM constitutive species and CAM inducible species under water stress or salinity have been recognized. Diurnal acidity cycle in Sedium caucasicum species which is considered endemic in Caucases occurs similar to that of in CAM plants regardless if they are grown in a greenhouse or naturally. The obtained results confirm that Sedum caucasicum is an obligate CAM plant. Keywords: CAM photosynthesis, Sedum caucasicum, titritable acidity, photochemical efficiency INTRODUCTION fixation and ∆H+ but no nocturnal stomatal aper- ture; and (d) idling, with small ∆H+ and stomatal Crassulacean acid metabolism (CAM) is a wa- closure during the entire day and night in severely ter-conserving mode of photosynthesis and one of stressed plants (Ana Herrera, 2009). On the basis of three photosynthetic pathways which occurs in 7% the magnitude of ∆H+ obligate CAM, can be strong of vascular plant species (Winter and Smith, 1996), and weak CAM form (Winter and Holtum, 2002; many of which dominate the plant biomass of arid, Silvera et al., 2005; Holtum et al., 2007). In faculta- marginal regions of the world. CAM is a modifica- tive species, CAM may be induced by factors such tion of the basic C3 pathway and represents CO2- as drought (Borland and Griffiths, 1990; Herrera et concentrating mechanisms that elevate CO2 around al., 1991; Olivares et al., 1984), salinity (Winter Rubisco and suppress photorespiration. This is and von Willert, 1972), photoperiod (Brulfert et al., achieved in two principal phases separated in time. 1988), high photosynthetic photon flux (PPF) At night, atmospheric CO2 is incorporated by phos- (Maxwell, 2002), nitrogen deficiency (Ota, 1988) phoenolpyruvate carboxylase (PEPC) via ox- and phosphorus deficiency (Paul and Cockburn, aloacetate into malic acid, which accumulates in the 1990), among others. large vacuoles of chloroplast-containing mesophyll Sedum is a large genus of flowering plants in cells. During the following light period, malic acid the family Crassulaceae, members of which are is released from the vacuoles and decarboxylated, commonly known as stonecrops. The genus has and the CO2 thus liberated is refixed by Rubisco been described as containing up to 600 spe- and reduced in the Calvin cycle (Osmond, 1978; cies updated to 470. There are 22 species of the ge- Winter & Smith, 1996). Decarboxylation of malate nus Sedium in the flora of Azerbaijan. Sedum cau- generates high intercellular CO2 and is associated casicum (Grossh.) Hylotelephium caucasicum, with stomatal closure, minimizing water loss in the grows on dry stony and limestone slopes in mixed middle of the day when evaporative demand is deciduous forests of the Caucasus (Figure 1). The highest. The net result of CAM is an improvement plant is herbaceous, perennial, with the aerial part in water use efficiency generally 6-fold higher than completely dying for the winter. Roots are thick- for C3 plants and 3-fold higher than for C4 plants ened, fusiform. Stems are simple, among several, under comparable conditions (Nobel 1996). raised, strong, straight, green or dark purple. Due to CAM may operate in different modes: (1) ob- the arrangement of leaves plants can use solar ener- ligate CAM, with high nocturnal acid accumulation gy with maximum efficiency. (∆H+) and CO2 fixation; (2) facultative or inducible Work presented in this paper aimed at investi- CAM, also known as C3-CAM, with a C3 form of gating the presence of CAM in S.caucasicum Inves- + CO2 fixation and nil (∆H ) in the non-induced state, tigations involved field and greeenhouse studies of and small nocturnal CO2 fixation and ∆H+ in the diurnal changes in titratable acidity. induced state; (c) CAM-cycling, with daytime CO2 26 Diurnal Changes Of The Titratable Acidity In A New CAM Plant Figure 1. S.caucasicum plants at anthesis under natural conditions. MATERIAL AND METHODS CAM at these 4 various stages of drought was as- sessed by measuring the overnight accumulation of The study site was located at the Lesser Cau- titratable acidity. Diurnal acidity changes were ob- casus Mountain (Tovuz region) in the north-west of served under natural as well as controlled condi- Azerbaijan (40°59′31.92″N, 45°37′44.04″E) and at tion. The quantity of titratable acid which was very the Main Caucasus Montain (Ismayilli region) in high in the early morning, decreased during the day the north-eastern part of Azerbaijan (40°51′11.16″ and reached a minimum value at 1700-1800. At the N, 48°23′35.16″E). early hours of night (at 2200) average acidity was Plant material. Mature Sedum caucasicum 1/3 of the acidity observed in the morning hours. plants were used as plant material. Total titratable acidity in flowers and green stems Titratable acidity was determined in leaves of under natural conditions was less than in leaves and Sedum caucasicum at different times of. Extracts it did not change pronouncedly during the day. De- were prepared by grinding 0.5 gr frozen tissue in spite the fact that, the quantity of titratable acidity distilled water and titrating them to a pH 7.0 end was less in juvenile leaves compared with young and point with 0.01 N NaOH (FJ.Castillo,1996) middle-aged leaves of the plants studied under both Determination of RWC. To obtain the leaf natural conditions, cyclic changes of acidity occurred RWC, fresh mass, mass after rehydration (equi- in juvenile leaves contrary to flowers and green librated at 100% RH, at 4°C in distilled water for stems of young and middle-aged ones (Table 1). 24 h in the dark) and dry mass were determined Acidity of old leaves, completed their vegeta- (Castillo,1996). tion, was relatively less than that of young and mid- RWC was calculated as follows: dle-aged leaves. It suggests that the rate of photo- RWC=(FW-DW)/(SW-DW)x100 synthesis and the intensity of metabolism are positi- where FW is the fresh weight, DW is the dry vely correlated with the titritable acidity. The quanti- weight, and SW is the mass after rehydration. ty of titritable acidity in S.caucasicum is much less Fluorescence induction was measured with in- in initial leaves, flowers (mature and immature) and tact leaves, at ambient temperature with a portable stems compared with leaves and this amount remains fluorimeter (Mini-PAM fluorometer, WALZ, Ger- unchanged during the day. This shows that CAM many). Fluorescence parameters were as follows: F0, photosynthesis does not occur in flowers, stem and initial fluorescence; Fm, maximum fluorescence, Fv, initial leaves of S.caucasicum. However, quantity of variable fluorescence (Fm-Fo; Bolhar-Norden-kampf acidity and its diurnal changes have similar patterns et al., 1989). Measured chlorophyll fluores-cence in young and middle-aged leaves. This confirms that parameters included Fv/Fm, and Φ PSII, reflecting there is no significant difference in the rate of CAM the efficiency of PSII antenna and the quantum yield photosynthesis between these leaves. of PSII, respectively. The titritable acidity in different aged leaves of the S.caucasicum plant cultivated in controlled RESULTS AND DISCUSSION conditions was lower in older leaves than in young and middle aged plants (Table 2). Thus, in young CAM in Sedum caucasicum L. (Crassulaceae) and middle aged leaves the quantity of titratable was analysed by studying diurnal acidity changes were ΔH+=120.0±0.2 μmol g-1 and ΔH+=124.0±0.2 depending on environmental factors both in natural μmol g-1 respectively, and ΔH+=106.0±0.15 μmol g-1 habitat of the plant and in phytotron. The extent of in older leaves. 27 Bayramov et al. Table 1. Diurnal changes in the titritable acidity in above-ground organs and leaf tiers of S.caucasicum under natural conditions. Plant organs 800 1200 1600 2000 2400 Initial leaf 44.0±0.2 42.0±0.1 38.0±0.2 40.0±0.1 44.0±0.1 1 leaf 114.0±0.1 46.0±0.02 26.0±0.01 20.0±0. 2 60.0±0.2 2 leaves 122.0±0.2 76.0±0.02 26.0±0.01 22.0±0. 2 70.0±0.3 3 leaves 126.0±0.25 88.0±0.03 24.0±0.01 20.0±0. 1 70.0±0.3 4 leaves 130.0±0.3 96.0±0.03 24.0±0.01 20.0±0.2 65.0±0.3 5 leaves 124.0±0.3 88.0±0.03 24.0±0.01 20.0±0.2 60.0±0.3 6 leaves 124.0±0.3 60.0±0.02 22.0±0.01 18.0±0.3 60.0±0.3 Flower 40.0±0.01 38.0±0.01 38.0±0.01 40.0±0.1 40.0±0.1 Stem 36.0±0.01 36.0±0.01 36.0±0.01 40.0±0.1 40.0±0.1 Note: Quantity of titritable acidity is expressed as µmol g-1 fresh weight Table 2.
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