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Induced Suppression of Crassulacean Acid Metabolism Characteristics in Five Woody Weeds*

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Induced Suppression of Crassulacean Acid Metabolism Characteristics in Five Woody Weeds* Indian J. Plant Physiol., Vol. XXVII No.1 pp. 64-73 (January 1984) PARAQUAT - INDUCED SUPPRESSION OF CRASSULACEAN ACID METABOLISM CHARACTERISTICS IN FIVE WOODY WEEDS* I. MADHtTSUDANA RAO", P.M. SWAMY AND V.S. RAMA DAS Department of Botany, Sri Venkateswara University. Tirupati 517502, Andbra Pradesb, India (Revised: February 22,1983) SUMMARY Five nonsucculent semi-arid scrub species, Carissa spinarum. Maha huxifolia. Flocourtia sepiaria, Gymnosporia emorginata and Dodonaea viscosa exhibi.ted scotoactive stomatal opening, nocturnal carbon assimila­ tion and dark acidification while day time CO. uptake was predominant in Chomelia asiatica. Foliar application of 100 mg 1-1 paraquat (I, 1'­ dimethlyl-4,4'-bipyridinium ion) completely inhibited the dark uptake of CO.. Phosphoenolypyruvate (PEP) carboxylase activity was suppres­ sed in the treated plants while Ribulose bisphosphate (RuBP) carboxy­ lase was relatively unaffected. The results indicated that tbe process of dark CO. uptake was sensitive to paraquat. INTRODUCTION Crassulacean acid metabolism (CAM) has been described in 17 flowering plant families, (13 dicotyledon and 4 monocotyledon families) 1 Gymnos­ perm (Welwitchia) and in two ferns (Kluge and Ting, 1978; Ting and Gibbs, 1982). Earlier workers had noticed the occurrence of CAM in some nonsuccu­ lent leaves (Mc William, 1970; Kluge et. al., 1973). We observed the occurrence of CAM characteristics in five nonsucculent scrub species, belonging to five taxonomically diverse dicotyledon famHies. Apocynacea, Ebenacae, Flacourti­ aceae Celastraceae and Sapindacea (Rao et. al., 1979). These are unproductive woody weeds, growing in a semi-arid climate. We observed earlier that paraquat treatment could bring the reversal of the scotoactive stomatal opening, exhibited by these plants, into photoactive ·The results form a part of the Ph.D. thesis of I.M.R. approved by Sri Venkateswara Univer. sity, Tirupati. ·*Present address: Department of Plant &. Soil Biology, University of California, 108, Hilgard Hall, Berkeley, CA, 94720 USA. SUPPRESSION OF CAM BY PARAQUAT 65 resulting in rapid desiccation and death of the foliage (Rao et. al., 1977). The above investigation prompted us to undertake a detailed study on the action of paraquat on the photosynthetic metabolism of these woody weeds. MATERIALS AND METHODS The scrub formation investigated, is situated in the Tirumala foot hills, near Tirupati, India. The scrub comprises of the following non-succulent woody weeds in a decreasing order of dominance: Carisso spinarum, L; Maba buxifolia, CI; Flacourtia sepiaria, Roxb; Chomelia aSiatica, O. Kze; Gymnosporia emarginata, Laws; and Dodonaea viscosa, L. Paraquat (1. l' - dimethyl-4,4' -bipyridinium ion) at 100 mg 1-1 was applied as foliar spray at 9 a.m. The width of the stomatal apertures was measured by obtaining silicone rubber and nail polish impressions on the abaxial leaf surfaces (Rao et. al.,1977). The titratable acidity of leaves was determined following Ting and Dugger (1968). Malic acid content was estimated by recording the reduction of NAD+ in the presence of malate dehydrogenase (photometrically) at 340 nm (Hohorst. 1970). The dark and light CO2 uptake was determined by using a HCOz feed­ ing technique (Berry et. al., 1970). PEP carboxylase and RuBP carboxylase enzymes were assayed according to the methods followed by Raghavendra and Das (1977). RESULTS The morphological and physical features of leaves of six weed species, growing under natural habitat are given in Table I. A striking result of the study of the diurnal rhythms of the size of the stomatal aperture was the occure­ nce of pronounced nocturnal opening (scotoactivity) of stomata in five of the six weed species investigated namely C. spinarum, M. buxifolia, F. sepiaria, D viscosa and G. emarginata. Foliar application of paraquat caused an appreci­ able reversal of the night opening behaviour of stomata into day opening- This was more pronounced in M. buxifolia, F. sepiaria and D. viscosa (Table 1). Studies on the diurnal variations in the titratable acidity and malic acid indicated high degree of dark acidification in five of the six weed species which exhibited scotoactive stomatal opening (Figs. I and 2). Consistent with the dark acidification, malic acid also accumulated in large quantities only during night (Fig. 2). Paraquat strongly inhibited the degree of dark acidification in all the five weed species. Paraquat caused an increase in the titrable acidity and malic acid content during day in D. viscosa and G. emarginala (Figs. 1 and 2). ~~c - :;-,--­ I •. "'11 66 J. MADHUSUDANA, P.M. SWAMY AND V.S. llAMA DAS Table I : Morphological and physical features of leaves under natural habitat Plant Species Average Stomatal Leaf Ratio of Dry wei&ht leaf area frequency Thickness dry weight to percml (cml) (No. per mm') (mm) fresh weight (mg) Apocynaceae: Carissa spinarum 3.10 652 0.45 0.70 17.20 Ebenaceae: Maba buxijolill O.g() 147 0.34 0.74 31.00 Flacourtiaceae: F[llcourtia sepiarill 1.82 716 0.20 0.86 20.12 CeJastraceae: Gymnosporill emarignala 1.35 450 0.48 0.78 n.2() Sapindaceae: Dodonaea viscosa 7.10 410 0.20 0.81 14.34 Rubiaceae: Chomelill asiatica 9.11 504 0.28 0.86 57.20 The occurrence of nocturnal carbon assimilation in addition to normal day time COl! uptake was noticed in five weed species which exhibited dark acidification while there was only day time COli uptake in C. asiatica. Paraquat remarkably inhibited COli uptake both during day time and also in darkness (Fig. 3). Studies on the carbon assimilation of leaves up to an extended period of 3 days after chemicaftreatment indicated a strong inhibition of carbon dioxide uptake with paraquat treatment in all the weed species tested (Table 2). The levels of PEP carboxylase activity were high in F. sepiaria, M. buxifolia and G. emarginata. Paraquat suppressed the PEP carboxylase activity in all the weed species up to 72 h after treatment, whereas RuBP carboxylase was relatively unaffected (Table 3 and 4). DISCUSSION The present investigation holds an important implication of the occurr­ ence ofsome CAM characteristics in five non-succulent scrub species belonging to five taxonomically diverse dicotyledon families of higher plants. The occurr­ ~. Table II: Effect. or"paraquat on stomatal diurnal rhythm upto 24 hours after treatment (Stomatal openin, in microns (p.m) ) . Plant Ipecies Treatment 12 Noon 3 p.m. 6 p.m. 9 p.m. 12 Midnight 3a.m 6a.m 9 a.m. Carissa spinaru", Control 2.0 4.8 5.8 6.0 8.0 4.0 5.4 5.8 paraquat 7.4 7.6 9.4 9.8 11.2 8.8 5.6 6.8 Maba buxifolia Control 2.4 4.8 1.6 8.0 8.2 6.0 2.8 3.0 Ul ~ paraquat 8.6 8.6 9.2 9.2 9.2 7.8 2.8 8.2 ." ~ EJ Flacourtia Control 1.8 4.6 0.8. 2.0 6.8 1.2 3.0 3.2 5"" sepiarie paraquat 6.2 9.2 8.4 4.~ 4.2 2.6 2.6 5.4 Z g GymnDsporia Control 0.8 1.8 2.8 4.0 4.2 3.1l 3.0 2.4 ("l > 'marz/nata paraquat 4.2 5.2 4.6 4.2 3.0 3.2 3.0 3.8 .ii: 2.6 -<= Dodonaea Control 3.4 1.8 2.8 6.0 2.4 2.2 3.5 ." > viscosa paraquat 9.8 9.6 3.8 3.8 3.6 2.8 2.2 2.8 )1;1 ~ Chomelia Control 2.4 3.0 1.9 0.0 0.0 0.0 0.8 2.2 >~ o-j asiatica paraquat 3.6 4.2 3.0 0.0 0.0 0.0 1.0 2.8 0­ -.....I .~ 1 ~. " 68 I. MADHUSUDANA. P.M. SWAMY ANDV.S.RAMA DAS O·itO C.SPINAfWM 0·60 F' - SE PIARIA 0.30 ... ::>"" 0 -'" (lJ ~ '­ 0-20 >- 0.4-0 :::> "!: ..... VI 0 c: ... (lJ .2! 0. 0-10 .5 E 0·20 (lJ .s;;; ~ -Cl' :::; I 0 0 M·BUXIFOLIA 0·4-0­ D.VISCOSA 0·50 ......s;;; 01 -0:; 0.30 ~ .r:. '"(lJ L. .'t­ -01 cT (lJ E 0 0·4-0 EMARGINATA • 0·12 0·04 3a.1II 9a.m Sam lp.m la.m TIlliE OF THE DAY Fig. 1. Influence of paraquat on the diurnal rhythm of titratable acidity of leaves (0-0 control; /::,.-/::,. paraquat). The diurnal change in light intensity (0-0) and temperature <II-B) in the field are also shown. ence of CAM characteristics was characterized by scotoactive stomatal beha­ viour. nocturnal carbon assimilation and high activity of PEP carboxylase. resulting in the dark acidification. These species thus form an addition to the r'~........... '.. "." SuPPRESSION OF CAM BY PARAQUAT I 360 420 , C. SPINA RUM F. SEPIARIA ~o 100 420 ~. BUXIFO,LIA 340 ao ~ ..,.:x iii '3: 180 :x 111 w ~ 100 -,- .. ~oo G. Et.4ARGINATA 220 III... 0s ::a.. no 160 60L-__~~____________~ fa.m. 1 p.m. 9 p.m. 3a.rn U.m. 9a.m. lp.m. 9p.m. la.lII. 9p.rn TIM. E 0 F T 4 E 1")'4 Y Fig. 2. Influence of paraquat on the diurnal variation in malic acid content of leaves (0-0 control; f:::.-f:::. paraquat). list of nonsucculent CAM plants already known (Me Williams. 1970; Kluge et. ai., 1973). Further this expanding list of nonsueerilent CAM plants s.upports the views of Kluge et. 01. (1973) who treats CAM as cellular succulence rather than the succulence of whole plant or plant organ. 70 I. MADHUSUDANA, P.M. SWAMY AND V.S. KAMA DAS C. SPIHARUM 2·0 0 '·2 .2..VISCOSA i.4 -I .s::. N 1· 6 I E '0 N 0 0·' u en E 0 3· 2 ~.
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