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1.  Water relations and photosynthetic capacity of two species of Calotropis in a tropical semi-arid ecosystem 
Annals of Botany  2010;107(3):397-405.
Background and Aims
Calotropis procera and Calotropis gigantea, originally from warm parts of Africa and Asia, are now pan-tropical and in ecological terms considered an indicator of overgrazed, disturbed lands; they grow successfully in dry areas. Variations in water relations, morphology and photosynthesis of the two species growing in the same habitat were studied to assess possible mechanisms of tolerance to drought and how these relate to their ecophysiological success. Also the hypothesis that their photosynthetic rate (A) under drought would be affected by stomatal and non-stomatal limitations was tested.
Water relations, gas exchange, water use efficiency (WUE), fluorescence parameters, pubescence and specific leaf area (SLA) of Calotropis procera and C. gigantea plants growing in the field were evaluated during the wet (WS) and dry (DS) seasons.
The xylem water potential (ψ) was similar in both species during the WS and DS; drought caused a 28 % decrease of ψ. In C. procera, A, stomatal conductance (gs) and carboxylation efficiency (CE) were higher in the WS with half the values of those during the DS, this species being more affected by drought than C. gigantea. A high δ13C of C. gigantea (–26·2 ‰) in the WS indicated a higher integrated WUE, in agreement with its lower gs. Leaves of C. gigantea were more pubescent than C. procera. Relative stomatal and non-stomatal limitation of A increased with drought in both species; no changes in maximum quantum yield of photosystem II (PSII; Fv/Fm) were observed. The decrease in the relative quantum yield of PSII (φPSII) and in the photochemical quenching coefficient (qP) was more pronounced in C. procera than in C. gigantea.
The photosynthetic capacity of C. procera was higher than that of C. gigantea. During the DS, A was regulated by stomatal and non-stomatal factors in a coordinated manner and drought did not cause chronic photoinhibition. A higher density of trichomes and leaf angle in C. gigantea may contribute to the maintenance of A and confer more efficient protection of photochemical activity in the DS. Ecophysiological traits such as high photosynthetic rate throughout the year even during the DS, and high WUE, highly pubescent leaves and low SLA observed in both species contribute to the establishment and growth of Calotropis in dry conditions.
PMCID: PMC3043925  PMID: 21149276
Photosynthesis; Calotropis procera; Calotropis gigantea; fluorescence; non-stomatal limitation; stomatal limitation; drought
2.  Causes of decreased photosynthetic rate and metabolic capacity in water-deficient leaf cells: a critical evaluation of mechanisms and integration of processes 
Annals of Botany  2009;103(4):561-579.
Water deficit (WD) decreases photosynthetic rate (A) via decreased stomatal conductance to CO2 (gs) and photosynthetic metabolic potential (Apot). The relative importance of gs and Apot, and how they are affected by WD, are reviewed with respect to light intensity and to experimental approaches.
Scope and Conclusions
With progressive WD, A decreases as gs falls. Under low light during growth and WD, A is stimulated by elevated CO2, showing that metabolism (Apot) is not impaired, but at high light A is not stimulated, showing inhibition. At a given intercellular CO2 concentration (Ci) A decreases, showing impaired metabolism (Apot). The Ci and probably chloroplast CO2 concentration (Cc), decreases and then increases, together with the equilibrium CO2 concentration, with greater WD. Estimation of Cc and internal (mesophyll) conductance (gi) is considered uncertain. Photosystem activity is unaffected until very severe WD, maintaining electron (e−) transport (ET) and reductant content. Low A, together with photorespiration (PR), which is maintained or decreased, provides a smaller sink for e−, causing over-energization of energy transduction. Despite increased non-photochemical quenching (NPQ), excess energy and e− result in generation of reactive oxygen species (ROS). Evidence is considered that ROS damages ATP synthase so that ATP content decreases progressively with WD. Decreased ATP limits RuBP production by the Calvin cycle and thus Apot. Rubisco activity is unlikely to determine Apot. Sucrose synthesis is limited by lack of substrate and impaired enzyme regulation. With WD, PR decreases relative to light respiration (RL), and mitochondria consume reductant and synthesise ATP. With progressing WD at low A, RL increases Ci and Cc. This review emphasises the effects of light intensity, considers techniques, and develops a qualitative model of photosynthetic metabolism under WD that explains many observations: testable hypotheses are suggested.
PMCID: PMC2707350  PMID: 19155221
Water stress; photosynthesis; photorespiration; stomata; ATP synthase; ATP; photoinhibition; electron transport; Rubisco; fluorescence; sucrose; mesophyll conductance
3.  Photosynthetic Responses of the Tropical Spiny Shrub Lycium nodosum (Solanaceae) to Drought, Soil Salinity and Saline Spray 
Annals of Botany  2003;92(6):757-765.
Water relations and photosynthetic characteristics of plants of Lycium nodosum grown under increasing water deficit (WD), saline spray (SS) or saline irrigation (SI) were studied. Plants of this perennial, deciduous shrub growing in the coastal thorn scrubs of Venezuela show succulent leaves which persist for approx. 1 month after the beginning of the dry season; leaf succulence is higher in populations closer to the sea. These observations suggested that L. nodosum is tolerant both to WD and salinity. In the glasshouse, WD caused a marked decrease in the xylem water potential (ψ), leaf osmotic potential (ψs) and relative water content (RWC) after 21 d; additionally, photosynthetic rate (A), carboxylation efficiency (CE) and stomatal conductance (gs) decreased by more than 90 %. In contrast, in plants treated for 21 d with a foliar spray with 35 ‰ NaCl or irrigation with a 10 % NaCl solution, ψ and RWC remained nearly constant, while ψs decreased by 30 %, and A, CE and gs decreased by more than 80 %. An osmotic adjustment of 0·60 (SS) and 0·94 MPa (SI) was measured. Relative stomatal and mesophyll limitations to A increased with both WD and SS, but were not determined for SI‐treated plants. No evidence of chronic photoinhibition due to any treatment was observed, since maximum quantum yield of PSII, Fv/Fm, did not change with either drought in the field or water or salinity stress in the glasshouse. Nevertheless, WD and SI treatments caused a decrease in the photochemical (qP) and an increase in the non‐photochemical (qN) quenching coefficients relative to controls; qN was unaffected by the SS treatment. The occurrence of co‐limitation of A by stomatal and non‐stomatal factors in plants of L. nodosum may be associated with the extended leaf duration under water or saline stress. Additionally, osmotic adjustment may partly explain the relative maintenance of A and gs in the SS and SI treatments and the tolerance to salinity of plants of this species in coastal habitats.
PMCID: PMC4243616  PMID: 14534200
Drought; Lycium nodosum; fluorescence; mesophyll limitation; saline stress; stomatal limitation; water deficit
4.  Operation of the Xanthophyll Cycle and Degradation of D1 Protein in the Inducible CAM plant, Talinum triangulare, under Water Deficit 
Annals of Botany  2003;92(3):393-399.
Changes in photochemical activity induced by water deficit were investigated in Talinum triangulare, an inducible CAM plant. The aim was to analyse the interactions between C3 photosynthesis, induction and activity of CAM, photosynthetic energy regulation and the mechanisms responsible for photoprotection and photoinhibition under water stress. Gas exchange, chlorophyll a fluorescence, titratable acidity, carotenoid composition and relative contents of the PSII reaction centre protein (D1) were measured. A decrease in xylem tension (ψ) from –0·14 to –0·2 MPa substantially decreased daytime net CO2 assimilation and daily carbon gain, and induced CAM, as shown by CO2 assimilation during the night and changes in titratable acidity; a further decrease in ψ decreased nocturnal acid accumulation by 60 %. Non-photochemical quenching of chlorophyll a fluorescence (NPQ) increased with water deficit, but decreased with a more severe drought (ψ below –0·2 MPa), when CAM activity was low. NPQ was lower at 0900 h (during maximum decarboxylation rates) than at 1400 h, when malate pools were depleted. Down-regulation of PSII activity related to the rise in NPQ was indicated by a smaller quantum yield of PSII photochemistry (ΦPSII) in droughted compared with watered plants. However, ΦPSII was larger at 0900 h than at 1400 h. The de-epoxidation state of the xanthophyll cycle increased with drought and was linearly related to NPQ. Intrinsic quantum yield of PSII (FV/FM) measured at dusk was also lower in severely stressed plants than in controls. Under maximum photosynthetic photon flux and high decarboxylation rates of organic acids, the D1 content in leaves of droughted plants showing maximal CAM activity was identical to the controls; increased drought decreased D1 content by more than 30 %. Predawn samples had D1 contents similar to leaves sampled at peak irradiance, with no signs of recovery after 12 h of darkness. It is concluded that under mild water stress, early induction of CAM, together with an increased energy dissipation by the xanthophyll cycle, prevents net degradation of D1 protein; when water deficit is more severe, CAM and xanthophyll cycle capacities for energy dissipation decline, and net degradation of D1 proceeds.
PMCID: PMC4257515  PMID: 12881404
CAM; D1 protein; photoinhibition; Talinum triangulare; water deficit; xanthophylls.

Results 1-4 (4)