Nitrogen Availability, Leaf Life Span And Nitrogen Conservation Mechanisms In Leaves Of Tropical Trees [disponibilidade De Nitrogênio, Longevidade Foliar E Mecanismos De Conservação De Nitrogênio Em Folhas De Espécies Arbóreas Tropicais]

Evergreen species of temperate regions are dominant in low-nutrient soils. This feature is attributed to more efficient mechanisms of nutrient economy. Nevertheless, the cashew (Anacardium occidentale-Anacardiaceae), a deciduous species, is native to regions in Brazil with sandy soil, whilst the annatto (Bixa orellana-Bixaceae), classified as an evergreen species native to tropical America, grows spontaneously in regions with more humid soils. Evergreens contain robust leaves that can resist adverse conditions for longer. The physical aspects of the leaves and mechanisms of nutrient economy between the two species were compared, in order to verify whether the deciduous species had more efficient mechanisms that might explain its occurrence in regions of low soil fertility. The mechanisms of nitrogen economy were also compared for the two species at available concentrations of this nutrient. The following were analysed: (i) leaf life span, (ii) physical leaf characteristics (leaf mass per area, and rupture strain), (iii) nitrogenous compounds (nitrogen, chlorophyll, and protein), (iv) nitrogen conservation mechanisms (nitrogen resorption efficiency, resorption proficiency, and use efficiency), and (v) nitrogen conservation mechanisms under different availability of this mineral. The higher values of leaf mass per area and leaf rupture strain found in A. occidentale were related to its longer leaf life span. A. occidentale showed lower concentrations of nitrogen and protein in the leaves than B. orellana. Under lower nitrogen availability, A. occidentale had higher nitrogen resorption proficiency, nitrogen use efficiency and leaf life span than B. orellana. These characteristics may contribute to the adaptation of this species to sandy soils with low nitrogen content.666812818Aerts, R., Nutrient use efficiency in evergreen and deciduous species from heathlands (1990) Oecologia, 84, pp. 391-397Aerts, R., Verhoeven, J.T.A., Whigham, D.F., Plant- mediated controls on nutrient cycling in temperate fens and bogs (1999) Ecology, 80, pp. 2170-2181Bradford, M.M., A rapid and sensitive method for the quantization of microgram quantities of protein utilizing the principle of protein-dye binding (1976) Analytical Biochemistry, 72, pp. 248-254Bremner, J.M., Nitrogen-Total (1996) Methods of Soil Analysis, pp. 1085-1121. , In: KLUTE, A.WEAVER, R.W.MICKELSON, S.H.SPARKS, D.L.BARTELS, J.M. (Ed.), Madison: SSSA, chap. 3, (Chemical Methods. Book Series, 5)Carrera, A.L., Sain, C.L., Bertiller, M.B., Patterns of nitrogen conservation in shrubs and grasses in the Patagonian Monte, Argentina (2000) Plant and Soil, 224, pp. 185-193Carrera, A.L., Bertiller, M.B., Sain, C.L., Mazzarino, M.J., Relationship between plant nitrogen conservation mechanisms and the dynamics of soil nitrogen in the arid Patagonian Monte, Argentina (2003) Plant and Soil, 255, pp. 595-604Castro-Díez, P., Puyravaud, J.P., Cornelissen, J.H.C., Leaf structure and anatomy as related to leaf mass per area variation in seedlings of a wide range of woody plant species and types (2000) Oecologia, 124, pp. 476-486Chabot, B.F., Hicks, D.J., The ecology of leaf life spans (1982) Annual Review of Ecology and Systematics, 13, pp. 229-259Cordell, S., Goldstein, G., Meinzer, F.C., Vitousek, P.M., Regulation of leaf life-span and nutrient-use efficiency of Metrosideros polymorpha trees at two extremes of a long chronosequence in Hawaii (2001) Oecologia, 127, pp. 198-206Dal Fabbro, I.M., Murase, H., Segerlind, L.J., Strain failure of apple, pear and potato tissue (1980) Summer Meeting of The American Society of Agricultural Engineers, , Saint Joseph, 1980. Proceedings. Saint JosephDiehl, P., Mazzarino, M.J., Funes, F., Fontenla, S., Gobbi, M., Ferrari, J., Nutrient conservation strategies in native Andean-patagonian forests (2003) Journal of Vegetation Science, 14, pp. 63-70Distel, R.A., Moretto, A.S., Didoné, N.G., Nutrient resorption from senescing leaves in two Stipa species native to central Argentina (2003) Austral Ecology, 28, pp. 210-215Feller, I.C., McKee, K.L., Whigham, D.F., O'Neill, J.P., Nitrogen vs. phosphorus limitation across an ecotonal gradient in a mangrove forest (2003) Biogeochemistry, 62, pp. 145-175Givnish, T.J., Adaptive significance of evergreen vs. deciduous leaves: Solving the triple paradox (2002) Silva Fenica, 36, pp. 703-743Haddad, C.R.B., Lemos, D.P., Mazzafera, P., Leaf life span and nitrogen in semideciduous forest tree species (Croton priscus and Hymenaea courbaril) (2004) Scientia Agricola, 61, pp. 462-465Hikosaka, K., Leaf canopy as a dynamic system: Ecophysiology and optimality in leaf turnover (2005) Annals of Botany, 95, pp. 521-533Hiscox, J.D., Israelstam, G.F., A method for the extraction of chlorophyll from leaf tissue without maceration (1979) Canadian Journal of Botany, 57, pp. 1332-1334Hoagland, D.R., Arnon, D.I., (1938) The Water Culture Method For Growing Plants Without Soil, , Davis: University of California Agricultural Experimental Station, (Bulletin, 347)Ishida, A., Diloksumoun, S., Ladpala, P., Staporn, D., Panuthai, S., Gamo, M., Yazaki, K., Puangchit, L., Contrasting seasonal leaf habits of canopy trees between tropical dry-deciduous and evergreen forests in Thailand (2006) Tree Physiology, 26, pp. 643-656Killingbeck, K.T., Nutrients in senesced leaves: Keys to the search for potential resorption and resorption proficiency (1996) Ecology, 77, pp. 1716-1727Lal, C.B., Anapurna, C., Raghubanshi, A.S., Singh, J.S., Effect of leaf habit and soil type on nutrient resorption and conservation in woody species of a dry tropical environment (2001) Canadian Journal of Botany, 79, pp. 1066-1075Lichtenthaler, B.K., Wellburn, A.R., Determination of total carotenoids and chlorophylls a and b of leaf extracts in different solvents (1983) Biochemical Society Transactions, 11, pp. 591-592Lima, A.L.S., Zanella, F., Schiavinato, M.A., Haddad, C.R.B., Nitrogenous compounds, phenolic compounds and physical aspects of leaves: Comparison of deciduous and semideciduous arboreal legumes (2006) Scientia Agricola, 63, pp. 40-45Lima, A.L.S., Zanella, F., Schiavinato, M.A., Haddad, C.R.B., N availability and mechanisms of N conservation in deciduous and semideciduous tropical forest legume trees (2006) Acta Botanica Brasilica, 20, pp. 625-632Lorenzi, H., (2000) Árvores brasileiras, p. 352. , 3 ed. Nova Odessa: Editora PlantarumMilla, R., Castro-Diez, P., Maestro-Martinez, M., Montserrat-Marti, G., Environmental constraints on phenology and internal nutrient cycling in the Mediterranean winter-deciduous shrub Amelanchier ovalis Medicus (2005) Plant Biology, 7, pp. 182-189Montgomery, D.C., (1992) Introduction to Linear Regression Analysis, p. 544. , 2 ed. New York: John Wiley, (Wiley Series in Probability and Mathematical Statistics)Morellato, L.P.C., Leitão-Filho, H.F., Reproductive phenology of climbers in a southeastern Brazilian forest (1996) Biotropica, 28, pp. 180-191Morellato, L.P.C., Talora, D.C., Takahasi, A., Bencke, C.C., Romera, E.C., Ziparro, V.B., Phenology of Atlantic rain forest trees: A comparative study (2000) Biotropica, 32, pp. 811-823Nelson, D.W., Sommers, L.E., Determination of the total nitrogen in plant material (1973) Agronomy Journal, 65, pp. 109-112Oleksyn, J., Reich, P.B., Zytkowiak, R., Karolewski, P., Tjoelker, M.G., Nutrient conservation increases with latitude of origin in European Pinus sylvestris populations (2003) Oecologia, 136, pp. 220-235Piedade, S., Fabbro, I.M.D., Vidal, B.C., Gamga, R., Ensaio uniaxial de tração de tendões de músculos grácil e semitendinoso (2001) Acta Ortopédica Brasileira, 9, pp. 5-11Prior, L.D., Eamus, D., Bowman, D.M.J.S., Leaf attributes in the seasonally dry tropics: A comparison of four habitats in northern Australia (2003) Functional Ecology, 17, pp. 504-515Pugnaire, F.I., Chapin, F.S., Controls over nutrient resorption from leaves of evergreen Mediterranean species (1993) Ecology, 74, pp. 124-129Pujol, B., Salager, J.-L., Beltran, M., Bousquet, S., McKey, D., Photosynthesis and leaf structure in domesticated cassava (Euphorbiaceae) and close wild relative: Have leaf photosynthetic parameters evolved under domestication? (2008) Biotropica, 40, pp. 305-312Read, J., Edwards, C., Sanson, G.D., Aranwela, N., Relationships between sclerophylly, leaf biomechanical properties and leaf anatomy in some Australian heath and forest species (2000) Plant Biosystems, 134, pp. 261-277Reich, P.B., Uhl, C., Walters, M.B., Ellsworth, D.S., Leaf lifespan as a determinant of leaf structure and function among 23 tree species in Amazonian forest communities (1991) Oecologia, 86, pp. 16-24Reich, P.B., Walters, M.B., Ellsworth, D.S., Leaf life-span in relation to leaf, plant, and stand characteristics among diverse ecosystems (1992) Ecological Monographs, 62, pp. 365-392Richardson, S.J., Peltzer, D.A., Allen, R.B., McGlone, M.S., Parfitt, R.L., Rapid development of phosphorus limitation in temperate rainforest along the Franz Josef soil chronosequence (2004) Oecologia, 139, pp. 267-276Sakata, T., Nakano, T., Iino, T., Yokoi, Y., Contrastive seasonal changes in ecophysiological traits of leaves of two perennial Polygonaceae herb species differing in leaf longevity and altitudinal distribution (2006) Ecological Research, 21, pp. 633-640(1990) Sas/stat User's Guide: Version 8, , SAS INSTITUTE, Cary: SAS Institute, CD-ROMSmart, C.M., Gene expression during leaf senescence (1994) New Phytologist, 126, pp. 419-448(2003) Keys to Soil Taxonomy, p. 331. , SOIL SURVEY STAFF, 9 ed. Washington, D.C.: Natural Resources Conservation Service, US Department of AgricultureTakashima, T., Hikosaka, K., Hirose, T., Photosynthesis or persistence: Nitrogen allocation in leaves of evergreen and deciduous Quercus species (2004) Plant and Cell Environment, 27, pp. 1047-1054Vitousek, P.M., Litterfall, nutrient cycling and nutrient limitation in tropical forests (1982) Ecology, 65, pp. 285-298Westoby, M., Falster, D.S., Moles, T., Vesk, P.A., Wright, I.J., Plant ecological strategies: Some leading dimensions of variation between species (2002) Annual Review of Ecology and Systematics, 33, pp. 125-159Wright, I.J., Westoby, M., Reich, P.B., Convergence towards higher leaf mass per area in dry and nutrient-poor habitats has different consequences for leaf lifespan (2002) Journal of Ecology, 90, pp. 534-543Wright, I.J., Westoby, M., Nutrient conservation, resorption and lifespan: Leaf traits of Autralian sclerophyll species (2003) Functional Ecology, 17, pp. 10-1

The Effect Of 24-epibrassinolide And Clotrimazole On The Adaptation Of Cajanus Cajan (l.) Millsp. To Salinity

The objective of this study was to evaluate the effects of one of brassinosteroids (24-epibrassinolide) and clotrimazole, (an inhibitor of brassinosteroid synthesis) on plant growth parameters, parameters related to leaf gas exchange (photosynthetic and transpiration rates; stomatal conductance; water use efficiency), photosynthetic pigment content and osmolyte (sugars and proline) content in Cajanus cajan exposed to salinity. Salt stress-caused by NaCl treatment-affected values of all parameters analyzed. The effects were ameliorated by 24-epibrassinolide and intensified by clotrimazole. The hormone increased fresh mass of the plant, shoot dry mass, leaf area, water content of leaves and roots, photosynthetic pigments, sugar concentration, photosynthetic rate, and water use efficiency. The effects of hormone were less evident in the absence of salt. However, under this condition the application of clotrimazole affected the values of parameters studied, indicating the importance of brassinosteroid synthesis for the normal development of the plant. © 2011 Franciszek Górski Institute of Plant Physiology, Polish Academy of Sciences, Kraków.33518871896Akintayo, E.T., Oshodi, A.A., Esuoso, K.O., Effects of NaCl, ionic strength and pH on the foaming and gelation of pigeon pea (Cajanus cajan) protein concentrates (1999) Food Chem, 66, pp. 51-56Ali, B., Hayat, S., Fariduddin, Q., Ahmad, A., 24-Epibrassinolide protects against the stress generated by salinity and nickel in Brassica juncea (2008) Chemosphere, 72, pp. 1387-1392Ali, Q., Athar, H.U.R., Ashraf, M., Modulation of growth, photosynthetic capacity and water relations in salt stressed wheat plants by exogenously applied 24-epibrassinolide (2008) Plant Growth Regul, 56, pp. 107-116Amzallag, G.N., Vaismam, J., Influence of brassinosteroids on initiation of the root gravitropic response in Pisum sativum seedlings (2006) Biol Plant, 50, pp. 283-286Ashraf, M., Harris, P.J.C., Potential biochemical indicators of salinity tolerance in plants (2004) Plant Sci, 166, pp. 3-16Babita, M., Maheswari, M., Rao, L.M., Shanker, A.K., Rao, D.G., Osmotic adjustment drought tolerance and yield in castor (Ricinus communis L.) hybrids (2010) Environ Exp Bot, 69, pp. 243-249Bates, L.S., Waldren, R.P., Rapid determination of free proline for water-stress studies (1973) Plant Soil, 39, pp. 205-207Bieleski, L.R., Turner, N.A., Separation and estimation of amino acids in crude plant extracts by thin-layer electrophoresis and chromatography (1966) Anal Biochm, 17, pp. 278-293Carillo, P., Mastrolonardo, G., Nacca, F., Fuggi, A., Nitrate reductase in durum wheat seedlings as affected by nitrate nutrition and salinity (2005) Funct Plant Biol, 32, pp. 209-219Chen, Z., Cuin, T.A., Zhou, M., Twomey, A., Naidu, B.P., Shiabala, S., Compatible solute accumulation and stress-mitigating effects in barley genotypes contrasting in their salt tolerance (2007) J Exp Bot, 58, pp. 4245-4255Corzo, O., Fuentes, A., Moisture sorption isotherms and modeling for pre-cooked flours of pigeon pea (Cajanus cajans L. millsp.) and lima bean (Canavalia ensiformis) (2004) J Food Engin, 65, pp. 443-448Dubois, M., Gilles, K.A., Hamilton, J.K., Rebers, P.A., Smith, F., Colorimetric method for determination of sugars and related substances (1956) Anal Chem, 28, pp. 350-356(2000) Global Network On Integrated Soil Management For Sustainable Use of Salt Affected Soils, , http://www.fao.org/ag/AGL/agll/spush/intro.htm, FAO, (accessed 22 December 2008)Flowers, T.J., Hajibagheri, M.A., Salinity tolerance in Hordeum vulgare: ion concentrations in root cells of cultivars differing in salt tolerance (2001) Plant Soil, 231, pp. 1-9Fujioka, S., Sakurai, A., Brassinosteroids (1997) Nat Prod Rep, 14, pp. 1-10Handel, E.V., Direct microdetermination of sucrose (1968) Anal Biochem, 22, pp. 280-283Hasegawa, P.M., Bressan, R.A., Zhu, J.K., Bohnert, H.J., Plant cellular and molecular responses to high salinity (2000) Annu Rev Plant Physiol Plant Mol Biol, 51, pp. 463-499Haubrick, L.L., Torsethaugen, G., Assmann, S.M., Effect of 24-epibrassinolide, alone and in concert with abscisic acid, on control of stomatal aperture and potassium currents of Viciafaba guard cell protoplasts (2006) Physiol Plant, 128, pp. 134-143Hiscox, J.D., Israelstam, G.F.A., A method for the extraction of chlorophyll from leaf tissue without maceration (1979) Can J Bot, 57, pp. 1332-1334Hoagland, D.R., Arnon, D.I., The water culture method for growing plants without soil (1938) Calif Agric Exp Sta Circ, p. 347Janeczko, A., Gullner, G., Skoczowski, A., Dubert, F., Barna, B.E., Effects of brassinosteroid infiltration prior to cold treatment on ion leakage and pigment contents in rape leaves (2007) Biol Plant, 51, pp. 355-358Kagale, S., Divi, U.K., Krochko, J.E., Keller, W.A., Krishna, P., Brassinosteroid confers tolerance in Arabidopsis thaliana and Brassinca napus to a range of abiotic stresses (2007) Planta, 225, pp. 353-364Khripach, V.A., Zhabinskii, V.N., Groot, A.E., (1999) Brassinosteroids-a New Class of Plant Hormones, , Academic Press, San DiegoKim, T.W., Chang, S.C., Lee, J.S., Takatsuto, S., Yokota, T., Kim, S.K., Novel biosynthetic pathway of castasterone from cholesterol in tomato (2004) Plant Physiol, 135, pp. 1231-1242Kim, Y.S., Joo, S.H., Hwang, J.Y., Park, C.H., Kim, S.K., Characterization of C29-brassinosteroids and their biosynthetic precursors in immature seeds of Phaseolus vulgaris (2006) Bull Korean Chem Soc, 27, pp. 1117-1118Lichtenthaler, B.K., Welburn, A.R., Determination of total carotenoids and chlorophylls a and b of leaf extracts in different solvents (1983) Biochem Soc Transact, 11, pp. 591-592Liska, A.J., Shevchenko, A., Pick, U., Katz, A., Enhanced photosynthesis and redox energy production contribute to salinity tolerance in Dunaliella as revealed by homology-based proteomics (2004) Plant Physiol, 136, pp. 2806-2817Lisso, J., Altmann, T., Mussig, C., Metabolic changes in fruits of the tomato d(x) mutant (2006) Phytochemistry, 67, pp. 2232-2238Morillon, R., Catterou, M., Sangwan, R.S., Sangwan, B.S., Lassalles, J.P., 24-epibrassinolide may control aquaporin activities in Arabidopsis thaliana (2001) Planta, 212, pp. 199-204Nakajima, N., Toyama, S., Effects of epibrassinolide on sugar transport and allocation to the epicotyl in cucumber seedlings (1999) Plant Prod Sci, 2, pp. 165-171Ogweno, J.O., Song, X.S., Shi, K., Hu, W.H., Mao, W.H., Zhou, Y.H., Yu, J.Q., Nogués, S., Brassinosteroids alleviate heat-induced inhibition of photosynthesis by increasing carboxylation efficiency and enhancing antioxidant systems in Lycopersicum esculentum (2008) J Plant Growth Regul, 27, pp. 49-57Ozdemir, F., Bor, M., Demiral, T., Türkan, I., Effects of 24-epibrassinolide on seed germination, seedling growth, lipid peroxidation proline content and antioxidative system of rice (Oryza sativa L.) under salinity stress (2004) Plant Growth Regul, 42, pp. 203-211Paul, M.J., Foyer, C.H., Sink regulation of photosynthesis (2001) J Exp Bot, 52, pp. 1383-1400Reggiani, R., Nebuloni, M., Mattana, M., Brambilla, I., Anaerobic accumulation of amino acids in rice roots: role of the glutamine synthetase/glutamate synthase cycle (2000) Amino Acids, 18, pp. 207-217Sasse, J.M., Physiological actions of brassinosteroids: an update (2003) J Plant Growth Regul, 22, pp. 276-288Shahbaz, M., Ashraf, M., Athar, H.U.R., Does exogenous application of 24-epibrassinolide ameliorate salt induced growth inhibition in wheat (Triticum aestivum L.)? (2008) Plant Growth Regul, 55, pp. 51-64Sharifi, M., Ghorbanli, M., Ebrahimzadeh, H., Improved growth of salinity-stressed soybean after inoculation with salt pre-treated mycorrhizal fungi (2007) J Plant Physiol, 164, pp. 1144-1151Tester, M., Davenport, R., Na+ tolerance and Na + transport in higher plants (2003) Ann Bot, 91, pp. 503-527Xia, X.J., Wang, Y.J., Zhou, Y.H., Tao, Y., Mao, W.H., Shi, K., Asami, T., Yu, J.Q., Reactive oxygen species are involved in brassinosteroid-induced stress tolerance in cucumber (2009) Plant Physiol, 150, pp. 801-814Yazici, I., Tuerkan, I., Sekman, A.H., Demiral, T., Salinity tolerance of purslane (Portulaca oleraceae L.) is achieved by enhanced antioxidative system, lower level of lipid peroxidation and proline accumulation (2007) Environ Exp Bot, 61, pp. 49-57Yu, J.Q., Huang, L.F., Hu, W.H., Zhou, Y.H., Mão, W.H., Ye, S.F., Nogués, S., A role for brassinosteroid in the regulation of photosynthesis in Cucumis sativus (2004) J Exp Bot, 55, pp. 1135-114