Nitrogen and Phosphorus Resorption Efficiency in Some Native Tropical Trees Planted on a Mine Spoil in Singrauli Coalfields, India

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1 Article International Journal of Environment and Bioenergy, 2014, 9(3): International Journal of Environment and Bioenergy Journal homepage: ISSN: Florida, USA Nitrogen and Phosphorus Resorption Efficiency in Some Native Tropical Trees Planted on a Mine Spoil in Singrauli Coalfields, India Arvind Singh Department of Botany, Banaras Hindu University, Varanasi , India arvindsingh_bhu@hotmail.com Article history: Received 26 May 2014, Received in revised form 25 July 2014, Accepted 31 July 2014, Published 4 August Abstract: A study was conducted to explore the nitrogen and phosphorus resorption efficiencies in eight native tropical trees planted on a mine spoil. Of the eight investigated species Acacia catechu, Albizia lebbeck, Dalbergia sissoo and Pongamia pinnata were represented by leguminous tree species while Azadirachta indica, Gmelina arborea, Tectona grandis and Terminalia bellerica were represented by non-leguminous tree species. All the tree species have shown higher nitrogen and phosphorus resorption efficiencies on nitrogen and phosphorus deficient mine spoil. Non-leguminous species had shown greater efficiency for nitrogen resorption than the leguminous species. However, no such trend emerged for phosphorus resorption efficiency between both groups of plants. Keywords: Mine Spoil, Nitrogen Resorption Efficiency, Phosphorus Resorption Efficiency, Tropical Trees 1. Introduction Nutrient resorption is a physiological process by which the plants use to withdraw nutrients from senescing leaves which are utilized later in the emergence of new structures. Nutrient resorption is a key component of nutrient conservation strategies and hence of productivity and elemental cycling

2 162 in ecosystems (Vergutz et al., 2012). The process of nutrient resorption reduces dependence of plants on soil nutrient supply. Nutrient resorption is one of the most important pathways employed by the plants to conserve the nutrients especially in nutrient deficient habitats. Several studies report that plants growing on less fertile habitats are more efficient in nutrient resorption (Stackhurski and Zimka, 1975; Boerner, 1984; Ralhan and Singh, 1987; Pugnaire and Chapin, 1993; Singh, 2004, 2007). However, other studies demonstrate greater resorption efficiency in more fertile habitats (Chapin and Kedrowski, 1983; Lajtha, 1987; Nambiar and Fife, 1987; Singh, 2005). Still some studies indicate no relationship between nutrient resorption and habitat fertility (Schlesinger et al., 1989; Chapin and Moilanen, 1991). The present study was performed on a mine spoil which is drastically disturbed and is poor in nutrients and microorganisms (Singh and Jha, 1993). Nitrogen and phosphorus are the two major limiting nutrients in mine spoils (Mays and Bengston, 1978). Therefore, the nitrogen and phosphorus resorption efficiencies were tested in some young native tropical trees planted for revegetation of mine spoil in a dry tropical environment. The main objective behind the study was to explore the effect of nitrogen and phosphorus poverty on nitrogen and phosphorus resorption efficiencies in native tropical trees planted on a mine spoil. Nutrient resorption efficiency is the percentage of the nutrient pool withdrawn from the foliage before leaf abscission. 2. Materials and Methods 2.1. Site Description The study was conducted at Jayant coal mine in Singrauli Coalfields, India (Fig. 1). The coalfields of Singrauli extend over 2200 km 2 (23 o o 12 NL; 81 o o 52 EL and elevation m msl), of which 80 km 2 lie in Sonebhadra District of Uttar Pradesh and rest in Singrauli District of Madhya Pradesh. The climate is tropical monsoonal and the year in divisible into a mild winter (November February) and hot summer (April - June) and a warm rainy season (July September). Mean monthly minimum temperature within annual cycle ranges from o C and mean monthly maximum from o C. The annual rainfall averages 1069 mm, 90% of which occurs during the period between June and September. Winter rains are negligible. The potential natural vegetation is a tropical deciduous forest Experimental Design and Methods Nursery raised 1 year old individuals of eight native tropical tree species: Acacia catechu, Albizia lebbeck, Azadirachta indica, Dalbergia sissoo, Gmelina arborea, Pongamia pinnata, Tectona

3 163 grandis and Terminalia bellerica were planted on fresh flat coal mine spoil in July 1993 at Jayant Project of Singrauli Coalfields, India. The seedlings were planted in 20 m x 20 m plots with a spacing of 2 m x 4 m. The within row spacing distance was 2 m whereas the between row spacing distance was 4 m. Three replicate plots were maintained for each tree species. The crops Pennisetum typhoides and Cajanus cajan were seeded in plantation plots annually in July from The non-legume crop P. typhoides was seeded in plantation plots of leguminous tree species while the legume crop Cajanus cajan was seeded in plantation plots of non-leguminous tree species. The texture of the spoil material was 80% sand, 10% silt and 10% clay. The total nitrogen (N) in spoil material ranged from to 0.039% and the total phosphorus (P) ranged from to %. Figure 1: Location of study site within Jayant coal mine of Singrauli Coalfields Green leaves were sampled in the first week of September 1996 and senesced leaves were sampled in the first week of December Senescent leaves were collected by lightly shaking the same branches from which green leaves were sampled. Leaf samples were collected from 5 marked individuals from middle of the crown for each species from the three replicate plots. The samples from 5 individuals were mixed from each replicate plots and brought to laboratory in polyethylene bags.

4 164 Blocks of 1cm 2 were cut, oven dried at 65 o C weighed and ground. Nitrogen was determined by microkjeldahl technique of Jackson (1958) and phosphorus was analyzed after digestion in mixture of HClO4, HNO3 and H2SO4 (1:5:1) using phosphomolybdic acid blue colour method of Jackson (1958). The nutrient mass was computed as the product obtained by multiplying dry mass (cm 2 ) of leaves by their nutrient concentration. Per cent resorption of nutrient was calculated using the formula: 100 x (nutrient mass cm -2 in mature leaf - nutrient mass cm -2 in senesced leaf) (nutrient mass cm -2 in mature leaf). 3. Results 3.1. Foliar Nitrogen Concentrations, Resorbed Nitrogen Pools and Nitrogen Resorption Efficiencies The foliar nitrogen concentration in mature and senesced leaves was greater in leguminous tree species than in non-leguminous tree species. In leguminous species the foliar nitrogen concentration in mature leaf ranged between 2.08 and 2.26% ( =2.18%) whereas it ranged between 1.17 and 1.97% ( =1.54%) in non-leguminous species (Table 1). Furthermore, the foliar nitrogen concentration in senesced leaves of leguminous species ranged between 1.14 and 1.45% ( =1.34%) while it ranged between 0.60 and 0.96% ( = 0.78%) in non-leguminous species (Table 1). Table 1. Nitrogen (N) concentration and mass in mature and senesced leaves, resorbed nitrogen pool and nitrogen resorption efficiency in native tropical trees planted on mine spoil in Singrauli Coalfields (Mean 1 SE) S. N. Tree species (Family) 1. Acacia catechu 2. Albizia lebbeck 3. Azadirachta indica (Meliaceae) 4. Dalbergia sissoo 5. Gmelina arborea (Verbenaceae) 6. Pongamia pinnata 7. Tectona grandis (Verbenaceae) 8. Terminalia bellerica (Combretaceae) Mature leaf Senesced leaf Resorbed pool (µg cm -2 ) Resorption efficiency (%) Concentration Mass Concentration Mass (%) (µg cm -2 ) (%) (µg cm -2 ) 2.26 ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± ± 0.49

5 165 No any consistent trend emerged for resorbed nitrogen pools between leguminous and nonleguminous tree species. In leguminous species the resorbed nitrogen pool ranged between and cm -2 ( = cm -2 ) whereas it ranged between and cm -2 ( = cm -2 ) in nonleguminous tree species (Table 1). The nitrogen resorption efficiency was greater in non-leguminous tree species than the leguminous tree species. In leguminous species the nitrogen resorption efficiency ranged between and 55.15% ( = 53.70%) while it ranged between and 69.19% ( = 62.37%) in nonleguminous species (Table 1) Foliar Phosphorus Concentrations, Resorbed Phosphorus Pools and Phosphorus Resorption Efficiencies No any consistent trend emerged for foliar phosphorus concentration between leguminous and non-leguminous tree species. However, the two non-leguminous tree species T. grandis and T. bellerica have lower foliar phosphorus concentration in mature leaf (Table 2). The foliar phosphorus concentration in mature leaf of leguminous species ranged between and 0.188% ( = 0.179%) while it ranged between and 0.179% ( =0.150%) in non-leguminous species. Moreover, the foliar phosphorus concentration in senesced leaf of leguminous species ranged between and 0.094% ( = 0.079%) whereas it ranged between and 0.103% ( = 0.090%) in non-leguminous tree species (Table 2). No any definite trend emerged for resorbed phosphorus pools between leguminous and nonleguminous species. The resorbed phosphorus pool in leguminous species ranged between 4.54 and 11 cm -2 ( = 7.68 cm -2 ) whereas it ranged between 5.96 and 9.14 cm -2 ( = 7.38 cm -2 ) in non-leguminous species (Table 2). No any consistent trend emerged for phosphorus resorption also between leguminous and nonleguminous tree species as two non-leguminous species A. indica and G. arborea have greater phosphorus resorption efficiency i.e. in range of leguminous species. The phosphorus resorption efficiency in leguminous species ranged between and 68.10%. ( = 66.61%) while it ranged between 42 and 65.51% ( = 53.83%) in non-leguminous species (Table 2).

6 166 Table 2. Phosphorus (P) concentration and mass in mature and senesced leaves, resorbed phosphorus pool and phosphorus resorption efficiency in native tropical trees planted on mine spoil in Singrauli Coalfields (Mean 1 SE) S. N. Tree species Mature leaf Senesced leaf Resorbed Resorption (Family) Concentration (%) Mass (µg cm -2 ) Concentration (%) Mass (µg cm -2 ) pool (µg cm -2 ) efficiency (%) 1. Acacia catechu ± ± ± ± ± ± Albizia lebbeck ± ± ± ± ± ± Azadirachta indica ± ± ± ± ± ± 0.23 (Meliaceae) 4. Dalbergia sissoo ± ± ± ± ± ± Gmelina arborea ± ± ± ± ± ± 1.15 (Verbenaceae) 6. Pongamia pinnata ± ± ± ± ± ± Tectona grandis ± ± ± ± ± ± 1.88 (Verbenaceae) 8. Terminalia bellerica (Combretaceae) ± ± ± ± ± ± Discussion The greater resorption efficiencies for both nitrogen and phosphorus in the present study in all the tree species clearly indicates that plant growing on nitrogen and phosphorus deficient habitats have greater efficiencies for nitrogen and phosphorus resorption. The result of the study was in conformity to the previous findings that plants growing on mine spoils have greater efficiencies for nitrogen and phosphorus resorption (Singh, 2004, 2005, 2007, 2011). Several workers have found that resorption process is more efficient in nutrient poor habitats than in nutrient rich habitats (Stackhurski and Zimka, 1975; Vitousek, 1982; Singh 2004, 2007). The greater nitrogen and phosphorus resorption efficiencies in the study suggest that effective resorption is a mechanism in plants to overcome the problem of nutrient poverty. In the study the tree species with lower nitrogen concentration in mature leaf have shown greater efficiencies for nitrogen resorption than those tree species having higher nitrogen concentration. Therefore the study suggests that plants with low nitrogen concentrations are more efficient in nitrogen resorption. This result conforms to previous findings by Stackhurski and Zimka (1975), Pugnaire and Chapin (1993) and Kobe et al. (2005). Contrary to the result of the present study other workers have found that plants with greater nitrogen concentration are more efficient in nitrogen resorption (Chapin and Kedrowski, 1983; Nambiar and Fife, 1987). Lajtha (1987), however, had

7 167 emphasized that plants of low nutrient status should have low resorption efficiency because most leaf nitrogen would be structurally bound and less accessible to hydrolysis and resorption. Pugnaire and Chapin (1993) found that proposed mechanism was correct but the ecological pattern was reverse. They reported that plants growing on infertile site with low nitrogen concentration had high proportion of soluble nitrogen. Similarly many other species have been reported to have a greater proportion of soluble nitrogen when grown at low nitrogen availability (Côté et al., 1989; Navari-Izzo et al., 1990). As a consequence, resorption efficiency depends on labile pool of the nutrients instead of bound in structural proteins. All the four leguminous species in present investigation have lower efficiencies for nitrogen resorption compared to non-leguminous species. This lower efficiency for nitrogen resorption in leguminous species may be owing to their nitrogen fixing attribute. Several studies suggest that nitrogen fixing plants are less efficient in nitrogen resorption (Stackhurski and Zimka, 1975; Dawson and Funk, 1981; Rodriguez - Barrueco et al., 1984; Côté and Dawson, 1986; Côté et al., 1989; Killingbeck, 1993a; Singh, 2004, 2005, 2011). On other hand side all leguminous species have shown higher efficiencies for phosphorus resorption. This affinity for phosphorus probably may be because phosphorus is being used during the process of nitrogen fixation. Several other studies also suggest that nitrogen fixing species are more efficient in phosphorus resorption (Chapin and Kedrowski 1983; Côté et al., 1989; Singh, 2004, 2005, 2011). Half or more of the maximum nitrogen and phosphorus content of a deciduous leaf is retranslocated to other plant parts before leaf abscission (Guha and Mitchell, 1966; Chapin and Kedrowski, 1983). All the tree species in the present study were deciduous thus showing high efficiencies for both nitrogen and phosphorus resorption. Greater percentage of nitrogen resorption have been reported from leaves of deciduous species than evergreen species (Ralhan and Singh, 1987; Aerts 1996; Yuan and Chen 2009b) which is an ecological need to compensate for the shorter life span of leaves and to produce heavy leaf crop annually (Chapin and Tryon, 1983). Greater phosphorus resorption efficiency has also been reported in deciduous species than in evergreen species (Huang et al., 2007). Actively growing leaves are very effective nutrient sinks (Kozlowski, 1971; Nambiar and Fife, 1987). Thus fast growing tree species are expected to be more efficient in nutrient resorption in comparison to slow growing tree species. In the present study the fast growing A. indica and G. arborea have greater nitrogen and phosphorus resorption efficiencies than slow growing T. grandis and T. bellerica.

8 Conclusion It can be concluded from the study that efficient nutrient resorption is strategies in plants to conserve nutrients when growing in low nutrient environment like mine spoil. The leguminous and non-leguminous species differ in nitrogen resorption efficiency. The non-leguminous species have a greater efficiency for nitrogen resorption while leguminous species have greater efficiency for phosphorous resorption. References Aerts, R. (1996). Nutrient resorption from senescing leaves of perennials: are there general patterns? J. Ecol., 84: Boerner, R.E.J. (1984). Foliar nutrient dynamics and nutrient use efficiency of four deciduous tree species in relation to site fertility. J. App. Ecol., 21: Chapin, F.S. III, and Kedrowski, R.A. (1983). Seasonal changes in nitrogen and phosphorus fractions and autumnal retranslocation in evergreen and deciduous taiga trees. Ecology, 64: Chapin, F.S.III, and Moilanen, L. (1991). Nutritional control over nitrogen and phosphorus resorption from Alaskan birch leaves. Ecology, 72: Chapin, F.S.III, and Tryon, P.R. (1983). Habitat and leaf habit as determinants of growth, nutrient absorption and nutrient use by Alaskan taiga forest species. Can. J. For. Res., 13(5): Côté, B., and Dawson, J.O. (1986). Autumnal changes in total nitrogen, salt extractable protein and amino acids in leaves and adjacent bark of black alder, eastern cottonwood and white bass wood. Physiol. Plant., 67: Côté, B., Vogel, C., and Dawson, J.O. (1989). Autumnal changes in tissue nitrogen of Autumn olive, Black alder, and Eastern cottonwood. Plant Soil, 118: Dawson, J.O., and Funk, D.T. (1981). Seasonal change in foliar nitrogen concentration of Alnus glutinosa. For. Sci., 27: Guha, M.M., and Mitchell, R.L. (1966). The trace and major element composition of the leaves of some deciduous trees II. Seasonal changes. Plant Soil, 24: Huang, J.,Wang, X., and Yan, E. (2007). Leaf nutrient concentration, nutrient resorption and litter decomposition in an evergreen broad-leaved forest in eastern China. For. Ecol. Manage., 239:

9 169 Jackson, M.L. (1958). Soil Chemical Analysis: Prentice-Hall, Englewood, Cliff, NJ. Killingbeck, K.T. (1993a). Inefficient nitrogen resorption in genets of the actinorhizal nitrogen fixing shrub Comptonia peregrina: physiological ineptitude or evolutionary trade off? Oecologia, 94: Kobe, R.K., Lepezyk, C.A., and Iyer, M. (2005). Resorption efficiency decreases with increasing green leaf nutrients in a global data set. Ecology, 86: Kozlowski, T.T. (1971). Growth and Development of Trees, Vol. 1 & 2, Academic Press, New York. Lajtha, K. (1987). Nutrient resorption efficiency and the response to phosphorus in desert shrub Larrea tridentata (DC.) Cov. Biogeochemistry, 4: Mays, D.A., and Bengston, G.W. (1978). Lime and fertilizer use in land reclamation in humid regions. In Schaller, F.W. and Sutton, P. (Eds). Reclamation of Drastically Disturbed Lands, CSSA and SSA, Madison, WI USA, pp Nambiar, E.K.S., and Fife, D.N. (1987). Growth and nutrient retranslocation in needles of radiata pine in relation to nitrogen supply. Ann. Bot. 60: Navari-Izzo, F., Quartacci, M.F., and Izzo, R. (1990). Water-stress induced changes in protein and free amino acids in field grown maize and sunflower. Plant Physio. Biochem. 28: Pugnaire, F.I., and Chapin, F.S.III (1993). Control over nutrient resorption from leaves of evergreen mediterranean species. Ecology, 74: Ralhan, P.K., and Singh, S.P. (1987). Dynamics of nutrient and leaf mass in central Himalyan forest trees and shrubs. Ecology, 68: Rodriguez - Barrueco, C., Miguel, C., and Subramaniam, P. (1984). Seasonal fluctuations of mineral concentration of alder (Alnus glutinosa (L.) Gaertn) from the field. Plant Soil, 78: Schlesinger, W.H., De Lucia, E.H., and Billings, W.D. (1989). Nutrient use efficiency of woody plants on contrasting soils in the Western Great Basin, Nevada. Ecology, 70: Singh, A. (2004). Effect of fertilization of N and P resorption efficiency of selected leguminous and nonleguminous tropical trees planted on mine spoil. J. Indian Inst. Sci., 84 (5): Singh, A. (2005). Influence of variation in site fertility on nitrogen and phosphorus resorption efficiency in young native tropical woody species planted on mine spoil. Indian For., 131 (11):

10 170 Singh, A. (2007). N and P retranslocation efficiency in three evergreen tree species planted in two different habitats of varying fertility status. Indian For., 133 (7): Singh, A. (2011). Influence of interplanted species on N and P resorption efficiency of companion species is mixed plantations of various species combinations raised on mine spoil. Indian J. For., 34 (1): Singh, J.S., and Jha, A.K. (1993). Restoration of degraded land: an overview. In Singh, J.S. (Ed.) Restoration of Degraded Land: Concepts and Strategies. Rastogi Publications, Meerut, India, pp Stackhurski, A., and Zimka, J.R. (1975). Methods for studying forest ecosystem: leaf area, leaf production and withdrawal of nutrients from leaves of trees. Ekol. Pols., 23: Vergutz, L., Manzoni, S., Porporato, A., Novais, R.F. and Jackson, R.B. (2012). Global resorption efficiencies and concentration of carbon and nutrients in leaves of terrestrial plants. Ecol. Monog., 82 (2): Vitousek, P.M. (1982). Nutrient cycling and nutrient use efficiency. Am. Nat. 199: Yuan, Z.Y., and Chen, H.Y.H. (2009b). Global trends in senesced-leaf nitrogen and phosphorus. Glob. Ecol. Biogeog., 18:

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