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Dinitrogen-Fixation by Three Neotropical Agroforestry Tree Species Under Semi-Controlled Field Conditions

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Dinitrogen-Fixation by Three Neotropical Agroforestry Tree Species Under Semi-Controlled Field Conditions Plant Soil (2007) 291:199–209 DOI 10.1007/s11104-006-9186-0 ORIGINAL PAPER Dinitrogen-fixation by three neotropical agroforestry tree species under semi-controlled field conditions Humberto A. Leblanc Æ Robert L. McGraw Æ Pekka Nygren Received: 19 October 2006 / Accepted: 15 December 2006 / Published online: 2 February 2007 Ó Springer Science+Business Media B.V. 2007 Abstract Cultivating dinitrogen-fixing legume E. poeppigiana, E. fusca, and V. guatemalensis trees with crops in agroforestry is a relatively were planted in the same field using the existing common N management practice in the Neotrop- cylinders. The 15N application was repeated at the ics. The objective of this study was to assess the rate of 20 kg [N] ha–1 15 days after planting and –1 N2 fixation potential of three important Neotrop- 10 kg [N] ha was added three months after ical agroforestry tree species, Erythrina poeppigi- planting. Trees were harvested 9 months after ana, Erythrina fusca, and Inga edulis, under planting in both years. The 15N content of semi-controlled field conditions. The study was leaves, branches, stems, and roots was deter- conducted in the humid tropical climate of the mined by mass spectrometry. The percentage Caribbean coastal plain of Costa Rica. In 2002, of atmospheric N fixed out of total N (%Nf) seedlings of I. edulis and Vochysia guatemalensis was calculated based on 15N atom excess in were planted in one-meter-deep open-ended leaves or total biomass. The difference between plastic cylinders buried in soil within hedgerows the two calculation methods was insignificant of the same species. Overall tree spacing was for all species. Sixty percent of I. edulis trees 1 · 4 m to simulate a typical alley-cropping fixed N2;%Nf was 57% for the N2-fixing 15 design. The N was applied as (NH4)2SO4 at trees. Biomass production and N yield were 15 10% N atom excess 15 days after planting at the similar in N2-fixing and non-N2-fixing I. edulis. rate of 20 kg [N] ha–1. In 2003, seedlings of No obvious cause was found for why not all I. edulis trees fixed N2. All E. poeppigiana and E. fusca trees fixed N ;%N was ca. 59% and & 2 f H. A. Leblanc ( ) 64%, respectively. These data were extrapolated EARTH University, 4442-1000 San Jose, Costa Rica to typical agroforestry systems using published e-mail: [email protected] data on N recycling by the studied species. Inga edulis may recycle ca. 100 kg ha–1 a–1 of N fixed R. L. McGraw from atmosphere to soil if only 60% of trees fix Department of Agronomy, University of Missouri, –1 –1 201 Waters Hall, Columbia, N2, E. poeppigiana 60–160 kg ha a , and E. MO 65211, USA fusca ca.80kgha–1 a–1. P. Nygren Keywords Erythrina fusca Á Erythrina Department of Forest Ecology, University of Helsinki, P.O. Box 27, poeppigiana Á Inga edulis Á Humid tropics Á 00014 Helsinki, Finland 15N dilution method 123 200 Plant Soil (2007) 291:199–209 Introduction common shade tree for perennial crops in Central and South America (Sa´nchez et al. 1993; Santana While nitrogen is seldom the limiting nutrient in and Rosand 1985). Green prunings of Erythrina natural tropical ecosystems (Martinelli et al. spp. are rich in N and form an excellent green 1999; Vitousek et al. 2002), serious N deficits manure. The amount of N supplied in prunings develop in many tropical agroecosystems because (Ramı´rez 1993; Russo and Budowski 1986)or of heavy N export in crop harvest (Lal 2004). litterfall (Escalante et al. 1984; Heuveldop et al. Cultivating dinitrogen-fixing legume trees in bio- 1988; Santana and Rosand 1985)ofErythrina spp. logical interaction with crops in tropical agrofor- is greater than the N removal in the harvest of estry systems is a relatively common N coffee berries or cacao (Theobroma cacao L.) management practice in the Neotropics, espe- pods. cially in production of perennial plantation crops Although both Inga and Erythrina spp. are like coffee and cacao (Beer et al. 1998). Legume widely used in agroforestry practices and their trees are also used for fodder production in positive effect on soil organic matter and N association with tropical grasses, green manure reserves have been shown in various studies (Beer production for annual crops in alley cropping, et al. 1998; Haggar et al. 1993; Hands 1998; living hedgerow fences and fence posts, support Leblanc and McGraw 2006), early attempts to for shade-tolerant climber crops, and timber quantify N2 fixation in agroforestry resulted in production. The beneficial effects of legume trees low estimates of annual N2 fixation rates for both on tropical soils include increased soil and Erythrina (Fassbender 1987; Lindblad and Russo microbial C and N content in comparison to 1986) and Inga spp. (Roskoski 1982; Roskoski annual cropping (Mazzarino et al. 1993; Sierra and Van Kessel 1985). These data were based on et al. 2002), and long-term accumulation of soil closed-chamber acetylene reduction assays, which organic matter and N (Dulormne et al. 2003; significantly underestimates the N2 fixation rate Haggar et al. 1993). Trees of legume genera (Minchin et al. 1983, 1986), or N balance com- Erythrina L. (Fabaceae: Papilionoideae) and Inga parisons of ecologically mismatching agroforestry Mill. (Mimosoideae) are common woody compo- systems. Although these early estimates were nents in agroforestry practices throughout Neo- inadequate, they started a vigorous debate on tropics. whether the true value of legume trees in agro- The genus Inga is composed of ca. 300 forestry is N2 fixation or beneficial modifications Neotropical tree species, many of which have of microenvironment that could be associated multiple uses on local farms (Lawrence et al. with any tree species (Beer 1988; Beer et al. 1998; 1995). Fruits of 50 Inga species are eaten and Inga Budowski et al. 1984; Kass et al. 1997). fuelwood is used domestically and marketed in Recent studies based on either natural abun- many Latin American countries. At least 33 Inga dance of the stable 15N isotope (Ladha et al. 1993; species are used as shade trees for perennial crops Nygren et al. 2000; Peoples et al. 1996) or label- in agroforestry. Inga edulis Mart. may be the most ling with 15N (Liyanage et al. 1994; Peoples et al. economically important species of this genus 1996; Sta˚hl et al. 2002, 2005) have shown consid- because it grows well on acidic soils, produces erably higher N2 fixation potential of several tree edible fruits, and is a common shade and fuel- species used in agroforestry. Several Inga spp. wood tree on Latin American farms (Hands 1998; showed N2 fixation potential according to a Le´on 1998). screening based on 15N natural abundance in a The genus Erythrina is pantropical with 70 rain forest (Roggy et al. 1999) and a freshwater species in the Neotropics (Neill 1993). Erythrina swamp forest (Koponen et al. 2003) in French poeppigiana (Walpers) O.F. Cook is the most Guiana. Percentage of N derived from atmo- popular shade tree for coffee (Coffea arabica L.) sphere out of total N in I. edulis determined by in Costa Rica (Ramı´rez 1993). Erythrina fusca 15N dilution varied from 10 to 52% in a pot Lour. (syn. E. glauca), which is the only naturally experiment, depending on growth medium pantropical species of the genus (Neill 1993), is a (Leblanc et al. 2005). Thus, it appears that N2 123 Plant Soil (2007) 291:199–209 201 fixation by important Erythrina and Inga spp. temperature is 25.1°C. The soil is classified as a should be reassessed using N isotopic methods. typic dystropept. The main problem of the 15N dilution method under field conditions is achieving uniform label- N2 fixation under semi-controlled conditions ling of the soil both spatially and temporarily, and selecting a suitable non-N2-fixing reference spe- Seeds of I. edulis, E. poeppigiana,andE. fusca cies (Chalk and Ladha 1999). This problem is were collected from trees at the EARTH Uni- especially important in the case of trees that may versity campus. The fresh pulp of the seeds of I. scavenge nutrients from a horizontally extensive edulis was removed by immersing them in water area and from several soil layers. Many agrofor- at ambient temperature for 1 h. After pulp estry trees have extensive root systems and it may removal, the seeds were inoculated with brady- be impossible to find areas without the presence rhizobia by immersing for 12 h in water contain- of legume roots within tens of meters from the ing macerated root nodules taken from mature I. nearest tree (Hauser 1993; Hauser and Gichuru edulis trees together with soil taken from beneath 1994; Sierra and Nygren 2006). This means that the same trees. No pulp removal was necessary 15N label should also be applied in large areas for seeds of the Erythrina spp. but otherwise they outside the study plots in order to be sure that N2- were treated and inoculated following the same fixing legumes and reference trees take up only procedure. 15N-labelled N from the soil. An alternative to The field experiment on I. edulis was con- overcome this problem is to apply 15N within an ducted in 2002–2003 and on Erythrina spp. in area restricted by root barriers (Sta˚hl et al. 2002, 2003–2004. Seeds of I. edulis and the non-N2- 2005). The use of barriers reduces loss of the 15N fixing control, Vochysia guatemalensis Donn. Sm, label in horizontal flow of soil solutes, growing of were planted in plastic bags and the seedlings tree roots out of the labelled area, and mixing of were grown in a greenhouse for 2 months.
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