Grain Legumes and Green Manures for Soil Fertility in ... - cimmyt
Grain Legumes and Green Manures for Soil Fertility in ... - cimmyt
Grain Legumes and Green Manures for Soil Fertility in ... - cimmyt
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Arbuscular mycorrhizal fungi (AMF) colonize roots<br />
of about 80% of terrestrial plant species <strong>and</strong> can <strong>in</strong>crease<br />
plant P uptake by <strong>in</strong>creas<strong>in</strong>g the surface uptake<br />
area (Koide 1991). Synergistic effects on legumes<br />
are frequently seen when both symbionts (the<br />
rhizobia <strong>and</strong> the fungi) are present (Goss <strong>and</strong> de<br />
Varennes, 2002; Sang<strong>in</strong>ga et ai., 1999; Fitter <strong>and</strong> Garbaye,<br />
1994). Both nodule number <strong>and</strong> dry weight<br />
usually <strong>in</strong>crease after mycorrhizal colonization<br />
(Reddy <strong>and</strong> Bagyraj, 1991), which is often expla<strong>in</strong>ed'<br />
by <strong>in</strong>creased P uptake by the fungi. While mycorrhizal<br />
fungi are components of most natural ecosystems,<br />
their abundance <strong>and</strong> efficacy can be severely<br />
retarded by common agricultural practices such as<br />
fallow<strong>in</strong>g, soil disturbance through till<strong>in</strong>g <strong>and</strong> weed<br />
management, <strong>and</strong> prolonged cultivation of non-host<br />
plants (Boswell et al. 1998; Kabir et al., 1997; Douds<br />
et al. 1995; Har<strong>in</strong>ikumar <strong>and</strong> Bagyraj, 1989).<br />
The objective of our work was to underst<strong>and</strong> the<br />
role of AMF <strong>for</strong> legume per<strong>for</strong>mance <strong>in</strong> subsistence<br />
farmers' fields, <strong>and</strong> to determ<strong>in</strong>e if an enhanced<br />
AMF abundance can promote nodulation <strong>and</strong> N2<br />
fixation. In this paper we discuss the results of two<br />
pot experiments. In Experiment 1, the effect of an<br />
altered AMF abundance on nodulation <strong>and</strong> shoot N<br />
content on groundnut was determ<strong>in</strong>ed by enhanc<strong>in</strong>g<br />
the AMF abundance through an <strong>in</strong>oculation<br />
with a common AMF, or by reduc<strong>in</strong>g the <strong>in</strong>digenous<br />
AMF abundance through a fungicide application.<br />
In Experiment 2, the abundance of <strong>in</strong>digenous<br />
fungi was enhanced <strong>and</strong> the effects on nodule number,<br />
nodule mass <strong>and</strong> shoot N content were determ<strong>in</strong>ed<br />
on lab lab bean. Groundnut was chosen s<strong>in</strong>ce<br />
it is a common legume grown by the subsistence<br />
farmer <strong>in</strong> Zimbabwe, <strong>and</strong> lablab bean because it is a<br />
green manure crop <strong>and</strong> there<strong>for</strong>e has a higher potential<br />
to improve soil fertility.<br />
Material <strong>and</strong> methods<br />
General<br />
The soil used <strong>in</strong> both Experiment 1 <strong>and</strong> Experiment<br />
2 was a Tsholotsho luvisol (from Simeon Moyo's<br />
farm) where legume P limitations had been demonstrated<br />
previously. The pH of the soil was 6.2 (1:2 V<br />
soil: V water), <strong>and</strong> available P was 1.2 ppm (Olsen).<br />
<strong>Soil</strong> was collected to a depth of 15 cm <strong>in</strong> December<br />
1999, <strong>for</strong> Experiment 1, <strong>in</strong> April 2001 <strong>for</strong> the <strong>in</strong>oculum<br />
production part of Experiment 2, <strong>and</strong> <strong>in</strong> November<br />
2001, <strong>for</strong> the <strong>in</strong>oculation part of Experiment<br />
2. For Experiment 1 the soil was collected r<strong>and</strong>omly<br />
from the field where plants were currently grown<br />
<strong>and</strong> no fertilizers had been added, <strong>and</strong> <strong>for</strong> Experiment<br />
2 from areas where maize had been grown the<br />
previous year without fertilizer additions.<br />
Experiment 1<br />
Groundnut (var. Falcon) was planted on December<br />
28 1999, <strong>in</strong> 1.6 L pots <strong>and</strong> grown <strong>for</strong> 6 weeks <strong>in</strong> soil<br />
amended with P [2 g s<strong>in</strong>gle superphosphate pot- l<br />
(19% P20S)], AMF (2000 spores pot l of Glomus <strong>in</strong>traradices,<br />
Schenk <strong>and</strong> Smith), a fungicide Benomyl<br />
(200 mL of 0.1% solution added one day prior to<br />
plant<strong>in</strong>g), or control consist<strong>in</strong>g of non-sterile soil.<br />
No additional fertilizers were added <strong>and</strong> the plants<br />
were watered as needed. At harvest, nodule numbers<br />
were determ<strong>in</strong>ed along with shoot N<strong>and</strong> P<br />
concentrations <strong>and</strong> AM colonization us<strong>in</strong>g st<strong>and</strong>ard<br />
procedures (Brundrett et ai., 1996; Watanabe <strong>and</strong><br />
Olsen, 1965; Jensen 1962).<br />
Data were analyzed us<strong>in</strong>g a one-way ANOV A <strong>and</strong><br />
trans<strong>for</strong>med where appropriate. When trans<strong>for</strong>mation<br />
failed to generate data that fulfilled the underly<strong>in</strong>g<br />
assumptions, data were analyzed us<strong>in</strong>g nonparametric<br />
tests.<br />
Experiment 2<br />
Production ofAMF <strong>and</strong> control <strong>in</strong>oculum<br />
Maize was planted <strong>in</strong> the soil <strong>in</strong> July 2001, <strong>and</strong><br />
grown <strong>for</strong> three months to enhance the abundance<br />
of <strong>in</strong>digenous AMF. Control pots consisted of<br />
maize grown <strong>in</strong> sterile soil that had been given<br />
spore wash<strong>in</strong>gs conta<strong>in</strong><strong>in</strong>g soil bacteria <strong>and</strong> fungi<br />
but lack<strong>in</strong>g AMF. The plants were fertilized six<br />
times with Peters fertilizer 15-0-15NK plus micronutrients<br />
(at a N concentration of 100 ppm) <strong>and</strong><br />
amended with 0.15 giL MgS04 <strong>and</strong> 5 mg PIL as<br />
KH2P04. After three months the maize plants were<br />
allowed to wilt, <strong>and</strong> dry soil <strong>and</strong> cut root pieces<br />
served as a source of <strong>in</strong>oculum, which consisted of<br />
AMF spores, external hyphae <strong>and</strong> colonized root<br />
pieces. The control roots were non-mycorrhizal.<br />
Inoculation experiment<br />
Lablab bean (var. Rongai) was planted on 3 November<br />
2002, <strong>in</strong> pots amended with either 200 mL control<br />
<strong>in</strong>oculum or 200 mL of AMF <strong>in</strong>oculum, <strong>and</strong><br />
grown <strong>for</strong> 7 weeks <strong>and</strong> watered as needed. At harvest,<br />
nodule number <strong>and</strong> weight were recorded,<br />
AM colonization determ<strong>in</strong>ed <strong>and</strong> shoot N<strong>and</strong> P<br />
concentration measured (Brundrett et ai., 1996; Watanabe<br />
<strong>and</strong> Olsen, 1965; Jensen 1962). Data were<br />
analyzed us<strong>in</strong>g a one tailed paired t-test where<br />
+AMF <strong>and</strong> control pairs shared the site orig<strong>in</strong> from<br />
the field.<br />
Results<br />
Experiment 1<br />
Enhanc<strong>in</strong>g AMF <strong>in</strong> the test luvisol significantly <strong>in</strong>creased<br />
AM colonization compared to the control<br />
(Table 1). This resulted <strong>in</strong> a doubl<strong>in</strong>g <strong>in</strong> nodule<br />
numbers <strong>and</strong> a significantly higher N content <strong>in</strong> the<br />
shoot. There was a strong beneficial effect of P on<br />
the number of nodules, which resulted <strong>in</strong> almost a<br />
doubl<strong>in</strong>g <strong>in</strong> N content <strong>in</strong> the shoot. However, <strong>in</strong>ter-<br />
44<br />
<strong>Gra<strong>in</strong></strong> <strong>Legumes</strong> <strong>and</strong> <strong>Green</strong> <strong>Manures</strong> <strong>for</strong> <strong>Soil</strong> <strong>Fertility</strong> <strong>in</strong> Southern Africa