Characterization of a New Defensin from Cowpea (Vigna unguiculata (L.) Walp
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Abstract Intercropping cotton (Gossypium
hirsutum L.) and cowpea (Vigna unguiculata (L.)
Walp) is one of the ways to improve food security
and soil fertility whilst generating cash income of
the rural poor. A study was carried out to find out
the effect of cotton–cowpea intercropping on
cowpea N2-fixation capacity, nitrogen balance
and yield of a subsequent maize crop. Results
showed that cowpea suppressed cotton yields but
the reduction in yield was compensated for by
cowpea grain yield. Cowpea grain yield was sig-
nificantly different across treatments and the
yields were as follows: sole cowpea (1.6 Mg ha–1),
1:1 intercrop (1.1 Mg ha–1), and 2:1 intercrop
(0.7 Mg ha–1). Cotton lint yield was also signifi-
cantly different across treatments and was sole
cotton (2.5 Mg ha–1), 1:1 intercrop (0.9 Mg ha–1)
and 2:1 intercrop (1.5 Mg ha–1). Intercropping
cotton and cowpea increased the productivity
with land equivalence ratios (LER) of 1.4 and 1.3
for 1:1 and 2:1 intercrop treatments, respectively.
There was an increase in percentage of N fixation
(%Ndfa) by cowpea in intercrops as compared to
sole crops though the absolute amount fixed
(Ndfa) was lower due to reduced plant popula-
tion. Sole cowpea had %Ndfa of 73%, 1:1 inter-
crop had 85% and 2:1 intercrop had 77% while
Ndfa was 138 kg ha–1 for sole cowpea, 128 kg ha–1
for 1:1 intercrop and 68 kg ha–1 for 2:1 intercrop
and these were significantly different. Sole cow-
pea and the intercrops all showed positive N
balances of 92 kg ha–1 for sole cowpea and 1:1
intercrop, and 48 kg ha–1 for 2:1 intercrop. Cow-
pea fixed N transferred to the companion cotton
crop was very low with 1:1 intercrop recording
3.5 kg N ha–1 and 2:1 intercrop recording
0.5 kg N ha–1. Crop residues from intercrops and
sole cowpea increased maize yields more than
residues from sole cotton. Maize grain yield was,
after sole cotton (1.4 Mg ha–1), sole cowpea
(4.6 Mg ha–1), 1:1 intercrops (4.4 Mg ha–1) and
2:1 intercrops (3.9 Mg ha–1) and these were sig-
nificantly different from each other. The LER,
crop yields, %N fixation and, N balance and
residual fertility showed that cotton–cowpea in-
tercropping could be a potentially productive
system that can easily fit into the current small-
holder farming systems under rain-fed conditions.
The fertilizer equivalency values show that sub-
stantial benefits do accrue and effort should be
directed at maximizing the dry matter yield of the
legume in the intercrop system while maintaining
L. Rusinamhodzi (&) Æ H. K. MurwiraTSBF-CIAT Zimbabwe, Box MP 228, Mt Pleasant,Harare, Zimbabwee-mail: [email protected]
J. NyamangaraDepartment of Soil Science and AgriculturalEngineering, University of Zimbabwe, Box MP167,Mt Pleasant, Harare, Zimbabwe
Plant Soil (2006) 287:327–336
DOI 10.1007/s11104-006-9080-9
123
ORIGINAL PAPER
Cotton–cowpea intercropping and its N2 fixation capacityimproves yield of a subsequent maize crop underZimbabwean rain-fed conditions
L. Rusinamhodzi Æ H. K. Murwira ÆJ. Nyamangara
Received: 24 January 2006 / Accepted: 4 July 2006 / Published online: 24 August 2006� Springer Science+Business Media B.V. 2006
or improving the economic yield of the compan-
ion cash crop.
Keywords Cotton cowpea intercropping Æ N2
fixation Æ N balance Æ maize yield Æ rain-fed
conditions
Introduction
Biological nitrogen fixation (BNF) in legumes is a
fundamental process for maintaining soil fertility
and the continued productivity of low-input
cropping systems. Carry-over of N from BNF, e.g.
in roots and stover, can supply the N demand of
subsequent non-N2 fixing crops (van Kessel and
Hartley 2000). BNF therefore can be used in
agricultural systems for replenishing N, which is
often the most limiting growth factor in tropical
soils (Jeranyama et al. 1998). The percentage N
fixation (%Ndfa) and the N contribution from
leguminous crops are influenced by crop species
and a number of environmental factors such as
soil type, water availability and temperature
(Jensen et al. 1997).
Most legumes in Zimbabwe are grown as in-
tercrops with cereals (Jeranyama et al. 1998), and
evidence shows that mycorrhizal connections be-
tween the intercropped components may provide
a route of N transfer (He et al. 2003). Although,
any such N benefit to an intercropped non-legume
species would only be significant under low-
yielding conditions (Hardy 1993). In addition,
when parts of the legume senesces and decom-
pose, soil N supply to the associated crop is im-
proved (Rao and Mathuva 2000).
Cowpea (Vigna unguiculata (L.) Walp) is a
grain legume grown in savanna regions of the
tropics and has a long history of cultivation in
Zimbabwe (Reid 1977). Spreading types are
predominant and their leaves, as well as, seeds,
are consumed as an important supplement to the
staple diet of maize. Cotton (Gossypium hirsutum
L.), on the other hand is an important cash crop
for farmers in the smallholder sector that repre-
sents 80% of national production and it is the
second largest foreign currency earner among
agricultural products after tobacco (Nicotiana
tabacum L.) in Zimbabwe (Cotton Research
Institute Report 1993–1994). It is also important
to other sub-Saharan African countries, for
example, in Tanzania cotton is currently rated
third, after cashew (Anacardium occidentale L.)
and coffee (Coffea arabica L.), in terms of foreign
exchange earnings (Myaka and Kabissa 1996).
Although measurements of %Ndfa in sole crop
cowpea are well documented, whether it is en-
hanced or suppressed in cotton–cowpea inter-
cropping systems is poorly understood. The aim
of this study was to quantify the N2 fixation
capacity of cowpea as a sole crop and in two in-
tercropping systems with cotton and its effects on
a subsequent maize crop. The amount of N fixed
and transferred from cowpea to cotton in the
intercropped systems was also quantified.
Materials and methods
Site description
The field experiment was conducted during the
2003/2004 and 2004/2005 cropping seasons at
Kadoma Cotton Research Institute (CRI), Zim-
babwe (29�53¢E, 18�19¢S, Altitude 1156 m). The
soils at CRI are well-drained, reddish brown fer-
siallitics (5E.2) (Zimbabwe), Ferralic Cambisol
(FAO) and Oxic Ustropept (USDA), (Thompson
and Purves 1978; Nyamapfene 1991). The area
was under soybean during 2001/2002 but was
left uncropped fallow during 2002/2003 season.
Soil was sampled randomly using a soil auger
(0–15 cm) and taken to the laboratory for basic
characterization. The land was ploughed during
the dry season and then tilled with a disk plough
just before the start of rains in 2003.
Experimental layout
Cowpea used in this experiment was the short
season, erect type variety called CBC2 released
by Agricultural Research and Extension (AREX)
which was derived from IT18 variety, while cotton
used was the high yielding, small spreading and
medium season variety, Albar SZ9134 (CRI Re-
port 1993–1994). The treatments were (i) sole
cowpea, (ii) sole cotton, (iii) one row of cotton
alternating with one row of cowpea, planted
simultaneously, and (iv) two rows of cotton
328 Plant Soil (2006) 287:327–336
123
alternating with a row of cowpea, planted simul-
taneously. These treatments were laid out in plots
(6 · 9 m) in a completely randomised design
(CRD) with four replicates. Cowpea in sole crop
was sown at a spacing of 0.5 m between rows and
0.15 m within the rows. In intercrops row spacing
for cowpea was adjusted to 1.0 m for the 1:1
(cotton: cowpea) intercrop treatment so that a
row of cotton could fit in-between and 2.0 m for
the 2:1 (cotton: cowpea) intercrop treatment so
that two rows of cotton could fit in-between but
the in-row spacing remained 0.15 m. Cotton was
sown at spacing of 1.0 m between rows and
0.30 m within rows in both sole and intercrops.
Within each main plot two microplots
(2 · 1 m) received 25 kg N ha–1 as (15NH4)2SO4,
with an enrichment of 10.38% 15N atom excess.
The N-fertilizer was mixed with sufficient sucrose
as a carbon source to give a C:N ratio of 10:1 in
solution, and dissolved in water to allow even
application to the microplots. The area outside
the microplots was fertilized with unlabelled
(15NH4)2SO4 also at 25 kg N ha–1.
Recommended rates for cotton for P
(45 kg P ha–1 as single superphosphate, SSP) and
K (25 kg K ha–1 as Muriate of Potash) and B
(0.5 kg B ha–1 as borax) were applied (Cotton
Training Center 2000).
Cowpea residual N fertility effect in the second
season (2004/2005) was determined by incorpo-
rating all residues and comparing the response of
subsequent maize crop. Aboveground cotton
residues except litter fallen before harvest were
not returned to the soil because the stems were
burnt to control the pink bollworm (Pectinophora
gossypiella). The yields from these plots were
then compared to an N fertilizer response curve
of 0, 30, 60 kg N ha–1 applied as ammonium ni-
trate to calculate N equivalencies. All the plots
received a basal application of 45 kg P ha–1 as
SSP.
Plant sampling and analysis
All plants were sampled (cut at 1 cm above the
ground) when cowpea had reached physiological
maturity and cowpea pods separated from the
leaves and stems. Cotton plants were sampled
at the same time with cowpea and at that time
cotton was between flowering and ball formation
and plant parts were not separated. The 15N
sampling of cowpea and cotton in the first season
were measured by removing the shoots from 1 m2
drying and weighing. The yield plots were 4 m2,
and everything from this was collected at harvest
for grain and dry matter determination. In the
second season, maize yield plots were 4 m2 and all
the above ground material was measured for
grain and dry matter.
The plant samples were first dried at 72�C,
ground in a Wiley mill to pass through a 0.5 mm
sieve and then ball-milled to < 150 microns. Total
N in plant samples was determined using a mod-
ified micro-Kjeldahl procedure. Plant samples
(0.1 g) were digested in 4.4 ml of digestion mix-
ture, which consists of H2O2, H2SO4 and Se cat-
alyst. After digestion, total N was determined
colorimetrically by spectrophotometry (Bremner
and Mulvaney 1982).
The 15N abundance in the ground plant mate-
rial was measured on an ANCA 20-20 GSL iso-
tope ratio mass spectrometer (PDZ Europa Ltd.,
Cheshire, UK) following dry combustion. 15N-la-
belled Plant Reference Material supplied by the
International Atomic Energy Agency (IAEA)
was used as standard.
Land equivalency ratio
The productivity and efficiency of intercropping
was measured by calculating the land equivalency
ratio (LER) (De Wit and Van den Bergh 1965).
The intercrop yields (kg ha–1) were divided by the
pure stand yields for each component crop in the
intercrop (Eq. 1). These two figures were then
added together, values greater than 1 show an
advantage while those less than 1 show a disad-
vantage of intercropping (Sullivan 1998).
LER ¼ Cottonintercrop
Cottonsolecropþ
Cowpeaintercrop
Cowpeasolecrop
" #ð1Þ
Biological N fixation
The %Ndfa (Eq. 2) was measured using the iso-
tope dilution (ID) method. The major assumptions
Plant Soil (2006) 287:327–336 329
123
are that the isotopic composition of the soil N and
its availability to both the fixing and non-fixing
crop is assumed to be similar, any dilution in plant15N isotopic composition is assumed to result from
BNF. The intensity of Ndfa (Eq. 3) is determined
by the extent to which plant 15N isotopic compo-
sition is diluted, this is obtained by comparing 15N
enrichment in N fixing plants and non-fixing le-
gume isolines or non-legumes (McAuliffe et al.
1958).
%Ndfa¼�1� %atom excess of fixing plant
%atom excess of non� fixing plant
��100
ð2Þ
Ndfa¼ Total legume N� % Ndfa
100ð3Þ
Nitrogen transfer
When a non-N fixing plant is cultivated in asso-
ciation with an N-fixing plant, N taken up by the
non N-fixing plant in the soil is derived from two
sources: X% (Eq. 4) coming from the Ndfa by
the legume and Y% coming directly from the
soil N (Kurdali et al. 1990). The d15N signatures
of plants relying solely on N2 fixation by bacte-
rial symbionts are similar to those of atmospheric
N2 (i.e. close to 0) as opposed to much wider
range of signatures of plants relying on soil N
sources (Tjepkema et al. 2000). The natural
abundance of the heavy isotope, 15N, is com-
monly expressed as a d15N value, denoting the
relative deviation (in 0/00) from the ratio 15N: 14N
in atmospheric N2. When discussing the 15N
enrichment of natural materials, that is their
natural abundance which is usually a small value
of atom % 15N excess, then the term d15N is used
(Giller 2001a).
Additions of atmospheric N2 or other gaseous
forms of N further modify d15N signatures of
mineral N and organic N pools in soils (Heaton
et al. 1997). The amount transferred (kg N ha–1)
is the calculated using the total N content
(kg N ha–1) of the cotton crop (Eq. 5).
Percentage Ntransferred ðX%Þ
¼ d15Nref:pl � d15Nassociation
d15Nref:pl � d15Nfixation
� � � 100 ð4Þ
Where d15Nfixation is obtained from cowpea,
d15N ref.pl is obtained from cotton sole crop,
d15Nassociation is obtained from cotton in inter-
crops.
Amount of Ntransferred ¼X% � Total N cotton plant
100
ð5Þ
Nitrogen balance
The difference between Ndfa and N exported in
seeds is the amount of N introduced to the soil
system and this has been referred as N balance
(Eq. 6) and this is measured in kg ha–1. The
assumption is that the legume residues will
decompose and N will be added to the soil. Root
fixed N was not measured in this study. Root N
is important in soil N balance calculations and
studies have shown that root N may be 25–41%
of total plant N (Russell and Fillery 1996;
Rochester et al. 1998; Unkovich and Pate 2000).
More recently, Peoples et al. (2001) used an
average of 33% for estimating root N for grain
legume crops in order to calculate soil N bal-
ance. Basing literature from similar studies, the
same figure (33%) was used for calculating N in
roots in this study.
Nitrogen balance ¼ Ndfa�GrainN ð6Þ
Fertilizer equivalence
Crop residue fertilizer equivalencies (FE) were
calculated by comparing residual plot yields fol-
lowing intercrop and residue incorporation, to a
mineral fertilizer response curve experiment in
the same site and season. The response curve
quadratic expression was used to calculate the FE
of the second season crop, by fitting the yield (y)
to calculate the corresponding rate of N applica-
tion to achieve the observed yield (Equation 7).
330 Plant Soil (2006) 287:327–336
123
To compare the FE, FE% (Eq. 8) was calculated
by dividing FE by the actual amount of N applied
in crop residues and expressed as a percentage
(Mutuo et al. 1999).
FE ¼ �b �ffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiffiðb2 � 4acÞ
p2a
ð7Þ
%FE ¼ FE
N applied� 100 ð8Þ
Statistical analysis
Analysis of variance (ANOVA) using Genstat
(2002) was used to identify treatment effects.
Variables on which statistical analyses were per-
formed are cowpea grain and total dry matter
yield, amount of N fixed (Ndfa), cotton lint and
total dry matter yield, LER, maize grain and total
dry matter yield. Treatment differences were
identified using standard error of differences
between means (SED).
Results
Soil characterization
Soil organic carbon (OC) was 1.5% and shows
that the soil has a good capacity to meet long-
term crop nutrient demands. The pH was greater
than 5 and provides a suitable environment for
crops that are commonly grown in Zimbabwe.
Available N of 28.4 mg kg–1 (Table 1) falls within
low category (15–30 mg kg–1), (Mashiringwani
1983) and more N can be released by minerali-
zation from OC (Cooper and Fenner 1981). The
sodium absorption ratio (SAR) of 13 is high but
the pH of 6.5 means that the soil is normal
according to the categories given by Brady (1990).
Yield of intercrops
Cowpea grain yield was higher in the sole
crop (1.4 Mg ha–1) than in the intercrops at
1.1 Mg and 0.7 Mg ha–1 for 1:1 and 2:1inter-
crop treatments, respectively. Intercrops were
more productive than sole crops as shown by
land equivalence values (LER) which were
greater than 1 in both cases (Table 2). The
same trend was observed for cotton lint yield
with sole cotton recording 2.5 Mg ha–1 but
the 2:1 intercrop treatment yielded higher
lint (1.5 Mg ha–1) than the 1:1 intercrop
(0.9 Mg ha–1) treatment (Table 2). These yields
were significantly different (P < 0.05). Both
crops recorded higher total aboveground dry
matter yield in sole crops compared to inter-
crops. Cowpea total dry matter yields were
4.7 Mg ha–1 for sole cowpea, 3.8 Mg ha–1 for
1:1 cotton–cowpea intercrops and 2.2 Mg ha–1
for 2:1 cotton–cowpea intercrops. Cotton yields
were 12.3 Mg ha–1 for sole cotton, 6.5 Mg ha–1
Table 1 Soil properties from the experimental site
Sand(%)
Clay(%)
Silt(%)
pH(CaCl2)
OC(%)
N (mg kg–1) AvailableP (%)
Ca(cmol+ kg–1)
Mg(cmol+ kg–1)
K(cmol+ kg–1)
Na(cmol+ kg–1)
Mineral Total
42 37 21 6.5 1.5 28.4 1880 0.03 8.4 7.7 0.3 3.7
Table 2 Effect of sole crops and intercrops on total dry matter (TDM), grain and lint yield, %Ndfa and Ndfa
Treatment CowpeaTDM(Mg ha–1)
CottonTDM(mg ha–1)
Cowpeagrain yield(mg ha–1)
Cotton lintyield (mg ha–1)
LandEquivalenceratio (LER)
%Ndfa Ndfa (kg ha–1)
Sole crop 4.7 12.3 1.4 2.5 1.0 73 1381:1 intercrop 3.8 6.5 1.1 0.8 1.4 85 1282:1 intercrop 2.2 9.5 0.6 1.5 1.3 77 68SED 0.3 0.8 0.2 0.4 0.01 3 27
Plant Soil (2006) 287:327–336 331
123
for 1:1 intercrop treatment and 9.5 Mg ha–1 for
2:1 intercrop (Table 2).
Biological nitrogen fixation
The amounts of Ndfa were significantly different
(P < 0.05) among treatments (Table 2). The
highest %Ndfa (85%) was recorded for 1:1 cot-
ton/cowpea intercrop, the lowest percentage
(73%) was recorded for sole cowpea treatment
and 2:1 cotton/cowpea intercrop treatment had
77%. The highest amount of Ndfa (138 kg N ha–1)
was however obtained from the sole cowpea crop,
the 1:1 cotton/cowpea intercrop had 128 kg N ha–1
and the 2:1 cotton/cowpea treatment had the
lowest amount of 68 kg N ha–1.
Nitrogen balance
The N balances of all the treatments were positive
(48–92 kg N ha–1) (Table 3). The 1:1 cotton/
cowpea intercrop treatment had the highest N
balance of all the treatments. Uptake from the
soil ranged between 15 and 38 kg N ha–1 (Ta-
ble 3). The amount of N transferred between
crops was insignificant with the 1:1 intercrop
having the highest amount of 3.5 kg N ha–1
while
the 2:1 intercrop recorded only 0.5 kg N ha–1.
The intercrop system’s planting arrangement and
fixation capability had an effect on the percentage
of N in the associated cotton crop coming from
N2-fixation by cowpea.
Yield of maize after intercrops
Previous crop residues had a significant effect on
the following maize crop yields, with maize fol-
lowing sole cotton having a significantly lower
yield than either sole crop cowpea or the two
intercrop treatments (P < 0.05) (Table 4).
Following sole cotton, maize yields had a
highly negative %FE (–27%) that was signifi-
cantly lower than the other three treatments. Sole
cowpea (27%), 1:1 intercrop (32%) and 2:1
intercrop (29%) showed no significant differences
between them.
Discussion
The %Ndfa by cowpea was higher in intercrop
treatments than in sole crop indicating a higher
productivity of the cotton/cowpea intercropping
system. This was due to increases in competition
for soil N that led to better nodulation. Marschner
(1995) reported that growing a non-legume with a
legume reduced the amounts of mineralized N in
the soil and legumes responded by fixing more N
Table 3 Total N in plant, N uptake from soil, Ndfa, N removed in grain and N balance in soil after intercrops as comparedto sole cowpea
Treatment Total N inplant (kg ha–1)
Ndfa(kg N ha–1)
N uptakefrom soil(kg N ha–1)
Grain Nremoved(kg N ha–1)
N balance(kg N ha–1)
Sole cowpea 189 138 51 46 921:1 Intercrop 150 128 22 36 922:1 intercrop 88 68 20 20 48SED 36 27 12 5 22
Table 4 Effect of residual fertility (N) on stover and grain yield of maize after growing cotton and cowpea intercrops aswell as sole crops as compared to three mineral fertilizer levels at CRI
Yield Previous crop Fertilizer level (kg N ha–1) SED
Cotton Cowpea 1:1 2:1 0 30 60
Grain (mg ha–1) 1.4 4.6 4.4 3.9 2.3 4.9 6.3 1.2Stover (mg ha–1) 4.3 8.6 8.1 7.3 5.9 8.8 11.2 1.6
332 Plant Soil (2006) 287:327–336
123
than they do in a pure stand, so long as the le-
gumes dominated the mixture. The Ndfa was
however lower in intercrop treatments than in
sole crop due to the lower density of the fixing
plant in intercrop than in sole cropping. The
greater density in cowpea sole crop increased the
dry matter yield per unit area and hence the total
amount of N fixed.
This study made use of the isotope dilution
method (ID). A great advantage with the use of15N isotope dilution technique to estimate N2
fixed is its ability to give an integrated estimate of
N fixation over a growing season or longer
(McAuliffe et al. 1958). It is the only method that
can distinguish between soil, fertilizer and fixed N
in field-grown crops. Estimates made using the
isotope dilution technique have established that
most legumes derive a large proportion of their N
from fixing atmospheric N2 and, in general, this is
in excess of 70–80% of their total N requirements
(Eaglesham et al. 1977) and results reported here
are close to that range. Errors often result from
differences in the pattern of N uptake between
reference and N2-fixing plants (Giller and Witty
1987). The addition of mineral N to the soil is
likely to disturb the process under measurement
as this added amount can constitute a significant
fraction of the mineral N available to the legume
(McNeill et al. 1996). The major problem with the
ID method is that 15N uptake is affected by var-
iation in rooting depth and timing of N uptake
between fixing and non-fixing crop (Giller,
2001a). Cowpea has a deep rooting system
(Giller, 2001a), and so is cotton but initial growth
is slow (Cotton Training Center 2000). Due to its
slow initial growth, the peak of 15N uptake could
have been different between cotton and cowpea
because available soil 15N rapidly declines due to
plant uptake, other losses and dilution due to
mineralization of N with smaller enrichment from
soil organic matter (Giller, 2001a). This could
have resulted in lower enrichment of 15N in the
cotton reference crop and an underestimate of
N2-fixation.
The LER obtained for cotton and cowpea in-
tercrops were all greater than one (>1.0). This
meant that the land requirement for the inter-
crops was lower than that for monocrops. Inter-
cropped legumes and cotton used plant growth
resources more efficiently than pure stands, thus
supporting results of Fukai and Trenbath (1993)
and Rao et al. (1987) on intercropping systems
with legumes and non-legumes. They attributed
the advantage of intercropping to different
aboveground and belowground growth habits and
morphological characteristics of the crops, and to
the higher efficiency in the utilization of water
and radiation energy. Cowpea rooting system is
more extensive than those of soybean (another
potential companion with cotton and also a crop
gaining prominence in Zimbabwe) and it can ex-
plore deep soil profiles for moisture (Miller 1998).
Cotton rooting on the other hand depends on
moisture availability. Under conditions of little
moisture, the main root system usually develops
to a considerable depth of up to 2.0 m (Monks
1999).
Although %Ndfa was high in the intercrop
systems (>75%), the associated cotton crop did
not get much of the N fixed by cowpea. Little
transfer was observed in both cases although it
was a little higher for 1:1 intercrop than for 2:1.
This could be explained by the fact that in 1:1
intercrop, the cotton crops are surrounded by two
rows of cowpea crops whereas in 2:1, two rows of
cotton will share a row of cowpea and hence
transfer is very low. Snoeck et al. (2000) were able
to show that under field conditions; roughly 30%
of N effectively fixed by a legume was transferred
to the associated non-legume. They reported that
N transfer was high because it was enhanced by
the interactions of roots and root exudates. The
low value (3%) for N transfer reported in this
study could be because the period of the study was
short (2 months) compared to at least two seasons
for Snoeck et al. (2000). The crops they used for
their study were coffee (Coffea arabica (L.) and
legumes, Flemingia macrophylla (Willd.) Merr.,
Desmodium intortum (Mill.) Urb., Leucaena leu-
cocephala (Lam.) De Wit., Leucaena diversifolia
(Schelecht.) Bentham, Calliandra calothyrsus
(Meissn.) and Erythrina abyssinica (Lam.).
The higher N balance (92 kg N ha–1) for the
cowpea sole crop and 1:1 intercrop could be ex-
plained by a higher amount of N fixation but far
much lower removal of N in the grain of
46 kg N ha–1 and 36 kg N ha–1, respectively. This
figure includes the contribution of roots to N
Plant Soil (2006) 287:327–336 333
123
balance and represents the maximum possible
amount of N added to the soil after growing sole
cowpea or 1:1 intercrops.
There is evidence of soil fertility improvement
in cropping systems that includes legumes such as
cowpea as shown by the yield of the succeeding
maize crop especially where cowpea plant density
was high (Giller 2001b). In rotation, legume res-
idues have a longer time to decompose as com-
pared to intercropping releasing more N that lead
to better yield of the following cereal (Jeranyama
et al. 2000). Increase in yield of crops following
cowpea has also been attributed to reduced
moisture stress in rotations (Rao and Mathuva
2000), reduction in Striga hermonthica (Reddy
et al. 1994) and also the ability of cowpea to
access sparingly soluble P (Vanlauwe et al. 2000).
However for the benefits to be fully realized the
legume should produce more residue and careful
management of the residue is required.
The fertilizer equivalence (FE) values for the
previous crop residues showed that cowpea resi-
dues contribute more to the N nutrition of the
following crop than cotton residues. Cadisch and
Giller (1997) reports that legumes are able to
provide more mineral N in the short term than
residues of other crops because of their better
chemical composition which has a major influence
on the rates of decomposition and N release. The
reduction in yield obtained in plots that had sole
cotton could be explained by the poor quality cot-
ton residues which caused immobilization of soil
available N (data not shown). The higher maize
grain yields from plots previously under intercrops
than from fallow indicated an obvious incentive to
integrate legumes in the existing cropping systems.
The positive effect of the previous cowpea ob-
served in these trials could have been partly due to
the previous cowpea absorbing less soil N than the
previous cotton, which is referred to as sparing ef-
fect (Peoples et al. 1995). Additionally, there was
probably additional N fixed and left in the soil for
the subsequent maize crop.
Conclusions
The intercropping of cotton with a short season
cowpea variety is a potentially productive system
as shown by high LER values. Nitrogen fixation
capacity of cowpea is enhanced in the intercrop
compared to sole crops. Nitrogen transfer from
cowpea to cotton does occur in cotton/cowpea
intercrops but it is of little significance to the
overall nutrition of the cotton crop. The positive
N balance in the intercrop indicated a further
advantage, and this N could be used to produce a
subsequent high yielding crop such as maize
which has high nutrient demands. This is shown
by the improvement in the grain and stover yield
of maize following intercrops without additional
N fertilizer. The benefits mentioned in this study
accrued without substantial additional costs ex-
cept for cowpea seed, farmers would be able to
save on N fertilizer input and should now divert
the little resources they have on securing P fer-
tilizer as it is the second most limiting element to
crop production.
Acknowledgements Special thanks to N. Mutasa andR. Dhliwayo from Kadoma Cotton Research Institute’sCrop Productivity Unit for letting me use their land andassistance with laying out the trials especially during thefirst season. Financial support from TSBF-CIAT throughthe IFAD Project is greatly appreciated.
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