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Nitrogen Transfer to Forage Crops from a Caragana Shelterbelt

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Caragana shelterbelts are a common feature of farms in the Northern Great Plains of North America. We investigated if nitrogen (N) from this leguminous shrub contributed to the N nutrition of triticale and oat forage crops growing adjacent to the shelterbelt row. Nitrogen transfer was measured using N-15 isotope dilution at distances of 2 m, 4 m, 6 m, 15 m and 20 m from the shelterbelt. At 2 m caragana negatively impacted the growth of triticale and oat. At 4 m from the shelterbelt productivity was maximum for both forage crops and corresponded to the highest amount of N originating from caragana. The amount of N transferred from caragana decreased linearly with distance away from the shelterbelt, but even at 20 m from the shelterbelt row measureable amounts of N originating from caragana were detectable in the forage biomass. At 4 m from the shelterbelt approximately 40% of the N in both oat and triticale was from caragana, and at 20 m from the shelterbelt approximately 20% of the N in oat and 8% of the N in triticale was from caragana.
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Forests 2015, 6, 1922-1932; doi:10.3390/f6061922
forests
ISSN 1999-4907
www.mdpi.com/journal/forests
Article
Nitrogen Transfer to Forage Crops from a Caragana Shelterbelt
Gazali Issah 1, Anthony A. Kimaro 2, John Kort 3 and J. Diane Knight 4,*
1 Western Applied Research Corporation (WARC), P.O. Box 89 Scott, SK S0K 4A0, Canada;
E-Mail: gazali.issah@warc.ca
2 World Agroforestry Centre, ICRAF-Tanzania Programme, Dar-es-Salaam, Tanzania;
E-Mail: A.Kimaro@cgiar.org
3 Agroforestry Development Centre, Agriculture and Agri-Food Canada (retired), P.O. Box 940,
Indian Head, SK S0G 2K0, Canada; E-Mail: korts@sasktel.net
4 Department of Soil Science, University of Saskatchewan, 51 Campus Drive, Saskatoon, SK S7N
5A8, Canada
* Author to whom correspondence should be addressed; E-Mail: diane.knight@usask.ca;
Tel.: +1-306-966-2703; Fax: +1-306-966-6881.
Academic Editor: Heinz Rennenberg
Received: 7 April 2015 / Accepted: 20 May 2015 / Published: 29 May 2015
Abstract: Caragana shelterbelts are a common feature of farms in the Northern Great
Plains of North America. We investigated if nitrogen (N) from this leguminous shrub
contributed to the N nutrition of triticale and oat forage crops growing adjacent to the
shelterbelt row. Nitrogen transfer was measured using 15N isotope dilution at distances of
2 m, 4 m, 6 m, 15 m and 20 m from the shelterbelt. At 2 m caragana negatively impacted
the growth of triticale and oat. At 4 m from the shelterbelt productivity was maximum for
both forage crops and corresponded to the highest amount of N originating from caragana.
The amount of N transferred from caragana decreased linearly with distance away from the
shelterbelt, but even at 20 m from the shelterbelt row measureable amounts of N
originating from caragana were detectable in the forage biomass. At 4 m from the
shelterbelt approximately 40% of the N in both oat and triticale was from caragana, and at
20 m from the shelterbelt approximately 20% of the N in oat and 8% of the N in triticale
was from caragana.
Keywords: agroforestry; oat; triticale; 15N isotope dilution; Saskatchewan
OPEN ACCESS
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1. Introduction
Field shelterbelts are a common feature on farms in the Northern Great Plains of North America.
The first shelterbelts in Canada were planted in 1903 as windbreaks to protect soils, crops and
farmyards [1]. In the USA, as part of the 1935 Soil Conservation Act, trees were planted in large
numbers as a means of soil erosion control in response to the devastating drought, windstorms and
resulting soil erosion that occurred during the Great Depression [2]. Shelterbelts consist of a variety of
coniferous and deciduous trees, with specific species adapted to the variety of soils and environments
encountered across the region. Planting of shelterbelts had a number of positive impacts including
protecting soil from wind erosion [3,4], positively influencing field hydrology through trapping
snow [5] and protecting crops from wind damage [3], among other beneficial impacts [6]. Shelterbelts
do not only serve as physical barriers to soil and snow movement; they are living barriers that separate
cropped fields and as such interact with the adjacent field crops in either a competitive or
complementary fashion [7]. Trees compete with adjacent field crops for above-ground and
below-ground resources. In the semi-arid climates of the Northern Great Plains, soil moisture is
frequently the limiting resource [7,8]. Furthermore, crop yields near shelterbelts can be reduced due to
allelopathy by the tree roots [9], nutrient leaching during snowmelt [10], shading [11] and changes to
microclimate [12]. Nonetheless, field shelterbelts that typically occupy 5% to 6% of the cropland can
produce economic returns to producers based entirely on the increased yields in sheltered areas [13]. In
temperate climates, yield increments for a number of field crops affected by shelterbelts ranged from
6% to 44% [6].
Caragana (Caragana arborescens Lam.) is a common shrub found in shelterbelts all across the
Canadian prairies. The species is one of the hardiest, small deciduous shrubs planted on the northern
Great Plains [12]. It readily adapts to sandy, alkaline soils in open (unshaded) environments [14].
Because it is a legume, caragana has the capacity to form root nodules and fix atmospheric
nitrogen (N) [14,15] and enhance N availability [16,17]. Biological N2 fixation accounted for up to
80% of N in caragana leaves in a soil-less system [15] and up to 65% in field soil [17,18]. An
intercropping study conducted in Saskatchewan, Canada provides indirect evidence for transfer of N
from caragana to willow (Salix miyabeana). Caragana enhanced the growth of willow at two sites, but
was outcompeted by willow at a third site [17]. At the first two sites, willow growth increased as a
function of the proportion of caragana in the mixed stand suggesting that increased N availability was
driving the relationship. Nitrogen transfer was not measured directly.
The objective of this study was to quantify the transfer of N from a caragana shelterbelt to adjacent
triticale (X Triticosecale Wittmack) and oat (Avena sativa L.) crops grown for forage in successive
years. The influence of distance from the shelterbelt was investigated to determine the extent of the
influence of the shelterbelt on N nutrition and forage quality of the adjacent field crops.
2. Experimental Section
2.1. Experimental Setup
The experiment was established adjacent to an existing caragana shelterbelt at the Agroforestry
Development Centre, Indian Head, SK, Canada (50°33′ N 103°39′ W) (Figure 1). The shelterbelt was
Forests 2015, 6
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planted on the nursery site in 1933 in a north-south orientation. Caragana was approximately 5 m tall
at the time of the experiment. The shelterbelt is a single row shelterbelt, characteristic of most
Canadian shelterbelt plantings [6]. The soil on site is an Orthic Oxbow with a loamy, morainal and
hummocky landform, with a slope of 2% to 5% [19]. Because the site was originally a nursery site,
historically the shelterbelt received periodic irrigation, fertilization and herbicide and pesticide
applications uncharacteristic of shelterbelts on farmland. However, no applications of fertilizers,
herbicides nor pesticides have been made for the last twenty-five years.
The shelterbelt is adjacent to an annually cropped field located on the eastern side of the shelterbelt.
In the spring of 2011, prior to establishing the experiment, soils were sampled from the site in 15 cm
increments to a 45 cm depth. Four soil cores were extracted at five distances from the shelterbelt row,
at 2 m, 4 m, 6 m, 15 m and 20 m for physiochemical characterization (Table 1). All analyses were
performed using standard protocols [20]. Electrical conductivity (EC) and pH were measured on 1:2
soil: water extracts. In 2011 the field was planted by staff at the Agroforestry Development Centre
with spring triticale (X Triticosecale Wittmack cv. AC Ultima) at a seeding rate of 90 kg ha1 and in
2012 the field was planted with Pinnacle oat (Avena sativa L.) at a seeding rate of 90 kg ha1. Total
precipitation during the 2011 growing season (May to September) was 290 mm and average
temperature was 15.3 °C. In the 2012 growing season (May to September) the total precipitation was
285 mm and temperature 15.1 °C. The long-term (1971 to 2000) averages for the area were 255 mm
precipitation and temperature of 15.9 °C.
Table 1. Physiochemical properties of Orthic Oxbow soil at distances perpendicular to a
caragana shelterbelt.
Distance (m)
pH
EC
(mS cm1)
P
NO3
S
OC
K+
Mg2+
Ca2+
mg kg1
g kg1
cmol (+) kg1
015 cm
2
8.04
0.26
22.04
8.27
5.66
7.55
0.70
1.86
9.83
4
8.05
0.17
26.21
2.40
5.15
7.05
0.62
1.66
9.29
6
8.06
0.19
21.34
1.52
4.47
7.00
0.52
1.98
9.50
15
8.01
0.22
27.23
2.23
7.65
7.20
0.63
2.21
7.75
20
7.93
0.26
31.94
2.99
9.00
7.45
0.65
2.44
7.27
1530 cm
2
8.21
0.20
6.40
1.50
4.18
4.50
0.36
2.28
9.07
4
8.31
0.16
6.31
1.02
4.25
5.10
034
2.06
9.32
6
8.37
0.15
11.48
1.25
4.29
4.95
0.33
2.11
8.86
15
8.17
0.20
9.50
1.17
6.55
5.90
0.34
2.35
7.79
20
8.26
0.23
8.51
0.87
11.16
5.15
0.29
2.91
8.21
3045 cm
2
6.40
0.13
5.36
0.22
3.37
2.45
0.23
2.58
6.13
4
8.49
0.18
9.77
0.29
4.18
3.00
0.31
3.10
8.34
6
8.45
0.20
6.09
0.67
4.11
3.55
0.24
2.70
8.27
15
8.44
0.21
5.64
0.49
7.80
3.45
0.23
2.75
7.90
20
8.56
0.29
5.16
0.46
nd
3.80
0.25
3.34
7.79
Mean values (n = 4); nd = no data.
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In each of the two years of the study four 15 m × 20 m plots were established perpendicular to the
shelterbelt on the eastern side of the shelterbelt (Figure 1). Within each plot, 0.6 m × 0.6 m subplots
were established at distances 2 m, 4 m, 6 m, 15 m and 20 m from the centre of the shelterbelt.
Above-ground biomass from the triticale and oat sub-plots was hand-harvested approximately 5 cm
above the soil surface using scissors, 9 and 10 weeks after planting, respectively. The time of sampling
corresponded to the mid to late milk stage of development (BBCH = 76) for each crop, after maximum
N uptake occurred. In cereal crops maximum N uptake typically occurs around 60 to 65 days after
emergence [21]. Four additional reference samples were collected from outside the main plots
approximately 50 m from the shelterbelt. Plants from these reference plots were assumed to be far
enough from the caragana so as not to be influenced by the caragana and not to have accessed N fixed
by the caragana. Leaves from caragana were collected from four randomly selected locations within
the canopy of the shelterbelt in the experimental plot. Delta 15N values from the reference crops and
the caragana were used in calculating N transfer from the caragana to the adjacent crops (Equation 1).
Plant samples were oven-dried at 60 °C to stable weight and ground using a Wiley mill (Thomas
Scientific, Swedesboro, NJ, USA), and re-ground to fine powder using a ball grinder (8000D
Mixer/Mill, SPEX Sample Prep® LLC., Metuchen, NJ, USA). Subsamples (ca. 1 mg) were weighed
using a micro-balance (Sartorius Microbalance, CPA2P, Bradford, MA, USA) and encapsulated in
8 mm × 5 mm tin capsules for analysis of N concentration and δ15N on a Costech ECS4010 elemental
analyzer (Costech Analytical Technologies Inc., Valencia, CA, USA) coupled to a Delta V Advantage
mass spectrometer (Thermo Scientific, Bremen, Germany) at the University of Saskatchewan
(Saskatoon, SK, Canada). The δ15N of the samples was used to calculate the percentage of N derived
from the atmosphere (% Ndfa) that was transferred from the caragana to the triticale and oats
(% Ntransfer) [21]:
   


  
  (1)
Where δ15Ncrop is the value for the reference triticale or oat not influenced by the caragana and
accessing only soil available N; δ15Ntree/crop is triticale or oat influenced by the caragana and assumed to
access N from soil available N and caragana fixed N; and δ15Ntree is the value for the caragana.
Amount of N transferred from caragana to triticale or oat was calculated as the product of %Ntransfer and
amount of N in the crop.
Statistical analyses were performed on %Ntransfer and amount of N transferred using SAS
version 9.2 [22]. Simple linear regression was performed on %Ntransfer, the amount of N transferred,
total N in the shoots of the forage crops and shoot biomass, as a function of distance from the
shelterbelt using PROC REG procedure in SAS. Regression analysis was performed with data from the
2 m distance included and excluded. Only regressions where the 2 m distance measurements were
excluded were significant (P < 0.05) and are reported.
Forests 2015, 6
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Figure 1. Diagram (not drawn to scale) of experimental plot layout showing replicates and
sampling subplots at 2 m, 4 m, 6 m, 15 m and 20 m from the caragana shelterbelt (60 cm ×
60 cm) within each 15 m swath.
3. Results and Discussion
Substantial N transfer from caragana to triticale and oat occurred in the two years of the study
(Figure 2A and 2D). Except at the 2 m distance, both percentage of N and the amount of N (g N m2)
originating from caragana in the triticale and oat biomass decreased linearly with increasing distance
from the shelterbelt (Figure 2A and 2D). Both triticale and oat productivity were severely limited 2 m
from the shelterbelt with both crops producing about 50% of the biomass produced 4 m from the
shelterbelt (Figure 2C and 2F). Productivity of both crops was maximum at 4 m (Figure 2C and 2F)
corresponding to the same distance at which the maximum amount of N was transferred (Figure 2A
and 2D). The amount of N transferred to triticale at 2 m was about 58% of the transfer at 4 m and for
oat only about 3.5% of the transfer at 4 m.
Low yields near shelterbelts have long been reported e.g., [11,23]. At the 2 m distance from the
shelterbelt competition for water, light and/or nutrients between the caragana and the crop plants were
probably responsible for the low yields of the crops. In temperate climates, competition for water is
generally more limiting than competition for light [8] and has been reported to decrease yields of crops
near tree rows [7]. Some early research focused on estimating the zone of influence of shelterbelts in
terms of the negative impact on yield. Read [23] estimated yield reductions occurred to 1 h, where h is
the height of the shelterbelt. Stoeckeler [11] suggested a range of 0.5 h to 1.5 h and Lyles et al. [24]
reported reduced yields at a distance of 2 h. In our study the negative influence on forage yield of the
approximately 5 m-tall shelterbelt was narrow at less than 4 m from the shelterbelt for both crops
(i.e., <1 h). Considering that caragana has a very vigorous and aggressive growth habit and can be
invasive in some conditions [14,25] this narrow zone of negative influence was somewhat surprising.
Forests 2015, 6
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The benefits from the N2-fixing capabilities of the plant might balance the negative impacts from
the invasive nature of the shrub. Indeed the vigorous root growth of caragana might be responsible for
the significant N transfer observed up to about 15 m in both crops. Considering tree root density
decreases with increasing distance from the shelterbelt row, this approximately 15 to 20 m zone of
influence in this study is assumed to be indicative of the zone where direct transfer of N takes place
through root to root contact or mycorrhizal associations, probably augmented by indirect transfer
through mineralization of below-ground roots and sloughed cells, nodules, and other organic
compounds excreted from roots, and possibly leaf litter fall.
Studies using 15N labeling have demonstrated net N transfer from legume tree saplings to grass
grown together in containers [26] and have provided evidence of N transfer via mycorrhiza between
N2-fixing and non-fixing tree species [2729]. Natural abundance methodologies have provided
similar estimates of N transfer between trees [3032]. The natural abundance method has an advantage
over 15N labeling studies in that N is not added to the soil or plant, enabling N transfer to be estimated
without disturbing the system. However the method relies on very small differences in natural
abundance values between the legume and non-legume plants and in the soil inorganic N pool. Any
soil processes like nitrification, denitrification and ammonium volatilization that discriminate against
15N can bias the estimates by altering the δ15N signature of the inorganic N pool accessed by the plants.
Furthermore, isotope ratio mass spectrometry (IRMS) analyses require a very small amount of an
organic sample (typically 1 mg10 mg) representative of the entire sample. It is essential that the
organic material sampled is finely ground and homogeneously mixed. Because of the small size of the
subsample for IRMS even small deviations in weighing samples can have a relatively large effect,
particularly on the percent N values. Nonetheless natural abundance studies have been used
successfully to estimate N transfer [3032]. In our study, differences in δ15N between the caragana and
triticale and oat are sufficiently large and consistent to provide a confident estimate of N
transfer (Table 2).
Table 2. Delta 15N values of leaves (caragana) or shoots (triticale and oats) where triticale
and oats are grown adjacent to a mature caragana shelterbelt row (distance 0) and sampled
at perpendicular distances away from the shelterbelt.
Caragana
Triticale
Oats
Distance (m)
δ15N (per mil)
Std Dev
δ15N (per mil)
Std Dev
δ15N (per mil)
Std Dev
0
4.55
0.44
-
-
-
-
2
-
-
7.03
0.65
8.175
1.20
4
-
-
7.38
0.44
5.845
0.75
6
-
-
8.22
0.78
6.268
0.18
15
-
-
8.99
1.91
6.285
0.19
20
-
-
10.83
1.19
7.935
1.18
P-value
0.0028
0.0047
The negative impact of the shelterbelt on forage crop yield 2 m from the shelterbelt was also
observed for the amount of N transferred (Figure 2A and 2D). Despite lower yields and lower amounts
of N transferred at 2 m compared to 4 m, in triticale approximately 60% of the N in the shoots was
transferred from caragana at 2 m, whereas in oat only about 5% of N in the shoots was transferred
Forests 2015, 6
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from caragana. It is clear that the two forage crops interacted differently in the immediate vicinity of
the shelterbelt trees. The concentration of N in the oat shoot biomass was higher at 2 m
(2.3 ± 0.23 mg N g1) compared to the further distances (1.5 ± 0.13 mg g1 averaged over the 4 to 20 m
distance) indicating a concentration of N because of low dry matter production but similarly indicating
that amounts of N present were sufficient to support the biomass produced at this distance. In contrast,
in triticale N concentration was constant across the sampling transect (2.0 ± 0.13 mg N g1). Initial soil
sampling in 2011 indicated that amounts of nitrate in the top 15 cm were higher 2 m from the
shelterbelt than elsewhere in the field (Table 1). Soils were not sampled in 2012, but it appears that the
levels of available N near the shelterbelt were sustained at high enough levels to fully supply the N
requirement of oat at this distance. Amounts of soil N are typically higher near the tree line than in
cropped areas in agroforestry systems [3335].
Even though oat did not appear to rely on direct transfer from caragana through root to root contact
or mycorrhizal connections, the lack of any apparent N transfer from caragana to the oat crop at the
2 m distance was unexpected. Oats at the 2 m distance should still be accessing N mineralized from
caragana litter fall. In addition to the 15N signature of organic matter inputs, changes in 15N natural
abundance of the available soil N can also occur through soil processes like nitrification, denitrification
and NH3 volatilization. Discrimination against 15N occurs in all processes causing a decrease
in δ15N associated with nitrification or an increase in δ15N associated with volatilization and
denitrification [36]. Soil conditions near the shelterbelts should be conducive for denitrification in that
snow melt in this snow accumulation zone could potentially provide the necessary anaerobic
conditions, and mineralization of leaf litter the necessary nitrate and soil organic carbon substrates
(Table 1). The high δ15N signature of the oat growing at the 2 m distance could be a result of a high
δ15N signature in the soil N because of denitrification and/or volatilization. Alternatively, snowmelt
and or/rainfall may have leached nitrate out of the rooting zone of the oat crop [10].
Despite N transfer to oat being negligible close to the caragana shelterbelt, the positive impact of
the caragana on N transfer was sustained over larger distances for oat than triticale (Figure 2D and
2A). Whereas the amount of N transferred to triticale at 20 m was only 9% of N transferred at 4 m, the
amount transferred to oat at 20 m was 22% of that transferred at 4 m. At 20 m approximately 20% of
the N in oat was from caragana, whereas in triticale, N from caragana constituted only about 8% of the
shoot N. The total amount of N in the shoot tissue of triticale (i.e., transferred N + soil derived N) was
about 1.3 times the amount in oat indicating a stronger sink for N (Figure 2B and 2E). It appears that
the stronger demand for N probably led to the supply of available N being used up faster in triticale
than oat.
There are few studies in the literature examining the distance to which N transfer occurs particularly
in mature shelterbelts. Daudin and Sierra [37] measured transfer between Gliricidia sepium and
Dichanhium aristatum (Poir) in a 16-year old tropical agroforestry system over a 5 m distance. They
measured a very similar pattern of decreasing N transfer to the grass with distance away from the tree
row, and a decreasing trend in total N uptake from the soil. In general % N derived from the tree was
relatively stable at around 50% over 3 m to 4 m, with percentages decreasing slightly at 5 m [37].
Isaac et al. [38] reported that N2-fixation in Acacia senegal decreased with the age of the tree. In
contrast, caragana appears to sustain relatively high N2-fixation in mature trees at levels that sustain
significant transfer to the adjacent crop.
Forests 2015, 6
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Figure 2. Nitrogen transfer characteristics and above-ground productivity in triticale
(AC) and oat (DF) as a function of perpendicular distance away from a caragana
shelterbelt. (A,D) are percentage of N in the forage crops transferred (dots) from caragana
and amount of N transferred (triangles); (B,E) are total N in the above-ground biomass and
(C,F) are above-ground biomass.
4. Conclusions
Significant amounts of N were transferred from a mature caragana shelterbelt to adjacent oat and
triticale forage crops. Close to the shelterbelt (<4 m) caragana negatively impacted the productivity of
the adjacent forage crop. However at 4 m from the shelterbelt row, forage biomass and N content were
at their maximum. Using the 15N dilution protocol significant amounts of N originating from the
caragana were detected in the shoot biomass of triticale and oat. Nitrogen transferred from caragana
represented approximately 40% of the total N in the shoot of both triticale and oat 4 m from the
shelterbelt. Both the percentage of N transferred and the amount of N transferred decreased linearly
with distance away from the shelterbelt. Measureable amounts of N originating from caragana were
detectable in the triticale and oat biomass 20 m from the shelterbelt row.
Acknowledgments
Funding from Agriculture and Agri-Food Canada (ADC) under the Research Affiliate Program
(RAP) and University of Saskatchewan is appreciated. Authors also acknowledge the support from
staff and the administration of the Agroforestry Development Centre, Indian Head, SK, Canada, during
fieldwork. Our thanks to the Stable Isotope Laboratory, University of Saskatchewan for the
15N analysis.
0
2
4
6
8
N transfer (g m-2)
0 5 10 15
Distance from shelterbelt (m)
20
0
20
40
60
80
N transfer (%)
Oat
N transfer amount r2=0.97, P=0.015
0 5 10 15
Distance from shelterbelt (m)
20
0
2
4
200
400
6
8
10
Total shoot N (g m-2)
0 5 10 15
Distance from shelterbelt (m)
20
0
100
300
500
600
Shoot biomass (g m-2)
0 5 10 15
Distance from shelterbelt (m)
20
0
2
4
200
400
6
8
10
Total shoot N (g m-2)
Oat
0 5 10 15
Distance from shelterbelt (m)
20
0
100
300
500
600
Shoot biomass (g m-2)
Oat
0 5 10 15
Distance from shelterbelt (m)
20
0
20
40
60
80
N transfer (%)
0
2
4
6
8
N transfer (g m-2)
N transfer amount r2=0.95, P=0.0002
%N transfer r2=0.97, P<0.0001 r2=0.77,P=0.122 r2=0.54, P=0.104
r2=0.86,P=0.002 r2=0.71, P=0.037
% N transfer r2=0.79, P=0.005
AB C
DEF
Forests 2015, 6
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Author Contributions
This manuscript reports on the results of one chapter of Gazali Issah’s MSc research supervised by
Anthony A. Kimaro and Knight. J. Kort (retired) was the scientist at the AAFC-Agroforestry
Development Centre. All authors provided input into the design and implementation of the experiment.
The manuscript was written by Gazali Issah and J. Diane Knight and edited by the other authors. All
authors contributed significantly to the research.
Conflicts of Interest
The authors declare no conflicts of interest.
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... For example, in temperate tree-based intercropping (TBI) systems, variable soil organic carbon (C) and soil N patterns from tree litter inputs were observed as far as~10 m from tree row (Bambrick et al. 2010). In shelterbelt systems, contributions from fixed N from a leguminous shrub (Caragana arborescens) declined from 40% to 20% of crop N with increasing distance (4 m to 20 m) from shrub row (Issah et al. 2015). Yet these important nutrient contributions overlap with a concentrated zone of competition; for example, in a TBI system, decline in some crop yields were observed within 2 m from the tree row (Thevathasan et al. 2012). ...
... Kurppa et al. (2010) used soil isotopic enrichment to investigate the transfer of N from leguminous shade trees, Gliricidia sepium and Inga edulis, to cocoa saplings under semi-controlled conditions in the field and found all transfer of fixed-N was attributed to belowground transfer. In Caragana arborescens shelterbelt systems, Issah et al. (2015) report 40% of annual crop N uptake was derived from fixed N using 15 N isotopic dilution. Leguminous tree and grass systems have shown a range of N transfer rates via root-root interactions. ...
... Rao and Giller (1993) showed a~3% contribution between Leucaena diversifolia and the grass Cenchrus ciliaris, while in a greenhouse study, Jalonen et al. (2009) showed the grass Dichantium aristatum absorbed 22% of N exuded from roots of the leguminous tree Gliricidia sepium with~4-14% of grass N derived from leguminous tree root exudates. While the study of N 2 fixation and N transfer within tropical agroforestry has predominately drawn upon methodologies originally developed for temperate agriculture, the unique spatial arrangement and morphological characteristics of crops and N 2 -fixing trees within agroforestry systems present unique challenges for conceptualizing, as well as (sampling and) measuring, the fate of fixed N. Importantly, field studies using natural abundance techniques have shown clear spatial variability in the magnitude of the contribution of fixed N to N acquisition in agroforestry systems (Issah et al. 2015;Meylan et al. 2017). Notable though, while the potential for N 2 -fixing trees and shrubs to enhance soil fertility within agroforestry systems is clear, the transfer of fixed-N to associated crops, however, does not always occur. ...
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Background Disentangling nutrient acquisition strategies between trees and crops is central to understanding positive nutrient interactions in agroforestry systems for improved low-input agriculture. However, as plants are responsive to a complex soil matrix at multiple scales, generalizable diagnostics across diverse agroforests remains challenging. Scope We synthesize research at various scales of the tree-crop interface that are cumulatively hypothesized to underpin nutrient acquisition strategies in agroforestry systems. These scales span the whole root system to fine-scale sites of acquisition actively engaged in biological and chemical interactions with soil. We target vertical and horizontal dimensions of acquisition patterns; localized root-soil dynamics including biological associations; root-scale plasticity for higher acquisition; and nutrient additions via biological nitrogen fixation and deep soil nutrient uplift. We consolidate methodological advances and the effects of environmental change on well-established nutrient interactions. Conclusions Root distribution patterns remain one of the most universal indicators of nutrient acquisition strategies in a range of agroforestry systems, while root functional traits are emerging as an effective root-scale indicator of nutrient acquisition strategy. We validate that in agroforestry systems crop root functional traits reveal bivariate trade-offs similar to, but weaker than, crops in monoculture, with mechanistic links to nutrient acquisition strategies. While interspecific root overlap may be associated with nutrient competition, clear cases of enhanced chemically and microbially meditated processes result in species- and management-specific nutrient facilitation. We argue for agroforestry science to use distinct and standardized nutrient acquisition indicators and processes at multiple scales to generate more nuanced, while also generalizable, diagnostics of tree-crop interactions. And extensive research is needed on how agroforestry practices stabilize key nutrient acquisition patterns in the face of environmental change.
... In addition to N fixation, legumes can also transfer a part of N they fixed to coexisting non-legumes (Murray et al., 2008;Issah et al., 2015). Nitrogen transfer mainly occurs via three pathways: i.e. mycorrhizal networks between legume and non-legume roots (Jalonen et al., 2009), direct uptake of N compounds exuded by legume roots (Ta et al., 1986;Jiang and Zhai 2000), or direct uptake of mineralized N derived from dead legume nodules and senescing root materials (Butler et al., 1959;Pirhofer-Walzl et al., 2012). ...
... Coexisting non-legume species can acquire N fixed by legumes mainly through mycorrhizal networks between their roots and legume roots (Jalonen et al., 2009), by absorption of mineralized N derived from dead root nodules (Butler et al., 1959;Pirhofer-Walzl et al., 2011), or by uptake of N compounds exuded directly by legume roots (Ta et al., 1986;Jiang and Zhai 2000). Therefore, the difference in N transfer could be ascribed to the root characteristics and regrowth strategies of plant species (Temperton et al., 2007;Pirhofer-Walzl et al., 2012), the distance between legumes and non-legumes (Issah et al., 2015) and the N status in soil (Murray et al., 2008). The root architectures and regrowth strategies of non-legume species (Pirhofer-Walzl et al., 2012) partly explain the reason for the difference in their N content and d 15 N when they growing with the legume shrubs and growing alone. ...
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Quantification of symbiotic nitrogen (N) fixation by legume and its N transfer to the neighboring plants is an important prerequisite to understand the N cycling in the N-limited grasslands. The objective of this study is to explore the symbiotic N 2-fixation by a legume shrub (Caragana microphylla) and its N transfer to coexisting non-legumes with the 15 N dilution technique. A paired experimental design (legume vs. non-legume quadrats) was used. A small amount (0.03 g N m À2) of 15 N-labeled KNO 3 (99.5% 15 N enrichment) was evenly distributed in these paired labeled quadrats, with only water additions as the control. One month after the 15 N addition, aboveground parts of C. microphylla and four non-legumes of reference plants (i.e. Cleistogenes squarrosa, Artemisia frigida, Artemisia capillaris and Leymus chinensis) in both labeled and controlled quadrats were harvested. Symbiotic N 2-fixation and transfer to non-legumes were estimated based on the aboveground biomass and its 15 N in these plant species. The N derived from the atmosphere (%Ndfa) of C. microphylla was estimated to range from 84 to 95%. According to the aboveground biomass of C. microphylla, symbiotic N 2-fixation was estimated to be 4.3 g m À2 yr À1 in the mixed C. microphylla community in the temperate grassland of northern China. The amount of N transfer from C. microphylla to their coexisting plants varied with plant species. About half of above N in C. squarrosa derived from C. microphylla, while L. chinensis showed almost no N derived from C. microphylla. Both A. frigida and A. capillaris showed about 36% and 37% of N derived from C. microphylla, respectively. Totally, about 17% of N fixed in the aboveground part of C. microphylla was transferred to its associated species. Considering the amount of N fixed and transferred by C. microphylla, we conclude that C. microphylla plays an important role in the N cycling of temperate grasslands in northern China. ã
... The transfer of plant N derived from N 2 fixation to associated crops was from zero percent to more than 30% in agroforestry systems (Munroe and Isaac 2014). For instance, in a shelterbelt agroforestry system in Saskatchewan, Canada, Issah et al. (2015) showed that crop N originating from leguminous shrubs was up to 40% and decreased with distance from the N 2 fixer shrub. Jalonen et al. (2009) showed that in Lesser Antilles, around 4-14% of the grass Dichantium aristatum's N nutrition was derived from root exudates from the leguminous tree, Gliricidia sepium. ...
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... Biomass production of alien competitor ( (Wu et al., 2017;Yu et al., 2018) and also transfer a part of N which fixed by itself to the coexisting non-legumes (Sierra and Daudin, 2010;Issah et al., 2015). Besides, N enrichment altered the ability of A. fatua and A. adsurgens in the hierarchy of competitive response when they competed with alien species M. sativa (Liu et al., 2013;Vasquez et al., 2008b). ...
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Plant competitive superiority has long been hypothesized to play a key role in the process of plant invasion, which is also affected by global change components such as anthropogenic nitrogen (N) deposition. To explore mutual interactions between alien and native plants at different N available levels, a pot experiment was carried out at a common garden in northern China. Two alien plants (Lolium perenne and Medicago sativa) and four native species (Agropyron cristatum, Elymus dahuricus, Astragalus adsurgens, and Avena fatua) were monocropped and intercropped alternately, under low (0.28 mg day⁻¹) and high (0.84 mg day⁻¹) N available levels. Results demonstrated N enrichment produced higher biomass of both alien and native plants, and species identity of the neighbor modified the growth responses of both native and alien plants to N enrichment. The alien grass L. perenne increased the biomass of native grass E. dahuricus at high N enrichment level. The alien legume M. sativa promoted the growth of native grasses A. cristatum and A. fatua at high N enrichment level, and promoted the growth of native grass E. dahuricus at low N enrichment level. The native legume A. adsurgens increased the growth of the alien grass L. perenne at low N enrichment level, and increased the growth of the alien legume M. sativa at high N enrichment level. Both alien and native legume species showed positive interactions and thus promoted plant growth. Considering the positive effects of the alien legume M. sativa on the growth of native plants and itself, the introduction of M. sativa could be beneficial for biomass production of plant communities and management of degraded temperate grasslands.
... Close proximity is achieved either through direct root contact or mycorrhizal hyphae connections [61]. However, Issah et al. [62] reported that maximum oat productivity was obtained when grown 4 m from caragana shelterbelt compared to 2 m from the shelterbelt. ...
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Below-ground transfer of nitrogen (N) fixed by legume trees to associated non-N2-fixing crops has received little attention in agroforestry, although the importance of below-ground interactions is shown in other ecosystems. We used 15N natural abundance to estimate N transfer from the legume tree Gliricidia sepium (Jacq.) Kunth ex Walp. to C4 grass Dichanthium aristatum (Poir.) C.E. Hubb. in a silvopastoral system, where N was recycled exclusively by below-ground processes and N2 fixation by G. sepium was the sole N input to the system. Finding a suitable reference plant, a grass without contact with tree roots or litter, was problematic because tree roots invaded adjacent grass monocrop plots and soil isotopic signature in soil below distant grass monocrops differed significantly from the agroforestry plots. Thus, we used grass cultivated under greenhouse conditions in pots filled with agroforestry soil as the reference. A model of soil 15N fractionation during N mineralization was developed for testing the reliability of that estimate. Experimental and theoretical results indicated that 9 months after greenhouse transplanting, the percentage of fixed N in the grass decreased from 35% to