Field Crops Research 287 (2022) 108637

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Field Crops Research

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Optimizing arrangement and density in maize and alfalfa intercropping and

the reduced incidence of the invasive fall armyworm (Spodoptera frugiperda)
in southern China
Kaixian Wu a, Chunhe Jiang b, Shiyong Zhou b, *, Hongli Yang c, *
a
Department of Resources and Environment, Moutai Institute, P.O. 564507, Renhuai, Guizhou Province, PR China
b
Faculty of Agronomy and Biotechnology, Yunnan Agricultural University, P.O. 650201, Kunming, Yunnan Province, PR China
c
Department of Brewing Engineering, Moutai Institute, P.O. 564507, Renhuai, Guizhou Province, PR China

A R T I C L E I N F O A B S T R A C T

Keywords: Maize and alfalfa intercropping has yield advantages and is an important cropping system. However, the row
Intercropping arrangement and density design of narrow-strip intercropping systems require improvement for the synergistic
Row arrangement production of maize and alfalfa with higher yield advantages. Moreover, whether this system is adequate for pest
Density
control is unclear. In this study, a three-year field experiment was conducted at Yunnan in southwestern China.
Invasive fall armyworm
Maize
Using monocultures of maize and alfalfa as controls (CK), the experimental design comprised intercropping with
Alfalfa three row arrangements: 2M2A (two maize rows with two alfalfa rows); 1M3A (one maize row with three alfalfa
rows); and 2M3A (two maize rows with three alfalfa rows) under low density with a replacement design and high
density with an additive design for the maize density. The growth and yield of maize and alfalfa were measured;
the total and partial land equivalent ratio, overyielding of intercropping, and the competition ratio of crops were
evaluated. Moreover, the incidence of the invasive fall armyworm (Spodoptera frugiperda, a major agricultural
pest) was investigated. Maize and alfalfa intercropping produced a yield advantage, with 1.31 being the highest
land equivalent ratio. The yield was affected by planting row arrangement and maize density. The land equiv­
alent ratio of 1M3A was significantly higher than that of 2M2A and 2M3A by 20% and 18%, respectively. A high
maize density with additive design increased the land equivalent ratio by 8% compared with a low maize density
with replacement design; the maize had a competitive advantage in intercropping; the row arrangement
significantly affected the alfalfa yield but not the maize yield, while a high maize density with additive design
slightly increased maize yield compared with a low maize density with replacement design, but had no signif­
icant effect on alfalfa yield. In addition, compared with sole maize cropping, intercropping significantly reduced
the incidence of the invasive fall armyworm by 80%. These findings suggested that increasing the alfalfa row
number and maize density improved the land equivalent ratio for narrow-strip intercropping, and that the 1M3A
arrangement under high maize density with an additive design synergistically produced maize and alfalfa.
Importantly, the results also suggest that maize and alfalfa intercropping has the potential to ecologically control
the incidence of the fall armyworm for the sustainable production of maize.

1. Introduction intercropping system pairing a perennial legume with an annual food

Intercropping perennial legume crops with annual grain crops is a
sustainable cropping system that has attracted increasing attention
(Hauggaard-nielsen et al., 2012; Wu. and Wu, 2014; Xu et al., 2022a).
Alfalfa is an important perennial legume crop that plays a key role in
forage and livestock production. Maize is the largest bulk food crop in
the world. The combination of alfalfa and maize is a typical
crop. This system has been widely practiced around the world, such as in
the United States (Smith and Carter, 1998), China (Sun et al., 2019),
Spain (Cela et al., 2011), and Denmark (Hassan et al., 2019), and has
yield, quality, and economic benefits (Smith and Carter, 1998; Zhang
et al., 2011; Sun et al., 2018). Moreover, alfalfa can provide high-quality
forage for livestock and is helpful in promoting more sustainable
livestock-crop systems (Herrero et al., 2010). However, systems for

* Corresponding authors.
E-mail addresses: zhoushiyong_ynau@outlook.com (S. Zhou), alve_0402@163.com (H. Yang).

https://doi.org/10.1016/j.fcr.2022.108637
Received 1 March 2022; Received in revised form 6 July 2022; Accepted 23 July 2022
Available online 1 August 2022
0378-4290/© 2022 Elsevier B.V. All rights reserved.
K. Wu et al.
intercropping maize with alfalfa need improved designs.
One of the most important intercropping design factors is the row
arrangement. Maize and alfalfa strip intercropping can be divided into
wide and narrow row arrangements. In this study, when the strip width
of both maize and alfalfa strips is less than about 1.40 m and the number
of alfalfa and maize rows does not exceed three and two, respectively,
the maize and alfalfa intercropping system is regarded as a relatively
typical example of narrow-strip intercropping. In narrow-strip inter­
cropping, the shading from maize hinders the growth and development
of alfalfa (Osterholz et al., 2018a; Grabber et al., 2021; Patel et al.,
2021). So, maize is usually planted as the main crop, while alfalfa is
mostly planted as a cover crop (Ghosheh et al., 2005; Osterholz et al.,
2018a; Grabber et al., 2021), resulting in the limited production of
forage from alfalfa. Although intercropped alfalfa and maize can be
seasonally staggered, with alfalfa harvested in spring and maize in
summer (Xu et al., 2022a), legume feed cannot be provided in summer
and the production potential of alfalfa may decrease due to the limited
moisture and low temperature in winter and spring. Consequently,
narrow-strip intercropping limits the application of maize and alfalfa
intercropping systems for crop–livestock system development. When the
strip width of intercropping is above 1.40 m, and there are more than
three rows of alfalfa, the combination of maize and alfalfa can be
considered as typical wide-strip intercropping. Many row arrangements
have been reported in wide-strip intercropping, and most have exhibited
yield advantages, including systems with one maize row with four al­
falfa rows (Grabber et al., 2021); two maize rows with three, four, and
five alfalfa rows (Zhang et al., 2013); three maize rows with three or five
alfalfa rows (Sun et al., 2018); four maize rows with four (Nasar et al.,
2020; Sun et al., 2020) or six alfalfa rows (Sun et al., 2019, 2020).
Wide-strip intercropping is conducive to mechanization, but the spatial
contact area between crops is reduced, which reduces the potential to
use interspecific facilitation. For example, in maize and soybean strip
intercropping systems, when the strip width exceeds 2 m, the inter­
cropping yield advantage decreases (Raza et al., 2020). Therefore, the
optimization of row arrangement based on narrow strip width to coor­
dinate the production of maize and alfalfa could be valuable for the
application of maize and alfalfa intercropping systems, especially in
mountainous areas where agricultural mechanization is difficult.
Another important intercropping design factor is the plant density of
the crops. In the intercropping theory, an additive design (where the
crop density in intercropped areas is higher than that in the same area
under sole cropping) and a replacement design (the crop density in
intercropped areas is equal to that at the same area under sole cropping)
are the two main density designs (Banik et al., 2006; Xu et al., 2008).
These schemes represent relatively high-density and low-density de­
signs, respectively. In maize and alfalfa intercropping, the maize density
may be important for yield advantage, because maize yield is sensitive to
density (Ren et al., 2016; Testa et al., 2016; Zhang et al., 2018). How­
ever, the intercropped maize density has been ignored in most of the
present studies in which maize and alfalfa intercropping was performed
using the replacement design (e.g., Zhang et al., 2013; Sun et al., 2018;
Nasar et al., 2020; Shao et al., 2020; Zhang et al., 2020). Few studies
explored the effect of maize density on yield (Grabber et al., 2021; Xu
et al., 2022b). Therefore, this raises the question of whether we can
further improve land productivity for maize and alfalfa narrow-strip
intercropping by increasing the maize planting density.
In addition to intercropping design, the interspecific facilitation be­
tween maize and alfalfa for yield advantage has attracted increasing
attention. First, alfalfa provides nitrogen for maize through nitrogen
fixation and transfer (Nasar et al., 2020; Shao et al., 2021), which re­
duces the nitrogen fertilizer demand of maize (Cela et al., 2011). Alfalfa
can also transfer potassium to maize through root contact and mycor­
rhizal fungi (Habben and Blevins, 1989). Second, the complementary
root distribution between maize and alfalfa increases the absorption and
utilization of nutrient resources in the soil profile (Zhang et al., 2013).
Moreover, the deep roots of alfalfa transfer water from the deep to
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Field Crops Research 287 (2022) 108637
shadow soil layer, which increases the soil water available to maize roots
(Corak et al., 1987), and improves the growth of maize under drought
conditions (Jiang et al., 2015; Sun et al., 2018; Xu et al., 2021), espe­
cially at the maize seedling stage (Hauggaard-nielsen et al., 2008). Al­
falfa can also improve soil quality by decreasing soil bulk density and
penetration resistance through the root residue (Latif et al., 1992). In
addition, the complementary of above-ground architecture and light
requirements can enhance the utilization of light resources (Smith and
Carter, 1998; Feng et al., 2020). However, although the role of crop
diversity in pest control has been widely confirmed in legume-cereal
intercropping systems (Pitan and Odebiyi, 2001; Tanyi et al., 2020),
whether maize and alfalfa intercropping can reduce insect pest damage
is still unclear.
The invasive fall armyworm (Spodoptera frugiperda) is a major
epidemic pest in agriculture. This insect has the characteristics of long-
distance migration, polyphagy, high fecundity, and strong adaptability
(Wan et al., 2021). The fall armyworm reduces the yield and quality of
maize, intensifies the use of pesticides, increases costs, and leads to
economic losses and the reduction of agricultural sustainability (De
Groote et al., 2020; Kassie et al., 2020). The fall armyworm entered
China from Southeast Asia in January 2019 (Huang et al., 2020). There
is an urgent need for ecological methods to control pests (Harrison et al.,
2019). Studies have reported that intercropped beans can control the fall
armyworm on gramineous crops (Hailu et al., 2018; Ndayisaba et al.,
2020; Tanyi et al., 2020). In addition, alfalfa is often used to inhibit pests
in diverse cropping systems (Pitan and Odebiyi, 2001; Ju et al., 2019;
Clemente-Orta et al., 2020). Therefore, we speculated that the maize
yield loss due to the fall armyworm may be reduced in maize and alfalfa
intercropping.
The main purpose of the present study was to determine whether
maize and alfalfa intercropping could produce over-yielding and could
inhibit the fall armyworm in mountainous areas of southern China,
where the fall armyworm has been present since 2019. To this end, we
conducted a three-year field experiment in Yunnan, southwest China.
The experiment was designed with different row arrangements (two
maize rows with three alfalfa rows; one maize row with three alfalfa
rows; two alfalfa rows with two maize rows) with two maize densities:
high density with an additive design and low density with a replacement
design. Our hypotheses were as follows. (1) Increasing the proportion of
alfalfa or the number of alfalfa rows and the maize density under
narrow-strip intercropping would enable the synergistic production of
maize and alfalfa with higher yield advantages. (2) Maize and alfalfa
intercropping would lower the incidence of the fall armyworm but this
function may depend on the row arrangement.
2. Materials and methods
2.1. Experimental site
Field experiments were conducted from August in 2018 to October in
2021 at the Daheqiao Agricultural Experimental Station, Yunnan Agri­
cultural University, Xundian, Yunnan Province, southwest China
(103◦ 16′ 41′ E, 25◦ 31′ 07′ N). This site is located at an altitude of 1860 m
with a mean annual sunshine duration of 2122 h, and a mean frost-free
period of 231 days. The daily air temperature and rainfall during the
experimental period are shown in Fig. 1. The main weather events
limiting crop growth in this district are drought in spring and winter.
The field soil was a silty clay loam (Chinese Classification) with the
potato as the previous crop. The soil (0–20 cm depth) had a pH of 7.91,
with total N, P, and K contents of 1.02 g, 1.74 g, and 9.91 g kg− 1,
respectively. Available N, P, and K were 43.42 mg, 22.18 mg, and 24.36
mg kg− 1, respectively. The soil organic matter content was 18.12 g kg− 1.
2.2. Experimental design
Three cropping systems (maize monoculture, alfalfa monoculture,
K. Wu et al.
Fig. 1. Daily air temperature and rainfall during the experiment
period (2018–2020).
and alfalfa and maize intercropping) were set up in the field experiment.
For intercropping, the narrow-strip design comprised three row ar­
rangements: two maize rows with two alfalfa rows (2M2A; the strip
width was 1.45 m, with the area proportions of maize and alfalfa being
0.528 and 0.472, respectively); one maize row with three alfalfa rows
(1M3A; the strip width was 1.10 m, with the area proportions of maize
and alfalfa being 0.308 and 0.692, respectively); and two maize rows
with three alfalfa rows (2M3A; the strip width was 1.70 m, with the area
proportions of maize and alfalfa being 0.528 and 0.472, respectively)
(Fig. 2). In order to explore the impact of different maize densities on
yield advantage, the three row arrangements for intercropping were
implemented under two maize densities: a low density with a replace­
ment design and a high density with an additive design (increasing 50%
density based on replacement design). Therefore, the density and plant
spacing for intercropped maize between row arrangements were varied
(Table 1). In particular, the density design for 1M3A was based on the
idea of merging two rows in 2M2A and 2M3A into one row. The spacings
between maize intraspecific rows, alfalfa intraspecific rows, and inter­
specific rows were 60 cm, 25 cm, and 30 cm, respectively. The density
of monoculture maize was 72495 ha− 1 with a plant spacing of 23 cm,
values that were determined according to local production practices
(Fig. 2). The field experiment had a total of eight treatments, each with
three replicates. The plots were arranged in a completely randomized
design with 6.0 m × 4.0 m areas.
The selected varieties of maize and alfalfa in the field experiment
were Yunrui 88 (a local maize variety) and Victoria (suitable for
planting in southern China). The alfalfa seed was sown on July 15, 2018,
3
Field Crops Research 287 (2022) 108637
after the harvest of the previous crop (potato). Before alfalfa sowing,
rotary tillage was conducted for all plots. The alfalfa was sown by hand
in soil ditches with a sowing rate of 15 kg ha− 1, 25 cm row spacing,
2–3 cm sowing depth, and topsoil cover. After alfalfa sowing, the plot
was covered with shading nets and watered until seedling emergence.
Before maize sowing, the soil of the maize planting area was tilled three
times using a rotary cultivator (tillage width and depth were 70 cm and
20 cm, respectively). The maize seed was sown in May of 2019, 2020,
and 2021, and plants were thinned at the third-leaf stage. During the
experimental periods in each year, nitrogen fertilizer (N:240 kg ha− 1)
and phosphate fertilizer (P2O5:80 kg ha− 1) were applied to each plot.
For maize, the nitrogen fertilizer (N) was applied by side dressing at the
seeding, jointing, and flowering stages at rates of 72 kg ha− 1,
48 kg ha− 1, and 120 kg ha− 1, respectively. The phosphate fertilizer was
applied once by side dressing at the seeding stage. For alfalfa, both the
nitrogen (N: 60 kg ha− 1) and phosphate (P2O5: 20 kg ha− 1) fertilizers
were applied to the alfalfa through broadcasting after cutting manage­
ment (four times per year). At the maize planting area, manual weeding
was carried out twice a year at the maize 4–5 leaves and 10–13 leaves
stages. In the alfalfa planting area, manual weeding was carried out once
after the emergence of alfalfa in 2018. To test whether intercropping
could control the invasive fall armyworm, no pesticides were used in any
of the plots during the experimental period.
2.3. Samples and measurements
2.3.1. Maize and alfalfa yield and growth parameters
The maize and alfalfa were harvested separately. For maize, four
rows were randomly selected at the crop maturity stage. The ear was
separated from the plant and dried in the sun to a constant weight. The
weight of maize grains was then determined after threshing; then the
yield was calculated. Eight maize plants within each plot were randomly
selected and cut at the stem base. The plant height and ear length, the
stem (at the middle part of the second internode), and the ear diameter
were measured using steel tapes and calipers. Then, each plant was
divided into the stem, leaf, kernel, and cob and dried at 60 ◦ C to constant
weight for measuring the leaf, stem, and grain biomass. The plant
biomass and the harvest index (grain to plant biomass ratio) were
calculated.
For alfalfa, from 22 April 2019, alfalfa was cut four times per year
(three times in 2021 due to drought) to measure the yield and biomass.
The first time was before maize sowing, ensuring that maize seedling
would not be affected. The other cutting events occurred from May to
November (rainy season) and were about 45 days apart. Alfalfa was not
cut from December to April due to the dry season; this ensured the
survival of alfalfa in winter. For cutting, three rows 0.5 m long (three
sides and inside rows in 1M3A and 2M3A, respectively) were randomly
selected in each plot for sample cutting. The stubble height was 5–6 cm.
The cut alfalfa was put into net bags and dried in the sun in order to
measure the total biomass; then, the yield was determined based on the
biomass. The other alfalfa rows in the plot were cut according to the
same standard in sample rows, and then the above-ground branches and
leaves were removed from the plot.
2.3.2. Investigation of the invasive fall armyworm
In 2019 and 2020, the fall armyworm invaded the experimental area.
We investigated and counted the maize plants infested by the fall
armyworm for each plot (except for the alfalfa monoculture). The in­
vestigations were conducted at the seedling, jointing, big bell mouth,
heading, and late flowering stages of maize. To evaluate the impact of
the fall armyworm on maize yield, ten maize plants infested or non-
infested in each maize monoculture plot were randomly selected to
measure their grain yield per plant (in 2020).
K. Wu et al. Field Crops Research 287 (2022) 108637

Fig. 2. Row arrangements of maize and alfalfa in field experiment: (a) 2M2A intercropping (two maize rows with two alfalfa rows); (b) 1M3A intercropping (one
maize row with three alfalfa rows); (c) 2M3A intercropping (two maize rows with three alfalfa rows); (d) sole maize; (e) sole alfalfa; and (f) the photo for
field experiment.

0 indicates a yield advantage for the intercropped crop.

Table 1
The interspecific interaction between maize and alfalfa was evalu­
Design of maize density and plant spacing for field experiment.
ated by the competition ratio (CR) (Zhang et al., 2011). CRma = (Y
Cropping Low density with replace High density with addition
intercrop M/(PM*Y sole crop M))/(Y intercrop A/(PA*Y sole crop A)), where Y
system design design
intercrop M and Y intercrop A denote the yields of intercropped maize and
Maize density Plant Maize density Plant alfalfa, respectively; Y sole crop M and Y sole crop A denote the yield of
(plants ha− 1) spacing (plants ha− 1) spacing
monoculture maize and alfalfa, respectively; PM and PA are the area
(m) (m)
proportions of maize and alfalfa, respectively. An CRma value greater
2M2A 44520 0.31 65700 0.21 than 1 indicates that the competitiveness of maize is greater than that of
intercropping
alfalfa. We also calculated the effect of intercropping on the growth of
1M3A 45480 0.20 60630 0.15
intercropping alfalfa by advantage of side row (ASR) = biomass at side row/biomass at
2M3A 37965 0.31 56055 0.21 the inside row in intercropping with three rows of alfalfa.
intercropping The incidence of pests is an important parameter used to evaluate the
Maize 72495 0.23 – –
occurrence and degree of damage by crop pests. We calculated the rate
monoculture
of fall armyworm infestation on maize (%) = the number of infested
maize plants/total number of maize plants in the plot (Tanyi et al.,
2.4. Data analysis 2020). We also calculated the incidence of fall armyworm at the plot
level (%) = the number of plots infested by fall armyworm/the total
The yield advantages of intercropping were evaluated by the land number of plots for a treatment.
equivalent ratio (LER), including partial LER (PLER) = Y intercrop /Y sole Data were analyzed using SPSS statistical software (version 21). One-
crop and total LER (TLER) = Y intercrop M/Y sole crop M + Y intercrop A/Y sole way analysis of variance (ANOVA) with treatments as fixed factors was
crop A (Rao and Willey, 1980). Here, Y intercrop M and Y intercrop A denoted performed to compare the differences in maize growth and yield, alfalfa
the yields of intercropped maize and alfalfa, respectively; Y sole crop M biomass and yield, LER, overyielding, and incidence of fall armyworm
and Y sole crop A denoted the yields of monoculture maize and alfalfa, between monoculture and intercropping over 3 years. Two-way ANOVA
respectively. An LER value greater than 1 indicates a yield advantage for with row arrangement and maize density as fixed factors was performed
intercropping. to compare the differences in maize growth and yield, alfalfa biomass
We also calculated the overyielding to quantitatively evaluate the and yield, LER, overyielding, and incidence of the fall armyworm be­
yield dominance for each crop. Overyielding (%) = (Y intercrop - P * Y sole tween different intercropping treatments over three years. One-way
crop)/(P * Y sole crop) * 100% (Li et al., 2011a, 2011b). Here, P is the area ANOVA was performed to compare the differences in maize yield be­
proportion of an intercropped crop. An overyielding value greater than tween the fall armyworm infested and non-infested maize plants, and on

4
K. Wu et al. Field Crops Research 287 (2022) 108637

ASR between 2M3A and 1M3A. The years in all ANOVAs were consid­ Table 2
ered as random variables. The significance of the correlation between Analyses of variance (ANOVA) for the yield and growth parameters of inter­
maize yield and incidence of fall armyworm was tested using Pearson cropped maize and alfalfa, and the incidence of invasive fall armyworm on
correlation (two-tailed tests). The normality and homogeneity of vari­ maize as affected by the intercropping row arrangements, maize density and
ances were previously verified; if necessary, natural logarithms (log) of years. LER: land equivalent ratio of intercropping; PLERm and PLERa: partial
land equivalent ratio of maize and alfalfa, respectively; Overyieldingm and
raw data were taken to meet homoscedasticity and normality
Overyieldinga: overyielding of maize and alfalfa, respectively; CRma: competi­
requirements.
tion ratio of maize to alfalfa; ASR: Advantage of side row for alfalfa in inter­
cropping. There was no significant interaction effect between row arrangement
3. Results and maize density on all parameters, so the P values were not showed.
* P < 0.05.
3.1. Yield of maize and alfalfa
Object Parameters Row Maize Year
arrangement density
Intercropping significantly reduced the yield of maize by
Maize Yield 0.448 0.060 0.093
28.60–47.11% (F = 11.616, P < 0.001) and that of alfalfa by Total biomass 0.074 0.061 0.041 *
36.58–59.19% (F = 38.486, P < 0.001) (Fig. 3). For intercropped maize, Harvest index 0.395 0.150 < 0.001 *
row arrangement had no significant effect on maize yield, but the high Ear length 0.186 0.132 0.013 *
maize density with additive design slightly increased maize yield by Ear diameter 0.080 0.124 0.003 *
Plant height 0.138 0.926 0.008 *
18% compared with the low maize density with replacement design (F =
Stem diameter 0.499 0.243 0.058
15.291, P = 0.060; Table 2, Fig. 3). For intercropped alfalfa, the yield Alfalfa Yield 0.015 * 0.352 0.035 *
was not significantly affected by maize density but was significantly Biomass 0.151 0.623 0.020 *
affected by the row arrangement. The alfalfa yield in 1M3A (one row of ASR 0.624 0.705 0.094
Maize and alfalfa LER 0.089 0.034 * 0.591
maize with three rows of alfalfa) was significantly higher than those in
PLERm 0.459 0.037 * 0.271
2M3A (two rows of maize with three rows of alfalfa) and 2M2A (two PLERa 0.008 * 0.294 0.722
rows of maize with two rows of alfalfa) by 29.8% and 52.7%, respec­ Overyieldingm 0.002 * 0.026 * 0.353
tively (Table 2, Fig. 3). There was no significant interaction effect among Overyieldinga 0.236 0.513 0.424
row arrangement, maize density, and year in either intercropped maize CRma < 0.001 * 0.078 0.183
Incidence of Seedling stage 0.183 0.845
or alfalfa yields (P > 0.05).
–
noctuid moth on Jointing stage 0.801 0.355 0.174
maize Trumpet stage 0.781 0.951 0.398
3.2. Biomass and growth of maize and alfalfa Heading stage 0.769 0.740 0.130
Flowering 0.615 0.804 0.173
stage
When compared to monoculture, intercropping significantly reduced
the maize plant biomass (under 1M3A, or maize density with additive
design; F = 8.869, P < 0.001), plant height (F = 29.641, P < 0.001), and found in 1M3A under the high maize density, and this treatment showed
stem diameter (F = 5.680, P < 0.001) but did not affect the ear length a yield advantage of intercropping (i.e., saving 31% land area). The
and diameter, the advantage of side row alfalfa (ASR) or the harvest lowest TLER value (0.99) was found in 2M2A with the low maize density
index (Table 2, Fig. 4). The alfalfa biomass in 1M3A was significantly and had no clear advantage of intercropping (Fig. 5). Row arrangement
lower than that in monoculture, 2M3A, or 2M2A (F = 38.486, did not significantly affect the TLER (Table 2, Fig. 5). The TLER in 1M3A
P < 0.001). In intercropping, neither the row arrangement nor the was significantly higher than that in 2M2A and 2M3A by 20.0% and
maize density significantly affected the biomass or other growth pa­ 18.8%, respectively. The high maize density with an additive design
rameters for both crops (Table 2, Fig. 4). There was no significant significantly increased the TLER compared with the low maize density
interaction effect among row arrangement, maize density, and year in with a replacement design by 8%.
both intercropped maize and alfalfa biomass and growth parameters Furthermore, low maize density with additive design significantly
(P > 0.05). increased the partial land equivalent ratio (PLERm) and the overyielding
of maize (Overyieldingm) but not those of alfalfa (PLERa and Over­
3.3. Intercropping advantage and interspecific relationship yieldinga) when compared to the low maize density with an additive
design. While the PLERa in 1M3A was significantly higher than that in
The highest total land equivalent ratio (TLER) value (1.31) was

Fig. 3. Yield of maize (a) and alfalfa (b) under different cropping systems (2M2A: two maize rows with two alfalfa rows intercropping; 1M3A: one maize row with
three alfalfa rows intercropping; 2M3A: one maize row with three alfalfa rows intercropping; SM: sole cropping) and maize density design (low density with replace
design; high density with addition design). Values (means ± S.D.) with same letters between treatments are not significant different at P < 0.05 by Duncan.

5
K. Wu et al. Field Crops Research 287 (2022) 108637

Fig. 4. Cropping systems (2M2A: two maize rows with two alfalfa rows intercropping; 1M3A: one maize row with three alfalfa rows intercropping; 2M3A: two maize
rows with three alfalfa rows intercropping; SM: sole cropping) and maize density design (low density with replace design; high density with addition design) on
biomass of maize (a) and alfalfa (b), harvest index (c), ear length (e), ear stem (f), plant height (g) and stem diameter (mm) of maize, advantage of side row (ASR) for
alfalfa in intercropping (d). Values (means ± S.D.) with same letters between treatments are not significant different at P < 0.05 by Duncan.

2M2A and 2M3A, but the Overyieldinga in 1M3A was significantly lower the CR in 1M3A and 2M3A was significantly higher than that in 2M2A
than that in 2M2A and 2M3A. All CR (competition ratio of maize to by 76.30% and in 2M3A by 116.36%. There was also no significant
alfalfa) values for the three arrangements were greater than 1, indicating interaction effect among row arrangement, maize density, and year in
that maize was the dominant crop in intercropping designs. Moreover, intercropping advantages and interspecific relationship parameters

6
K. Wu et al. Field Crops Research 287 (2022) 108637

Fig. 5. Effect of cropping systems (2M2A: two maize rows with two alfalfa rows intercropping; 1M3A: one maize row with three alfalfa rows intercropping; 2M3A:
one maize row with three alfalfa rows intercropping; SM: sole cropping) and maize density design (low density with replace design; high density with addition
design) on (a) total land equivalent ratio (TLER), partial land equivalent ratio of (b) maize (PLERm) and (c) alfalfa (PLERm), (d) relative competition of maize to
alfalfa in intercropping (CRma), Overyield of (e) maize (Overyieldingm) and (f) alfalfa (Overyieldinga). Values (means ± S.D.) with same letters between treatments
are not significant different at P < 0.05 by Duncan.

(P > 0.05).
3.4. Incidence of invasive fall armyworm
The investigation showed that the incidence of invasive fall army­
worm on maize was increased from seedling to heading stages in 2019
and 2020 (P < 0.05). However, intercropping compared with mono­
culture significantly reduced the incidence of fall armyworm on maize
during the investigation period (Table 3). The highest incidence of
invasive fall armyworm on maize in monoculture was 15%, while that in
intercropping was only 3%. Meanwhile, intercropping significantly
reduced the incidence of fall armyworm at the maize flowering stage by
80% in the two years. In addition, the incidence of fall armyworm at the
plot level in both 2019 and 2020 was 100% (n = 3) at the flowering
stage in monoculture, while in intercropping in 2019 and 2020 this was
only 66% and 25% (n = 18), respectively. Therefore, intercropping
reduced the average incidence of fall armyworm at the plot level by
53%. Importantly, the controlling effect of intercropping on the inci­
dence of fall armyworm was not significantly affected by row arrange­
ment or maize density (Table 2). Unexpectedly, although the fall
armyworm reduced the yield of plants in monoculture by 12.7%, the
yield difference between infected maize plants (189.33 ± 16.19 g
plant− 1) and non-infected plants (217.07.33 ± 12.93 g plant− 1) was not
significant (F = 5.376, P = 0.081). In addition, there was no significant
correlation between the incidence of fall armyworm and maize yield (for
monoculture: r = − 0.103, P = 0.847; for intercropping with high maize
density: r = − 0.269, P = 0.280; for intercropping with low maize den­
sity: r = 0.043, P = 0.866; for all intercropping treatments: r = 0.457,
P = 0.002).
4. Discussion
4.1. Effect of row arrangement and maize density on crop yield and
interspecific relationships
The TLER (total land equivalent ratio) values in maize and alfalfa
intercropping were greater than 1, suggesting that maize–alfalfa inter­
cropping had a yield advantage (Smith and Carter, 1998; Zhang et al.,
2013; Xu et al., 2022a). Importantly, the experiments demonstrated that
one maize row with three alfalfa rows (1M3A), as a type of narrow-strip
intercropping, had the highest TLER and synergistically produced maize
and alfalfa. These findings could be great significance for the develop­
ment of crop–livestock systems. To our knowledge, few studies have
reported the 1M3A row arrangement. Compared to 2M2A (two maize
rows with two alfalfa rows) and 2M3A (two maize rows with three al­
falfa rows), 1M3A (one maize row with three alfalfa rows) increased the
planting area of alfalfa by 78.8% and 46.6%, respectively. This increased
alfalfa production and reduced the light competition from maize

Table 3
Analyses of variance (ANOVA) for the incidence (%) of invasive fall armyworm on maize as affected by the cropping system in 2019 and 2020. Values are means ± S.D.
* P < 0.05.
Years Cropping systems Growth stage of maize

Seedling Jointing Trumpet Heading Flowering

2019 Sole cropping – 6.4 ± 1.3c 10.8 ± 1.2b 14.8 ± 1.8a 15.1 ± 2.2a
Intercropping – 0.5 ± 0.8b 1.3 ± 1.5b 3.0 ± 2.6a 3.3 ± 2.8a
2020 Sole cropping 1.7 ± 0.9c 5.2 ± 1.0b 8.2 ± 2.9a 8.3 ± 2.0a 8.3 ± 2.0a
Intercropping 0.2 ± 0.6a 0.4 ± 0.9a 1.1 ± 2.7a 1.3 ± 3.1a 1.3 ± 3.1a
Average Sole cropping – 5.8 ± 1.3c 9.5 ± 2.5b 11.6 ± 3.8a 11.7 ± 4.1a
Intercropping – 0.5 ± 0.8b 1.2 ± 2.2b 2.2 ± 3.0a 2.3 ± 3.1a
P-value Cropping system (C) < 0.001 * < 0.001 * < 0.001 * 0.003 * 0.002 *
Year (Y) – 0.103 0.324 0.017 * 0.008 *
C*Y – 0.705 0.064 0.013 * 0.016 *

7
K. Wu et al.
(Osterholz et al., 2018a; Berti et al., 2021). Notably, in this study, there
was no significant difference in alfalfa biomass between 2M2A or 2M3A
and monoculture, but the biomass of 1M3A was significantly lower than
that of monoculture, indicating that the increased alfalfa yield in the
1M3A arrangement was mainly due to the increase in planting propor­
tion. In narrow-strip intercropping systems, increasing the number of
rows and the planting proportion of alfalfa with 1M3A may have many
other benefits, such as reducing the aggravation of alfalfa diseases
caused by high temperature and humidity in summer, which in turn
affect the feed quality (Berti et al., 2021), as well as controlling soil
erosion (Osterholz et al., 2018b; Gamble et al., 2022) and nutrient loss
(Basso and Ritchie, 2005; Osterholz et al., 2021). In addition, the 1M3A
row arrangement could reduce the area of tillage needed for maize,
which would reduce both tillage and labor input so as to improve the
sustainable development of agriculture.
Increasing the row number or strip width of alfalfa will result in the
reduction of the maize planting area. In order to maintain and improve
maize production, we increased the maize density with additive design
to supplement the reduced planting area. The results showed that
increasing maize density slightly improved maize yield when inter­
cropped with alfalfa, even at an initial density of 45480 plants ha− 1
accounting for 27% of the field in 1M3A. A recent study also reported
that increasing maize density could significantly improve the maize
yield and LER in maize and alfalfa intercropping (Xu et al., 2022b).
Compared to sole cropping, there may be more space available for
increasing density under intercropping because there is less light
competition than under monoculture (Wang et al., 2017). In contrast to
Grabber et al. (2021), who reported that increased maize density was
not conducive to the establishment and growth of alfalfa seedling when
one maize row was intercropped with two alfalfa rows, the present study
found that the yield and growth of matured alfalfa were not sensitive to
increased maize density. This difference could be related to the greater
interspecific row spacing and strip width of alfalfa in this study and
showed that increasing maize density in narrow-strip intercropping
could be a feasible way to improve the productivity of maize and alfalfa
intercropping systems. At present, most studies designed a low density
intercropped maize with the replacement design in wide-strip inter­
cropping (Zhang et al., 2013, 2020; Sun et al., 2018; Nasar et al., 2020;
Shao et al., 2020); so, the potential yield advantages in maize–alfalfa
intercropping systems may be underestimated. Nevertheless, we
emphasize that further increasing the maize density in the 1M3A row
arrangement should be undertaken cautiously because a plant spacing
less than 15 cm can intensify intraspecific competition. Excessive
intraspecific competition cannot ensure a sufficient plant emergence
rate and may increase the weak plants (Rossini et al., 2011; Xia et al.,
2019). In addition, the design in the present study did not consider
complete addition (the total density in intercropping was equal to that in
monoculture) due to the workload constraints. The present study also
did not consider different densities for monoculture. Therefore,
increasing intercropped maize density needs to be further explored in
future research.
Under narrow-strip intercropping systems, increasing the alfalfa
width and maize density may intensify the competition between plants.
Although many forms of interspecific facilitation between maize and
alfalfa have been reported (Jellum and Kuo, 1996; Sun et al., 2020), the
competition for resources cannot be ignored. Studies have shown that
increasing the application of nitrogen (Nasar et al., 2020) or phosphorus
and potassium (Jellum and Kuo, 2013) fertilizers can often significantly
improve the advantages of intercropping. In this study, the biomass of
intercropping alfalfa and maize was lower than that of monoculture,
indicating that there was a net competitive effect between the two crops.
The biomass of alfalfa was not affected by maize density, and the edge
effect of alfalfa was not obvious, while the alfalfa biomass of 1M3A and
monoculture was significantly lower than that of 2M2A and 2M3A. This
suggested that alfalfa was subjected to greater intraspecific competition
with the increase of planting proportion, especially the competition for
8
Field Crops Research 287 (2022) 108637
water. Water deficiency is an important limiting factor for alfalfa pro­
ductivity (Ottman et al., 1996), and the experimental site used in the
present study was dry in winter and spring (Fig. 1). For maize, the
biomass in 1M3A was not only lower than that in monoculture maize but
was also lower than that in 2M2A and 2M3A. Moreover, higher maize
density decreased the maize biomass. Consequently, it is necessary to
further reduce the competition between maize and alfalfa, especially in
the 1M3A row arrangement.
From the perspective of relative competitiveness, the results showed
that maize had a competitive advantage in 1M3A, which was not in
accordance with studies based on wide-strip intercropping (Zhang et al.,
2011; Sun et al., 2019) but was consistent with studies based on
narrow-strip maize–alfalfa intercropping (Berti et al., 2021; Xu et al.,
2022a). Therefore, further improvements in maize yield in 1M3A could
be achieved through reducing intraspecific competition. The intraspe­
cific competition in intercropped maize was likely to have risen from
underground interactions rather than aboveground interactions because
the light resources on the vertical row side were sufficient. Thus,
appropriate water and fertilizer management is necessary to further
improve the yield advantage. In addition, the 1M3A row arrangement
may be more suitable in soils with suitable water and fertilizer condi­
tions. The interspecific facilitation between maize and alfalfa can also be
improved to increase maize yield. For example, the lateral root growth
of alfalfa could be limited by deep tillage or vertical tillage at the maize
planting area, thereby reducing water and fertilizer competition (Jellum
and Kuo, 1996; Sun et al., 2018) or allelopathic effects (Li et al., 2005)
exerted by alfalfa on maize, which could promote the release and
transfer of nutrients to maize from alfalfa root residues (Muhammad
et al., 2007; Shao et al., 2020) and stimulate the root growth of alfalfa,
improving the spatial complementary of the root growth of maize and
alfalfa (Shao et al., 2021). Less tillage in the maize planting area may
also improve water use (Ghaffarzadeh et al., 1997). To improve alfalfa
performance, it will be necessary to reduce the shading effect from
maize, such as through selecting the dwarf maize varieties, increasing
the 5–10 cm interspecific row spacing, and applying growth regulators.
4.2. Ecological control of the invasive fall armyworm
The fall armyworm (Spodoptera frugiperda) can pose a serious threat
to maize production in terms of reducing yield and quality, aggravating
the use of pesticides, increasing costs, and leading to economic losses
and the reduction of agricultural sustainability (Kassie et al., 2020).
Unexpectedly, the fall armyworm in our study had a limited effect on
maize yield in monoculture. The reason may be partly attributed to the
short duration of pest infestation, because the incidence of the fall
armyworm did not increase after the maize heading stage (Table 3). The
intercropping near the monoculture plot (Clemente-Orta et al., 2020)
and the pesticide application near our experimental site may reduced
the pest effect of fall armyworm and its duration. Although the maize
yield did not be significantly influenced by the fall armyworm, this study
found that intercropped alfalfa effectively reduced the incidence of the
fall armyworm on maize by 80%. Presently, the main measure adopted
for controlling the fall armyworm is the use of chemical pesticides
(Harrison et al., 2019), an effective method but that has high cost and
negative impacts on the environment (Tambo et al., 2020). The reduced
incidence of the fall armyworm in our study suggested that maize and
alfalfa intercropping can control the infestation of the invasive fall
armyworm, which provides a new potential intercropping system for
low-cost and ecological control of the fall armyworm, and may be sig­
nificant for the comprehensive management of the fall armyworm
(Harrison et al., 2019).
Harrison et al. (2019) reviewed the possible mechanisms of inter­
cropping for reducing fall armyworm infestation. Importantly, legumes
(alfalfa) can provide a habitat for many natural enemies of insect pests
on gramineous crops (maize) (Madeira et al., 2019; Clemente-Orta et al.,
2020; Udayakumar et al., 2021). Indeed, alfalfa has been demonstrated
K. Wu et al.
to provide a habitat for insect natural enemies (Madeira et al., 2019;
Clemente-Orta et al., 2020). However, legumes have been shown to
release bioactive volatiles that can effectively inhibit, poison, and dispel
adult pests or inhibit oviposition, resulting in a "pull-push" effect (Ju
et al., 2019). For example, Lysimachia and lychee, as "avoidance trap"
crops, reduced the occurrence of Spodoptera litura on maize by 168%
(Yeboah et al., 2021). Using Pennisetum as an "avoidance trap" crop
lowered S. litura on maize by 30% (Khan et al., 2018). For controlling the
fall armyworm, studies have reported that intercropped legumes (dwarf
kidney beans and climbing kidney beans) (Tanyi et al., 2020; Udaya­
kumar et al., 2021) and soybeans and peanuts (Hailu et al., 2018)
reduced the occurrence of the fall armyworm by 50% and 40%,
respectively. The control effects in these studies were lower than in our
study, suggesting that legume crop types may play a role in the control
effect for the fall armyworm. It was worth noting that if the pest exhibits
a negative effect on alfalfa, such as aphids and thrips (Kaplin et al.,
2020), the benefit from alfalfa on maize for controlling the fall army­
worm could be neutralized. This possible issue needs to be considered in
future research. In addition to the effects from alfalfa, the changed maize
quality in intercropping may have substantially affected and hindered
the pest. For example, the nitrogen content of maize has been found to
increase when intercropped with alfalfa (Nasar et al., 2021; Xu et al.,
2022b), and studies have shown that the host nutritional status can
affect the growth and reproduction of fall armyworm (Real-Santillán
et al., 2019; Ribeiro et al., 2020).
5. Conclusion
Our study showed that increasing the proportion or row number of
intercropped alfalfa and increasing the maize density enhanced the yield
advantage under narrow-strip intercropping. Importantly, the arrange­
ment of one maize row with three alfalfa rows (1M3A) under a high
maize density with an additive design could promote the synergistic
production of maize and alfalfa and benefit the application of
crop–livestock systems, especially in mountainous area where agricul­
tural mechanization is limited and soil and water loss is serious. The
intercropped alfalfa effectively reduced the incidence of the fall army­
worm on maize, providing an ecological method to control invasive fall
armyworm on maize and other cereals. To improve maize and alfalfa
intercropping using the 1M3A arrangement under high maize density
with additive design, the plant interactions, including interspecific and
intraspecific as well as the aboveground and belowground competition,
need to be elucidated. In addition, efforts should be made to explore the
mechanism of alfalfa and maize intercropping to control fall armyworm.
In practice, the design (the 1M3A row arrangement with high maize
density design) could increase the water consumption and caution
should be used when applying this method in dry areas. Moreover, the
design will not be recommended to adopt in the area where the maize
producing is dominant due to the reduced maize yield. The range of
density that maize can be grown under varying environmental condi­
tions needs to be further clarified. It is suggested that researchers
develop feasible measures (such as determining the optimal tillage
depth, water and nutrient management, and harvest technology) to
reduce the intraspecific competition in maize and the underground
competition between alfalfa and maize.
Funding
This study was financially supported by the National Natural Science
Foundation of China [32160492] and [31401336], and by the Tech­
nology Research and Development Program of ZunYi (HZ[2021]324).
Declaration of Competing Interest
The authors have no relevant financial or non-financial interests to
disclose.
9
Field Crops Research 287 (2022) 108637
Acknowledgments
We thank Xiaoyun Zhang, Chao Wang for their assistance during the
experiment. We also thank LetPub (www.letpub.com) for its linguistic
assistance during the preparation of this manuscript.
Authors’ contributions
Kaixian Wu contributed to the study conception and design. Material
preparation, data collection were performed by Kaixian Wu, Shiyong
Zhou, Chunhe Jiang. Analysis were performed by Hongli Yang. The first
draft of the manuscript was written by Kaixian Wu. Manuscript was
reviewed and edited by Shiyong Zhou and Hongli Yang. All authors
commented on previous versions of the manuscript and approved the
final manuscript.
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