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Interspecific interaction and productivity in a dryland wheat/alfalfa strip

intercropping

Article in Field Crops Research · April 2024

DOI: 10.1016/j.fcr.2024.109335

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Inte rspe cific inte raction and prod uctivity in a dryland whea t/alfalfa
strip inte rcropp ing

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Qianhu Ma a, Yuhuan Wu a, Yanan Liu a, Yuying Shen a, b , Zikui Wang a, b , ⁎

a Stat e Key Labo rato ry of He rb ag e Im pr ov em en t an d Gras slan d Ag ro −e co sy st em s, Co lleg e of Past or al Ag ricu lture Sc ie nc e an d Tech no lo gy , Lanz ho u
Un iver sity, 768 Jiayug ua n We st Rd, Lanz ho u 730020, Ch ina
b Na tion al Fiel d Sc ie nt ific Ob se rv atio n an d Re se arch Stat io n of Gras slan d Ag ro −Eco sy st em s, Qi ng yang 745004, Ch ina

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ARTICLE INFO ABSTRACT

Keywo rd s: Co nt ex t: Inte rcrop ping pa sture with gr ain crop s provide s a strateg y fo r diversifying crop ping syste ms
Bo rde r row effect and es tablish ing a fo unda tion fo r mixed crop - livestoc k farming syste ms .
Dryland farming Ob je ct ives : Investiga ting the effects of strip de sign and nitroge n (N) ap plication on inte rsp ecific inte rac-
Grain and pa sture crop s inte rcrop ping
tion , crop yield, N us e efficien cy (NUE), and land eq uivalent ratio (LER) in dryland whea t/alfalfa inte rcrop -
Land eq uivalent ratio
ping .
Nitroge n us e efficien cy
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Me th od s: We co ndu cted a fo ur-year field expe rime nt, which te sted three inte rcrop s, name ly, 2- row
whea t with 1- row alfalfa (I21), 4- row whea t with 2- row alfalfa (I42), and 8- row whea t with 4- row alfalfa
(I84), as well as so le whea t (SW) and alfalfa (SA). N rate s of 150 (HN) and 75 kg ha −1 (LN) were ap plied fo r
whea t.
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Re su lts: Winte r whea t exhibited ea rlier sp ring gree n- up than alfalfa and ma intained do minance du ring
co - growth. Over the fo ur se aso ns , I21, I42, and I84 resu lted in whea t yield (bas ed on the whea t strip
area ) increa se s of 19.0%, 16.7%, and 24.0%, resp ec tively, unde r HN , and 34.1%, 31.9%, and 43.4% un-
de r LN. Alfalfa growth was co nstrained du ring co - growth bu t reco vered afte r the whea t harvest. Al-
falfa in I42 and I84 ma intained fo rag e yield (bas ed on the alfalfa strip area ) as high as SA. Inte rcrop -
ping increa se d whea t NU E unde r bo th N co ndition s bu t did no t sig nificantly affect alfalfa NU E. I84 at-
tained the high es t LER of 1.01–1.14 and 1.11–1.25 unde r HN and LN resp ec tively. Ne t reven ue of I84
EC

was lower than SA bu t three time s high er than SW .

Co nc lusion : Wide - strip whea t/alfalfa inte rcrop with low- N sh owed the be st pe rfo rma nce, which is
prom ising in diversifying farming syste m and improving ag ricultural su stainab ility.

1. Introd uction mu ltane ou s cultivation of two or mo re crop s in the sa me field, is a

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low−inpu t crop ping syste m that ma ximize s reso urce utilization ,

Ag ricultural inte nsification has long be en rega rded as the pri- en hances produ ctivity, and improves syste m resilien ce
ma ry ap proa ch to ad dres s fo od se curity (Sn ap p et al., 2010). Ho w- (Rase du zzam an and Jens en , 2017; Rodrigue z et al., 2020; Tesh ita et
ever, co nce rns ab ou t the su stainab ility of inte nsive ag riculture al., 2023). Ce real/legu me inte rcrop ping , in pa rticular, offers the
have be en raised du e to its de trime ntal effects on ag ricultural pro- op po rtunity to op timize nitroge n (N) utilization , ma intain high
du ction syste ms , biodiversity, and exce ssive reliance on chem ical yield and qu ality levels, and minimize the ne ed fo r exce ssive fertil-
inpu ts su ch as fertilizers and pe sticide s (Fras ier et al., 2017; Sn ap p izer N inpu ts, thus redu cing po te ntial en vironme ntal impa cts as so -
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et al., 2010; Wang et al., 2023a). To en hance ag ricultural su stain- ciated with inte nsive ag ricultural syste ms (Pelzer et al., 2012; Wang
ab ility and fo od se curity, the transfo rma tion from the prevailing et al., 2021).
ne ar-mo no culture crop ping syste m to a diversified crop ping sys- The co nvention al crop ping syste ms practice d in the Loes s
te m is recog nize d as a prom ising ag ricultural practice (Zo u et al., Platea u of China predo minantly rely on co ntinuou s whea t
2021; Stratton et al., 2022). Inte rcrop ping , which involves the si- (Triticum ae stivu m L.) cultivation and whea t-ma ize (Ze a ma ys L.)

⁎ Co rres po nd ing author at: State Key Labo rato ry of He rbag e Improvem en t and Gras sland Ag ro−e co system s, Co lleg e of Pastoral Ag riculture Science and

Techno logy, Lanzho u University, 768 Jiayug uan We st Rd, Lanzho u 730020, China.
E- ma il ad dr es s: wzk@ lzu.ed u.cn (Z. Wang ).

https://doi.org/10.1016/j.fcr.2024.109335
Received 15 November 2023; Received in revised form 27 January 2024; Accepted 3 March 2024
0378-4290/© 20XX

Note: Low-resolution images were used to create this PDF. The original images will be used in the final composition.
Q. Ma et al. Field Crops Research xxx (xxxx) 109335

rotation (Wang et al., 2020c). Ho wever, thes e practice s, includ ing me nts ob se rved in bo rde r rows, co mpa red to inne r rows or
co ntinuou s crop ping , su mme r-autumn fallowing, and exce ssive mo no crop s, can be attributed to reso urce co mpleme ntarity, which
fertilizer ap plication , have led to se vere en vironme ntal challeng es lead s to weaker co mpe tition from ne ighbo ring crop s of a differen t
su ch as so il po llution (Zhang et al., 2022), so il erosion (Yin et al., sp ec ies when averag ed over time , althou gh it ma y be strong er
2020), and de ep so il de siccation (Ali et al., 2021). He nce, there is an du ring sp ecific pe riod s of co - growth (Wang et al., 2017, 2023b).
urge nt ne ed to sh ift away from the current mo no culture pa tte rns. Go u et al. (2016) co ndu cted a stud y that de mo nstrate d su bstantial
Inte rcrop ping of whea t and alfalfa em erge s as a valuab le so lution yield increa se s in the bo rde r rows of whea t in a whea t/ma ize inte r-
to mitiga te the en vironme ntal prob lems as so ciated with inte nsive crop ping syste m. Similarly, in an oa t (Aven a sa tiva L.)/co mmo n
cultivation , as it offers po te ntial be nefits of en hanced so il fertility vetch inte rcrop ping syste m, Li et al. (2021) fo und that oa ts in the
(Kermah et al., 2017), redu ce d pe st and disea se incide nce (Co ng , bo rde r row yielde d 41%- 52% high er co mpa red to the inne r row.

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2020), and weed su ppres sion (Jung ers et al., 2021). Mo reover, the Ho wever, ne g ative bo rde r row effects can also oc cur in inte rcrop -
sig nificant biolog ical N 2 fixation capa city of alfalfa se rves as an al- ping syste ms . Inte ns e inte rsp ecific co mpe tition within inte rcrop -
te rnative N so urce fo r the crop s, redu cing the de pe nde nce on syn- ping syste ms ma y inhibit the bo rde r row effect of at leas t on e crop ,

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thet ic fertilizer ap plication s (Su n et al., 2018). Co nse qu en tly, inte r- thereb y su ppres sing the yield ad vantage as so ciated with inte r-
crop ping alfalfa (Me d icag o sa tiva L.) with locally ad ap te d gr ain crop ping (Vand erme er, 1989). The sp ecific bo rde r row effects in in-
crop s like whea t would no t on ly provide ad equ ate fo rag e fo r live- te rcrop ping syste ms involving pe rennial legu me pa sture and ce -
stoc k bu t also offer an en vironme ntally su stainab le and prom ising real crop s rema in unclea r.He nce, we hypo thesize that the yield ad -
ag ricultural syste m fo r the region al de velop me nt. vantage of the do minant sp ec ies is ma inly co ntributed by the bo r-
Qu antifying inte rsp ecific co mpe titivene ss is crucial fo r accu- de r rows, and the ne g ative bo rde r row effects on the su bdo minant
rate ly as se ss ing the produ ctivity of ce real/legu me inte rcrop sys- sp ec ies co uld be mitiga te d throug h ap propriate field ma nag e-

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te ms , as inte rsp ecific co mpe tition se rves as the prima ry driver of me nt te chniqu es su ch as N ap plication and strip de sign .
yield ad vantage (Willey and Rao, 1980; Luo et al., 2021). It is co m- We co ndu cted a 4- year field expe rime nt on whea t and alfalfa
mo nly as su me d that ce reals po sse ss a co mpe titive ad vantage over mo no cultures and inte rcrop ping syste ms in this stud y, with the
legu me s, en ab ling them to do minate inte rcrop ping syste ms and ob jectives of : (1) me asu ring growth dynamics of whea t and alfalfa
acce ss mo re reso urce s (Ne ug sc hwandt ne r and Kaul, 2015; Zhang plants and elucida ting the co mpe titive and co mpleme ntary effects
et al., 2021). The pe rfo rma nce of the do minant crop directly influ- of whea t and alfalfa unde r varying strip widt hs and N ap plication
en ce s the overall pe rfo rma nce of the inte rcrop ping syste m (Hu et levels; (2) investiga ting the bo rde r row effects in inte rcrop ping and
al., 2016). For instance , Dh ima et al. (2007) ob se rved that the yield as se ss ing whea t gr ain yield and alfalfa fo rag e yield, and (3) qu an-
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ad vantage in whea t/co mmo n vetch (Vicia sa tiva L.) inte rcrop ping tify N us e efficien cy and land us e ad vantage of whea t/alfalfa inte r-
was prima rily attributed to the su pe rior co mpe titivene ss of whea t. crop ping over so le planting syste ms .
Zhang et al. (2022) high ligh te d the en hanceme nts in yield and wa-
te r us e efficien cy in ma ize/so ybe an (Glycine ma x (L.) Me rr) inte r- 2. Ma terial and Me thod s
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crop ping resu lting from root plas ticity and the te mpo ral and sp a-
tial co mpleme ntarities be twee n the two sp ec ies. Perennial and an- 2.1. Expe rime nt al site
nual crop s exhibit sig nificant dispa rities in te rms of ph ysiolog ical
ec olog y, root sp atial distribution , and nutrition al requ ireme nts Field expe rime nts were co ndu cted at the Qing yang Expe rime n-
(Jeuffroy and Ne y, 1997), su g ge sting the po te ntial fo r ec olog ical tal Station of Lanzho u University (35º 40´N, 107º 51´E, a.s.l 1298 m)
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co mpleme ntarity be twee n thes e type s of crop s. Ho wever, limited from Se pte mbe r 2017 to Se pte mbe r 2021. The expe rime ntal sta-
rese arch has be en co ndu cted on the inte rcrop ping of annual ce re- tion is situated in the ce ntral pa rt of the Loes s Platea u, Gans u
als and pe rennial legu me fo rag es . province , China. The region expe rien ce s a long −term averag e an-
N availab ility plays a sig nificant role in reg ulating inte rsp ecific nual te mpe rature of 10.1°C (from 1981−2020) and an averag e an-
co mpleme ntarity and/or co mpe tition within ce real/legu me inte r- nual rainfall of 568.2 mm . Fig. 1 illustrates the mo nthly to tal pre-
crop syste ms (Zhang and Li, 2003). Hu et al. (2017) disco vered that cipitation and averag e air te mpe rature reco rded du ring the expe r-
low N inpu ts inte nsify inte rsp ecific co mpe tition and en hance ime nt. From whea t so wing to alfalfa harvest in the fo ur growing
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yields throug h the facilitation effect de rived from co mpe tition . se aso ns , the annual precipitation rang ed from 425.1 to 766.0 mm .
Co nversely, exce ssive ap plication of N fertilizer in annual ce real/ So me ba sic so il prop erties were me asu red prior to so wing in 2017.
legu me inte rcrop ping exacerba te s the inhibito ry effect, po sing The so il in the 0–100 cm profile is a silty loam with a high silt co n-
threats to bo th ag ricultural produ ctivity and en vironme ntal su s- te nt of 73.8%- 80.5% and clay co nte nt of 6.6%- 8.8% (Cao et al.,
tainab ility. Ad dition ally, strip widt h is a critical facto r influen cing 2021). So il orga nic carbo n was me asu red us ing H 2SO 4- K2Cr 2O 7 ox-
the pe rfo rma nce of strip inte rcrop ping (Ma hallati et al., 2015; Raza ida tion and titration me thod (Stanislaw and Da vid, 1973), had a
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et al., 2020a). Previou s stud ies have de mo nstrate d that wide r st rip co nte nt of 6.74 g kg −1 ; to tal nitroge n co nte nt was me asu red with
widt hs op timize land us e efficien cy (Raza et al., 2020b; Zhang et Kjelda hl me thod (Brem ne r, 1965), had a co nte nt of 0.69 mg g −1 ;
al., 2022). Ho wever, increa sing strip widt h has also be en ob se rved availab le nitroge n was me asu red with mo lybde num antimo ny co l-
to weaken ligh t co mpleme ntarity be twee n sp ec ies, thereb y redu c- orime tric me thod (Broa db en t, 1981), had a co nte nt of
ing the inte rcrop ad vantage (Raza et al., 2021; Wang et al., 2020a). 23.21 mg kg −1 ; and ph os ph orus was me asu red with Olse n me thod
Ma hallati et al. (2015) repo rted an 18.1% redu ction in the land (Olse n, 1954), had a co nte nt of 1.48 mg kg −1 .
eq uivalent ratio (LER) when increa sing the inte rcrop row ratio from
2:2–5:5 in a ma ize/be an (Ph as e o lus vu lga ris L.) inte rcrop syste m. 2.2. Expe rime nt al de sign
Ne vertheless , the sp ecific trad e- offs as so ciated with N fertilization
and strip de sign in pe rennial legu me pa sture and annual ce real in- The expe rime nt co mprised five differen t planting co mbina-
te rcrop ping syste ms have no t be en thorou gh ly investiga te d. tion s, includ ing so le whea t (SW) and so le alfalfa (SA) as co ntrol
In strip inte rcrop ping , the bo rde r rows of ad jace nt crop strips trea tme nts, as well as whea t/alfalfa inte rcrop ping trea tme nts with
play a crucial role in facilitating inte rsp ecific inte raction s, ma king varying strip widt hs . The inte rcrop ping trea tme nts co nsisted of 2-
the bo rde r row effect a sig nificant facto r in en hancing inte rcrop row whea t and 1- row alfalfa inte rcrop (I21), 4- row whea t and 2-
produ ctivity (Mu nz et al., 2014; Wang et al., 2017). Yield incre- row alfalfa inte rcrop (I42), and 8- row whea t and 4- row alfalfa inte r-

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Q. Ma et al. Field Crops Research xxx (xxxx) 109335

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2245px x 1976px

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Fig. 1. The ma ximu m, minimu m, and me an air te mpe rature (a) as well as the rainfall and so lar radiation (b) reco rded du ring the fo ur se aso ns sp anning
from Se pte mbe r 2017 to Se pte mbe r 2021 at the Qing yang Expe rime ntal Station in Gans u, China.
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crop (I84). The expe rime ntal plot s were arrang ed in an ea st-west 2.3. Me as urem en ts an d calculat io ns
row sp acing co nfig uration . Local whea t variet y ‘T ong ma i 6’ and
alfalfa variet y ‘L on g do ng ’ were se lected fo r this expe rime nt. 2.3.1. Ab ov eg ro un d bi om as s
Wh ea t rows were sp aced 15 cm ap art, while alfalfa rows were The ab ovegrou nd bioma ss of whea t and alfalfa was as se ss ed at
sp aced 30 cm ap art. In the inte rcrop plot s, the row sp acing of bo th reg ular inte rvals of 15–20 da ys from em erge nce until the en d of
crop s was the sa me as in the mo no crop , and the inte rcrop rows of the growing se aso n. For bo th crop s, 0.5- me te r-long se ction s of
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whea t and alfalfa were se t at 22.5 cm ap art (Fig. 2). Two N fertilizer rows were sa mpled within ea ch plot . In so le- crop pe d plot s, three
trea tme nts, name ly high nitroge n (HN) and low nitroge n (LN), rows were chose n fo r sa mpling, while in inte rcrop ping plot s, on e
were em ployed fo r whea t in the expe rime nt. In the HN trea tme nt, co mplet e strip of whea t and alfalfa was se lected . The sa mpled
whea t rece ived a ba sa l ap plication of 75 kg N ha −1 and 150 kg plants were placed in an oven at 105 °C fo r 30 minutes to halt bio-
P 2O 5 ha −1 be fo re so wing, alon g with an ad dition al 75 kg N ha −1 log ical activity, fo llowed by drying at 75 °C until reaching a co n-
to pdres sing at the jointing stag e in ea ch se aso n. In the LN trea t- stant weight.
me nt, the N fertilizer ap plication fo r whea t was redu ce d to
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37.5 kg N ha −1 at so wing and 37.5 kg N ha −1 fo r to pdres sing. Un- 2.3.2. Wh ea t gr ain yiel d an d alfalfa fo rage yiel d
de r bo th HN and LN, alfalfa was fertilized with 75 kg N ha −1 and At crop harvesting, all plants within three on e− me te r−long
50 kg P 2O 5 ha −1 be fo re so wing in 2018, while 50 kg P 2O 5 ha −1 was rows were ma nually cut at grou nd level in the so le crop plot s. In in-
ap plied be fo re regrowth in the sp ring of 2019, 2020, and 2021. The te rcrop ping plot s, three on e- me te r-long strips co mprising bo th
so wing, harvesting, and fertilizing da te s are prese nted in Table 1. whea t and alfalfa were harvested , and the plants in the bo rde r
The dime nsion s of the SW , SA , and I21 plot s were 3×6 m 2. There rows and inne r rows were me asu red se parate ly. Wh ea t gr ains were
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were five co mplet e inte rcrop ping strips (includ ing on e whea t strip air dried until their wate r co nte nt reache d ap proxima te ly 14%. Ini-
and on e alfalfa strip ea ch) in I21 plot as ea ch strip was 1.2- m- wide . tially, the ea r de nsity of whea t within the en tire sa mpling area was
At leas t three co mplet e inte rcrop ping strips sh ou ld be includ ed in de te rmine d. Su bse qu en tly, te n ea rs were co unte d to de te rmine
the wide - strip inte rcrop ping plot s to ma ke the so il and plant sa m- the averag e numbe r of gr ains pe r ea r, and all the harvested sa m-
pling represe ntative (Li et al., 2020; Wu et al., 2023). On e co mplet e ples were thresh ed to extract gr ains . Finally, all whea t gr ains were
strip was 1.2 and 2.4 m fo r I42 and I84 resp ec tively, fo ur and three weighe d to de te rmine the gr ain yield. The thou sa nd - gr ain weight
strip were de sign ed in ea ch plot , so the area fo r I42 plot me asu red of whea t was calculate d by dividing the to tal numbe r of gr ains by
4.8×6 m 2, and that fo r I84 plot me asu red 7.2×6 m 2. The expe ri- their to tal weight within the en tire sa mpling area . For alfalfa hay
me ntal plot s did no t rece ive irriga tion , and ot he r field ma nag e- yield, fres h weight me asu reme nts were taken, and su bse qu en tly,
me nt practice s fo llowed local produ ction proto co ls. the sa mples were su bjected to oven - drying at 105 °C fo r 30 min-
utes to halt biolog ical activity fo llowed by drying at 75 °C until a
co nstant weight was achieved .

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Q. Ma et al. Field Crops Research xxx (xxxx) 109335

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2245px x 1980px

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Fig. 2. The sc he ma tic represe ntation of the whea t/alfalfa inte rcrop ping syste m unde r variou s strip widt h trea tme nts. SW , so le whea t; SA , so le alfalfa. I21,
I42, and I84 de pict inte rcrop s co nsisting of two rows of whea t inte rcrop pe d with on e row of alfalfa, fo ur rows of whea t inte rcrop pe d with two rows of al-
falfa, and eigh t rows of whea t inte rcrop pe d with fo ur rows of alfalfa, resp ec tively.
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Table 1
So wing, harvesting and fertilizing da te s (dd/mm /yyyy) of whea t and alfalfa in ea ch growing se aso n from 2017 to 2021.
Growing se as on Wh ea t Alfalfa

So wing Harves ting Top dres sing So wing

1st cut 2nd cut 3nd cut Top dres sing
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2017−18 27/09/2017 27/06/2018 19/04/2018 11/04/2018

29/7/2018 01/10/2018 07/08/2018
2018−19 23/09/2018 29/06/2019 20/04/2019 01/06/2019 21/7/2019 10/09/2019 20/04/2019
2019−20 21/09/2019 26/06/2020 21/04/2020 23/05/2020 08/07/2020 10/09/2020 21/04/2020
2020−21 27/09/2020 27/06/2021 20/04/2021 29/05/2021 08/07/2021 29/08/2021 20/04/2021

2.3.3. Inte rs pe cific co mp et itio n the inte rcrop ping syste m is mo re produ ctive than the resp ec tive
The inte rsp ecific co mpe tition is qu antified us ing the co mpe ti- mo no crop s. Co nversely, an LER value less than 1 sig nifies an inte r-
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tion ratio (CR), which se rves as a me tric fo r evaluating the co mpe t- crop disad vantage . The pa rtial land eq uivalent ratio (pLER) fo r
itive streng th of differen t sp ec ies in inte rcrop ping syste ms (Willey whea t (pLERW ) and alfalfa (pLERA) was calculate d to as se ss the
and Rao, 1980). The CR reflec ts the exte nt to which on e crop co m- co ntribu tion of ea ch crop to the overall produ ctivity of the inte r-
pe te s with the ot he r in an inte rcrop pe d en vironme nt. It is de te r- crop ping syste m. LERs are calculate d us ing the fo llowing eq ua-
mine d as : tion s:
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Image (1) Image (2)

Image (3)
where CR W- A is the co mpe titive ratio of whea t relative to alfalfa;
Image (4)
BYIW and BYIA are the bioma ss yield of whea t and alfalfa in inte r-
crop ping , resp ec tively, while BYSW and BYSA are the bioma ss yield where YIW and YIA are the yields of whea t and alfalfa in the in-
of whea t and alfalfa in mo no culture; P W and P A are the propo rtion s te rcrop ping syste m, resp ec tively, and YSW and YSA are the yields of
of the land area oc cupied by whea t and alfalfa in the inte rcrop - whea t and alfalfa in so le crop s.
ping , resp ec tively. A CR value grea te r than 1.0 indicate s that whea t
exhibits co mpe titive ad vantage over alfalfa in the inte rcrop ping 2.3.5. N us e indi cato rs
syste m. Su b- sa mp les fo r de te rmination of N were grou nd and pa ss ed
throug h a 1 mm sieve, and then the N co nce ntration in the sa m-
2.3.4. Land eq uivale nt ratio ples was de te rmine d us ing the Kjelda hl me thod (Brem ne r, 1965).
The land eq uivalent ratio (LER) is a me tric us ed to as se ss the N us e efficien cy (NUE) is calculate d as a qu otien t be twee n crop
produ ctivity and efficien cy of inte rcrop ping syste ms and evaluate yield (Y, whea t gr ain yield or alfalfa bioma ss yield) and N up take
the utilization of land reso urce s (Li et al., 2001). An LER value (de Wit, 1953):
grea te r than 1.0 indicate s an inte rcrop ad vantage , indicating that

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Q. Ma et al. Field Crops Research xxx (xxxx) 109335

2.4. Stat istical an alys is

Image (5)
All statistical analyse s were carried ou t us ing SPSS 25.0 (SPSS
Partial facto r produ ctivity of ap plied N (PFPN ) was calculate d as
Inc., Chicag o, IL, USA). Analysis of variance (ANO VA) was us ed to
a qu otien t be twee n crop yield (Y, whea t gr ain yield or alfalfa bio-
as se ss trea tme nt effects on bioma ss yield, gr ain yield, whea t yield
ma ss yield) and the am ou nt of N ap plication (N ap p , Shi et al., 2017):
co mpo ne nts, co mpe titive ratio, NU E, PFP N , NER, LER, and ne t rev-
(6) en ue . Leas t sig nificant differen ce s (LSD ) were us ed to se parate
Image
trea tme nt differen ce s in me ans at 0.05 level. The univariate Ge n-
N eq uivalent ratio (NER) is de fine d as the am ou nt of nitroge n eral Linea r Mo de l proce du re was us ed to as se ss the effects of in-
fertilizer us ed by mo no crop ping crop s to achieve the sa me yield as te rcrop ping trea tme nt, nitroge n fertilizer level and their inte rac-

F
in the inte rcrop ping syste m (Li et al.,2020): tion .

Image (7) 3. Results

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where N IW and N IA are N fertilizer inpu t pe r unit area of whea t 3.1. Plan t gr ow th an d inte rs pe cific co mp et itio n
and alfalfa in the inte rcrop ping syste m, resp ec tively. N SW and N SA
are N fertilizer inpu t pe r unit area of whea t and alfalfa, resp ec - Inte rcrop ping had a sig nificant po sitive impa ct on the growth
tively. The NER value >1 indicate s that inte rcrop ping sa ves N fer- of whea t, as evide nced by the high er bioma ss yield of inte r-
tilizer. crop pe d whea t co mpa red to so le whea t across mu ltiple sa mpling
da te s in all fo ur se aso ns , rega rdless of N ap plication (Fig. 3). In the
2.3.6. Ne t re ve nu e

PR
initial year of alfalfa es tablish me nt (2018), whea t in the I21 and I42
To calculate the ne t reven ue of ea ch trea tme nt, the prices of inte rcrop ping co nfig uration s expe rien ce d the grea tes t facilitation
whea t gr ain and alfalfa fo rag e were de te rmine d by the actual ma r- unde r HN and LN co ndition s, resp ec tively. Su bse qu en tly, in the
ket price (Table S1). The co st of se ed , fertilizer, tillag e, so wing and fo llowing three years, whea t in the I84 inte rcrop ping co nfig uration
co nstant and harvesting were es tima te d ba se d on local rate s (Raza exhibited the high es t de gree of facilitation unde r bo th N ap plica-
et al., 2021; Mu et al., 2023), which were sh own in Table S2. Total tion s.
reven ue was calculate d by mu ltiplying the me asu red yield with the Wh ea t exhibited do minance in the co mpe tition with alfalfa
yearly local ma rket prices of whea t gr ain and alfalfa fo rag e. Ne t du ring the co - growth pe riod , as indicate d by the CR W- A value s
D
reven ue was de te rmine d by su btracting all inpu ts from the gros s co nsiste ntly exce ed ing 1.0 across the fo ur se aso ns , irresp ec tive of
reven ue (Raza et al., 2021). N ap plication levels (Table 2). Co nse qu en tly, the growth of alfalfa
was co nstrained (Fig. 4). The es tablish me nt of alfalfa face d sig nifi-
cant limitation s in the initial se aso n of 2018, bu t it gr ad ually reco v-
TE
EC
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2245px x 2110px
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Fig. 3. Plant growth of whea t in differen t se aso ns : (a) 2017–18, (b) 2018–19, (c) 2019–20, and (d) 2020–21. HN , high N fertilization ; LN, low N fertilization ;
SW , so le whea t; I21, I42, and I84 represe nt inte rcrop s with two- row whea t and on e- row alfalfa, fo ur-row whea t and two- row alfalfa, and eigh t-row whea t
and fo ur-row alfalfa, resp ec tively.

5
Q. Ma et al. Field Crops Research xxx (xxxx) 109335

Table 2 3.2. Wh ea t yiel d an d th e bo rd er ro w ef fe ct s

Co mpe titive ratio of whea t relative to alfalfa du ring the co - growth pe -
riod in whea t/alfalfa inte rcrop ping . I21, I42, and I84 represe nt inte rcrop s Inte rcrop ping had a po sitive impa ct on whea t gr ain yield (Fig.
with two- row whea t and on e- row alfalfa, fo ur-row whea t and two- row 5). On averag e across the fo ur se aso ns , whea t yield in I21, I42, and
alfalfa, and eigh t-rows whea t and fo ur-row alfalfa, resp ec tively. Differen t I84 exhibited improveme nts of 19.0%, 16.7%, and 24.0%, resp ec -
lowercase lette rs indicate sig nificant differen ce s am on g differen t plant-
tively, co mpa red to so le whea t unde r HN . The yield ad vantage was
ing pa tte rns at α =0.05.
even mo re prono unce d unde r LN, with improveme nts of 34.1%,
Growing planting High N Low N P value
se as on pa ttern ap plication ap plication
31.9%, and 43.4% in I21, I42, and I84, resp ec tively.
2017–18 I21 4.36a 5.89a Planting pa ttern The bo rde r row effect played a crucial role in the yield increa se
<0.001 ob se rved in I42 exce pt fo r 2019–20, whea t in bo rde r row ac-

F
I42 3.72b 5.44b N ap plication<0.001 co unte d fo r 70.1% and 86.4% of the yield improveme nt (with a half
I84 2.08c 2.75c Planting pa ttern ×N whea t strip area ) unde r HN and LN resp ec tively over the fo ur se a-
ap plication <0.001
so ns . In I84, bo th bo rde r and inne r rows ou t yielde d the so le whea t

OO
2018–19 I21 1.17b 1.23a Planting pa ttern=
0.018 in all of the crop se aso ns , althou gh the bo rde r row ou t yielde d the
I42 1.09b 1.61a N ap plication=0.028 inne r rows on ly in the first se aso n 2017–18. The yield ad vantage of
I84 1.53a 1.68a Planting pa ttern ×N whea t in I21 over so le whea t gr ad ually diminishe d with the du ra-
ap plication =0.166
tion of inte rcrop ping .
2019–20 I21 1.12b 1.76a Planting pa ttern
<0.001
I42 1.08b 1.02b N ap plication=0.002 3.3. Wh ea t yiel d co mp on en ts an d th e bo rd er ro w ef fe ct s
I84 1.55a 1.52a Planting pa ttern ×N

PR
ap plication=0.012 The ea r de nsity of whea t was sig nificantly influen ce d by inte r-
2020–21 I21 1.35a 1.57a Planting pa ttern
crop ping , with increa se s of 8.6%, 13.5%, and 15.2% in I21, I42, and
<0.001
I42 1.15b 1.06c N ap plication=0.010 I84, resp ec tively, unde r HN over the fo ur se aso ns ; unde r LN, it in-
I84 1.16b 1.35b Planting pa ttern ×N crea se d by 21.3%, 23.3%, and 26.6% (Fig. 6a). Co mpa red to the in-
ap plication=0.008 ne r row in inte rcrop ping strip, whea t ea r de nsity in I42- B increa se d
by 10.2% and 21.1% unde r HN and LN, resp ec tively. Positive bo r-
ered afte r the harvest of whea t. In the su bse qu en t se aso ns , al- de r row effect on plant ea r de nsity in I84 was ob se rved on ly in the
thou gh the growth of inte rcrop pe d alfalfa was so me what su p- initial year. The effects of inte rcrop ping on whea t gr ain numbe r
D
pres se d, it rema ined co mpa rable to that of mo no culture alfalfa, pe r ea r we re mo stly insign ificant (Fig. 6b). Similarly, inte rcrop ping
with the exce ption of the final se aso n of 2020–21, where inte r- also led to an increa se in the 1000- gr ain weight of whea t, with av-
crop pe d alfalfa exhibited lower bioma ss co mpa red to mo no cul- erag e improveme nts of 12.3%, 11.4%, and 10.8% in I21, I42, and
I84, resp ec tively, unde r HN , and 6.6%, 6.7%, and 6.3% unde r LN
TE

ture alfalfa.
(Fig. 6c). The bo rde r row effects on 1000- gr ain weight were on ly
evide nt in the initial year.
EC
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2495px x 1746px
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Fig. 4. Plant growth of alfalfa at differen t growth stag es in: (a) 2017–18, (b) 2018–19, (c) 2019–20, and (d) 2020–21. HN , high N fertilization ; LN, low N fer-
tilization ; SA , so le alfalfa; I21, I42, and I84 represe nt inte rcrop s with two- row whea t and on e- row alfalfa, fo ur-row whea t and two- row alfalfa, and eigh t-
row whea t and fo ur-row alfalfa, resp ec tively.

6
Q. Ma et al. Field Crops Research xxx (xxxx) 109335

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2495px x 1822px

OO
PR
Fig. 5. Wh ea t gr ain yield in so le and inte rcrop ping syste ms in: (a) 2017–18, (b) 2018–19, (c) 2019–20, and (d) 2020–21. SW , so le whea t; I21, I42, and I84
represe nt inte rcrop s with two- row whea t and on e- row alfalfa, fo ur-row whea t and two- row alfalfa, and eigh t-rows whea t and fo ur-row alfalfa, resp ec -
D
tively. I42- I and I42- B me an inne r row and bo rde r rows in I42 resp ec tively, I84- I and I84- B me an inne r and bo rde r rows in I84. N, P and N×P sh ow the ef-
fects of N ap plication , planting pa tte rn and their inte raction . Differen t lette rs indicate sig nificant differen ce s am on g differen t inte rcrop ping trea tme nts
at α =0.05. “⁎ ” and “⁎ ⁎” indicate sig nificant and very sig nificant differen ce s at α =0.05 and α =0.01 resp ec tively, and ns me ans no n- sign ificant dif-
feren ce at α = 0.05.
TE

3.4. Fo rage yiel d of alfalfa an d th e bo rd er ro w ef fe ct s NU E. Unde r HN co ndition s, alfalfa NU E in I21, I42 and I84 was im-
proved by 1.8–8.9% co mpa red to SA , while unde r LN co ndition s,
In the initial year of es tablish me nt, inte rcrop ping had a sig nifi- the co rresp on ding improveme nts rang ed in 0.2–7.0%. There was
cant ne g ative impa ct on the bioma ss yield of alfalfa du ring the co - no bo rde r row effect of I84 on alfalfa NU E.
growth pe riod (P<0.05, Table 3). Ho wever, afte r whea t harvest, the Inte rcrop ping sh owed N fertilization us e ad vantage co mpa red
EC

bioma ss yield of alfalfa in the inte rcrop ping trea tme nts reco vered . to so le crop ping syste ms , as all of the inte rcrop ping trea tme nts
Ultima te ly, fo rag e yield in I21, I42 and I84 was redu ce d by 63.0%, achieved NER value s exce ed ing 1.0 (Table 5). I84 ob tained the
45.8% and 1.6% co mpa red to so le alfalfa unde r HN , resp ec tively, high es t NER value s in the fo ur years, with averag es of 1.48 and
by 66.8%, 39.2% and 3.4% unde r LN, resp ec tively. In the su bse - 1.68 unde r HN and LN, resp ec tively. Mo reover, redu cing N fertil-
qu en t three years, inte rcrop ping co ntinue d to redu ce the bioma ss izer ap plication increa se d the NER of ea ch trea tme nt. The NER in-
yield of alfalfa du ring the co - growth pe riod . Ho wever, in the I42 crea se d by an averag e of 11.8%, 12.6% and 12.4% unde r LN in I21,
RR

and I84 trea tme nts, the bioma ss yield of alfalfa reco vered afte r I42 and I84, resp ec tively.
whea t harvest, while I21 further redu ce d the bioma ss yield. Ulti-
ma te ly, throug hou t the growing se aso n, all inte rcrop ping trea t- 3.6. LER
me nts resu lted in redu ce d bioma ss yield of alfalfa. We sh ou ld no te
that alfalfa in I42 and I84 slightly improved alfalfa yield or ma in- Inte rcrop ping sh owed land us e ad vantage co mpa red to so le
tained a yield as high as so le alfalfa unde r bo th N trea tme nts. crop ping syste ms , as all of the inte rcrop ping trea tme nts achieved
LER value s exce ed ing 1.0, exce pt fo r I21 and I42 unde r bo th N
CO

3.5. N us e ef ficien cy rate s in 2018 and I21 unde r HN in 2021 (Table 6). I84 ob tained the
high es t LER value s throug hou t the fo ur se aso ns , averag ed at 1.10
Inte rcrop ping had a po sitive impa ct on whea t NU E (Table 4). and 1.20 unde r high and low N rate s, resp ec tively. Mo reover, co m-
Across the fo ur se aso ns , the NU E of whea t in I21, I42, and I84 im- pa red to high N rate , the LER unde r low N increa se d on averag e by
proved by 3.9%, 2.2%, and −1.2%, resp ec tively, co mpa red to SW 6.1%, 12.1% and 9.7% in I21, I42 and I84, resp ec tively. Wh ea t
unde r HN co ndition s. The NU E ad vantage was even mo re pro- sh owed inte rcrop ping ad vantage in all fo ur years, with its pa rtial
no unce d unde r LN co ndition s, with improveme nts of 13.8%, 4.1%, LER exce ed ing its planting pe rce ntag e (0.5).
and 14.2% in I21, I42, and I84, resp ec tively. Inte rcrop ping also sig -
nificantly increa se d the PFP N of whea t; ho wever, increa sing N ap - 3.7. Ne t re ve nu e
plication sig nificantly redu ce d the PFP N . The bo rde r row effect
co ntributed su bstantially to the increa se of whea t PFP N ob se rved In the initial year of es tablish me nt, on ly I84 sig nificantly im-
in I42. proved the ne t reven ue co mpa red to SA unde r bo th HN and LN
In the initial year of es tablish me nt, inte rcrop ping had a sig nifi- (Fig. 7). Co mpa red with SW , I21 and I42 redu ce d ne t reven ue while
cant ne g ative impa ct on alfalfa NU E (Table 4). Ho wever, in the su b- I84 improved ne t reven ue unde r bo th N rate s. In the su bse qu en t
se qu en t three years, inte rcrop ping ma rginally improved alfalfa three years, unde r bo th HN and LN, inte rcrop ping sig nificantly im-

7
Q. Ma et al. Field Crops Research xxx (xxxx) 109335

F
OO
2911px x 2863px

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D
TE
EC

Fig. 6. Ear de nsity (a), gr ain numbe r pe r ea r (b), and 1000- gr ain weight (c) of whea t in so le and inte rcrop ping syste ms unde r differen t N rate s in 2017–18,
2018–19, 2019–20 and 2020–21. SW , so le whea t; I21, I42, and I84 represe nt inte rcrop s with two- row whea t and on e- row alfalfa, fo ur-row whea t and two-
row alfalfa, and eigh t-rows whea t and fo ur-row alfalfa, resp ec tively. I42- I and I42- B me an inne r row and bo rde r rows in I42 resp ec tively, I84- I and I84- B
me an inne r and bo rde r rows in I84. N, P and N×P sh ow the effects of N ap plication , planting pa tte rn and their inte raction . Differen t lette rs indicate sig -
RR

nificant differen ce s am on g differen t inte rcrop ping trea tme nts at α =0.05. “⁎ ” and “⁎ ⁎” indicate sig nificant and very sig nificant differen ce s at α =
0.05 and α =0.01 resp ec tively, and ns me ans no n- sign ificant differen ce at α =0.05.

proved ne t reven ue co mpa red to SW . Co mpa red with SA , inte r- ea rlier planting and faster re- gree ning in the sp ring co mpa red to
crop ping had a lower ne t reven ue . On averag e across the fo ur se a- alfalfa. Early crop s us ually have a grea te r plant he ight and are ad -
so ns , the ne t reven ue in I21, I42, and I84 improved by 134.6%, vantage ou s in ligh t co mpe tition . Ad dition ally, the ea rly de velop -
CO

169.4%, and 169.3%, resp ec tively, co mpa red to SW unde r HN , and me nt of the root syste m allows fo r grea te r acce ss to reso urce s,
by 240.8%, 344.8%, and 329.3% unde r LN. I84 and I42 sh owed even from ne ighbo ring plant strips . This me chanism is co mmo n in
co mpa rable ne t reven ue (P>0.05), which were also no t sig nifi- relay inte rcrop ping . For exam ple, previou s stud ies on whea t/ma ize
cantly differen t be twee n N levels (P>0.05). relay inte rcrop ping syste ms have repo rted that the ea rlier es tab-
lish me nt of whea t provide s a co mpe titive ad vantage fo r en viron-
4. Discussion me ntal reso urce s over ma ize, resu lting in sig nificant su ppres sion
on ma ize ea rly growth (Li et al., 2001; Ma et al., 2020). Se con dly,
4.1. Inte rs pe cific co mp et itio n an d yiel d pe rfor ma nc e in wh ea t an d ce real crop s are se nsitive to N and their growth can be facilitate d
alfalfa st rip inte rcro pp ing by the su pply of N from legu me crop s. This is su ppo rted by previ-
ou s stud ies have sh own that ce real- legu me inte rcrop ping effec-
The strong er co mpe titive crop s do minate the inte rcrop ping tively improves the yield of ce real co mpo ne nts (Li et al., 2021;
syste m and have acce ss to mo re reso urce s (Ne ug sc hwandt ne r and Wang et al., 2020b; Latati, Do kukin, Ao uiche, Rebo uh, Takouachet ,
Kaul, 2015). In ou r st ud y, winte r whea t do minate d the ec olog ical Hafnao ui, Hamd ani, Bacha, Ou nane , 2019). The co mpe tition fo r re-
niche in the co - growth pe riod and was mo re co mpe titive than al- so urce s be twee n the two co mpo ne nts in inte rcrop ping de pe nd s
falfa. The co mpe titive ad vantage of whea t was prima rily du e to on the availab ility of reso urce s and the growth of the crop s (Co rre-

8
Q. Ma et al. Field Crops Research xxx (xxxx) 109335

Table 3 The yield ad vantage of inte rcrop pe d whea t was ma inly de -

Alfalfa bioma ss yield of differen t trea tme nts be fo re and afte r whea t har- pe nd s on the su bstantial increa se in ea r de nsity and 1000- gr ain
vest and to tal alfalfa bioma ss yield throug hou t the growing se aso n in weight. Wh ea t returns gree n ea rlier than alfalfa, resu lting in inte r-
2017–18, 2018–19, 2019–20 and 2020–21. SA , so le alfalfa; I21, I42, and crop pe d whea t be ing ab le to inte rce pt mo re ligh t than so le whea t,
I84 represe nt inte rcrop s with two rows of whea t inte rcrop pe d with on e which co ntributes to whea t tillering and thus improves the whea t
row of alfalfa, fo ur rows of whea t inte rcrop pe d with two rows of alfalfa,
ea r de nsity (Go u et al., 2016; Mu et al., 2023). The radiation ad van-
and eigh t rows of whea t inte rcrop pe d with fo ur rows of alfalfa; Differen t
tage of inte rcrop pe d whea t in bo rde r rows is mo re ob viou s, as it
lowercase lette rs indicate sig nificant differen ce s am on g differen t inte r-
crop ping trea tme nts at α =0.05, resp ec tively. captures mo re ligh t and increa se s ph oto synthesis, which in turn
Growing Planting High N Low N
lead s to an increa se in the ea r de nsity of whea t (Wang et al., 2017).
se as on pa ttern Be fo re Afte r Total Be fo re Afte r Total The increa se in 1000- gr ain weight in inte rcrop pe d whea t is du e to

F
whea t whea t whea t whea t the ad vantage of utilizing natural reso urce s su ch as so il wate r and
harves t harves t harves t harves t radiation du ring the reprodu ctive growth (Zhang et al., 2022). Af-
2017−18 SA 2.2a 3.3ab 5.4a 2.2a 3.3b 5.5a te r the first cut of alfalfa, the co mpe titive ab ility of whea t is further

OO
I21 0.6c 1.4c 2.0b 0.4c 1.4c 1.8c
improved (Table 2), lead ing to a high er growth rate of whea t, ac-
I42 0.7c 2.3b c 2.9b 0.5c 2.9b 3.4b
I84 1.2b 4.2a 5.3a 1.0b 4.4a 5.4a ce lerating the transfer of dry ma tte r to the gr ain, and thus increa s-
2018−19 SA 10.0a 5.0a 15.0ab 10.8a 2.8c 13.6a ing the thou sa nd gr ain weight.
I21 9.9a 5.2a 15.1b 10.5ab 4.3b 14.8a Stud ies have sh own that inte rcrop ping ce real and legu me
I42 10.8a 5.0a 15.9a 9.7ab 6.4a 16.1a crop s do es no t en hance the yield of legu me crop s in mo st case s
I84 8.0b 5.8a 13.8b 8.3b 5.8a 14.1a (Ne ug sc hwandt ne r and Kaul, 2015; Raza et al., 2021). In ou r st ud y,
2019−20 SA 11.5a 5.8b 17.3ab 10.8a 5.6a 16.5ab
we also ob se rved that inte rcrop pe d alfalfa expe rien ce d su ppres -
I21 10.9a 4.6c 15.4b 7.9b 5.8a 13.7c

PR
I42 12.0a 7.3a 19.3a 12.4a 5.4a 17.8a sion du e to the strong co mpe titive ab ility of whea t du ring the co -
I84 9.5a 6.4ab 15.9b 8.7b 6.6a 15.3bc growth pe riod , resu lting in lower yields . Ho wever, fo llowing the
2020−21 SA 8.8a 5.0a 13.8a 9.6a 4.4a 14.0a harvest of whea t, inte rcrop pe d alfalfa had ap proxima te ly 70 da ys
I21 7.3b 3.3c 10.6c 6.9b 2.7b 9.6b of reco very growth, du ring which it co uld ab so rb reso urce s from
I42 8.2ab 3.7b c 11.9bc 10.3a 4.1a 14.4a the whea t strips . This reco very pe riod ultima te ly led to an increa se
I84 9.1a 4.3ab 13.4ab 8.7a 4.5a 13.2a
in the bioma ss yield of alfalfa. Althou gh the alfalfa yields sh owed
so me co mpe nsa tion du ring the growth reco very pe riod fo llowing
He llou et al., 2006). As discuss ed by Vand erme er (1989), when two whea t harvest, they did no t fully reco ver from the initial yield re-
D
sp ec ies co mpe te fo r a reso urce , su ch as wate r or N, ad ding mo re du ction . Ho wever, we revealed that the bioma ss yield of alfalfa in
of that reso urce ma y redu ce the inte nsity of co mpe tition . This was the I42 and I84 trea tme nts were co mpa rable or even high er than
also de mo nstrate d in ou r st ud y, as low N ap plication sig nificantly that of SA , this migh t be caus e increa sing the strip widt h of alfalfa
improved the co mpe titivene ss of whea t ag ains t alfalfa, ultima te ly redu ce d ligh t, wate r and nitroge n co mpe tition from ne ighbo ring
TE

increa sing the growth rate and yield ad vantage of inte rcrop pe d whea t du ring co - growth pe riod while increa se d facilitation from
whea t over so le whea t. whea t strip afte r whea t harvest. Similar resu lts were repo rted by Li

Table 4
N us e efficien cy (NUE) and pa rtial facto r produ ctively of ap plied N (PFPN ) of differen t trea tme nts in 2017–18, 2018–19, 2019–20 and 2020–2021. SW ,
EC

so le whea t; SA , so le alfalfa; I42- I, I42 inne r rows; I42- B, I42 bo rde r rows; I84- I, I84 inne r rows; I84- B, I84 bo rde r rows; Differen t lowercase lette rs indi-
cate sig nificant differen ce s am on g differen t inte rcrop ping trea tme nts at α =0.05, resp ec tively.
Growing se as on s Trea tm en ts Wh ea t Alfalfa
High N Low N High N Low N
NU E PFP N NU E PFP N NU E PFP N NU E PFP N
2017–18 SW 34.9b 25.0a 52.1d 23.6b - - - -
SA - - - - 33.32a 72.65b 32.24a 147.70a
RR

I21 41.7a 24.6a 64.6c 22.3b 27.90 cd 26.78d 26.49c 49.05d

I42- I 36.4a 24.5a 68.0c 24.4b 27.08d 39.01 cd 27.80c 89.79c
I42- B 41.8a 24.2a 72.0bc 24.2b
I84- I 41.0a 25.8a 78.0ab 31.9a 32.48ab 91.82a 31.42ab 166.66a
I84- B 44.7a 26.9a 85.6a 30.6a 29.99b c 49.93c 29.05b c 118.73b
2018–19 SW 33.4bc 25.6a 49.0d 25.9ab - - - -
SA 34.8a - 37.0a -
I21 37.4b 24.2a 64.3b 27.4a 35.4a - 38.1a -
CO

I42- I 38.5a 25.5a 64.5b 26.7a 36.1a - 37.1a -

I42- B 39.3a 23.5a 76.2a 25.8ab
I84- I 38.0ab 21.8ab 59.8bc 23.8ab 35.3a - 37.1a -
I84- B 31.1c 18.8b 54.3 cd 21.6b 35.5a - 37.0a -
2019–20 SW 26.6c 21.8b 43.4b 23.8b - - - -
SA - - - - 26.3a - 27.3c -
I21 35.1ab 28.6a 64.7a 29.6ab 27.6a - 29.0ab -
I42- I 29.0c 22.2b 43.5b 17.5c 28.6a - 29.2a -
I42- B 35.5ab 26.6ab 68.6a 30.9a
I84- I 38.1a 23.2ab 69.8a 28.9ab 28.5a - 28.9ab -
I84- B 31.5bc 25.0ab 66.5a 27.6ab 28.3a - 27.7bc -
2020–21 SW 27.9a 30.1a 49.6 cd 32.7a - - - -
SA - - - - 30.9a - 30.2a -
I21 31.5a 27.7a 65.5ab 42.7a 29.4a - 30.0a -
I42- I 31.3a 31.4a 44.1d 35.3a 29.6a - 28.8a -
I42- B 34.2a 31.0a 75.7a 36.9a
I84- I 38.0a 29.8a 74.9a 37.4a 29.6a - 29.2a -
I84- B 31.3a 31.0a 57.8bc 35.6a 29.4a - 30.0a -

9
Q. Ma et al. Field Crops Research xxx (xxxx) 109335

Table 5 ma king the bo rde r row effect a sig nificant facto r in en hancing in-
N eq uivalent ratio (NER) of differen t trea tme nts in 2017–18, 2018–19, te rcrop produ ctivity (Clém en t et al., 2022; Go u et al., 2016; Wang
2019–20 and 2020–2021. SW , so le whea t; SA , so le alfalfa; I21, I42, and et al., 2017). This stud y also de mo nstrate d that in whea t/alfalfa in-
I84 represe nt inte rcrop s with two rows of whea t inte rcrop pe d with on e te rcrop ping syste ms , favorab le ea rly growth co ndition s fo r whea t
row of alfalfa, fo ur rows of whea t inte rcrop pe d with two rows of alfalfa, resu lted in sig nificant yield increa se s in the bo rde r rows. This bo r-
and eigh t rows of whea t inte rcrop pe d with fo ur rows of alfalfa. Differen t
de r row ad vantage co ntributed sig nificantly to the overall yield of
lowercase lette rs indicate sig nificant differen ce s am on g differen t inte r-
inte rcrop pe d whea t. For exam ple, in I42, whea t in the bo rde r row
crop ping trea tme nts at α =0.05, resp ec tively.
acco unte d fo r 53.3–126.8% of the yield improveme nt while oc cu-
Growing se as on s Trea tm en ts NER
pying on ly 50% of the land area . Similar resu lts were fo und in
High N Low N
2017–18 I21 1.56b 1.57c whea t/faba be an, whea t/pe a, and whea t/ma ize strip inte rcrop ping

F
I42 1.66b 1.95b syste ms (Pelzer et al., 2012; Go u et al., 2016; Xiao et al., 2018). In
I84 2.17a 2.50a ad dition , we fo und that the bo rde r row effect of inte rcrop pe d
2018–19 I21 1.12a 1.31ab whea t was mo re prono unce d unde r LN co ndition s. This co nfirmed

OO
I42 1.17a 1.44a
the N facilitation from alfalfa to whea t in a whea t/alfalfa inte rcrop -
I84 1.09a 1.19b
2019–20 I21 1.32a 1.49a
ping syste m. On on e hand , bo rde r row whea t co mpe te s fo r so il ni-
I42 1.21a 1.29b troge n from the alfalfa rows, as whea t has a do minant po sition
I84 1.37a 1.59a (Wang et al., 2021); and on the ot he r hand , the strong co mpe tition
2020–21 I21 1.13a 1.32a from whea t migh t have promo te d biolog ical N 2 fixation by alfalfa,
I42 1.17a 1.21a which further provide d nitroge n to whea t bo rde r row (Me i et al.,
I84 1.30a 1.42a
2021; Wang et al., 2020b).

PR
For alfalfa, the bo rde r row also sh owed a sig nificantly high er
Table 6 bioma ss yield than the inne r row in I84. Alfalfa had ap proxima te ly
Land eq uivalent ratio (LER) of differen t trea tme nts in 2017–18, 2018–19, 70 da ys of reco very time afte r whea t harvest, alfalfa in the bo rde r
2019–20 and 2020–2021. SW , so le whea t; SA , so le alfalfa; I21, I42, and row can inte rce pt mo re ligh t than in the inne r row and utilize so il
I84 represe nt inte rcrop s with two rows of whea t inte rcrop pe d with on e
reso urce s in the ba re pa th (Qian et al., 2018). In co ntras t to the
row of alfalfa, fo ur rows of whea t inte rcrop pe d with two rows of alfalfa,
bo rde r row effects on whea t, the bo rde r row effects on alfalfa yield
and eigh t rows of whea t inte rcrop pe d with fo ur rows of alfalfa. Differen t
lowercase lette rs indicate sig nificant differen ce s am on g differen t inte r- were mo re sig nificant unde r HN . This was be caus e, firstly, whea t
crop ping trea tme nts at α =0.05, resp ec tively. exerte d less co mpe tition on alfalfa unde r HN , and se con dly, alfalfa
D
Growing trea tm en ts High N Low N migh t be ab le to utilize mo re residual so il N in the whea t strip un-
se as on s pLERW pLERA LER
pLERW pLERA LER de r HN (Xu et al., 2021).
2018 I21 0.60a 0.18b 0.78b 0.62b 0.17c 0.79c
4.3. N us e ef ficien cy
TE

I42 0.56a 0.27b 0.83b 0.67b 0.30b 0.98b

I84 0.60a 0.49a 1.09a 0.77a 0.48a 1.25a
2019 I21 0.56a 0.50a 1.06ab 0.66ab 0.55a 1.20ab The nitroge n utilization of inte rcrop pe d whea t has be en im-
I42 0.58a 0.53a 1.11a 0.72a 0.59a 1.31a
proved , lead ing to its increa se d yield ad vantage . The reaso ns fo r
I84 0.54a 0.46b 1.01b 0.60b 0.52a 1.11b
2020 I21 0.66a 0.45b 1.11a 0.75a 0.42c 1.16b the high NU E of whea t are: firstly, alfalfa has a de ep - root ing sys-
I42 0.61a 0.56a 1.16a 0.65b 0.54a 1.19ab te m, while whea t has a sh allow- root syste m, utilizing the su bsu r-
EC

I84 0.68a 0.46b 1.14a 0.79a 0.47b 1.26a face sp atial ec olog ical niches and improving N availab ility fo r
2021 I21 0.56a 0.38b 0.95b 0.66a 0.34b 1.00b whea t (Su n et al., 2019; Xu et al., 2021). Furthermo re, alfalfa N 2 fix-
I42 0.59a 0.43b 1.02ab 0.60a 0.51a 1.12ab ation can improve the so il N availab ility in the sh allow so il layer, fa-
I84 0.65a 0.48a 1.14a 0.71a 0.47a 1.18a
cilitating whea t N us e. In ad dition to sp atial N facilitation from al-
falfa to whea t strip, te mpo ral ec olog ical niche se paration is also a
et al. (2021) in an oa t/co mmo n vetch inte rcrop ping syste m, which ma jor reaso n to improve NU E of whea t (Hu et al., 2020; Ma hm ou d
sh owed that the bioma ss yield of inte rcrop pe d co mmo n vetch in- et al., 2022), as whea t re- gree ning oc curs ea rlier and alfalfa co m-
RR

crea se d with strip widt h. pe te s weakly fo r so il nitroge n. Finally, the high er wate r availab ility
du e to root co mpleme ntary and ligh t availab ility du e to strip co n-
4.2. Strip de sign an d bo rd er ro w ef fe ct s fig uration de sign also co ntributed to high N co nversion efficien cy
(Wang et al., 2020b; Xiao et al., 2018). For alfalfa, NU E was also im-
Strip widt h ma nag eme nt is critical fo r achieving high er crop proved exce pt fo r the first year of es tablish me nt, du e to sp atial
yields in strip inte rcrop ping syste ms (Raza et al., 2020a). Unde r no and te mpo ral co mpleme ntary inte raction on wate r, ligh t and ni-
wate r-limite d co ndition s, narrow strips increa se inte rsp ecific inte r-
CO

troge n utilization . Me anwhile, we disco vered that the NER of

action , facilitating the co mpleme ntary utilization of ligh t and re- whea t/alfalfa inte rcrop ping was grea te r than 1.0, su g ge sting that
su lting in increa se d inte rcrop ping yield ad vantage (Mu nz et al., whea t/alfalfa inte rcrop ping sa ved N fertilizer inpu t. The NU E and
2014; Wang et al., 2015a). Ho wever, unde r dryland co ndition s, a PFP N of whea t and alfalfa, and the NER of inte rcrop ping all de -
narrow strip pa tte rn redu ce s the inte raction sp ace be twee n two crea se d with increa sing N ap plication , in ag reeme nt with the stud -
crop s, limiting the inte rcrop ping ad vantage . In co mpa riso n, wide r ies of Kermah et al. (2017), Frazir et al. (2017), Me i et al. (2021), and
st rips can facilitate the exte nsion of crop root s in bo rde r rows to Wang et al. (2021), who proved that ce real/legu me inte rcrop ping
ot he r crop strips , thus ob taining mo re wate r and nutrien ts to im- improved N us e ad vantage over mo no cultures unde r low nitroge n
prove growth and yield (Ma hm ou d et al., 2022; Wang et al., 2020a). co ndition s.
Therefo re, we ob se rved that the yield ad vantage of whea t in I21
gr ad ually diminishe d with the du ration of inte rcrop ping , while, the 4.4. Land us e ad vant ag e an d ne t re ve nu e
yield ad vantage of whea t in I42 and I84 increa se d as the inte rcrop -
ping du ration progres se d. Co nse qu en tly, wide r st rips sh owed Unde r high N ap plication , whea t/alfalfa strip inte rcrop ping
grea te r ad vantage s fo r whea t produ ction . sh owed land us e ad vantage with LER rang ed in 0.78–1.16. We no te
In wide - strip inte rcrop ping , the bo rde r rows of ad jace nt crop that the LER value s in this stud y are mu ch high er than the pu b-
strips play a crucial role in facilitating inte rsp ecific inte raction s,

10
Q. Ma et al. Field Crops Research xxx (xxxx) 109335

F
OO
2245px x 2210px

PR
D
TE

Fig. 7. Ne t reven ue in so le and inte rcrop ping syste ms in: (a) 2017–18, (b) 2018–19, (c) 2019–20, (d) 2020–21 and (e) 4- year me an. SW , so le whea t; I21, I42,
and I84 represe nt inte rcrop s with two- row whea t and on e- row alfalfa, fo ur-row whea t and two- row alfalfa, and eigh t-rows whea t and fo ur-row alfalfa, re-
sp ec tively. N, P and N×P sh ow the effects of N ap plication , planting pa tte rn and their inte raction . Differen t lette rs indicate sig nificant differen ce s am on g
differen t inte rcrop ping trea tme nts at α =0.05. “⁎ ” and “⁎ ⁎” indicate sig nificant and very sig nificant differen ce s at α =0.05 and α =0.01 resp ec tively,
and ns me ans no n- sign ificant differen ce at α =0.05.
EC

lish ed da ta fo r whea t/alfalfa inte rcrop ping unde r irriga te d co ndi- similar to the resu lts of Su n et al. (2018) and Wu et al. (2023). De -
tion s (Mu et al., 2023), indicating that whea t/alfalfa inte rcrop ping sp ite the high er ne t reven ue from so le alfalfa, it canno t me et the
pe rfo rmed be tte r unde r rainfed co ndition s. Wh en N ap plication de ma nd s fo r grain produ ction to feed the po pulation . Further-
was redu ce d, the LER value s further improved to 0.79–1.31. So me mo re, alfalfa yields and ec ono mic returns will de crea se afte r 4–6
ot he r st ud ies also fo und that grea te r produ ctivity of inte rcrop pe d years of planting (Ali et al., 2021), so it is unsu itab le fo r long - te rm
syste ms relative to so le crop s of te n resu lt from low N co ndition s planting. The 4- year averag e ne t reven ue of I84 was high er than
(Sa rr et al., 2008; Rodrigue z et al., 2020). Am on g differen t co nfig u- that of the narrow- strip pa tte rns. Ad dition ally, the I84 wide - strip
RR

ration s, the LER value of I84 exce ed ed 1.0 rega rdless of the year pa tte rn is mo re me chanically acce ssible in practice . Therefo re, we
and N fertilizer, indicating that I84 had the high es t stability of in- su g ge st farmers to cultivate I84 unde r low- nitrog en co ndition s to
te rcrop do minance. This was be caus e the wide r st rip de sign in- su pply fo rag e fo r livestoc k, redu ce their reliance on N fertilizer, en -
crea se d the inte raction sp ace be twee n whea t and alfalfa, improv- hance their field produ ctivity and ne t reven ue .
ing the inte rsp ecific facilitation be twee n the sp ec ies (Hu et al.,
2016; Li et al., 2021). The pLER of whea t was high er than its plant- 5. Co nclusions
CO

ing ratio of 0.5 in fo ur years, co nfirming that whea t was the do mi-
nant sp ec ies in the inte rcrop ping syste m; me anwhile, alfalfa was a This stud y investiga te d the yield, nitroge n us e and ec ono mic
weaker co mpe tito r in inte rcrop ping with the low pLER value s of pe rfo rma nce of a dryland whea t/alfalfa inte rcrop ping throug h a 4-
less than 0.5 unde r mo st circumstance s. yr field expe rime nt. Dynamics of inte rsp ecific inte raction be twee n
Wh en co nsidering whet he r farmers en ga ge in inte rcrop ping whea t and alfalfa were clarified , we fo und winte r whea t ma in-
practice s, it is impo rtant to co nside r facto rs be yond simply crop tained do minance throug hou t the co - growth pe riod in the inte r-
yields , su ch as the ec ono mic be nefits involved . The profitab ility of crop ping syste m, alfalfa was co nstrained bu t reco vered afte r the
thes e practice s ultima te ly de cide s the farmers’ level of inte res t in whea t harvest. Wh ea t yield in inte rcrop ping increa se d by 19.0%-
ad op ting inte rcrop ping (Raza et al., 2021). Ou r find ings su g ge st 24.0% unde r HN , and by 34.1%- 43.4% unde r LN. Similarly, whea t
that increa sing N fertilizer ap plication did no t have a sig nificant NU E in inte rcrop ping increa se d by −1.1–3.9% unde r HN , and by
impa ct on the ne t reven ue , which indicate s that inte rcrop ping 4.1–14.2% unde r LN. The averag e land eq uivalent ratio over the
have the po te ntial to redu ce the ap plication of N fertilizer. The av- fo ur-year rang ed in 0.98–1.10 unde r HN co ndition s, and in
erag e ne t reven ue of inte rcrop ping was 134.6%- 344.8% high er 1.04–1.20 unde r LN. Co mbination I84 achieved the high es t LER
than that of so le whea t. Ho wever, co mpa red to so le alfalfa, inte r- and ec ono mic profit. Therefo re, the I84 pa tte rn with low N ap plica-
crop ping had a lower ne t reven ue du e to the high alfalfa hay price, tion alleviated inte rsp ecific co mpe tition and increa se d syste m

11
Q. Ma et al. Field Crops Research xxx (xxxx) 109335

profit, which was su g ge sted to be ap plied in ou r st ud y area and Hu, F., Gan, Y., Chai, Q., Feng , F., Zhao , C., Yu, A., Mu , Y., Zhang , Y., 2016. Bo os ting
system prod uctivity throug h the improved co ordination of inte rspe cific
similar dryland en vironme nts. co mp et ition in ma ize/pe a strip inte rcropp ing . Field Crop s Res 198, 50–60.
Hu, F., Zhao , C., Feng , F., Chai, Q., Mu , Y., Zhang , Y., 2017. Improving N ma nage me nt
Uncited refe rences throug h inte rcropp ing alleviates the inhibito ry effect of mine ral N on
no du lation in pe a. Plant So il 412, 235–251.
Jeuffroy, M ., Ne y, B., 1997. Crop ph ysiology and prod uctivity. Field Crop . Res. 53,
(Broa db en t, 1981; Clém en t et al., 2022; Wang et al., 2020a,a,c, 3–16.
2021, 2023b) Jung ers, J.M ., Yang , Y., Fernande z, C.W., Isbe ll, F., Lehm an, C., Wyse , D., Shea ffer, C.,
2021. Diversifying bioe ne rgy crop s increa se s yield and yield stab ility by
redu cing weed ab unda nce. Sci. Ad v. 7. . g8 531.
CRediT authorship contribution statem ent Kermah, M ., Franke, A.C., Ad jei- Ns iah, S., Ahiabo r, B.D.K., Ab aido o, R.C., Giller, K.E.,
2017. Ma ize- gr ain legu me inte rcropp ing fo r en hanced reso urce us e efficien cy

F
Qianhu Ma : Da ta curation , Investiga tion , Writing – original and crop prod uctivity in the Guinea sa vann a of no rthe rn Ghana. Field Crop . Res.
213, 38–50.
draft, Writing – review & ed iting. Yuhuan Wu : Investiga tion , Writ- Latati, Do kukin, Ao uiche, Rebo uh, Takouachet , Hafnao ui, Hamd ani, Bacha, Ou nane ,
ing – original draft. Zikui Wang : Co nce ptualization , Da ta curation , 2019. Sp ec ies inte ractions improve ab ove- grou nd biom as s and land us e

OO
Fund ing acqu isition , Su pe rvision , Writing – original draft, Writing efficien cy in inte rcropp ed whea t and chickpea unde r low so il inpu ts. Ag rono my
9, 765.
– review & ed iting. Yanan Liu: Co nce ptualization , Investiga tion . Li, C., Ho ffland , E., Kuyper, T.W., Yu, Y., Zhang , C., Li, H., van de r We rf, W., et al., 2020.
Yuying Shen: Co nce ptualization , Su pe rvision , Writing – review & Synd rome s of prod uction in inte rcropp ing impa ct yield ga ins. Na t. Plants 6 (6),
ed iting. 653–660.
Li, L., Su n, J., Zhang , F., Li, X., Yang , S., Reng el, Z., 2001. Wh ea t/ma ize or whea t/
so ybea n strip inte rcropp ing: I. Yield ad vantag e and inte rspe cific inte ractions on
De claration of Co mp eting Interest nutrients. Field Crop s Res 71, 123–137.
Li, R., Zhang , Z., Tang , W., Huang , Y., Na n, Z., 2021. Effect of row co nfiguration on

PR
yield and radiation us e of co mm on vetch−oa t strip inte rcropp ing on the
The authors de clare that they have no known co mpe ting finan-
Qing hai- Tibe tan platea u. Eur. J. Ag ron. 128, 126290.
cial inte res ts or pe rso nal relation sh ips that co uld have ap pe ared to Luo, C., Zhu, J., Ma , L., Guo, Z., Do ng , K., Do ng , Y., 2021. Effects of nitrog en regu lation
influen ce the work repo rted in this pa pe r. and strip inte rcropp ing on faba be an biom as s, nitrog en accumu lation and
distribu tion , and inte rspe cific inte ractions . Crop Sci. 61, 4325–4343.
Ma , L., Li, Y., Wu , P., Zhao , X., Gao, X., Chen , X., 2020. Reco very growth and wate r us e
Data Availability of inte rcropp ed ma ize fo llowing whea t harves t in whea t/ma ize relay strip
inte rcropp ing . Field Crop . Res. 256, 107924.
Da ta will be ma de availab le on requ es t. Ma hallati, M .N., Kooc he ki, A., Mo nd ani, F., Feizi, H., Am irmo radi, S., 2015.
De te rmination of op tima l strip widt h in strip inte rcropp ing of ma ize (Zea ma ys
D
L.) and be an (Phase olus vulgaris L.) in No rthe as t Iran. J. Clea n. Prod . 106 (3),
Acknowledg em ents 343–350.
Ma hm ou d, R., Casa de ba ig, P., Hilgert, N., Alletto, L., Fres chet , G.T., de Ma zancou rt,
C., Gaud io, N., 2022. Sp ec ies choice and N fertilization influe nce yield ga ins
This stud y was jointly su ppo rted by the Na tion al Key R& D Pro-
throug h co mp leme ntarity and se lection effects in ce real- legu me inte rcrops .
gr am of China (2022YFD1300803), the Na tion al Na tural Scien ce
TE

Ag ron. Su stain. De v. 42.

Founda tion of China (31871560, 32001403, 32101849), the Ma jor Me i, P.P., Wang , P., Yang , H., Gui, L.G., Christie, P., Li, L., 2021. Ma ize/faba be an
inte rcropp ing with rhizob ial inoc ulation in a reclaime d de se rt so il en hances
Scien ce and Techno log y Project of Gans u Province (21ZD 4NA 012),
prod uctivity and symb iotic n- 2 fixation and redu ce s ap pa rent n loss es . So il . Res.
and the Funda me ntal Rese arch Fund s fo r the Ce ntral Universities . (213- ), 213.
(lzujbky- 2022- ey22). Mu , L., Su , K., Zhou , T., Yang , H., 2023. Yield pe rforma nce, land and wate r us e,
ec on om ic profit of irriga te d sp ring whea t/alfalfa inte rcropp ing in the inland arid
area of no rthwes te rn China. Field Crop s Res 303, 109116.
EC

Ap pe ndix A. Supp orting info rmation Mu nz, S., Grae ff-Hö nn inge r, S., Lizaso , J.I., Chen , Q., Claupe in, W., 2014. Mo de ling
ligh t availability fo r a su bo rdinate crop within a strip–inte rcropp ing system .
Su ppleme ntary da ta as so ciated with this article can be fo und in Field Crop s Res 155, 77–89.
Ne ug sc hwandt ne r, R.W., Kaul, H., 2015. Nitrog en up take, us e and utilization
the on line version at do i:10.1016/j.fcr.2024.109335. efficien cy by oa t–pe a inte rcrops . Field Crop . Res. 179, 113–119.
Olse n, S., 1954. Estima tion of Available Phos ph orus in So ils by Extraction with
Refe rences So dium Bicarbon ate. No . 939. US De pa rtme nt of Ag riculture, Printing Of fice ,
Wash ingt on , DC .
RR

Pelzer, E., Bazo t, M ., Ma kowski, D., Co rre- He llou , G., Na ud in, C., Al Rifaï, M ., Barang er,
Ali, G., Wang , Z., Li, X., Jin, N., Chu, H., Jing , L., 2021. De ep so il wate r de ficit and
E., Be do us sa c, L., Biarnè s, V., Bo uche ny, P., Carrou ée , B., Do rvillez, D., Foissy, D.,
reco very in alfalfa fields of the Loes s Platea u of China. Field Crop . Res. 260,
Gaillard, B., Guichard, L., Ma ns ard, M ., Om on , B., Prieur, L., Yvergn iaux, M ., Justes ,
107990.
E., Jeuffroy, M ., 2012. Pea- whea t inte rcrops in low- inpu t co nd itions co mb ine
Brem ne r, J.M., 1965. Total nitrog en . In: No rman, A.G. (Ed.), Me thod s of So il Analysis.
high ec on om ic pe rforma nces and low en vironm en tal impa cts. Eur. J. Ag ron. 40,
Part 2. Chem ical and Microb iological Prop erties . Am erican So ciet y of
39–53.
Ag rono my, So il Science So ciet y of Am erica, pp . 1149–1178.
Qian, X., Zang , H., Xu, H., Hu, Y., Ren, C., Guo, L., Wang , C., Ze ng , Z., 2018. Relay strip
Broa db en t, F.E., 1981. Me thod olog y fo r nitrog en transforma tion and ba lance in so il.
inte rcropp ing of oa t with ma ize, su nflower and mu ng be an in se mi- arid region s
Plant So il 1981, 383–399.
of No rthe as t China: Yield ad vantag es and ec on om ic be ne fits. Field Crop . Res.
CO

Cao, Q., Wang , Z., Yang , X., Shen , Y., 2021. The effects of co cksfoo t co ver crop on so il
223, 33–40.
wate r ba lance, evap ot rans piration pa rtitioning , and system prod uction in an
Rase du zzam an, M ., Jens en , E.S., 2017. Do es inte rcropp ing en hance yield stab ility in
ap ple orchard on the Loes s Platea u of China. So il Tillag e Res. 205, 104788.
arab le crop prod uction ? A me ta- analysis. Eur. J. Ag ron. 91, 25–33.
Clém en t, C., Sleide rink, J., Svane, S.F., Sm ith, A.G., Diam anto po ulos , E., De sb røll, D.B.,
Raza, M .A., Feng , L.Y., We rf, W., Iqba l, N., Khan, I., Khan, A., Din, A.M .U., Na ee m, M .,
Thorup - Kriste ns en , K., 2022. Co mp aring the de ep root growth and wate r up take
Me raj, T.A., Hass an, M .J., Khan, A., Lu, F.Z., Liu, X., Ahme d, M ., Yang , F., Yang , W.,
of inte rmed iate whea tg rass (Kernza® ) to alfalfa. Plant So il 472, 369–390.
2020b. Op timu m strip widt h increa se s dry ma tter, nutrient accumu lation , and
Co ng , W., 2020. China’ s inte rcropp ing lead s to high er grain yield ga ins. Sci. China
se ed yield of inte rcrops unde r the relay inte rcropp ing system . Food Energy
Life Sci. 63, 1612–1614.
Se cur. 9.
Co rre- He llou , G., Fustec , J., Croz at, Y., 2006. Inte rspe cific Co mp et ition fo r So il N and
Raza, M .A., Cui, L., Qin, R., Yang , F., Yang , W., 2020a. Strip- widt h de te rmines
its Inte raction with N 2 Fixation , Leaf Expa ns ion and Crop Growth in Pea- Barley
co mp et itive streng ths and gr ain yields of inte rcrop sp ec ies in relay
Inte rcrops . Plant So il 282, 195–208.
inte rcropp ing system . Sci. Rep. 10.
Dh ima, K.V., Lithou rgidis, A.S., Vasilakoglou , I.B., Do rdas , C.A., 2007. Co mp et ition
Raza, M .A., Gul, H., Wang , J., Yasin, H.S., Qin, R., Bin Khalid, M .H., Na ee m, M ., Feng ,
indice s of co mm on vetch and ce real inte rcrops in two se ed ing ratio. Field Crop s
L.Y., Iqba l, N., Gitari, H., Ahma d, S., Battag lia, M ., Ansa r, M ., Yang , F., Yang , W.,
Res 100, 249–256.
2021. Land prod uctivity and wate r us e efficien cy of ma ize- so ybea n strip
Fras ier, I., No elleme yer, E., Am iotti, N., Qu irog a, A., 2017. Vetch- rye biculture is a
inte rcropp ing system s in se mi- arid area s: a case stud y in Punjab Province ,
su stainable alte rnative fo r en hanced nitrog en availability and low leaching
Pakistan. J. Clea n. Prod . 308, 127282.
loss es in a no −till co ver crop system . Field Crop s Res 214, 104–112.
Rodrigue z, C., Carlss on , G., Englund, J., Flöh r, A., Pelzer, E., Jeuffroy, M ., Ma kowski, D.,
Go u, F., van Itte rsum , M .K., Wang , G., van de r Putten , P.E.L., van de r We rf, W., 2016.
Jens en , E.S., 2020. Grain legu me - ce real inte rcropp ing en hances the us e of so il-
Yield and yield co mp on en ts of whea t and ma ize in whea t-ma ize inte rcropp ing
de rived and biolog ically fixed nitrog en in te mp erate ag roec os yste ms . A me ta-
in the Ne therland s. Eur. J. Ag ron. 76, 17–27.
analysis. Eur. J. Ag ron. 118, 126077.

12
Q. Ma et al. Field Crops Research xxx (xxxx) 109335

Sa rr, P.S., Khou ma , M ., Se ne , M ., Guisse , A., Badiane, A.N., Yama kawa, T., 2008. Effect system resilien ce . Ag ric. Wate r Ma nag. 276, 108067.
of pe arl millet-co wpea crop ping system s on nitrog en reco very, nitrog en us e Wang , Z., Zhao , X., Wu , P., Gao, Y., Yang , Q., Shen , Y., 2017. Bo rder row effects on ligh t
efficien cy and biolog ical fixation us ing the 15N tracer te chniqu e. So il Sci. Plant inte rcep tion in whea t/ma ize strip inte rcropp ing system s. Field Crop . Res. 214,
Nu tr. 54, 142–147. 1–13.
Shi, Y., Gao, S., Zhou , D., Liu, M ., Wang , J., Knop s, J.M .H., Mu , C., 2017. Fall nitrog en Wang , Z., Do ng , B., Stom ph , T.J., Evers, J.B., L. Van De r Putten , P.E., Ma , H., Miss ale, R.,
ap plication increa se s se ed yield, fo rage yield and nitrog en us e efficien cy mo re van de r We rf, W., 2023b. Temp oral co mp leme ntarity drives sp ec ies
than sp ring nitrog en ap plication in Leymus chinen sis, a pe renn ial gr as s. Field co mb inab ility in strip inte rcropp ing in the Ne therland s. Field Crop . Res. 291,
Crop s Res 214, 66–72. 108757.
Sn ap p, S.S., Blackie, M .J., Gilbert, R.A., Be zner-Kerr, R., Kanyam a- Phiri, G.Y., 2010. Wang , X., Gao, Y., Zhang , H., Shao , Z., Su n, B., Gao, Q., 2020b. Enhancem en t of
Biod iversity can su pp ort a gree ne r revolution in Africa. Proc . Na tl. Acad . Sci. 107, rhizos ph ere citric acid and de crea se of NO 3− /NH 4+ ratio by root inte ractions
20840–20845. . (USA). facilitate N fixation and transfer. Plant So il 447, 169–182.
Stanislaw, J., Da vid, S., 1973. A co mp arative stud y of titrimet ric and gr avimet ric Willey, R.W., Rao, M .R., 1980. A co mp et itive ratio fo r qu antifying co mp et ition
me thod s fo r the de te rmination of orga nic carbon in so il. J. Sci. Fd. Ag ric. 24, be twee n inte rcrops . Exp. Ag r. 117–125.

F
1085–1090. de Wit, C.T., 1953. A Physical Theo ry on Placem en t of Fertilizers. Wage ning en
Stratton , A.E., Co min, J.J., Sidd ique , I., Zak, D.R., Filipini, L.D., Lucas, R.R., Bles h, J., University.
2022. Asse ss ing co ver crop and inte rcrop pe rforma nce alon g a farm Wu , Y., Ma , Q., Ma , L., Wang , Z., 2023. Inte rspe cific co mp et ition and prod uctivity in a
ma nage me nt gr ad ient. Ag ric. Ecos yst. Environ. 332, 107925. dryland silage ma ize/alfalfa inte rcropp ing system . Food Energy Se cur 12.

OO
Su n, B., Gao, Y., Yang , H., Zhang , W., Li, Z., 2019. Performa nce of alfalfa rather than Xiao , J., Yin, X., Ren, J., Zhang , M ., Tang , L., Zhen g , Y., 2018. Co mp leme ntation drives
ma ize stimulates system ph os ph orus up take and overyielding of ma ize/alfalfa high er growth rate and yield of whea t and sa ves nitrog en fertilizer in whea t and
inte rcropp ing via change s in so il wate r ba lance and root mo rpho logy and faba be an inte rcropp ing . Field Crop s Res 221, 119–129.
distribu tion in a ligh t cherno ze mic so il. Plant So il 439, 145–161. Xu, R., Zhao , H., Liu, G., You, Y., Ma , L., Liu, N., Zhang , Y., 2021. Effects of nitrog en and
Su n, T., Li, Z., Wu , Q., Shen g , T., Du , M ., 2018. Effects of alfalfa inte rcropp ing on crop ma ize plant de ns ity on fo rage yield and nitrog en up take in an alfalfa- silage
yield, wate r us e efficien cy, and overall ec on om ic be ne fit in the Co rn Be lt of ma ize relay inte rcropp ing system in the No rth China Plain. Field Crop s Res 263,
No rthe as t China. Field Crop . Res. 216, 109–119. 108068.
Tesh ita, A., Feng , Y., Qian, R., Wang , X., Khan, W., Gao, Y., 2023. Alfalfa and ma ize Yin, W., Chai, Q., Zhao , C., Yu, A., Fan, Z., Hu, F., Fan, H., Guo, Y., Co ulte r, J.A., 2020.
inte rcropp ing en hances so il ne ma to de structure and fo od web co mp lexity in Wate r utilization in inte rcropp ing: A review. Ag ric. Wate r Ma nag. 241, 106335.

PR
low- nitrog en so ils. Ap pl. So il Ecol. 186, 104809. Zhang , F., Li, L., 2003. Us ing co mp et itive and facilitative inte ractions in inte rcropp ing
Vand erme er J., 1989. The Ecolog y of Inte rcropp ing. Camb ridg e University pres s, system s en hances crop prod uctivity and nutrient-us e efficien cy. Plant So il 248,
Camb ridg e, UK, 237 p. 305–312.
Wang , R., Su n, Z., Zhang , L., Yang , N., Feng , L., Bai, W., Zhang , D., Wang , Q., Evers, J.B., Zhang , W.P., Gao, S.N., Li, Z.X., Xu, H.S., Yang , H., Yang , X., Fan, H.X., Su , Y., Fornara, D.,
Liu, Y., Ren, J., Zhang , Y., van de r We rf, W., 2020a. Bo rder-row prop ortion Li, L., 2021. Shifts from co mp leme ntarity to se lection effects ma intain high
de te rmines streng th of inte rspe cific inte ractions and crop yields in ma ize/ prod uctivity in ma ize/legu me inte rcropp ing system s. J. Ap pl. Ecol. 58,
pe anut strip inte rcropp ing . Field Crop s Res 253, 107819. 2603–2613.
Wang , Z., Jiang , H., Shen , Y., 2020c. Forage prod uction and so il wate r ba lance in oa t Zhang , Y., Do ng , X., Yang , X., Mu nyam pirwa, T., Shen , Y., 2022. The lagg ing
and co mm on vetch so le crop s and inte rcrops cultivated in the su mm er-autumn mo veme nt of so il nitrate in co mp arison to that of so il wate r in the 500- cm so il
fallow se as on on the Chines e Loes s Platea u. Eur. J. Ag ron. 115, 126042. profile. Ag ric. Ecos yst. Environ. 326, 107811.
D
Wang , X., Liu, X., Zhao , Y., Wang , J., 2021. Nitrog en utilization and inte rspe cific Zhang , Y., Su n, Z., Su , Z., Du , G., Bai, W., Wang , Q., Wang , R., Nie, J., Su n, T., Feng , C.,
feed ba ck characte ristics of inte rcropp ed alfalfa/oa t with differen t root ba rriers. Zhang , Z., Yang , N., Zhang , X., Evers, J.B., van de r We rf, W., Zhang , L., 2022. Root
Acta Prataculturae Sin. 30, 73–85. plas ticity and inte rspe cific co mp leme ntarity improve yields and wate r us e
Wang , X., Chen , Y., Yang , K., Du an, F., Liu, P., Wang , Z., Wang , J., 2021. Effects of efficien cy of ma ize/so ybea n inte rcropp ing in a wate r-limite d co nd ition. Field
legu me inte rcropp ing and nitrog en inpu t on ne t gree nhou se ga s ba lances , Crop . Res. 282, 108523.
TE

inte ns ity, carbon fo ot print and crop prod uctivity in swee t ma ize crop land in Zo u, X., Shi, P., Zhang , C., Si, T., Wang , Y., Zhang , X., Yu, X., Wang , H., Wang , M ., 2021.
So uth China. J. Clea n. Prod . 314, 127997. Rotation al strip inte rcropp ing of ma ize and pe anuts has mu ltiple be ne fits fo r
Wang , Z., Zhao , X., Wu , P., He , J., Chen , X., Gao, Y., Cao, X., 2015a. Radiation ag ricultural prod uction in the no rthe rn ag ropa storal ec ot on e region of China.
inte rcep tion and utilization by whea t/ma ize strip inte rcropp ing system s. Ag ric. Eur. J. Ag ron. 129, 126304.
For. Me te orol. 204, 58–66.
Wang , S., Xion g , J., Yang , B., Yang , X., Du , T., Stee nhuis, T.S., Sidd ique , K.H.M ., Kang ,
S., 2023a. Diversified crop rotation s redu ce grou nd wate r us e and en hance
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