Paper · 2026 · Journal of Applied Ecology

Intercropping and cover cropping differentially alter maize secondary metabolism and affect herbivore resistance

Juan Pablo Jordán; André Kessler

8
claims
16
verbatim quotes
2
citations
3
research areas
✓
Open access

Abstract

Abstract Diversified agricultural systems such as intercropping and cover cropping provide a wide range of emergent properties that can positively impact agricultural performance metrics. Both cropping systems by leveraging distinct ecological processes such as associational resistance and plant–soil feedbacks can facilitate soil health while increasing pest resistance and crop productivity. However, the extent to which the cropping system in general and the companion plant species in particular induce changes to plant metabolism as a mechanism mediating plant resistance remains understudied and limits a wider application of companion cropping technologies. To close this gap, we use microcosm experiments that simulate intercropping and cover cropping practices by manipulating plant neighbourhood and soil conditioning to investigate how three different companion legume species change plant secondary metabolite profiles and the resulting effects on herbivore resistance of maize plants. By utilizing a conventional maize variety (genetically modified; bt‐transformed), we further test the potential for such ecosystem services to persist under industrialized agricultural systems. The non‐volatile secondary metabolite profiles of maize leaves and roots differed across intercropping and cover cropping systems, while volatile organic compound emissions remained largely unchanged. Defence‐related compounds such as benzoxazinoids were generally upregulated under intercropping, while the companion plant identity further shaped the maize defence profile with bean intercropping specifically upregulating benzoxazinoid compounds. Resistance bioassays with the generalist herbivore Spodoptera frugiperda revealed that intercropping significantly reduces leaf damage relative to cover cropping. Among the intercrops, beans increase larval mortality while alfalfa decreases larval mortality on maize plants. Synthesis and applications . We demonstrate that intercropping and cover cropping can shape the overall secondary metabolite profiles of maize and mediate herbivore resistance. These findings underscore that crop diversification strategies can specifically modulate the expression of chemical defences linked to herbivore resistance. In practical terms, this means that diversified cropping systems can be optimized to enhance constitutive levels of pest resistance, potentially reducing the reliance on insecticides, increasing sustainability and supporting yield. The fact that positive resistance effects persist even when using pest‐resistant GM crops suggests companion cropping practices as a valid tool to increase the sustainable management of insect pests.

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Claims 8

Intervention effectobservedoriginal resultmeasured

Claim · model-written summary

Intercropping and cover cropping differed in non-volatile secondary metabolite profiles of maize leaves and roots.

Taxon
maizeZea mayst0
Treatment
intercropping and cover cropping systemse1
Outcome
non‐volatile secondary metabolite profiles of maize leaves and rootse1
Direction
differede1

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

t0

“maize secondary metabolism”

✓ located verbatim in the source · title, characters 54–80 · show in text ↑
e1

“The non‐volatile secondary metabolite profiles of maize leaves and roots differed across intercropping and cover cropping systems”

✓ located verbatim in the source · abstract, characters 1193–1322 · show in text ↑
Observationobservedoriginal resultno effectmeasured

Claim · model-written summary

Volatile organic compound emissions remained largely unchanged across cropping systems.

Taxon
maizeZea mayst0
Outcome
volatile organic compound emissionse2
Direction
remained largely unchangede2

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

t0

“maize secondary metabolism”

✓ located verbatim in the source · title, characters 54–80 · show in text ↑
e2

“while volatile organic compound emissions remained largely unchanged”

✓ located verbatim in the source · abstract, characters 1324–1392 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Intercropping generally upregulated defence-related benzoxazinoids in maize.

Taxon
maizeZea mayst0
Treatment
intercroppinge3
Outcome
benzoxazinoidse3
Direction
upregulatede3

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

t0

“maize secondary metabolism”

✓ located verbatim in the source · title, characters 54–80 · show in text ↑
e3

“Defence‐related compounds such as benzoxazinoids were generally upregulated under intercropping”

✓ located verbatim in the source · abstract, characters 1394–1489 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Bean intercropping specifically upregulated benzoxazinoid compounds in maize.

Taxon
maizeZea mayst0
Treatment
bean intercroppinge4
Outcome
benzoxazinoid compoundse4
Direction
upregulatinge4

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

t0

“maize secondary metabolism”

✓ located verbatim in the source · title, characters 54–80 · show in text ↑
e4

“bean intercropping specifically upregulating benzoxazinoid compounds”

✓ located verbatim in the source · abstract, characters 1572–1640 · show in text ↑
Intervention effectobservedoriginal resultpositivesignificant (as stated)measured

Claim · model-written summary

Intercropping significantly reduced leaf damage by Spodoptera frugiperda relative to cover cropping.

Taxon
maizeZea mayst0
Treatment
intercroppinge5
Interactor
Spodoptera frugiperdaSpodoptera frugiperdae5herbivore
Outcome
leaf damagee5
Direction
reducese5
Comparator
cover croppinge5

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗
Spodoptera frugiperda
Spodoptera frugiperda · InteractorUSGS Bee Inventory and Monitoring Lab from Beltsville, Maryland, USA · Public domain · Wikimedia Commons ↗

Verbatim

t0

“maize secondary metabolism”

✓ located verbatim in the source · title, characters 54–80 · show in text ↑
e5

“Resistance bioassays with the generalist herbivore Spodoptera frugiperda revealed that intercropping significantly reduces leaf damage relative to cover cropping”

✓ located verbatim in the source · abstract, characters 1642–1803 · show in text ↑
Intervention effectobservedoriginal resultpositivemeasured

Claim · model-written summary

Beans increased larval mortality on maize.

Taxon
maizeZea mayst0
Treatment
beanse6
Outcome
larval mortalitye6
Direction
increasee6

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

t0

“maize secondary metabolism”

✓ located verbatim in the source · title, characters 54–80 · show in text ↑
e6

“Among the intercrops, beans increase larval mortality while alfalfa decreases larval mortality on maize plants”

✓ located verbatim in the source · abstract, characters 1805–1915 · show in text ↑
Intervention effectobservedoriginal resultnegativemeasured

Claim · model-written summary

Alfalfa decreased larval mortality on maize.

Taxon
maizeZea mayst0
Treatment
alfalfae6
Outcome
larval mortalitye6
Direction
decreasese6

Not reportedComparator

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

t0

“maize secondary metabolism”

✓ located verbatim in the source · title, characters 54–80 · show in text ↑
e6

“Among the intercrops, beans increase larval mortality while alfalfa decreases larval mortality on maize plants”

✓ located verbatim in the source · abstract, characters 1805–1915 · show in text ↑
Recommendationauthor interpretationoriginal resultbasis n/a

Claim · model-written summary

Companion cropping may be a valid tool to increase the sustainable management of insect pests in maize.

Taxon
maizeZea mayst0
Outcome
sustainable management of insect pestse7

Organisms pictured · Representative pictures of the matched species, not figures from the paper.

Zea mays
Zea mays · TaxonAkwugo · CC BY-SA 4.0 · Wikimedia Commons ↗

Verbatim

t0

“maize secondary metabolism”

✓ located verbatim in the source · title, characters 54–80 · show in text ↑
e7

“companion cropping practices as a valid tool to increase the sustainable management of insect pests”

✓ located verbatim in the source · abstract, characters 2582–2681 · show in text ↑

Medicago sativa · pest · Zea mays (maize)Positive and negative results

4 papers: 3 positive, 1 negative or no effect

3 in the opposite direction

  1. positive
  2. positive
  3. positive
See all 4 papers →

intercropping · Spodoptera frugiperda · Zea mays (maize)Positive and negative results

7 papers: 6 positive, 1 negative or no effect

5 papers in the same direction as this paper · 1 in the opposite direction

  1. no effect
  2. positive
  3. positive

3 papers more in this group

See all 7 papers →

Claims are grouped by organism, treatment and what was measured or targeted. A paper counts as positive when any of its claims in the group is positive.