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Intercropping improves resource utilization. Under wide–narrow-row maize (Zea mays) intercropping, maize plants are subjected to weak unilateral illumination and exhibit high photosynthetic performance. However, the mechanism regulating photosynthesis under unilateral weak light remains unknown. We investigated the relationship between photosynthesis and sugar metabolism in maize under unilateral weak light. Our results showed that the net photosynthetic rate (P n) of unshaded leaves increased as the level of shade on the other side increased. On the contrary, the concentration of sucrose and starch and the number of starch granules in the unshaded leaves decreased with increased shading due to the transfer of abundant C into the grains. However, sink loss with ear removal reduced the P n of unshaded leaves. Intense unilateral shade (40% to 20% normal light), but not mild unilateral shade (60% normal light), reduced grain yield (37.6% to 54.4%, respectively). We further found that in unshaded leaves, Agpsl , Bmy , and Mexl-like expression significantly influenced sucrose and starch metabolism, while Sweet13a and Sut1 expression was crucial for sugar export. In shaded leaves, expression of Sps1 , Agpsl , and Sweet13c was crucial for sugar metabolism and export. This study confirmed that unshaded leaves transported photosynthates to the ear, leading to a decrease in sugar concentration. The improvement of photosynthetic performance was associated with altered sugar transport. We propose a narrow-row spacing of 40 cm, which provides appropriate unilateral shade and limits yield reduction. [ABSTRACT FROM AUTHOR]

Soybean in relay intercropping is initially exposed to a shade environment, followed by exposure to full sunlight after the harvesting of primary crops, e.g., maize. Therefore, soybean's ability to acclimate to this changing light environment determines its growth and yield formation. However, the changes in soybean photosynthesis under such light alternations in relay intercropping are poorly understood. This study compared the photosynthetic acclimation of two soybean varieties with contrasting shade tolerance, i.e., Gongxuan1 (shade-tolerant) and C103 (shade-intolerant). The two soybean genotypes were grown in a greenhouse under full sunlight (HL) and 40% full sunlight (LL) conditions. Subsequently, after the fifth compound leaf expanded, half of the LL plants were transferred to a high-sunlight environment (LL-HL). Morphological traits were measured at 0 and 10 days, while chlorophyll content, gas exchange characteristics and chlorophyll fluorescence were assayed at 0, 2, 4, 7 and 10 days after transfer to an HL environment (LL-HL). Shade-intolerant C103 showed photoinhibition 10 days after transfer, and the net photosynthetic rate (Pn) did not completely recover to that under a high light level. On the day of transfer, the shade-intolerant variety, C103, exhibited a decrease in net photosynthetic rate (Pn), stomatal conductance (Gs) and transpiration rate (E) in the low-light (LL) and low-light-to-high-light (LL-HL) treatments. Additionally, intercellular CO2 concentration (Ci) increased in low light, suggesting that non-stomatal factors were the primary limitations to photosynthesis in C103 following the transfer. In contrast, the shade-tolerant variety, Gongxuan1, displayed a greater increase in Pn 7 days after transfer, with no difference observed between the HL and LL-HL treatments. Ten days after transfer, the shade-tolerant Gongxuan1 exhibited 24.1%, 10.9% and 20.9% higher biomass, leaf area and stem diameter than the intolerant C103. These findings suggest that Gongxuan1 possesses a higher capacity to adapt to variations in light conditions, making it a potential candidate for variety selection in intercropping systems. [ABSTRACT FROM AUTHOR]

Intercropping improves land-use efficiency under conditions of limited land and resources, but no information is currently available pertaining to land-use efficiency and silage quality based on whole-plant utilization. Therefore, a two-year field experiment was conducted with the following conditions: three maize–soybean strip intercropping patterns (SIPs), comprising two maize rows along with two, three, or four soybean rows (2M2S, 2M3S, and 2M4S, respectively); and two sole cropping patterns of maize (SM) and soybean (SS). The aim was to evaluate the biomass yield and silage quality under each condition. Our results showed that all SIPs had a land equivalent ratio (LER) of over 1.6 based on both fresh and dry matter yield, and a higher whole plant yield, compared to sole cropping. Specifically, 2M3S exhibited the highest whole crop dry matter LER (1.8–1.9) and yield (24.6–27.2 t ha−1) compared to SM and SS (20.88–21.49 and 3.48–4.79 t ha−1, respectively). Maize–soybean mixed silages also showed better fermentation quality with higher lactic acid content (1–3%) and lower ammonia-N content (2–8%) compared to SS silages, and higher crude protein content (1–1.5%) with lower ammonia-N content (1–2%) compared to SM silage. Among the intercropping patterns, 2M3S had the highest fermentation quality index V-score (92–95). Consequently, maize–soybean strip intercropping improved silage quality and biomass yield, with 2M3S being recommended, due to its highest LER and biomass yield, and most optimal silage quality. [ABSTRACT FROM AUTHOR]

Recovery growth is common in relay intercropping systems and is a useful practice to increase total productivity of these systems. A field experiment was conducted during 2014–2016 to investigate the phenotypic and physiological traits related to light interception and utilization changes and cultivars difference of soybean during the recovery growth period. Two cropping patterns (maize–soybean relay strip intercropping (RSI), analogous soybean monoculture (CK)) and two soybean cultivars (shade‐tolerant Nandou12 and shade non‐tolerant Jiuyuehuang) were evaluated. During the co‐growth period, the growth of both two soybean cultivars was suppressed and showed a series of shade avoidance traits. After maize harvest, the LAI and leaflet number of intercropped soybean exhibited rapid growth, the LAI of Nandou12 (3.09) and Jiuyuehuang (1.32) recovered to 79% and 43% of corresponding CK level, respectively. Subsequently, final biomass of Nandou12 (3972.7 kg ha−1) and Jiuyuehuang (2263.3 kg ha−1) recovered 85.7% and 52% of corresponding CK level, respectively. Leaf blade thickness of Nandou12 recovered to the corresponding CK level, while Jiuyuehuang was not. Pn recovered to 84.5% and 82.8% of sole Pn at 80 days after soybean sowing in 2015 and to 72% and 91.2% at 66 days after soybean sowing in 2016 for Nandou12 and Jiuyuehuang, respectively. The above recovery process promoted the recovery of pod formation and grain yield. The yield of Nandou12 (1159.5 kg ha−1) has no significant difference between the CK level (1325.5 kg ha−1) but significantly higher than that of Jiuyuehuang (566.8 kg ha−1). Taken together, some recovery growth did occur in Jiuyuehuang, but it did not completely compensate for the growth reduction that occurred during the co‐growth period. Shade‐tolerant soybean cultivar has more stronger recovery growth ability in LAI, leaflet number, leaf thickness, and photosynthesis after maize harvest, which promoted the recovery growth of aboveground biomass and grain yield. [ABSTRACT FROM AUTHOR]

Insufficient and unbalanced biomass supply inhibited soybean [ Glycine max (L.) Merr.] yield formation in the maize-soybean relay strip intercropping (IS) and monoculture soybean (SS). A field experiment was conducted to explore the soybean yield increase mechanism of DA-6 in IS and SS treatments. In this 2-year experiment, compact maize "Denghai 605" and shade-tolerant soybean "Nandou 25" were selected as cultivated materials. DA-6 with four concentrations, i.e., 0 mg/L (CK), 40 mg/L (D40), 60 mg/L (D60), and 80 mg/L (D80), were sprayed on soybean leaves at the beginning of flowering stage of soybean. Results showed that DA-6 treatments significantly (p < 0.05) increased soybean grain yield, and the yield increase ratio was higher in IS than SS. The leaf area index values and net photosynthesis rate of IS peaked at D60 and were increased by 32.2–49.3% and 24.1–27.2% compared with the corresponding CK. Similarly, DA-6 treatments increased the aboveground dry matter and the amount of soybean dry matter accumulation from the R1 stage to the R8 stage (VDMT) and highest at D60 both in IS and SS. D60 increased the VDMT by 29.0–47.1% in IS and 20.7–29.2% in SS. The TR G at D60 ranged 72.4–77.6% in IS and 61.4–62.5% in SS. The MDA content at D60 treatment was decreased by 38.3% in IS and 25.8% in SS. The active grain-filling day in IS was about 7 days longer than in SS. In D60 treatment, the Vmean and Vmax increased by 6.5% and 6.5% in IS and 5.7% and 4.3% in SS compared with the corresponding CK. Although the pod number and hundred-grain weight were significantly (p < 0.05) increased by DA-6 treatments, the grains per pod were maintained stable. The pod number and hundred-grain weight were increased by 30.1–36.8% and 4.5–6.7% in IS and 6.3–13% and 3.6–5.6% in SS. Thus, the grain yield at D60 was increased by 36.7–38.4% in IS and 21.7–26.6% in SS. DA-6 treatments significantly (p < 0.05) increased soybean grain yield and peaked D60 treatments both in IS and SS. [ABSTRACT FROM AUTHOR]

Mildew severely reduces soybean yield and quality, and pods are the first line of defence against pathogens. Maize‐soybean intercropping (MSI) reduces mildew incidence on soybean pods; however, the mechanism remains unclear. Changing light (CL) from maize shading is the most important environmental feature in MSI. We hypothesized that CL affects isoflavone accumulation in soybean pods, affecting their disease resistance. In the present study, shading treatments were applied to soybean plants during different developmental stages according to various CL environments under MSI. Chlorophyll fluorescence imaging (CFI) and classical evaluation methods confirmed that CL, especially vegetative stage shading (VS), enhanced pod resistance to mildew. Further metabolomic analyses and exogenous jasmonic acid (JA) and biosynthesis inhibitor experiments revealed the important relationship between JA and isoflavone biosynthesis, which had a synergistic effect on the enhanced resistance of CL‐treated pods to mildew. VS promoted the biosynthesis and accumulation of constitutive isoflavones upstream of the isoflavone pathway, such as aglycones and glycosides, in soybean pods. When mildew infects pods, endogenous JA signalling stimulated the biosynthesis of downstream inducible malonyl isoflavone (MIF) and glyceollin to improve pod resistance. Changing light from maize shading is the most important environmental feature in maize‐soybean intercropping, which reduces mildew incidence on soybean (Glycine max L. Merr.) pods.Weak to bright light conditions promotes constitutive isoflavone biosynthesis in soybean pods, and provides sufficient precursors for downstream inducible malonylated isoflavones and glyceolin to improve pod resistance.JA signalling and isoflavone have a synergistic effect on the enhanced resistance of changing light treated pods to mildew. [ABSTRACT FROM AUTHOR]

China purchases around 62% of the soybean that is traded internationally and is currently only 14.3% self-sufficient, which strains the global food supply. A key question is to what extent China can produce soybeans to meet its demand. Here we used 3 years of data for 26 soybean cultivars across 146 locations in China, combined with APSIM modeling, to quantify soybean yield spatial variations and the impact of varietal and climate changes on soybean yield potential. We further analyzed the key drivers of national soybean production and explored a potential solution to increase national soybean production with maize-soybean strip intercropping system based on data from 32 locations in the past 10 years. The results showed that the contrasting climate conditions across regions in China resulted in diverse maturity types of soybean cultivars, ranging from MG000 to MGⅧ, with grain yield ranging from 1.4 t ha−1 to 4.5 t ha−1. The key climate drivers are temperature at high latitude cool regions and radiation at low latitude hot and humid regions. The key physiological trait for increasing yield is grain number that is closely related to post-anthesis growth. There exists an obvious yield gap between potential yield and actual yield. At national level, total soybean productivity was limited by planting area due to competing maize crop. It remains difficult for soybean to replace maize due to economic competitiveness. However, maize-soybean strip intercropping offers a sustainable solution to produce an additional soybean crop with around half of the yield of a sole soybean while maintaining the current maize yield. This would enable to potentially produce additional 67.4 million tons of soybean yield each year, with minimum reduction of maize yield (land equivalent ratio of maize =0.96). Fully closing the gap between actual unit yield and potential yield level would only increase national soybean production by 8 million tons. Further increase in production rely on increasing soybean planting area is unrealistic due to the more profitable maize crop. Maize-soybean strip intercropping offers a sustainable solution to potentially produce additional 67.4 million tons of soybean yield while maintaining the current maize yield. • We used APSIM NG model to quantify the impact of varietal and climate changes on soybean yield potential. • Grain yield of soybean ranging from 1.4 t ha-1–4.5 t ha-1 across regions in China. • The key climate drivers and physiological trait for increasing soybean yield have been identified. • Maize-soybean strip intercropping would enable to potentially produce additional 67.4 million tons of soybean yield. [ABSTRACT FROM AUTHOR]

Multi-species intercropping is a sustainable agricultural practice worldwide used to utilize resources more efficiently. In intercropping systems, short crops often grow under vegetative shade of tall crops. Soybean, one important legume, is often planted in intercropping. However, little is known about the mechanisms of shade inhibition effect on leaf size in soybean leaves at the transcriptome level. We analyzed the transcriptome of shaded soybean leaves via RNA-Seq technology. We found that transcription 1085 genes in mature leaves and 1847 genes in young leaves were significantly affected by shade. Gene ontology analyses showed that expression of genes enriched in polysaccharide metabolism was down-regulated, but genes enriched in auxin stimulus were up-regulated in mature leaves; and genes enriched in cell cycling, DNA-replication were down-regulated in young leaves. These results suggest that the inhibition of higher auxin content and shortage of sugar supply on cell division and cell expansion contribute to smaller and thinner leaf morphology, which highlights potential research targets such as auxin and sugar regulation on leaves for crop adaptation to shade in intercropping. [ABSTRACT FROM AUTHOR]

BACKGROUND RESULTS CONCLUSION Relay intercropping of maize and soybean can improve land productivity. However, the mechanism behind N2O emissions in this practice remains unclear. A two‐factor randomized block field trial was conducted to reveal the mechanism of N2O emissions in a full additive maize–soybean relay intercropping. Factor A was three cropping systems – that is, monoculture maize (Zea mays L.), monoculture soybean (Glycine max L. Merr.) and maize–soybean relay intercropping. Factor B was different N supply, containing no N, reduced N and conventional N. Differences in N2O emissions, soil properties, rhizosphere bacterial communities and yield advantage were evaluated.The land equivalent ratio was 1.55–2.44, and the cumulative N2O emission (CEN2O$$ \mathrm{C}{\mathrm{E}}_{{\mathrm{N}}_2\mathrm{O}} $$) was notably lower by 60.2% in intercropping than in monoculture, respectively. Reduced N declined CEN2O$$ \mathrm{C}{\mathrm{E}}_{{\mathrm{N}}_2\mathrm{O}} $$ without penalty on the yield advantages. The relay intercropping shifted soil properties – for example, soil organic matter, total N, NH4+$$ {\mathrm{NH}}_4^{+} $$ and protease activity – and improved the soil microorganism community – for example, Proteobacteria and Acidobacteria. Intercropping reduced CEN2O$$ \mathrm{C}{\mathrm{E}}_{{\mathrm{N}}_2\mathrm{O}} $$ by directly suppressing nirS‐ and amoA‐regulated N2O generation during soil N cycling, or nirS‐ and amoA‐mediated soil properties shifted to reduce CEN2O$$ \mathrm{C}{\mathrm{E}}_{{\mathrm{N}}_2\mathrm{O}} $$ indirectly. Reduced N directly reduced CEN2O$$ \mathrm{C}{\mathrm{E}}_{{\mathrm{N}}_2\mathrm{O}} $$ by decreasing soil N content and reducing soil microorganism activities to alleviate N2O produced in soil N cycling.Conducting a full additive maize–soybean relay intercropping with reduced nitrogen supply provides a way to alleviate N2O emissions without the penalty on the yield advantage by changing rhizosphere bacterial communities and soil N cycling. © 2024 Society of Chemical Industry. [ABSTRACT FROM AUTHOR]

• We implemented the light interception model for strip intercropping into APSIM-Classic. • A quasi-Bayesian optimisation approach was used to derive parameter by constraining the model to LAI, biomass and yield. • Key model parameters were changed with row configuration, indicating a need to improve modelling physiology of intercrops. • A potential application is to optimize row configurations in strip intercropping across climates. Intercropping of two or more crop species increases the efficiency of resource use and often produces a greater yield per unit land area. The relative efficiency of intercropping depends on row configuration, but there is a shortage of modelling-based evaluation of alternative intercropping options due to the inadequacy of standard process-based crop models to simulate resource capture, growth and yield formation when the canopy is spatially structured in strips. We implemented a light interception model for strip crops into the APSIM Classic model and combined it with a quasi-Bayesian approach to derive the model parameters to simulate crop growth and grain yield in maize-soybean strip intercropping. We used 4 years of field data for 5 different row configurations to derive key model parameters for simulation of light interception, LAI dynamics, biomass growth and grain yield of maize and soybean intercrops. Key model parameters (e.g. RUE, k etc.) were found to change with row-spacing and configuration, posing challenges to simulate different configurations with a single parameter set. The potential ranges of these key parameters were derived by constraining the model to observed data. The model can be potentially used to evaluate impact of planting configurations on productivity of strip intercropping systems, but the variability of key model parameters among configuration treatments calls for further in-depth research to improve modelling physiology of strip intercrops. [ABSTRACT FROM AUTHOR]

Relay intercropping soybean [ Glycine max (L.) Merr.] with cereal provides an approach to increase land productivity and improve N fixation. To explore the physiological mechanisms underlying the N fixation, leaf and nodule traits, biomass and N accumulations, nodule sucrose content, paraffin section and transmission electron microscope observations, and nodule antioxidant capacity were compared in sole and relay intercropped soybean with N levels. The growth of relay intercropped soybean was suppressed during the coexistence duration, and N addition promotes leaf growth and suppressed nodule development. After maize harvest, a boosted recovery of intercropped soybean than sole cropping notably enhanced sucrose by 20.4% allocated to the nodule. Sufficient sucrose partition delayed nodule senescence by increasing antioxidant capacity in intercropped soybeans contrast in the sole cropping. The delayed nodule senescence of intercropped soybean continuously provides ureides to the aboveground. The ureide-N in total N of intercropped soybean was 37.3% notably higher than sole cropping at the R5 stage. Finally, the adverse effects of shade on soybean growth and N fixation during the coexistence duration were compensated through the recovery growth, which benefits obtaining yield advantage of the intercropped soybean. • A boosting recovery growth increasing nodule sucrose partition in relay intercropping. • A higher sucrose partition to nodules delayed nodule senescence. • The delayed nodule senescence improves shoot ureides accumulation in relay intercropping. [ABSTRACT FROM AUTHOR]

The maize-soybean intercropping system has become increasingly popular in many areas of the world, particularly in China, due to its high productivity and the harvest of two different grains. While efforts have been made to maintain the yield of the taller maize crop, there is limited understanding of how the morphology, growth and yield of the lower soybean crop changes in response to the shading by maize. We therefore conducted a three-year field experiment from 2013 to 2015 to investigate the changes in light environment, growth of individual organs, biomass, and grain yield of soybean under two intercropping patterns (1M1S, one row of maize with one row of soybean; 2M2S, two rows of maize with two rows of soybean) as compared to monoculture. Our results showed that at soybean flowering stage, the R:FR ratio at the top of soybean canopy was reduced 17–21% more than the photosynthetically active radiation (PAR) under intercropping compared to monoculture, with 15–19% more reduction under 1M1S than 2M2S. This led to increased internode lengths, plant height and specific leaf area (SLA), but reduced branching of soybean plants under intercropping. These morphological changes enabled the crop to intercept relatively more light and the shading also increased the light use efficiency (LUE) of soybean. However, these positive responses were not able to compensate the effect of reduced leaf area (due to smaller leaf size and less branching) and total light interception, leading to reduced biomass and grain. The reduction in grain yield was mainly caused by the reduced number of grains (particularly on the middle nodes) produced by the intercropped soybean plants, while the grain size remained unchanged. The data and results of this study may be used to develop and parameterize crop models for simulating development and growth of soybean crop in response to changes in the light environment under intercropping. [ABSTRACT FROM AUTHOR]

Maize–soybean strip intercropping can effectively alleviate arable land competition, improve land output, and be conducive to the sustainable development. The key to achieve these advantages is to stabilize the maize yield. Suitable narrow row spacing could maximize the benefit of maize–soybean strip intercropping, and an increasing bandwidth with suitable narrow row spacing is beneficial to the mechanized production of soybean. In practice, to ensure the maize yield in additive maize strip intercropping, maize plant spacing decreases and intraspecific competition intensifies as the bandwidth increases. However, interspecific interactions have been the primary focus of studies because of the better rhizosphere nitrogen fixation ability of soybean and the activation of maize rhizosphere nutrients. Therefore, it is particularly urgent to identify the trade-offs between intraspecific competition and interspecific interactions in maize and the regulatory effects on maize yield. Field experiments were conducted from 2017 to 2020 to analyze the results of interindividual competition in maize and interactions with soybeans under different bandwidths. It was found that maize biomass and yield at the maturity stage decreased by 8.0% and 6.4%; 20.5% and 19.4%; and 28.7% and 44.1% from 2 m to 2.8 m, respectively, but the biomass proportion in ears showed no significant difference. Intraspecific competition of maize was weakened by special promotion in intercropping, but stable yield was achieved only at 2 m. In response to suitable competition, maize roots can flourish only on the basis of ensuring aboveground biomass. The results revealed that the primary negative determinant of maize yield was intraspecific aboveground competition and strengthening intraspecific competition reduced biomass accumulation without altering the allocation ratio to the ear. We concluded that maize yield was guaranteed at 2 m due to the special strip planting design compensated for the enhanced intraspecific competition of maize aboveground parts in the additive maize–soybean strip intercropping system. • Lower intraspecific competition was a driver of maize yield under different bandwidths. • The aboveground part of maize contributed more to the yield advantage than the underground part. • Decrease in biomass, not the proportion of partitioning to the ear, affected the maize yield. [ABSTRACT FROM AUTHOR]