Intercropping is an important agronomic practice adopted to increase crop production and resource efficiency in areas with intensive agricultural production. Two sequential field trials were conducted in 2015–2016 to investigate the effect of shading on the morphological features, leaf structure, and photosynthetic characteristics of soybean in a maize-soybean relay-strip intercropping system. Three treatments were designed on the basis of different row configurations A1 (“50 cm + 50 cm” one row of maize and one row of soybean with a 50 cm spacing between the rows), A2 (“160 cm + 40 cm” two rows of maize by wide-narrow row planting, where two rows of soybean were planted in the wide rows with a width of 40 cm, and with 60 cm row spacing was used between the maize and soybean rows), and CK (sole cropping of soybean, with 70 cm rows spacing). Results showed that the photosynthetically active radiation transmittances of soybean canopy at V5 stage under A2 treatment (31.1%) were considerably higher than those under A1 (8.7%) treatment, and the red-to-far-red ratio was reduced significantly under A1 (0.7) and A2 (1.0) treatments compared with those under CK (1.2). By contrast with CK, stem diameter, total aboveground biomass, chlorophyll content and net photosynthetic rate decreased significantly except plant height under A1 and A2. The thickness of palisade tissue and spongy tissue of soybean leaf under A1 and A2 were significantly reduced at V5 stage compared with CK. The leaf thicknesses under A1 and A2 were lower than those in CK by 39.5% and 18.2%, respectively. At the R1 stage of soybean (after maize harvest), the soybean plant height, stem biomass, leaf biomass and petiole biomass under A1 and A2 treatments were still significantly lower than those under CK, but no significant differences were observed in Chl a/b, P_n, epidermis thickness and spongy tissue thickness of soybean leaves in A2 compared with CK. In addition, the soybean yields (g plant^-1) under A1 and A2 were 54.69% and 16.83% lower than those in CK, respectively. These findings suggested that soybean plants can regulate its morphological characteristics and leaf anatomical structures under different light environments.

The intensity and quality (red to far-red (R/Fr) ratio) of light directly affect growth of plant under shading. Gibberellins (GAs) and auxin [indole-3-acetic acid (IAA)] play important roles in mediating the shading adaptive responses of plants. Thus, the intensity and quality of the uncoupling light from shading were assessed to identify the influence of each component on the morphology and matter distribution of the leaf, stem, and petiole. This assessment was based on the changes in endogenous Gibberellin 1 (GA1) and IAA levels. Soybean plants were grown in a growth chamber with four treatments [normal (N), N+Fr, low (L), and L+Fr light]. Results revealed that the reductions in photosynthetically active radiation (PAR) and R/Fr ratio equally increased height and stem mass fractions (SMFs) of the soybean seedling. The light intensity significantly influenced the dry mass per unit area and mass fraction of soybean leaves, whereas the light quality regulated the petiole elongation and mass fraction. Low R/Fr ratio (high Fr light) increased the soybean biomass by improving the photosynthetic assimilation rate and quantum yield of photosystem II. In addition, the IAA and GA1 levels in the leaf, stem, and petiole did not reflect the growth response trends of each tissue toward light intensity and quality; however, trends of the IAA-to-GA1 content ratios were similar to those of the growth and matter allocation of each soybean tissue under different light environments. Therefore, the response of growth and matter allocation of soybean to light intensity and quality may be regulated by the IAA-to-GA1 content ratio in the tissues of the soybean plant.

Intercropping and close planting are important cultivation methods that increase soybean yield in agricultural production. However, plant shading is a major abiotic stress factor that influences soybean growth and development. Although shade affects leaf morphological parameters and decreases leaf photosynthesis capacity, information on the responses of soybean leaf photosynthesis to shading at proteomic level is still lacking. Compared with leaves under normal light (CK) treatment, leaves under shading treatment exhibited decreased palisade and spongy tissue thicknesses but significantly increased cell gap. Although shade increased the number of the chloroplast, the thickness of the grana lamella and the photosynthetic pigments per unit mass, but the size of the chloroplast and starch grains and the rate of net photosynthesis decreased compared with those of under CK treatment. A total of 248 differentially expressed proteins, among which 138 were upregulated, and 110 were downregulated, in soybean leaves under shading and CK treatments were detected via isobaric tags for relative and absolute quantification labeling in the three biological repeats. Differentially expressed proteins were classified into 3 large and 20 small groups. Most proteins involved in porphyrin and chlorophyll metabolism, photosynthesis-antenna proteins and carbon fixation in photosynthetic organisms were upregulated. By contrast, proteins involved in photosynthesis were downregulated. The gene family members corresponding to differentially expressed proteins, including protochlorophyllide reductase (Glyma06g247100), geranylgeranyl hydrogenase (Ggh), LHCB1 (Lhcb1) and ferredoxin (N/A) involved in the porphyrin and chlorophyll metabolism, photosynthesis-antenna proteins and photosynthesis pathway were verified with real-time qPCR. The results showed that the expression patterns of the genes were consistent with the expression patterns of the corresponding proteins. This study combined the variation of the soybean leaf structure and differentially expressed proteins of soybean leaves under shading. These results demonstrated that shade condition increased the light capture efficiency of photosystem II (PSII) in soybean leaves but decreased the capacity from PSII transmitted to photosystem II (PSI). This maybe the major reason that the photosynthetic capacity was decreased in shading.

Intercropping advantage occurs only when each species has adequate time and space to maximize cooperation and minimize competition between them. A field experiment was conducted for two consecutive years between 2013 and 2014 to investigate the effects of maize and soybean relay strip intercropping systems on the uptake and utilization of nitrogen, phosphorus, and potassium. The treatments included “40:160” (T1, maize narrow and wide row spacing of 40 and 160 cm, where two rows of soybean with a 40 cm row were planted in the wide rows. The area occupation ratio of maize and soybean both were 50% of the every experimental block), “80:120” (T2, maize narrow and wide row spacing of 80 and 120 cm, the soybean planting was the same as T1 treatment. The area occupation ratio of maize and soybean were 60% and 40% of the every experimental block), “100:100” (T3, one row of maize and one row of soybean with a 100-cm row. The area occupation ratio of maize and soybean was the same as T1 treatment), sole cropping of maize (CK1, The area occupation ratio of maize was 100% of the every experimental block), and sole cropping of soybean (CK2, The area occupation ratio of soybean was 100% of the every experimental block). The results show that, compared with the sole cropping system (sole maize), the economic yields in T1, T2, and T3 treatments increased by 761, 536, and 458 kg·ha⁻¹, respectively, and the biological yields increased by 2410, 2127, and 1588 kg·ha⁻¹. The uptake and utilization of nitrogen, phosphorus, and potassium in T1, T2, and T3 treatments were significantly higher than those in sole crops, and the nutrient advantage is mainly due to nutrient uptake rather than nutrient use efficiency. The land equivalent ratio values in T1, T2, and T3 treatments were 1.43, 1.32, and 1.20, respectively. In particular, the economic and biological yield in T1 treatment exhibited potential as an intercropping pattern.