Intercropping improves resource utilization. Under wide-narrow-row maize 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 under unilateral weak light. Our results showed that the net photosynthetic rate (Pn) 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 ¹³C into the grains. However, sink loss with ear removal reduced the Pn of unshaded leaves. Intense unilateral shade (40% to 20% normal light) reduced grain yield (37.6% to 54.4%) but not mild unilateral shade (60% normal light). We further found that in unshaded leaves, Agpsl, Bmy, and Mexl-like significantly influenced sucrose and starch metabolism, while Sweet13a and Sut1 were crucial for sugar export. In shaded leaves, Sps1, Agpsl, and Sweet13c were 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 that provides appropriate unilateral shade and limits yield reduction.

The border row crop in strip intercropped maize is often exposed to heterogeneous light conditions, resulting in increased photosynthesis and yield decreased. Previous studies have focused on photosynthetic productivity, whereas carbon allocation could also be one of the major causes of decreased yield. However, carbon distribution remains unclear in partially shaded conditions. In the present study, we applied heterogeneous light conditions (T), and one side of plants was shaded (T-30%), keeping the other side fully exposed to light (T-100%), as compared to control plants that were exposed entirely to full-light (CK). Dry weight, carbon assimilation, ¹³C abundance, and transport tissue structure were analyzed to clarify the carbon distribution in partial shading of plants. T caused a marked decline in dry weight and harvest index (HI), whereas dry weight in unshaded and shaded leaves did not differ. Net photosynthesis rate (Pn), the activity of sucrose phosphate synthase enzymes (SPS), and sucrose concentration increased in unshaded leaves. Appropriately, 5.7% of the ¹³C from unshaded leaves was transferred to shaded leaves. Furthermore, plasmodesma density in the unshaded (T-100%) and shaded (T-30%) leaves in T was not significantly different but was lower than that of CK. Similarly, the vascular bundle total area of T was decreased. ¹³C transfer from unshaded leaves to ear in T was decreased by 18.0% compared with that in CK. Moreover, ¹³C and sucrose concentration of stem in T were higher than those in CK. Our results suggested that, under heterogeneous light, shaded leaves as a sink imported the carbohydrates from the unshaded leaves. Ear and shaded leaf competed for carbohydrates, and were not conducive to tissue structure of sucrose transport, resulting in a decrease in the carbon proportion in the ear, harvest index, and ear weight.