Plants are adapted to sense numerous stress stimuli and mount efficient defense responses by directing intricate signaling pathways. They respond to undesirable circumstances to produce stress-inducible phytochemicals that play indispensable roles in plant immunity. Extensive studies have been made to elucidate the underpinnings of defensive molecular mechanisms in various plant species. Transcriptional factors (TFs) are involved in plant defense regulations through acting as mediators by perceiving stress signals and directing downstream defense gene expression. The cross interactions of TFs and stress signaling crosstalk are decisive in determining accumulation of defense metabolites. Here, we collected the major TFs that are efficient in stress responses through regulating secondary metabolism for the direct cessation of stress factors. We focused on six major TF families including AP2/ERF, WRKY, bHLH, bZIP, MYB, and NAC. This review is the compilation of studies where researches were conducted to explore the roles of TFs in stress responses and the contribution of secondary metabolites in combating stress influences. Modulation of these TFs at transcriptional and post-transcriptional levels can facilitate molecular breeding and genetic improvement of crop plants regarding stress sensitivity and response through production of defensive compounds.

Different planting patterns affect the light interception of intercrops under intercropping conditions. Here we revealed that narrow-wide-row relay-intercropping improves the light interception across maize leaves in wide rows (60cm) and narrow rows (40cm), accelerated the biomass production of intercrop-species and compensated the slight maize yield loss by considerably increasing the soybean yield. In a two-year experiment, maize was planted with soybean in different planting patterns (1M1S, 50:50cm and 2M2S, 40:60cm) of relay-intercropping, both planting patterns were compared with sole cropping of maize (M) and soybean (S). As compared to M and 1M1S, 2M2S increased the total light interception of maize leaves in wide rows (WR) by 27% and 23%, 20% and 10%, 16% and 9% which in turn significantly enhanced the photosynthetic rate of WR maize leaves by 7% and 5%, 12% and 9%, and 19% and 4%, at tasseling, grain-filling and maturity stage of maize, respectively. Similarly, the light transmittance at soybean canopy increased by 218%, 160% and 172% at V 2 , V 5 and R 1 stage in 2M2S compared with 1M1S. The improved light environment at soybean canopy in 2M2S considerably enhanced the mean biomass accumulation, and allocation to stem and leaves of soybean by 168%, and 131% and 207%, respectively, while it decreased the mean biomass accumulation, and distribution to stem, leaves and seed of maize by 4%, and 4%, 6% and 5%, respectively than 1M1S. Compared to 1M1S, 2M2S also increased the CR values of soybean (by 157%) but decreased the CR values of maize (by 61%). Overall, under 2M2S, relay-cropped maize and soybean produced 94% and 69% of the sole cropping yield, and the 2M2S achieved LER of 1.7 with net income of 1387.7 US $ ha -1 in 2016 and 1434.4 US $ ha -1 in 2017. Our findings implied that selection of optimum planting pattern (2M2S) may increase the light interception and influence the light distribution between maize and soybean rows under relay-intercropping conditions which will significantly increase the intercrops productivity. Therefore, more attention should be paid to the light environment when considering the sustainability of maize-soybean relay-intercropping via appropriate planting pattern selection.

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Strip width management is a critical factor for producing higher crop yields in relay intercropping systems. A 2-year field experiment was carried out during 2012 and 2013 to evaluate the effects of different strip width treatments on dry-matter production, major-nutrient (nitrogen, phosphorus, and potassium) uptake, and competition parameters of soybean and maize in relay intercropping system. The strip width (SW) treatments were 0.40, 0.40, and 0.40 m (SW1); 0.40, 0.40, and 0.50 m (SW2); 0.40, 0.40, and 0.60 m (SW3); and 0.40, 0.40, and 0.70 m (SW4) for soybean row spacing, maize row spacing, and spacing between soybean and maize rows, respectively. As compared to sole maize (SM) and sole soybean (SS), relay-intercropped maize and soybean accumulated lower quantities of nitrogen, phosphorus, and potassium in all treatments. However, maize in SW1 accumulated higher nitrogen, phosphorus, and potassium than SW4 (9%, 9%, and 8% for nitrogen, phosphorus, and potassium, respectively). Soybean in SW3 accumulated 25% higher nitrogen, 33% higher phosphorus, and 24% higher potassium than in SW1. The improved nutrient accumulation in SW3 significantly increased the soybean dry matter by 19%, but slightly decreased the maize dry matter by 6% compared to SW1. Similarly, SW3 increased the competition ratio value of soybean (by 151%), but it reduced the competition ratio value of maize (by 171%) compared to SW1. On average, in SW3, relay-cropped soybean produced 84% of SS seed yield and maize produced 98% of SM seed yield and achieved the land equivalent ratio of 1.8, demonstrating the highest level in the world. Overall, these results suggested that by selecting the appropriate strip width (SW3; 0.40 m for soybean row spacing, 0.40 m maize row spacing, and 0.60 m spacing between soybean and maize rows), we can increase the nutrient 2 of 14 | RAZA et Al.

Drastic changes in the climate and ecosystem due to natural or anthropogenic activities have severely affected crop production globally. This concern has raised the need to develop environmentally friendly and cost-effective strategies, particularly for keeping pace with the demands of the growing population. The use of nanobiofertilizers in agriculture opens a new chapter in the sustainable production of crops. The application of nanoparticles improves the growth and stress tolerance in plants. Inoculation of biofertilizers is another strategy explored in agriculture. The combination of nanoparticles and biofertilizers produces nanobiofertilizers, which are cost-effective and more potent and eco-friendly than nanoparticles or biofertilizers alone. Nanobiofertilizers consist of biofertilizers encapsulated in nanoparticles. Biofertilizers are the preparations of plant-based carriers having beneficial microbial cells, while nanoparticles are microscopic (1–100 nm) particles that possess numerous advantages. Silicon, zinc, copper, iron, and silver are the commonly used nanoparticles for the formulation of nanobiofertilizer. The green synthesis of these nanoparticles enhances their performance and characteristics. The use of nanobiofertilizers is more effective than other traditional strategies. They also perform their role better than the common salts previously used in agriculture to enhance the production of crops. Nanobiofertilizer gives better and more long-lasting results as compared to traditional chemical fertilizers. It improves the structure and function of soil and the morphological, physiological, biochemical, and yield attributes of plants. The formation and application of nanobiofertilizer is a practical step toward smart fertilizer that enhances growth and augments the yield of crops. The literature on the formulation and application of nanobiofertilizer at the field level is scarce. This product requires attention, as it can reduce the use of chemical fertilizer and make the soil and crops healthy. This review highlights the formulation and application of nanobiofertilizer on various plant species and explains how nanobiofertilizer improves the growth and development of plants. It covers the role and status of nanobiofertilizer in agriculture. The limitations of and future strategies for formulating effective nanobiofertilizer are mentioned.

Planting arrangements affect radiation use efficiency (RUE) and competitiveness of intercrop species in intercropping systems. Here, we reveal that narrow‐wide‐row planting arrangement in maize‐soybean relay‐intercropping system increases the dry matter and competitiveness of soybean, increased the RUE of maize and soybean, and compensates the yield loss of maize by substantially increasing the yield of soybean. In this field study, maize was planted with soybean in different planting arrangements (P1, 20:180, P2, 40:160; P3, 60:140, and P4, 80:120) of relay intercropping, all the relay‐intercropping treatments were compared with sole crops of maize (SM) and soybean (SS). Results showed that P1 improved the total RUE 3.26 g/MJ (maize RUE + soybean RUE) of maize and soybean in relay‐intercropping system. Compared to P4, treatment P1 increased the soybean competition ratio (CR) values (by 55%) but reduced the maize CR values (by 29%), which in turn significantly improved the yield of soybean by maintaining the maize yield. Generally, in P1, soybean produced 82% of SS yield, and maize produced 88% of SM yield, and it achieved the land equivalent ratio of 1.7. These results suggest that by maintaining the appropriate planting distances between maize and soybean we can improve the competitiveness and yield of intercrop species in relay‐intercropping system.

Drought stress causes heavy damages to crop growth and productivity under global climatic changes. Transcription factors have been extensively studied in many crops to play important roles in plant growth and defense. However, there is a scarcity of studies regarding WRKY transcription factors regulating drought responses in maize crops. Previously, ZmWRKY79 was identified as the regulator of maize phytoalexin biosynthesis with inducible expression under different elicitation. Here, we elucidated the function of ZmWRKY79 in drought stress through regulating ABA biosynthesis. The overexpression of ZmWRKY79 in Arabidopsis improved the survival rate under drought stress, which was accompanied by more lateral roots, lower stomatal aperture, and water loss. ROS scavenging was also boosted by ZmWRKY79 to result in less H2O2 and MDA accumulation and increased antioxidant enzyme activities. Further analysis detected more ABA production in ZmWRKY79 overexpression lines under drought stress, which was consistent with up-regulated ABA biosynthetic gene expression by RNA-seq analysis. ZmWRKY79 was observed to target ZmAAO3 genes in maize protoplast through acting on the specific W-boxes of the corresponding gene promoters. Virus-induced gene silencing of ZmWRKY79 in maize resulted in compromised drought tolerance with more H2O2 accumulation and weaker root system architecture. Together, this study substantiates the role of ZmWRKY79 in the drought-tolerance mechanism through regulating ABA biosynthesis, suggesting its broad functions not only as the regulator in phytoalexin biosynthesis against pathogen infection but also playing the positive role in abiotic stress response, which provides a WRKY candidate gene to improve drought tolerance for maize and other crop plants.

Without developing new agronomic practices, present rates of improvement in seed yields of cereal crops globally are insufficient to fulfil the estimated increasing food demand for 2050 and beyond. Intercropping is one of the agricultural practices that can lead to greater crop yields. However, there exists leaf redundancy for maize in intercropping systems, and the top canopy leaves shade more competent leaves at middle strata of maize plants. Therefore, this work aimed to elucidate the effect of leaf excision treatments in maize to understand the optimum leaf area of maize plants under a maize-soybean relay-intercropping system (MS R ) and a sole cropping system (SM). The effects of four-leaf excision treatments (T 1 , 0; T 2 , 2; T 3 , 4; T 4 , 6 leaves excised from the top of maize plants until 7 days after silking) on light interception, leaf area index (LAI), photosynthetic characteristics, total biomass accumulation at blistering stage (BS), dough stage (DS) and physiological maturity (PM), and seed yield of maize were investigated through field experiments for 2 years under MS R and SM. Results showed that, under MS R and SM, as compared to control (T 1 ), optimum excision of leaves (T 2 ) from the top of maize plants significantly improved the light interception (by 25, 18 and 16% at BS, DS and PM, respectively) to lower strata leaves and accelerated the biomass partitioning to maize seeds (by 13 and 12% at DS and PM, respectively). Importantly, plants under T 2 exhibited higher green leaf area than control, that is, excision the top two leaves led to an increase in LAI at PM by 10%, suggesting that leaf senescence under T 2 was delayed which enhanced the photosynthetic rate at PM by 7% in 2017 and 6% in 2018. Relative to T 1 , maize under T 2 produced 19 and 13% higher maize yield under MS R and SM, respectively, and relaycropped maize had 90% of SM seed yield. These results suggest that by manipulating the canopy structure of maize plants we can enhance the biomass accumulation and seed yield of maize crops under MS R and SM. K E Y W O R D Sbiomass, leaf excision, light interception, relay-intercropping, seed yield Muhammad A. Raza and Ling Y. Feng are co-first authors.