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Receptor-like kinases (RLKs) can participate in multiple signalling pathways and are considered one of the most critical components of the early events of intercellular signalling. As an RLK, the ERECTA family (ERf), which comprises ERECTA (ER), ERECTA-Like1 (ERL1), and ERECTA-Like2 (ERL2) in Arabidopsis, regulates multiple signalling pathways in plant growth and development. Despite its indispensability, detailed information on ERf-manipulated signalling pathways remains elusive. In this review, we attempt to summarize the essential roles of the ERf in plant organ morphogenesis, including shoot apical meristem, stem, and reproductive organ development.

Intensively farmed crops used to experience numerous environmental stresses. Among these, shade and drought significantly influence the morpho-physiological and biochemical attributes of plants. However, the interactive effect of shade and drought on the growth and development of soybean under dense cropping systems has not been reported yet. This study investigated the interactive effect of PEG-induced osmotic stress and shade on soybean seedlings. The soybean cultivar viz., C-103 was subjected to PEG-induced osmotic stress from polyethylene glycol 6000 (PEG-6000) under shading and non-shading conditions. PEG-induced osmotic stress significantly reduced the relative water contents, morphological parameters, carbohydrates and chlorophyll contents under both light environments. A significant increase was observed in osmoprotectants, reactive oxygen species and antioxidant enzymes in soybean seedlings. Henceforth, the findings revealed that, seedlings grown under non-shading conditions produced more malondialdehyde and hydrogen peroxide contents as compared to the shade-treated plants when subjected to PEG-induced osmotic stress. Likewise, the shaded plants accumulated more sugars and proline than non-shaded ones under drought stress. Moreover, it was found that non-shaded grown plants were more sensitive to PEG-induced osmotic stress than those exposed to shading conditions, which suggested that shade could boost the protective mechanisms against osmotic stress or at least would not exaggerate the adverse effects of PEG-induced osmotic stress in soybean seedlings.

Plants and crops are widely suffered by shade stress in the natural communities or in the agricultural fields. The three main phytohormones auxin, gibberellins (GAs) and brassinosteroids (BRs) were found essential in shade avoidance in Arabidopsis. However, their relationship have been seldom reported in plant shade avoidance control. Here, we report our investigation of the crosstalk of auxin, GAs and BRs in shade-induced hypocotyl elongation of soybean. Exogenous feeding of indol-3-acetic acid (IAA), GA3 or 24-epibrassinolide (EBL) distinctly promoted hypocotyl elongation in the white light, while the potent biosynthesis inhibitors of GA3, IAA, BRs severely diminished shade-induced hypocotyl elongation. Synergistic treatment of their biosynthesis inhibitors showed that GA3 fully, while EBL slightly, restored the hypocotyl elongation that was efficiently repressed by IAA biosynthesis inhibitor, GA3 and IAA dramatically suppressed the hypocotyl growth inhibition by BR biosynthesis inhibitor in the shade, whereas both IAA and EBL feeding cannot suppress the elongation inhibition by GA biosynthesis inhibitor. Further analyses revealed that shade remarkably upregulated expression of key genes of IAA, GA and BR biosynthesis in the soybean hypocotyls, and GA biosynthesis genes were effectively blocked by IAA, GA and BR biosynthesis inhibitors in the shade. Taken together, these results suggest that GAs modulate shade-induced hypocotyl elongation downstream of mutual promotion of auxin and BRs in soybean.

Soybean-based intercropping is important for sustainable agricultural practice on a regional and global scale. However, most soybean varieties use shade avoidance strategy to acquire more light absorption when suffered in canopy shade in intercropping systems, thus reduced the yield of the whole population on a farmland. The mechanisms underlying early response of soybean in shade avoidance is still largely unknown. Here we report our identification of differentially accumulated proteins in shade-sensitive soybean seedlings by global quantitative proteome analysis under white light (WL) and shade conditions. By using Tandem Mass Tag (TMT) labelling and HPLC fractionation followed by high-resolution LC-MS/MS analysis, 29 proteins were found up-regulated and 412 proteins were found down-regulated in soybean seedlings by 2-h shade stress than that by 2-h WL treatment. Multiple differentially expressed proteins are enriched in carbohydrate metabolic process especially in the biosynthetic pathways of cell wall polysaccharides in soybean seedlings by shade stress comparing to those in WL growth conditions. Physiological assays showed that saccharides were rapidly declined in shoot apex of soybean seedlings under a short-term shading. Our results would provide new insights into the mechanisms of shade avoidance responses in soybean seedlings.

The receptor-like kinase ERECTA (ER) plays vital roles in plant adaptation under environmental stress including shade avoidance in Arabidopsis. In our previous study, we identified four ER paralogues in soybean (GmERs) that showed high similarities to Arabidopsis ER. Each GmER was predicted to generate diverse alternative splicing variants. However, whether soybean GmERs contribute to shade avoidance is unknown. Here we report our characterization of GmERs in response to shading. Promoter::GUS staining analysis shows that expression of GmER paralogous genes was differentially induced under shade stress. Further analyses show that GmERa.1 and GmERa.2 exhibit a larger distinction in length than the other GmER variants. GmERa.2 has the shortest length of amino acid with only 15 leucine-rich repeats which is the part of the extracellular domain of GmERa.1. Overexpression of GmERa.2 fully rescued the hypocotyl length, leaf area and petiole length, and the sensitivity of the hypocotyl of Arabidopsis mutant er-3 to shading, suggesting that the truncated extracellular domain of GmERa might contribute importantly to shade avoidance.

Shade and drought are considered the most drastic abiotic stress factors which restrict normal crop growth especially for those high-density and intercropped crops in arid and semi-arid areas. However, relationship and mechanisms of the interactive effects of these two stress factors on crops are elusive. Here, we report our identification of relationship between shade and drought and their interactive effects on soybean plants. Our quantitative-real time PCR (q-PCR) results showed that shade significantly up-regulated Abscisic Acid (ABA) biosynthesis (NCED3 and AAO3) and signaling (ABI4 and ABI5) genes. Moreover, shade massively enhanced the ABA contents. Based on these results, we subjected soybean seedlings to pretreatment of shade (Pre-SH) and normal light (Pre-NL) followed by polyethylene glycol 6000 induced drought stress. Drought resulted in the reduction of relative water content, morphophysiological and biochemical parameters under both light regimes. However, plants grown under Pre-SH treatment protected their cells from damaging effects of drought by scavenging more reactive oxygen species (ROS) through enhancement of its defense system as compared to Pre-NL plants. Hence, our findings demonstrated that plants grown under Pre-NL conditions were more affected with drought stress than those of Pre-SH ones which suggested that pretreatment of shade can enhance the drought resistance of soybean seedlings.