# Regulatory Mechanisms Underlying Stem Strength and Toughness in Dicotyledonous Plants: Implications for Soybean Breeding

**Authors:** Ye Zhang, Elshan Musazade, Javaid Akhter Bhat, Songling Xie, Yaohua Zhang, Weitao Xu, Xianzhong Feng, Suxin Yang

PMC · DOI: 10.3390/cimb48020189 · Current Issues in Molecular Biology · 2026-02-07

## TL;DR

This review explores how stem strength and toughness in dicot plants, especially soybean, are regulated and how this knowledge can improve breeding for lodging resistance.

## Contribution

The paper integrates anatomical, structural, and genetic insights from model species to provide a translational framework for soybean improvement.

## Key findings

- Stem mechanical performance is influenced by vascular tissue development and secondary cell wall composition.
- Conserved regulatory pathways from model species can be applied to enhance soybean lodging resistance.
- Anatomical and genetic mechanisms are key to improving stem strength and yield stability in soybean.

## Abstract

Soybean (Glycine max) is a globally important crop valued for its high seed oil and protein content. However, lodging remains a major abiotic constraint that causes substantial yield losses. Lodging resistance is primarily determined by stem strength and toughness, which are governed by stem anatomical organization, vascular tissue development, and the composition and architecture of secondary cell walls (SCWs). This review synthesizes current knowledge on anatomical, structural, and genetic factors that are implicated in stem mechanical performance in dicotyledonous plants, with particular emphasis on vascular cambium activity, xylem and phloem differentiation, and the biosynthesis of major SCW components, including cellulose, hemicellulose, and lignin. These processes collectively determine stem rigidity, flexibility, and resistance to mechanical stress. By integrating insights from model species, especially Arabidopsis thaliana, and non-soybean dicots, this review highlights conserved regulatory pathways controlling stem development and SCW formation that are directly relevant to soybean improvement. The synthesis provides a translational framework for understanding how conserved anatomical and genetic mechanisms can be leveraged to enhance soybean stem strength, toughness, and lodging resistance. Overall, this review provides a conceptual foundation for future functional studies and breeding strategies to improve soybean yield stability and adaptability across diverse agronomic conditions.

## Linked entities

- **Species:** Glycine max (taxon 3847), Arabidopsis thaliana (taxon 3702)

## Full-text entities

- **Genes:** LOG3 (lysine decarboxylase family protein) [NCBI Gene 818298] {aka LONELY GUY 3, T2N18.3, T2N18_3}, MYB3 (myb domain protein 3) [NCBI Gene 838870] {aka ARABIDOPSIS THALIANA  MYB DOMAIN PROTEIN 3, ARABIDOPSIS THALIANA MYB DOMAIN PROTEIN 3, ATMYB3, T22J18.19, T22J18_19, TRANSCRIPTION FACTOR MYB3}, BXL2 (beta-xylosidase 2) [NCBI Gene 837940] {aka ATBXL2, BETA-XYLOSIDASE 2, T14P4.8, T14P4_8, beta-xylosidase 2}, NAC15 (NAC domain-containing protein) [NCBI Gene 100170744] {aka GmNAC065}, NAC012 (NAC domain containing protein 12) [NCBI Gene 840171] {aka ANAC012, F6N18.15, F6N18_15, NAC SECONDARY WALL THICKENING PROMOTING 3, NAC domain containing protein 12, NST3}, MYB54 (MYB transcription factor MYB54) [NCBI Gene 778030], IRX14 (Nucleotide-diphospho-sugar transferases superfamily protein) [NCBI Gene 829842] {aka AP22.51, AP22_51, irregular xylem 14}, CESA4 (cellulose synthase A4) [NCBI Gene 834426] {aka IRREGULAR XYLEM 5, IRX5, MRH10.14, MRH10_14, NWS2, cellulose synthase A4}, KNAT1 (homeobox knotted-like protein) [NCBI Gene 826364] {aka BP, BP1, BREVIPEDICELLUS, BREVIPEDICELLUS 1, F9M13.2, F9M13_2}, IRX1 (cellulose synthase family protein) [NCBI Gene 827612] {aka ATCESA8, CELLULOSE SYNTHASE 8, CESA8, F28A21.190, F28A21_190, IRREGULAR XYLEM 1}, VND7 (vascular related NAC-domain protein 7) [NCBI Gene 843524] {aka ANAC030, Arabidopsis NAC domain containing protein 30, F17M19.8, F17M19_8, vascular related NAC-domain protein 7}, MYB83 (myb domain protein 83) [NCBI Gene 819997] {aka AtMYB83, myb domain protein 83}, PXY (Leucine-rich repeat protein kinase family protein) [NCBI Gene 836269] {aka MCI2.4, MCI2_4, PHLOEM INTERCALATED WITH XYLEM, TDIF receptor, TDR}, CLV3 (CLAVATA3) [NCBI Gene 817267] {aka AtCLV3, CLAVATA3, F12K2.17, F12K2_17}, IRX9 (Nucleotide-diphospho-sugar transferases superfamily protein) [NCBI Gene 818285] {aka IRREGULAR XYLEM 9, T2N18.15, T2N18_15}, GUT2 (Exostosin family protein) [NCBI Gene 839635] {aka ATGUT1, F17L21.23, F17L21_23, IRX10}, GH3 [NCBI Gene 100811309], MYB [NCBI Gene 547568], VRN2 [NCBI Gene 100816782], IRX3 (Cellulose synthase family protein) [NCBI Gene 831608] {aka ATCESA7, CELLULOSE SYNTHASE  CATALYTIC SUBUNIT 7, CELLULOSE SYNTHASE CATALYTIC SUBUNIT, CELLULOSE SYNTHASE CATALYTIC SUBUNIT 7, CESA7, IRREGULAR XYLEM 3}, WOX4 (WUSCHEL related homeobox 4) [NCBI Gene 841113] {aka F2G19.11, F2G19_11, WUSCHEL related homeobox 4}, LOG4 (Putative lysine decarboxylase family protein) [NCBI Gene 824513] {aka LONELY GUY 4}, NAC101 (NAC-domain protein 101) [NCBI Gene 836359] {aka ANAC101, MMI9.6, MMI9_6, NAC-domain protein 101, VASCULAR-RELATED NAC-DOMAIN 6, VND6}, WOX14 (WUSCHEL related homeobox 14) [NCBI Gene 838660] {aka ATWOX14, F2D10.19, F2D10_19, WUSCHEL RELATED HOMEOBOX 14, WUSCHEL related homeobox 14}, ubiquitin [NCBI Gene 547906], HP6 (histidine phosphotransfer protein 6) [NCBI Gene 844350] {aka AHP6, F18B13.18, F18B13_18, histidine phosphotransfer protein 6}, FRA8 (Exostosin family protein) [NCBI Gene 817357] {aka F24D13.10, F24D13_10, FRAGILE FIBER 8, IRREGULAR XYLEM 7, IRX7}, MP (Transcriptional factor B3 family protein / auxin-responsive factor AUX/IAA-like protein) [NCBI Gene 838573] {aka ARF5, AUXIN RESPONSE FACTOR 5, F6F9.10, F6F9_10, IAA24, MONOPTEROS}, MYB46 (myb domain protein 46) [NCBI Gene 831127] {aka ATMYB46, MYB DOMAIN PROTEIN 46, T24H18.40, T24H18_40, myb domain protein 46}, MYB52 (MYB transcription factor MYB52) [NCBI Gene 778032], SS (sucrose synthase) [NCBI Gene 547508] {aka N-100}, CLE41 (CLAVATA3/ESR-RELATED 41) [NCBI Gene 822075] {aka CLAVATA3/ESR-RELATED 41}, CLE44 (CLAVATA3/ESR-RELATED 44) [NCBI Gene 826934] {aka CLAVATA3/ESR-RELATED 44}, MYB20 (myb domain protein 20) [NCBI Gene 842938] {aka AtMYB20, T6J19.5, T6J19_5, myb domain protein 20}
- **Diseases:** dwarfism (MESH:D004392), vascular abnormalities (MESH:D014652), growth retardation (MESH:D006130), injury to (MESH:D014947), dwarf (MESH:D004393), SND (MESH:D056988)
- **Chemicals:** UDP (MESH:D014530), vegetable oils (MESH:D010938), Hemicellulose (MESH:C007916), BRs (MESH:D060406), gibberellin (MESH:D005875), xylan (MESH:D014990), water (MESH:D014867), polymers (MESH:D011108), glucomannan (MESH:C022901), carbon (MESH:D002244), nitrogen (MESH:D009584), polysaccharide (MESH:D011134), thermospermine (MESH:C021097), Auxin (MESH:D007210), JA (MESH:C011006), H (MESH:D006859), IAA (MESH:C030737), Cellulose (MESH:D002482), sphingolipid (MESH:D013107), GA (MESH:D005708), p-coumaryl alcohol (MESH:C495469), sucrose (MESH:D013395), CK (MESH:D003583), Lignin (MESH:D008031), carbohydrate (MESH:D002241), UDP-glucose (MESH:D014532), coniferyl alcohol (MESH:C010559), potassium ion (MESH:D011188), sinapyl alcohol (MESH:C496130), AUX (-)
- **Species:** Glycine max (soybean, species) [taxon 3847], Medicago sativa (alfalfa, species) [taxon 3879], Oryza sativa (Asian cultivated rice, species) [taxon 4530], Sorghum bicolor (broomcorn, species) [taxon 4558], Homo sapiens (human, species) [taxon 9606], Arabidopsis thaliana (mouse-ear cress, species) [taxon 3702]

## Full text

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## Figures

7 figures with captions in the complete paper: https://tomesphere.com/paper/PMC12939125/full.md

## References

117 references — full list in the complete paper: https://tomesphere.com/paper/PMC12939125/full.md

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Source: https://tomesphere.com/paper/PMC12939125