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Rice Science ›› 2024, Vol. 31 ›› Issue (6): 659-672.DOI: 10.1016/j.rsci.2024.08.002

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  • 收稿日期:2024-05-13 接受日期:2024-08-21 出版日期:2024-11-28 发布日期:2024-12-10

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. [J]. Rice Science, 2024, 31(6): 659-672.

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链接本文: http://www.ricesci.org/CN/10.1016/j.rsci.2024.08.002

               http://www.ricesci.org/CN/Y2024/V31/I6/659

图/表 5

Table 1. Phenotyping for biochemical estimation.
Biochemical Staining Wet estimation technique Reference
Non-reducing sugars Iodine staining Anthrone test-glucose de Bruyn et al, 1968
Pectin Ruthenium red staining; 0.02% unesterified pectin; de-esterified with 0.1 mol/L Na2CO3 overnight at 4o followed by staining Carbazole method (galacturonic is standard); FTIR; HPLC Hou et al, 1999; Kyriakidis and Psoma, 2001
Suberin Sudan red staining Vapor osmometry; FTIR Lopes et al, 2000; Rao, 2011
Lignin Weisner staining (Phloroglucinol staining); Toluidine staining Karlson lignin; thioglycolic acid; acetyl bromide; DFRC, FTIR, HPLC, AIL, and ASL Sjöberg et al, 2004; Liljegren, 2010; Pradhan Mitra and Loqué, 2014; Dampanaboina et al, 2021a
Silicon Digestion with octyl-alcohol, H2O2, and NaOH, and determination by ICP-OES; FTIR Kraska and Breitenbeck, 2010
Potassium Digestion with H2O2 and H2SO4 followed by measured through plasma spectrometer Yue et al, 2023
Cellulose Calcofluor staining by the use of fluorescent microscope Updergraff’s method; modified Updergraff’s method; FTIR Bauer and Ibáñez, 2014; Wu et al, 2017; Dampanaboina et al, 2021b

Table 1. Phenotyping for biochemical estimation.

Biochemical Staining Wet estimation technique Reference
Non-reducing sugars Iodine staining Anthrone test-glucose de Bruyn et al, 1968
Pectin Ruthenium red staining; 0.02% unesterified pectin; de-esterified with 0.1 mol/L Na2CO3 overnight at 4o followed by staining Carbazole method (galacturonic is standard); FTIR; HPLC Hou et al, 1999; Kyriakidis and Psoma, 2001
Suberin Sudan red staining Vapor osmometry; FTIR Lopes et al, 2000; Rao, 2011
Lignin Weisner staining (Phloroglucinol staining); Toluidine staining Karlson lignin; thioglycolic acid; acetyl bromide; DFRC, FTIR, HPLC, AIL, and ASL Sjöberg et al, 2004; Liljegren, 2010; Pradhan Mitra and Loqué, 2014; Dampanaboina et al, 2021a
Silicon Digestion with octyl-alcohol, H2O2, and NaOH, and determination by ICP-OES; FTIR Kraska and Breitenbeck, 2010
Potassium Digestion with H2O2 and H2SO4 followed by measured through plasma spectrometer Yue et al, 2023
Cellulose Calcofluor staining by the use of fluorescent microscope Updergraff’s method; modified Updergraff’s method; FTIR Bauer and Ibáñez, 2014; Wu et al, 2017; Dampanaboina et al, 2021b
Fig. 1. Mechanics of lodging resistance in rice. A, Section modulus (a1 and b1, the major and minor outer diameters; a2 and b2, the major and minor inner diameters). B, Universal Testing Machine (UTM), used to measure bending stress or load and deflection. C, Plant lodging tester, not as bulky as the UTM used to measure bending stress or load and deflection and elastic modulus. D and E, Deflection of a weak culm is more (D), but a stiff culm is less (E). F and G, Bending moment at breaking (BMB) with leaf sheath (MWLS, F) and without leaf sheath (MWOLS, G) used to measure degree of leaf sheath reinforcement (DLSR) to evaluate the contribution of leaf sheath for lodging resistance. H, Understanding bending-type lodging, less plant height leads to reduced lodging index. I, Prostrate tester utilized to measure bending resistance of the canopy (Pushing resistance reading/tiller number per plant) and culm strength. Δσ, Change in stress; Δε, Change in strain; YM, Young modulus; FL, Load × fulcrum distance; FR, Flexural rigidity; SM, Section modulus; τ, Torque; F, Force; r, Distance from the pivot point; θ, The angle between the force and the lever arm; LI, Lodging index; PH, Plant height; FW, Fresh weight of culm; BS, Bending stress; CS, Culm strength; PR, Pushing resistance; PRL, Pushing resistance of lower internode; TN, Tiller number. The rice image is generated using Meta AI (Llama 3.1).

Fig. 1. Mechanics of lodging resistance in rice. A, Section modulus (a1 and b1, the major and minor outer diameters; a2 and b2, the major and minor inner diameters). B, Universal Testing Machine (UTM), used to measure bending stress or load and deflection. C, Plant lodging tester, not as bulky as the UTM used to measure bending stress or load and deflection and elastic modulus. D and E, Deflection of a weak culm is more (D), but a stiff culm is less (E). F and G, Bending moment at breaking (BMB) with leaf sheath (MWLS, F) and without leaf sheath (MWOLS, G) used to measure degree of leaf sheath reinforcement (DLSR) to evaluate the contribution of leaf sheath for lodging resistance. H, Understanding bending-type lodging, less plant height leads to reduced lodging index. I, Prostrate tester utilized to measure bending resistance of the canopy (Pushing resistance reading/tiller number per plant) and culm strength. Δσ, Change in stress; Δε, Change in strain; YM, Young modulus; FL, Load × fulcrum distance; FR, Flexural rigidity; SM, Section modulus; τ, Torque; F, Force; r, Distance from the pivot point; θ, The angle between the force and the lever arm; LI, Lodging index; PH, Plant height; FW, Fresh weight of culm; BS, Bending stress; CS, Culm strength; PR, Pushing resistance; PRL, Pushing resistance of lower internode; TN, Tiller number. The rice image is generated using Meta AI (Llama 3.1).

Fig. 2. Comparisons for understanding lodging resistance. A, High section modulus, i.e., less area in the medullary region/lacuna, generally leads to lodging resistance, while low section modulus, i.e., more area in the medullary region, leads to lodging susceptibility. B, Dark pink phloroglucinol stain suggests the presence of more lignin, while light stain signifies the presence of less lignin. C, Powdered X-ray diffraction (XRD) is used to determine cellulose crystallinity. A sharp peak at 22.5º indicates highly crystalline cellulose. Peaks at 22.5º and 18.0º on the X axis are checked for a rough estimation of cellulose crystallinity (sengal method). D, Genotype with high bending stress is observed withstand a load of 18.6 N before breaking, and the genotype with comparatively lower bending stress is observed withstand a load of 6.2 N load before breaking (y-axis, Load; x-axis, Time in seconds).

Fig. 2. Comparisons for understanding lodging resistance. A, High section modulus, i.e., less area in the medullary region/lacuna, generally leads to lodging resistance, while low section modulus, i.e., more area in the medullary region, leads to lodging susceptibility. B, Dark pink phloroglucinol stain suggests the presence of more lignin, while light stain signifies the presence of less lignin. C, Powdered X-ray diffraction (XRD) is used to determine cellulose crystallinity. A sharp peak at 22.5º indicates highly crystalline cellulose. Peaks at 22.5º and 18.0º on the X axis are checked for a rough estimation of cellulose crystallinity (sengal method). D, Genotype with high bending stress is observed withstand a load of 18.6 N before breaking, and the genotype with comparatively lower bending stress is observed withstand a load of 6.2 N load before breaking (y-axis, Load; x-axis, Time in seconds).

Table 2. Genes associated with lodging resistance.
Gene Chr. Marker Mapping population Remark Reference
qCD1.1, qCS1.1 1 Id10019973-Id1006772 Swarna × Moroberekan♂ (BIL) Lead to an increase in culm diameter and culm strength Yadav et al, 2017
qLR1, qLR8 1 BIN1-161-BIN1-162
BIN8-32-BIN8-33
93-11 × O. longistaminata♂ (BIL) An increase in CD and breaking strength pleiotropic effect qLR1 and qLR8 is seen for lodging resistance Long et al, 2020
OsCYPq1 1 RM12285-RM212 Cheongcheong♂ × Nagdong (DH) The candidate gene identified controls basal internode length and mutations in it led to reduced internode elongation and also showed pleiotropic effect by increasing yield Zhao et al, 2021
Gnla 1 N321-RM10316 R498♂ × R3551
(RIL and CSSL)
Increase in CD and crown root development hence can also deal with root lodging Tu et al, 2022
SCM1 1 RM562-RM5 Habataki♂ × Sasanishiki
(CSSL)
Increases culm wall thickness Yang et al, 2023
qSCM4 2 SSR1-RM5511 LTH♂ × Shennong265
(F2 and RIL)
Increase in CWT, CD, and culm folding resistance Yang et al, 2023
SCM3 3 RM15761-RM15782
SNPV318_03-RM15767
Koshihikari × Chugoku 117♂ (BIL) Improves CD, SM, and BMB Yano et al, 2015
prl4 4 RM5749 Nona Bokra♂ × Koshihikari (CSSL) Pushing resistance, and accumulates NSC in basal culm Kashiwagi, 2022
prl5 5 C1081 Nipponbare × Kasalath♂
(NIL)
Delays leaf senescence, PR, NSC content, starch reaccumulation, and high YM Kashiwagi et al, 2006
SCM2 6 RM20546-RM20562 Sasanishiki × Habataki♂ (CSSL and NIL) Increases CWT and affects yield because it increases the number of spikelets per panicle Ookawa et al, 2010a
qTyM6, qTyH6 6 RM3343-20318 Cheongcheong♂ × Nagdong (DH) Resistance reported in Typhoons, as it reduces plant height and lowers plant thrust resistance Zhao et al, 2023
sdm8 8 C10122 Kasalath♂ × Nipponbare (CSSL) Increases in CD, CWT, culm stiffness, and heavy lower part of the plant Kashiwagi et al, 2008
BSUC11 11 G44 and G257 Kasalath♂ × Koshihikari (CSSL) Increases content of holocellulose in upper culm; thickening of cortical fiber tissue in internode 1 Kashiwagi et al, 2016

Table 2. Genes associated with lodging resistance.

Gene Chr. Marker Mapping population Remark Reference
qCD1.1, qCS1.1 1 Id10019973-Id1006772 Swarna × Moroberekan♂ (BIL) Lead to an increase in culm diameter and culm strength Yadav et al, 2017
qLR1, qLR8 1 BIN1-161-BIN1-162
BIN8-32-BIN8-33
93-11 × O. longistaminata♂ (BIL) An increase in CD and breaking strength pleiotropic effect qLR1 and qLR8 is seen for lodging resistance Long et al, 2020
OsCYPq1 1 RM12285-RM212 Cheongcheong♂ × Nagdong (DH) The candidate gene identified controls basal internode length and mutations in it led to reduced internode elongation and also showed pleiotropic effect by increasing yield Zhao et al, 2021
Gnla 1 N321-RM10316 R498♂ × R3551
(RIL and CSSL)
Increase in CD and crown root development hence can also deal with root lodging Tu et al, 2022
SCM1 1 RM562-RM5 Habataki♂ × Sasanishiki
(CSSL)
Increases culm wall thickness Yang et al, 2023
qSCM4 2 SSR1-RM5511 LTH♂ × Shennong265
(F2 and RIL)
Increase in CWT, CD, and culm folding resistance Yang et al, 2023
SCM3 3 RM15761-RM15782
SNPV318_03-RM15767
Koshihikari × Chugoku 117♂ (BIL) Improves CD, SM, and BMB Yano et al, 2015
prl4 4 RM5749 Nona Bokra♂ × Koshihikari (CSSL) Pushing resistance, and accumulates NSC in basal culm Kashiwagi, 2022
prl5 5 C1081 Nipponbare × Kasalath♂
(NIL)
Delays leaf senescence, PR, NSC content, starch reaccumulation, and high YM Kashiwagi et al, 2006
SCM2 6 RM20546-RM20562 Sasanishiki × Habataki♂ (CSSL and NIL) Increases CWT and affects yield because it increases the number of spikelets per panicle Ookawa et al, 2010a
qTyM6, qTyH6 6 RM3343-20318 Cheongcheong♂ × Nagdong (DH) Resistance reported in Typhoons, as it reduces plant height and lowers plant thrust resistance Zhao et al, 2023
sdm8 8 C10122 Kasalath♂ × Nipponbare (CSSL) Increases in CD, CWT, culm stiffness, and heavy lower part of the plant Kashiwagi et al, 2008
BSUC11 11 G44 and G257 Kasalath♂ × Koshihikari (CSSL) Increases content of holocellulose in upper culm; thickening of cortical fiber tissue in internode 1 Kashiwagi et al, 2016
Fig. 3. Pleiotropic action of SCM3, SCM2 and depiction of other genes involved in lodging resistance. Gn1a is expressed in culm, roots, inflorescence meristem, and leaves (grey dots). The null allele gn1a in the roots leads to well-developed crown roots, and this allele leads to an increase in grain number. The Gn1a allele SCM1 increases culm diameter and strength. The allele of APO1 (SCM2) increases spikelet number and culm diameter (blue dots). SCM3 decreases tiller number but increases spikelet number (pink dots). The prl4 allele leads to an increase in the bending resistance of the lower internode. GA-inactive (#), GA-active (*), and GA-inactive GA (##) [conversions from active gibberellic acid (GA) to inactive GA alleles SD1 (GA20 oxidase) and SBI (GA2 oxidase)] lead to a reduction in GA levels, leading to dwarfism, prl5 is a gene associated with late senescence, and the orange boxes represent genes involved in lignin and cellulose biosynthesis.The rice image is generated using Meta AI (Llama 3.1).

Fig. 3. Pleiotropic action of SCM3, SCM2 and depiction of other genes involved in lodging resistance. Gn1a is expressed in culm, roots, inflorescence meristem, and leaves (grey dots). The null allele gn1a in the roots leads to well-developed crown roots, and this allele leads to an increase in grain number. The Gn1a allele SCM1 increases culm diameter and strength. The allele of APO1 (SCM2) increases spikelet number and culm diameter (blue dots). SCM3 decreases tiller number but increases spikelet number (pink dots). The prl4 allele leads to an increase in the bending resistance of the lower internode. GA-inactive (#), GA-active (*), and GA-inactive GA (##) [conversions from active gibberellic acid (GA) to inactive GA alleles SD1 (GA20 oxidase) and SBI (GA2 oxidase)] lead to a reduction in GA levels, leading to dwarfism, prl5 is a gene associated with late senescence, and the orange boxes represent genes involved in lignin and cellulose biosynthesis.The rice image is generated using Meta AI (Llama 3.1).

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