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Acta Agronomica Sinica ›› 2019, Vol. 45 ›› Issue (7): 1002-1016.doi: 10.3724/SP.J.1006.2019.84143


Cloning and expression analysis of sugarcane lipoxygenase gene ScLOX1

SUN Ting-Ting1,WANG Wen-Ju1,LOU Wen-Yue1,LIU Feng1,ZHANG Xu1,WANG Ling1,CHEN Yu-Feng1,QUE You-Xiong1,2,XU Li-Ping1,2,LI Da-Mei1,2,SU Ya-Chun1,2,*()   

  1. 1 Key Laboratory of Sugarcane Biology and Genetic Breeding (Fujian), Ministry of Agriculture / Fujian Agriculture and Forestry University, Fuzhou 350002, Fujian, China
    2 Key Laboratory of Crop Genetics and Breeding and Comprehensive Utilization, Ministry of Education / Fujian Agriculture and Forestry University, Fuzhou 350002, Fujian, China;
  • Received:2018-11-05 Accepted:2019-01-29 Online:2019-07-12 Published:2019-03-22
  • Contact: Ya-Chun SU E-mail:syc2009mail@163.com
  • Supported by:
    This study was supported by the National Natural Science Foundation of China(31501363);the Research Funds for Distinguished Young Scientists in Fujian Provincial Department of Education(SYC-2017);the Research Funds for Distinguished Young Scientists in Fujian Agriculture and Forestry University(xjq201630);the China Agriculture Research System(CARS-17)


LOX, which belongs to the lipoxygenase superfamily, is an important factor for fat oxidation and widely involved in the regulation of plant growth and development and the resistance to external stimuli. In this study, based on sugarcane (Saccharum spp.) transcriptome database, we first cloned a full-length cDNA sequence of ScLOX1 gene (GenBank accession number: MK106188) from ROC22 bud by RT-PCR. Bioinformatics analysis showed that the cDNA length of ScLOX1 gene was 2813 bp which had a 2664 bp length of open reading frame, encoding 887 amino acids. The theoretical isoelectric point, instability coefficient, and average hydrophilicity of the ScLOX1 protein were 6.23, 39.77, and -0.437, respectively. There were no signal peptide and transmembrane structure, but the PLAT_LH2 and lipoxygenase active sites in ScLOX1 protein. The similarity of amino acid sequences between ScLOX1 and Sorghum bicolor LOX (XP_002466613.1) was 95.96%. The protein encoded by ScLOX1 gene was predicted to be an acid-stable, hydrophilic, and non-secreted protein which belongs to the type I non-traditional 9-LOX. qRT-PCR results showed that ScLOX1 was specifically expressed in sugarcane bud tissue. The expression level of ScLOX1 gene was transiently increased in the smut-resistant sugarcane variety Yacheng05-179 but significantly decreased in the smut-susceptible sugarcane variety ROC22 after inoculated with Sporisorium scitamineum. When leaves of Nicotiana benthamiana were transiently overexpressed ScLOX1 gene and inoculated with tobacco pathogens Fusarium solani var. coeruleum and Ralstonia solanacearum, respectively, the results of phenotypic observation, 3,3’-diaminobenzidine (DAB) staining and expression analysis of tobacco immune-related genes revealed that the overexpression of ScLOX1 gene could enhance the defense of N. benthamiana to the F. solani var. coeruleum, but had no significant difference with the control on the defense effect against R. solanacearum. In addition, the expression level of ScLOX1 was down-regulated by methyl jasmine and salicylic acid, but up-regulated by abscisic acid, sodium chloride and polyethylene glycol. The above results provide references for further study on the function of sugarcane ScLOX1 gene.

Key words: sugarcane, lipoxygenase, bioinformatics, real-time flourescent quantitative PCR, biotic and abiotic stresses

Table 1

Sequence and purpose of the primers used in this study"

Primer name
Primer sequence (5'-3')
M13-F TGTAAAACGACGGCCAGT Transient overexpression vector construction
M13-R CAGGAAACAGCTATGACC Transient overexpression vector construction
NtHSR201-F CAGCAGTCCTTTGGCGTTGTC qRT-PCR after transient overexpression
Primer name
Primer sequence (5'-3')
NtHSR201-R GCTCAGTTTAGCCGCAGTTGTG qRT-PCR after transient overexpression
NtHSR203-F TGGCTCAACGATTACGCA qRT-PCR after transient overexpression
NtHSR203-R GCACGAAACCTGGATGG qRT-PCR after transient overexpression
NtHSR515-F TTGGGCAGAATAGATGGGTA qRT-PCR after transient overexpression
NtHSR515-R TTTGGTGAAAGTCTTGGCTC qRT-PCR after transient overexpression
NtPR-1a/c-F AACCTTTGACCTGGGACGAC qRT-PCR after transient overexpression
NtPR-1a/c-R GCACATCCAACACGAACCGA qRT-PCR after transient overexpression
NtPR2-F TGATGCCCTTTTGGATTCTATG qRT-PCR after transient overexpression
NtPR2-R AGTTCCTGCCCCGCTTT qRT-PCR after transient overexpression
NtPR3-F CAGGAGGGTATTGCTTTGTTAGG qRT-PCR after transient overexpression
NtPR3-R CGTGGGAAGATGGCTTGTTGTC qRT-PCR after transient overexpression
NtEFE26-F CGGACGCTGGTGGCATAAT qRT-PCR after transient overexpression
NtEFE26-R CAACAAGAGCTGGTGCTGGATA qRT-PCR after transient overexpression
NtAccdeaminase-F TCTGAGGTTACTGATTTGGATTGG qRT-PCR after transient overexpression
NtAccdeaminase-R TGGACATGGTGGATAGTTGCT qRT-PCR after transient overexpression
NtEF-1α-F TGCTGCTGTAACAAGATGGATGC qRT-PCR after transient overexpression
NtEF-1α-R GAGATGGGGACAAAGGGGATT qRT-PCR after transient overexpression

Fig. 1

Nucleotide acid sequence and deduced amino acid sequence of ScLOX1 gene The sequences underlined are cloning primers; * indicates stop codon."

Fig. 2

Third structure prediction of ScLOX1 protein"

Fig. 3

Conserved domains of amino acid sequences of ScLOX1"

Fig. 4

Phylogenetic tree estimated by the ScLOX1 and those proteins from other plants ScLOX1 is marked with a black triangle. Zea mays LOXs: ZmLOX1 (AF271894), ZmLOX3 (AF329371), ZmLOX6 (DQ335764), ZmLOX10 (DQ335768), ZmLOX11 (DQ335769); Oryza sativa LOXs: OsLOX1 (DQ389164), r9-LOX1 (AB099850); Lycopersicum esculentum LOXs: TomLOXA (U09026), TomLOXB (U09025), TomLOXC (U37839), TomLOXD (U37840), TomLOXE (AY008278), TomLOXF (FJ617476); Arabidopsis thaliana LOXs: AtLOX1 (NM_104376), AtLOX2 (AY062611), AtLOX3 (AJ249794), AtLOX4 (NM_105911); Nicotiana tabacum LOXs: NaLOX1 (X84040), NaLOX2 (AY254348), NaLOX3 (AY254349); Solanum tuberosum LOXs: StLOX1 (X95513), StLOX2 (X96405), StLOX3 (X96406); Glycine max LOXs: GmLOX9 (EU003576), GmLOX10 (EU003577); Arachis hypogaea LOX: PnLOX2 (DQ068249); Pisum sativum LOX: LOXN2 (AJ749702); Prunus persica LOXs: PpLOX1 (EU883638), PpLOX2 (FJ029110), PpLOX3 (FJ032015), PpLOX4 (EF568783); Vitis vinifera LOXs: VvLOXA (FJ858255), VvLOXC (FJ858256), VvLOXO (FJ858257); Olea europaea LOXs: OeLOX (EU678670), Oep1LOX2 (EU513352), Oep2LOX2 (EU513353); Capsicum annuum LOXs: CaLOX1 (FJ377708), CaLOX2 (JQ219046); Populus deltoids LOXs: PdLOX1 (DQ131178), PdLOX2 (DQ131179); Prunus dulcis LOXs: LOX1:Pd:1 (AJ404331), LOX1:Pd:2 (AJ418043); Phaseolus vulgaris LOX: PvLOX6 (EF196866); Brassica oleracea LOX: BoLOX (EF123056); Caragana jubata LOX: CjLOX (EF530043); Camellia sinensis LOX: CasLOX1 (EU195885); Corylus avellana LOX: CaLOX (AJ417975)."

Fig. 5

Amino acids sequence aligenment of ScLOX1 and other plant LOXs The substrate-binding domain (Domains I and II) and C-terminus conserved regions are shown by text boxes. An arrow indicates a conserved amino acid residue related to iron binding. The plus sign indicates TH(V)/R(K), a motif proposed to be essential for determining oxygen-adding positional specificity. OsLOX1: Oryza sativa (DQ389164); ZmLOX1: Zea mays (AF271894); HvLOXC: Hordeum vulgare (L37358); SbLOX4: Sorghum bicolor (XP_ 002466613); SiLOX4: Setaria italica (XP_004982082)."

Fig. 6

Expression pattern of ScLOX1 gene in different sugarcane tissues of ROC22 Using GAPDH as an internal reference gene. Data points are means ± SE (n = 3)."

Fig. 7

Gene expression level of ScLOX1 in the interactions of different sugarcane genotypes and smut pathogen Using GAPDH as an internal reference gene. Bars superscripted by different lowercase letters are significantly different at P < 0.05. Data points are means ± SE (n = 3). Yacheng05-179 is a smut-resistant sugarcane variety. ROC22 is a smut-susceptible sugarcane variety."

Fig. 8

Transient overexpression of ScLOX1 in Nicotiana benthamiana leaves A: RT-PCR analysis of ScLOX1 gene in the N. benthamiana leaves after infiltration by Agrobacterium strain GV3101 carrying vector 35S::00 or 35S::ScLOX1 for 1 d. B: DAB staining with N. benthamiana leaves 2 d after 35S::ScLOX1-containing Agrobacterium strain infiltration. (1) and (2) represent images taken by SONY camera and microscope, respectively. C: Relative expression level of the tobacco immunity-associated marker genes in 35S::ScLOX1-transiently expressing leaves after infiltration for 1 d. D and F: The results of disease symptoms and DAB staining of N. benthamiana leaves after inoculation with 35S::ScLOX1 1 d and then infection with Ralstonia solanacearum and Fusarium solani var. coeruleum for 1 d and 7 d, respectively. (1) and (2) represent images taken by SONY camera and microscope, respectively. E and G: Analysis of the immunity related marker genes in the N. benthamiana leaves by R. solanacearum and F. solani var. coeruleum for 1 d and 7 d post inoculation. The tobacco immunity-associated marker genes including the HR marker genes NtHSR201, NtHSR203 and NtHSR515, the SA pathway related genes NtPR-1a/c, the JA pathway-associated genes NtPR2 and NtPR3, and the ET synthesis depended genes NtEFE26 and NtAccdeaminase. Using NtEF-1α as an internal reference gene. Bars superscripted by different lowercase letters are significantly different at P < 0.05. Data points are means ± SE (n = 3). The vectors of 35S::00 and 35S::ScLOX1 are indicated by a and b, respectively."

Fig. 9

Relative expression level of ScLOX1 gene under the stress of different plant hormones Using GAPDH as an internal reference gene. Bars superscripted by different lowercase letters are significantly different at P < 0.05. Data points are means ± SE (n = 3)."

Fig. 10

Expression of ScLOX1 gene under the stresses of PEG and NaCl Using GAPDH as an internal reference gene. Bars superscripted by different lowercase letters are significantly different at P < 0.05. Data points are means ± SE (n = 3)."

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