A comparative study of the evolution and codon usage bias in WOX gene family of three Camellia sinensis cultivars

doi:10.12302/j.issn.1000-2006.202108049

JOURNAL OF NANJING FORESTRY UNIVERSITY ›› 2022, Vol. 46 ›› Issue (2): 71-80.doi: 10.12302/j.issn.1000-2006.202108049

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A comparative study of the evolution and codon usage bias in WOX gene family of three Camellia sinensis cultivars

WANG Zhanjun¹^,²(), WU Ziqi¹(), WANG Zhaoxia¹, OU Zulan¹, LI Jie¹, CAI Qianwen¹, XU Zhongdong¹, ZHANG Zhaoliang²^,^*()

1. College of Life Science, Hefei Normal University, Hefei 230601, China
2. State Key Laboratory of Camellia sinensis Plant Biology and Utilization, Anhui Agricultural University, Hefei 230036, China

Received:2021-08-29 Accepted:2021-11-29 Online:2022-03-30 Published:2022-04-08
Contact: ZHANG Zhaoliang E-mail:wangzhanjunhxj@163.com;1293014657@qq.com;zhlzhang@ahau.edu.cn

Abstract

Abstract:

【Objective】The WOX gene family plays an important molecular regulatory role in model plant tissue culture. Phylogeny studies and research on the codon usage bias in the WOX gene family in non-model plants are helpful for exploring genetic evolution and gene expression characteristics, which serve as a guideline for establishing a transgenic system.【Method】In this study, bioinformatics were used to identify the WOX genes among all coding sequences of three Camellia sinensis cultivars. The evolutionary patterns were analyzed using ClustalX 2.1 and MEGA X software. The codon usage index of the coding sequence of the WOX gene was analyzed, and the codon usage pattern and the source of codon variation were explored using Perl language, Codon W1.4.2, and SPSS 23.0. The main factors affecting codon usage bias were analyzed based on the results of ENc-GC_3s, a PR2-plot analysis, and a neutral analysis.【Result】A total of 42 WOX members were identified in this study, of which 11 were CSA, 18 were CSS and 13 were DASZ. The phylogenetic analysis revealed that the evolution of WOX genes of the three Camellia sinensis cultivars is highly conserved. The codon bias analysis revealed that the codons of WOX genes preferred to use A/T and end with A/T in the three Camellia sinensis cultivars; the codon usage patterns of WOX genes in CSA and DASZ are similar, indicating that they are closely related. When CSA and CSS are used as transgenic receptors, individual codons in WOX genes of three Camellia sinensis cultivars need to be optimized. Nicotiana tabacum is the best heterologous expression receptor for WOX genes of three Camellia sinensis cultivars. ENc-GC_3s mapping the PR2 analysis, and the neutrality analysis indicate that the natural selection is the main reason for the codon usage bias of WOX genes in the three Camellia sinensis cultivars.【Conclusion】This study showed that WOX genes of three Camellia sinensis cultivars are evolutionarily conserved, codons prefer to use A/T and end with A/T, and the codon usage bias is mainly due to the natural selection. The results reveal the optimization information of codons when transforming tea plants, and tobacco is the best heterologous expression receptor.

Key words: Camellia sinensis, WOX gene family, evolutionary analysis, codon usage bias, transgene expression

CLC Number:

Q786
S794.4

WANG Zhanjun, WU Ziqi, WANG Zhaoxia, OU Zulan, LI Jie, CAI Qianwen, XU Zhongdong, ZHANG Zhaoliang. A comparative study of the evolution and codon usage bias in WOX gene family of three Camellia sinensis cultivars[J]. JOURNAL OF NANJING FORESTRY UNIVERSITY, 2022, 46(2): 71-80.

Figures/Tables 6

Table 1

The information of WOX gene families in three Camellia sinensis cultivars"

基因名称 gene name	基因ID gene ID	氨基酸数量 amino acid quantity	等电点 isoelectric point	分子质量/ku molecular weight	不稳定系数 instability coefficient	脂溶性指数 aliphatic index	亲水性 hydrophilicity	亚细胞定位 subcellular localization
CsaWUS	CSA016122.1	292	5.91	32 238.09	78.16	43.42	-1.001	细胞核
CsaWOX1a	CSA009146.1	326	8.45	37 982.71	64.95	60.18	-0.913	细胞核
CsaWOX1b	CSA021658.1	125	9.26	14 771.54	72.38	66.96	-1.003	细胞核
CsaWOX1c	CSA017712.1	197	9.94	22 734.74	64.44	60.00	-1.076	细胞核
CsaWOX3	CSA034617.1	195	8.77	22 724.70	63.00	52.00	-0.835	细胞核
CsaWOX4	CSA021808.1	231	9.30	26 338.67	58.94	61.21	-0.905	细胞核
CsaWOX5	CSA035970.1	289	9.01	32 485.00	55.84	77.27	-0.522	细胞核
CsaWOX8a	CSA024131.1	285	5.54	32 067.08	56.18	68.07	-0.698	细胞核
CsaWOX8b	CSA032010.1	296	6.06	33 503.17	58.29	53.34	-0.963	细胞核
CsaWOX9	CSA016734.1	287	9.10	31 373.27	52.17	55.71	-0.574	细胞核
CsaWOX11	CSA027532.1	275	5.63	29 686.28	67.31	72.62	-0.286	细胞核
CssWUS	TEA008786.1	297	6.00	32 659.55	79.08	44.34	-0.986	细胞核
CssWOX1a	TEA025042.1	271	7.71	31 693.58	66.28	61.22	-0.913	细胞核
CssWOX1b	TEA006406.1	361	6.47	40 838.38	57.69	60.30	-0.804	细胞核
CssWOX2a	TEA014651.1	220	5.41	23 858.01	51.73	57.68	-0.771	细胞核
CssWOX2b	TEA032867.1	256	6.22	27 928.90	59.25	59.06	-0.702	细胞核
CssWOX3a	TEA018015.1	172	9.27	19 461.41	69.67	63.49	-0.553	细胞核
CssWOX3b	TEA012714.1	195	8.77	22 724.70	63.00	52.00	-0.835	细胞核
CssWOX4a	TEA029435.1	214	8.96	24 865.11	48.74	59.25	-1.007	细胞核
CssWOX4b	TEA022717.1	231	9.30	26 334.68	58.61	61.21	-0.910	细胞核
CssWOX4c	TEA007699.1	316	8.96	36 018.54	60.02	59.91	-0.736	细胞核
CssWOX8a	TEA027235.1	288	5.53	32 224.21	56.12	67.36	-0.695	细胞核
CssWOX8b	TEA017348.1	288	6.06	32 745.31	57.53	53.47	-0.984	细胞核
CssWOX9a	TEA033789.1	279	8.85	31 274.79	61.41	55.91	-0.834	细胞核
CssWOX9b	TEA029851.1	459	8.93	49 770.50	74.52	56.91	-0.550	细胞核
CssWOX11	TEA027449.1	212	5.09	22 791.41	79.30	67.12	-0.395	细胞核
CssWOX13a	TEA021703.1	410	6.84	46 513.77	50.56	76.49	-0.594	细胞核
CssWOX13b	TEA020072.1	706	7.84	81 150.51	49.65	80.88	-0.630	细胞核
CssWOX13c	TEA007801.1	287	7.72	32 557.99	58.76	74.36	-0.630	细胞核
DaszWOX1a	GWHPABKB016733	326	8.45	38 010.72	65.95	60.18	-0.915	细胞核
DaszWOX1b	GWHPABKB013155	916	6.51	102 942.02	48.18	76.12	-0.433	细胞核
DaszWOX2a	GWHPABKB014366	118	9.72	13 333.89	62.71	54.58	-0.840	细胞核
DaszWOX2b	GWHPABKB001060	254	6.53	27 823.75	59.93	59.53	-0.731	细胞核
DaszWOX4a	GWHPABKB015660	226	9.10	26 222.65	52.45	60.00	-0.977	细胞核
DaszWOX4b	GWHPABKB020125	234	9.30	26 676.05	57.99	61.67	-0.909	细胞核
DaszWOX5	GWHPABKB000744	157	9.62	17 769.22	53.75	75.03	-0.569	细胞核
DaszWOX8a	GWHPABKB020522	286	5.54	32 080.10	57.59	67.83	-0.698	细胞核
DaszWOX8b	GWHPABKB001976	287	6.07	32 259.75	61.39	53.66	-0.959	细胞核
DaszWOX9	GWHPABKB030319	354	8.63	38 729.50	49.81	59.18	-0.459	细胞核
DaszWOX11	GWHPABKB008034	197	5.25	21 261.58	82.03	63.35	-0.488	细胞核
DaszWOX13a	GWHPABKB017606	252	7.05	28 500.77	59.30	61.11	-0.928	细胞核
DaszWOX13b	GWHPABKB010750	183	9.10	20 592.04	61.42	56.50	-0.949	细胞核

Table 1

Fig.1

Table 2

Table 3

URSCU and fRFSC of codons of WOX gene families in three Camellia sinensis cultivars"

氨基酸 amino acid	密码子 codon	U_RSCU			f_RFSC			氨基酸 amino acid	密码子 codon	U_RSCU			f_RFSC
氨基酸 amino acid	密码子 codon	CSA	CSS	DASZ	CSA	CSS	DASZ	氨基酸 amino acid	密码子 codon	CSA	CSS	DASZ	CSA	CSS	DASZ
Phe	TTT	1.09	0.98	1.09	54.26	49.21	54.35	Ala	GCA	1.47	1.35	1.15	36.84	33.86	28.78
	TTC	0.91	1.02	0.91	45.74	50.79	45.65		GCG	0.18	0.29	0.43	4.51	7.17	10.73
Leu	TTA	0.78	0.70	0.84	13.07	11.60	13.95	Tyr	TAT	1.10	1.03	1.20	54.93	51.75	59.77
	TTG	1.53	1.59	1.60	25.57	26.52	26.75		TAC	0.90	0.97	0.80	45.07	48.25	40.23
	CTT	1.43	1.39	1.42	23.86	23.20	23.64		TAA	1.36	1.50	1.15	45.46	50.00	38.46
	CTC	1.16	1.23	1.21	19.32	20.44	20.16	TER	TAG	0.55	0.33	0.00	18.18	11.11	0.00
	CTA	0.48	0.55	0.47	7.95	9.12	7.75		TGA	1.09	1.17	1.85	36.36	38.89	61.54
	CTG	0.61	0.55	0.47	10.23	9.12	7.75	His	CAT	0.99	0.77	1.04	49.44	38.73	52.07
	ATT	1.18	1.22	1.37	39.39	40.66	45.61		CAC	1.01	1.23	0.96	50.56	61.27	47.93
Ile	ATC	0.93	0.98	0.98	31.06	32.60	32.75	Gln	CAA	1.15	1.20	1.21	57.50	60.12	60.32
	ATA	0.89	0.80	0.65	29.55	26.74	21.64		CAG	0.85	0.80	0.79	42.50	39.88	39.68
Met	ATG	1.00	1.00	1.00	100.00	100.00	100.00	Asn	AAT	1.10	1.06	1.18	55.13	52.94	59.19
Val	GTT	1.64	1.24	1.53	40.94	30.98	38.26		AAC	0.90	0.94	0.82	44.87	47.06	40.81
	GTC	0.54	0.74	0.51	13.39	18.43	12.76	Lys	AAA	0.78	0.71	0.67	38.96	35.42	33.50
	GTA	0.57	0.45	0.51	14.17	11.37	12.76		AAG	1.22	1.29	1.33	61.04	64.58	66.50
	GTG	1.26	1.57	1.45	31.50	39.22	36.22	Asp	GAT	0.98	1.11	1.21	49.02	55.61	60.54
Ser	TCT	1.40	1.45	1.59	23.31	24.09	26.59		GAC	1.02	0.89	0.79	50.98	44.39	39.46
	TCC	0.77	1.01	0.76	12.90	16.85	12.60	Glu	GAA	1.14	1.13	1.03	56.78	56.43	51.70
	TCA	1.33	1.11	1.38	22.22	18.48	23.01		GAG	0.86	0.87	0.97	43.22	43.57	48.30
	TCG	0.41	0.46	0.43	6.81	7.61	7.12	Cys	TGT	0.89	1.03	1.08	44.68	51.65	54.17
	AGT	1.18	1.03	1.04	19.71	17.21	17.26		TGC	1.11	0.97	0.92	55.32	48.35	45.83
	AGC	0.90	0.95	0.81	15.05	15.76	13.42	Trp	TGG	1.00	1.00	1.00	100.00	100.00	100.00
Pro	CCT	1.05	1.26	1.33	26.35	31.40	33.18	Arg	CGT	0.78	0.83	0.89	13.02	13.85	14.85
	CCC	0.77	0.69	0.73	19.16	17.36	18.18		CGC	0.85	0.85	0.71	14.20	14.19	11.79
	CCA	1.75	1.62	1.47	43.71	40.50	36.82		CGA	0.43	0.45	0.52	7.10	7.43	8.73
	CCG	0.43	0.43	0.47	10.78	10.74	11.82		CGG	0.43	0.41	0.71	7.10	6.76	11.79
Thr	ACT	1.47	1.47	1.66	36.69	36.87	41.49		AGA	1.74	1.78	1.60	28.99	29.73	26.64
	ACC	0.90	1.02	0.94	22.49	25.42	23.58		AGG	1.78	1.68	1.57	29.59	28.04	26.20
	ACA	1.30	1.21	1.05	32.54	30.17	26.20	Gly	GGT	1.14	1.17	1.26	28.43	29.14	31.52
	ACG	0.33	0.30	0.35	8.28	7.54	8.73		GGC	0.93	0.99	1.00	23.35	24.86	24.90
Ala	GCT	1.47	1.48	1.64	36.85	37.06	40.98		GGA	1.18	1.13	0.95	29.44	28.29	23.74
	GCC	0.87	0.88	0.78	21.80	21.91	19.51		GGG	0.75	0.71	0.79	18.78	17.71	19.84

Table 3

Table 4

A comparison of codon usage frequency between WOX gene families in three C. sinensis cultivars and six plants"

氨基酸 amino acid	密码子 codon	CsaWOX						CssWOX						DaszWOX
氨基酸 amino acid	密码子 codon	CSA	CSS	A	N	P	O	CSA	CSS	A	N	P	O	CSA	CSS	A	N	P	O
Phe	TTT	0.95	1.03	0.83	0.72	0.70	1.39	0.87	0.93	0.79	0.68	0.66	1.31	0.96	1.03	0.90	0.79	0.76	1.51
	TTC	1.06	0.97	0.74	0.85	0.87	0.68	1.18	1.08	0.86	0.98	1.01	0.79	1.06	0.97	0.80	0.92	0.95	0.74
Leu	TTA	1.06	1.07	0.64	0.61	0.55	1.34	0.94	0.95	0.60	0.57	0.51	1.26	1.14	1.14	0.75	0.71	0.64	1.55
	TTG	0.96	0.98	0.77	0.72	0.63	1.09	1.00	1.02	0.84	0.79	0.68	1.19	1.00	1.03	0.87	0.81	0.71	1.23
	CTT	1.04	1.22	0.62	0.62	0.52	0.98	1.01	1.18	0.64	0.64	0.53	1.01	1.03	1.20	0.67	0.67	0.55	1.06
	CTC	1.28	1.22	0.75	0.98	0.86	0.47	1.35	1.29	0.84	1.10	0.96	0.52	1.33	1.27	0.85	1.11	0.97	0.53
	CTA	0.77	0.75	0.50	0.53	0.41	0.65	0.88	0.86	0.61	0.64	0.50	0.78	0.75	0.73	0.53	0.56	0.43	0.68
	CTG	0.81	0.83	0.65	0.63	0.44	0.31	0.72	0.74	0.61	0.59	0.41	0.29	0.61	0.63	0.54	0.52	0.36	0.25
	ATT	0.83	0.92	0.86	0.67	0.63	1.30	0.85	0.95	0.94	0.73	0.69	1.43	0.96	1.07	0.95	0.74	0.70	1.44
Ile	ATC	1.12	0.96	0.79	1.05	0.96	0.75	1.17	1.01	0.88	1.17	1.07	0.84	1.18	1.01	0.80	1.06	0.97	0.76
	ATA	1.21	1.18	1.10	0.99	0.93	1.58	1.09	1.07	1.06	0.95	0.89	1.51	0.88	0.87	0.77	0.69	0.65	1.11
Met	ATG	1.00	1.00	1.32	1.30	1.38	1.36	1.00	1.00	1.09	1.07	1.14	1.13	1.00	1.00	1.11	1.08	1.15	1.14
Val	GTT	1.07	1.19	0.68	0.69	0.76	1.19	0.81	0.90	0.53	0.54	0.60	0.93	1.00	1.11	0.73	0.74	0.81	1.27
	GTC	0.76	0.70	0.47	0.55	0.54	0.30	1.05	0.96	0.67	0.77	0.76	0.43	0.73	0.66	0.51	0.59	0.58	0.33
	GTA	0.92	0.87	0.65	0.56	0.63	0.94	0.74	0.70	0.53	0.46	0.52	0.78	0.83	0.78	0.66	0.58	0.64	0.97
	GTG	1.09	1.05	0.82	0.85	0.84	0.59	1.36	1.31	1.05	1.09	1.08	0.75	1.26	1.21	1.07	1.12	1.10	0.77
Ser	TCT	0.94	1.10	0.92	1.16	1.13	1.82	0.97	1.13	0.96	1.21	1.18	1.91	1.07	1.25	1.01	1.28	1.24	2.01
	TCC	0.93	0.85	1.14	1.26	1.49	0.79	1.22	1.10	1.52	1.66	1.97	1.04	0.91	0.83	1.08	1.19	1.41	0.74
	TCA	0.99	0.98	1.21	1.25	1.12	1.78	0.82	0.82	1.02	1.06	0.94	1.50	1.03	1.02	1.21	1.25	1.12	1.78
	TCG	0.78	0.72	0.73	1.28	1.35	0.55	0.87	0.80	0.82	1.45	1.53	0.62	0.82	0.75	0.74	1.29	1.37	0.56
	AGT	1.17	1.23	1.40	1.47	1.30	2.22	1.02	1.08	1.24	1.30	1.15	1.97	1.02	1.08	1.18	1.25	1.10	1.88
	AGC	1.14	0.98	1.32	1.50	1.32	0.93	1.19	1.02	1.40	1.59	1.40	0.99	1.01	0.87	1.14	1.29	1.14	0.81
Pro	CCT	0.76	0.91	0.84	0.84	0.98	1.15	0.91	1.09	1.11	1.11	1.30	1.53	0.96	1.15	1.03	1.03	1.20	1.41
	CCC	1.13	0.97	2.15	1.73	2.19	0.94	1.02	0.88	2.17	1.74	2.21	0.95	1.07	0.92	1.98	1.59	2.02	0.87
	CCA	1.21	1.19	1.61	1.31	1.56	1.83	1.12	1.10	1.67	1.35	1.61	1.89	1.02	1.00	1.32	1.08	1.28	1.50
	CCG	0.87	0.74	0.75	1.28	1.60	0.36	0.87	0.74	0.83	1.42	1.78	0.40	0.95	0.81	0.79	1.37	1.71	0.38
Thr	ACT	1.07	1.26	1.26	1.09	1.53	2.08	1.07	1.27	1.38	1.19	1.67	2.27	1.20	1.42	1.43	1.23	1.73	2.36
	ACC	1.00	0.93	1.31	1.39	1.63	0.91	1.13	1.06	1.61	1.71	2.00	1.11	1.04	0.98	1.38	1.46	1.71	0.95
	ACA	0.99	0.93	1.25	1.13	1.28	1.69	0.92	0.86	1.26	1.13	1.29	1.70	0.80	0.75	1.00	0.91	1.03	1.36
	ACG	0.82	0.70	0.65	1.11	1.13	0.44	0.75	0.64	0.64	1.09	1.12	0.43	0.87	0.74	0.68	1.17	1.20	0.46
Ala	GCT	0.96	1.10	0.62	0.56	0.79	0.89	0.97	1.10	0.60	0.54	0.77	0.87	1.07	1.22	0.78	0.71	1.00	1.13
	GCC	1.11	1.01	1.00	0.83	1.05	0.34	1.12	1.02	0.97	0.80	1.02	0.33	0.99	0.91	1.02	0.84	1.07	0.34
	GCA	1.13	1.09	1.00	0.76	0.87	1.01	1.04	1.00	0.89	0.67	0.77	0.90	0.88	0.85	0.89	0.67	0.77	0.90
	GCG	0.48	0.41	0.24	0.37	0.58	0.08	0.76	0.64	0.36	0.57	0.89	0.12	1.13	0.96	0.64	1.00	1.56	0.22
Tyr	TAT	0.94	0.95	0.95	0.78	0.85	1.39	0.88	0.90	0.92	0.76	0.83	1.35	1.02	1.04	0.94	0.77	0.84	1.37
	TAC	1.09	1.06	0.83	0.84	1.20	0.75	1.16	1.14	0.92	0.93	1.33	0.83	0.97	0.95	0.67	0.68	0.97	0.61
	TAA	1.68	1.84	1.98	1.62	4.45	2.54	1.85	2.02	1.82	1.49	4.11	2.35	1.42	1.55	1.46	1.20	3.29	1.88
TER	TAG	0.82	0.86	1.42	1.42	1.78	0.89	0.50	0.53	0.73	0.73	0.91	0.46	0.00	0.00	0.00	0.00	0.00	0.00
	TGA	0.72	0.67	1.19	1.42	2.03	1.19	0.77	0.72	1.06	1.28	1.82	1.06	1.21	1.14	1.75	2.10	3.01	1.75
His	CAT	0.80	0.83	1.14	1.17	0.99	1.39	0.63	0.65	0.89	0.91	0.77	1.08	0.84	0.88	1.20	1.24	1.04	1.47
	CAC	1.33	1.24	1.84	1.84	1.93	1.16	1.61	1.50	2.22	2.22	2.33	1.40	1.26	1.18	1.75	1.75	1.84	1.11
Gln	CAA	1.00	0.94	2.11	1.98	1.96	3.03	1.04	0.98	1.93	1.81	1.79	2.77	1.05	0.99	2.02	1.89	1.87	2.90
	CAG	1.00	1.10	1.99	2.02	1.72	1.45	0.94	1.03	1.63	1.65	1.41	1.19	0.94	1.02	1.70	1.72	1.46	1.24
Asn	AAT	0.90	0.95	1.37	1.09	1.11	2.03	0.86	0.91	1.33	1.06	1.07	1.96	0.97	1.02	1.56	1.24	1.25	2.30
Asn	AAC	1.16	1.07	1.19	1.39	1.60	1.35	1.22	1.12	1.26	1.47	1.68	1.42	1.06	0.97	1.14	1.34	1.53	1.29
Lys	AAA	0.82	0.77	0.69	0.66	0.63	1.33	0.74	0.70	0.67	0.63	0.60	1.29	0.70	0.66	0.56	0.53	0.51	1.08
	AAG	1.17	1.24	1.02	1.00	1.03	1.04	1.24	1.31	1.15	1.12	1.16	1.16	1.27	1.35	1.05	1.03	1.06	1.07
Asp	GAT	0.72	0.75	0.49	0.48	0.43	0.70	0.82	0.85	0.62	0.61	0.54	0.89	0.89	0.92	0.64	0.63	0.56	0.93
	GAC	1.60	1.48	1.08	1.10	1.29	0.66	1.40	1.29	1.05	1.07	1.26	0.64	1.24	1.15	0.89	0.90	1.07	0.54
氨基酸 amino acid	密码子 codon	CsaWOX						CssWOX						DaszWOX
氨基酸 amino acid	密码子 codon	CSA	CSS	A	N	P	O	CSA	CSS	A	N	P	O	CSA	CSS	A	N	P	O
Glu	GAA	1.08	1.00	1.17	1.12	0.99	1.86	1.07	1.00	1.14	1.09	0.97	1.82	0.98	0.91	1.05	1.00	0.89	1.67
	GAG	0.91	1.00	0.95	1.04	0.94	0.79	0.92	1.00	0.94	1.03	0.93	0.78	1.02	1.11	1.05	1.14	1.04	0.87
Cys	TGT	0.81	0.87	0.71	0.76	0.67	1.21	0.93	1.00	0.82	0.88	0.77	1.38	0.98	1.05	0.98	1.05	0.92	1.65
	TGC	1.24	1.14	1.29	1.29	1.04	0.75	1.08	1.00	1.12	1.12	0.90	0.65	1.03	0.95	1.21	1.21	0.97	0.70
Trp	TGG	1.00	1.00	1.31	1.34	1.18	1.19	1.00	1.00	1.27	1.30	1.14	1.15	1.00	1.00	1.26	1.29	1.14	1.14
Arg	CGT	1.12	1.55	0.87	1.04	1.06	1.09	1.19	1.65	0.83	1.00	1.01	1.04	1.27	1.77	0.99	1.19	1.21	1.24
	CGC	1.66	1.97	2.25	2.19	1.90	0.53	1.66	1.97	2.02	1.97	1.70	0.48	1.38	1.64	1.87	1.82	1.58	0.44
	CGA	0.56	0.53	0.68	0.81	0.78	0.67	0.59	0.55	0.64	0.76	0.73	0.63	0.69	0.65	0.83	0.99	0.96	0.82
	CGG	0.68	0.69	0.87	1.15	0.75	0.32	0.65	0.66	0.74	0.99	0.64	0.27	1.13	1.14	1.45	1.92	1.25	0.53
	AGA	0.94	0.81	0.92	1.09	0.89	1.66	0.97	0.83	0.85	1.00	0.82	1.53	0.87	0.75	0.84	1.00	0.82	1.53
	AGG	1.14	1.18	1.62	1.46	1.40	1.11	1.08	1.12	1.38	1.24	1.19	0.95	1.01	1.05	1.43	1.29	1.24	0.99
Gly	GGT	0.96	1.14	0.90	0.89	1.11	1.35	0.99	1.17	0.84	0.83	1.03	1.26	1.07	1.26	0.96	0.96	1.18	1.44
	GGC	1.31	1.20	1.78	1.46	1.61	0.56	1.39	1.28	1.73	1.42	1.56	0.54	1.39	1.28	1.83	1.50	1.65	0.57
	GGA	0.95	0.86	0.85	0.89	0.91	1.30	0.92	0.82	0.75	0.78	0.80	1.14	0.77	0.69	0.66	0.69	0.71	1.01
	GGG	0.87	0.88	1.29	1.25	1.15	0.77	0.82	0.83	1.11	1.08	0.98	0.66	0.92	0.93	1.31	1.28	1.17	0.78

Table 4

Fig.2

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碱基组成 base composition	CSA	CSS	DASZ
GC₁	46.369	46.228	47.543
GC₂	45.577	44.789	45.697
GC₃	46.050	47.303	45.087
平均average	45.999	46.107	46.109

A comparative study of the evolution and codon usage bias in WOX gene family of three Camellia sinensis cultivars

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