香橼与佛手F1杂种鉴定及果形差异分析

doi:10.16420/j.issn.0513-353x.2022-0745

摘要/Abstract

摘要：

以香橼（Citrus medica L.）为母本，佛手（C. medica var. L. sarcodactylis Swingle）为父本进行授粉杂交，将得到的种子进行培育获得19份材料。从佛手和香橼转录组数据筛选出SSR位点设计TP-M13-SSR分子标记引物，对19份材料进行基因分型。每对核心引物获得17 ~ 32个等位基因，平均等位基因数为24.188个。观测杂合度变化为0.188 ~ 1.000，多态性指数为0.944。19个F₁代株系中17个确定为真杂种，杂交率为89.47%。通过影像分析F₁代果形，指状果与柑果果形分离比为2︰3。佛手和香橼花芽石蜡切片观察发现，形成指状或柑果果形差异起始于现蕾期至现瓣期的心皮隆起。对现蕾期至花后3 d的佛手、香橼花芽转录组信息与心皮特征进行相关性分析，分离得到一批雌蕊形成相关基因；在即将开花的佛手、香橼及F₁代雌蕊中进行qRT-PCR验证，分析这些基因与果形差异的关联性，WOX1基因在现蕾期和现瓣期的表达与指状特征相关。YABBY家族基因可与花发育C类和E类基因及WOX家族协同作用调控佛手指状果形建成。

关键词: 香橼, 佛手, SSR, 指状果, 雌蕊形成, WOX1

Abstract:

In this study，19 F₁ hybrids derived from the cross between Citrus medica L. and C. medica var. L. sarcodactylis Swingle were cultured as materials. SSR loci were selected from the transcriptome data of fingered citron and citron. Based on this，TP-M13-SSR primers were designed and then 19 F₁ hybrids were genotyped. A total of 17 to 32 alleles were obtained from 16 SSR primers，and the average number of alleles was 24.188. The heterozygosity was ranged from 0.188 to 1.000，and the polymorphism index was about 0.944. The results showed that 17 out of 19 F₁ lines were true hybrids，and the hybridization rate was 89.47%. Those materials were phenotyped by morphological photography analysis and the ratio of finger fruit to hesperidium is 2︰3. Paraffin section analysis of early flower buds of fingered citron and citron found the difference between their fruit shape initiating from the carpel bulges since the visible bud stage to the petals emerging stage. From the visible bud stage to 3 d after anthesis of fingered citron and citron，the correlation analysis between the transcriptome data and carpel feature was done，and a number of genes associated with gynoecium formation were identified. These genes were validated in the pistils of citron，fingered citron，and hybrid F₁ lines during the pre-anthesis stage. The results showed that the expression of WOX1 was related to the character of finger fruit from the visible bud stage to the petals emerging stage. The YABBY family coordinated with the classes C，E genes of the ABC model of flower development and WOX family to regulate the fruit shaping in fingered citron.

Key words: citron, fingered citron, SSR, finger fruit, gynoecium formation, WOX1

梅煜琳, 徐劲剑, 蒋梦玥, 尉俊海, 宗宇, 陈文荣, 廖芳蕾, 郭卫东. 香橼与佛手F₁杂种鉴定及果形差异分析[J]. 园艺学报, 2023, 50(7): 1402-1418.

MEI Yulin, XU Jinjian, JIANG Mengyue, YU Junhai, ZONG Yu, CHEN Wenrong, LIAO Fanglei, GUO Weidong. Identification of F₁ Hybrids and Analysis of Fruit Shape Difference Between Fingered Citron and Citron[J]. Acta Horticulturae Sinica, 2023, 50(7): 1402-1418.

图/表 13

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引物序号 Primer code	重复单元 Repeat motif	引物序列（5′-3′） Primer sequence		产物大小/bp Product size
Z1	(AT)*8	F：<M13>-TAACCTAGCATCCTCCAC	R：TATGCGGGTAAATCTCGT	184
Z5	(TC)*10	F：<M13>-CATTGATTCTTTCCTTCG	R：AAAATTAGCCAGAAACGT	247
Z7	(TA)*9	F：<M13>-TGATCTCCTCTGGCGAATG	R：CCTCCTCGTCACCATCCA	277
Z15	(TC)*6	F：<M13>-TCCCCTTCATTTATCCTT	R：TTTTGTTGATGGCGTTGG	172
Z17	(CT)*8	F：<M13>-ATGCAAGCCGCTTTTCTC	R：ACCCAAGATTCTGGTCAGG	203
Z24	(AT)*18	F：<M13>-ACCAGTCCTACGCATCAC	R：CCCGATACCCTGGATTTT	260
Z52	(TA)*12	F：<M13>-CTCCCACTGTCCCTTCGT	R：GCTTGCTGAGCGGATTTA	186
Z54	(AT)*14	F：<M13>-CATTAGAGTTCATTGGGAATA	R：TACCTTACGTTTTGTCGC	254
Z92	(CT)*12	F：<M13>-TACGGTATCAATTCCTTC	R：GGCATTATCAGACCCAAA	312
Z124	(AG)*23	F：<M13>-CCGATCATCAGGGACTACTA	R：CATCTCCAGCACCATTCTT	316
Z131	(CT)*9	F：<M13>-TATAAGGCAAGTGGGGTGA	R：GTGGCAACAAGAATACAAGAT	161
Z171	(TA)*17	F：<M13>-CTAAAGCGATCTTGACATA	R：GGACGGGACTTACTACCAG	252
Z178	(AT)*11	F：<M13>-GAGAACTTTTGTTTATTGGAG	R：TTCTTTCATGCTTCCGTTC	165
Z198	(TAA)*13	F：<M13>-GTAACTGGTGGATTTGTCG	R：CCATAGTCGGACCCTCTTT	201
Z199	(TAA)*13	F：<M13>-GTAACTGGTGGATTTGTCG	R：CCATAGTCGGACCCTCTTT	220
Z204	(TG)*9	F：<M13>-TGCTGGAAAGGAACGAAAC	R：CATTGCATGGCTGCTCATA	244
M13通用引物 <FAM>或<HEX>-TGTAAAACGACGGCCAGT
M13 universal primer

引物序号 Primer code	重复单元 Repeat motif	引物序列（5′-3′） Primer sequence		产物大小/bp Product size
Z1	(AT)*8	F：<M13>-TAACCTAGCATCCTCCAC	R：TATGCGGGTAAATCTCGT	184
Z5	(TC)*10	F：<M13>-CATTGATTCTTTCCTTCG	R：AAAATTAGCCAGAAACGT	247
Z7	(TA)*9	F：<M13>-TGATCTCCTCTGGCGAATG	R：CCTCCTCGTCACCATCCA	277
Z15	(TC)*6	F：<M13>-TCCCCTTCATTTATCCTT	R：TTTTGTTGATGGCGTTGG	172
Z17	(CT)*8	F：<M13>-ATGCAAGCCGCTTTTCTC	R：ACCCAAGATTCTGGTCAGG	203
Z24	(AT)*18	F：<M13>-ACCAGTCCTACGCATCAC	R：CCCGATACCCTGGATTTT	260
Z52	(TA)*12	F：<M13>-CTCCCACTGTCCCTTCGT	R：GCTTGCTGAGCGGATTTA	186
Z54	(AT)*14	F：<M13>-CATTAGAGTTCATTGGGAATA	R：TACCTTACGTTTTGTCGC	254
Z92	(CT)*12	F：<M13>-TACGGTATCAATTCCTTC	R：GGCATTATCAGACCCAAA	312
Z124	(AG)*23	F：<M13>-CCGATCATCAGGGACTACTA	R：CATCTCCAGCACCATTCTT	316
Z131	(CT)*9	F：<M13>-TATAAGGCAAGTGGGGTGA	R：GTGGCAACAAGAATACAAGAT	161
Z171	(TA)*17	F：<M13>-CTAAAGCGATCTTGACATA	R：GGACGGGACTTACTACCAG	252
Z178	(AT)*11	F：<M13>-GAGAACTTTTGTTTATTGGAG	R：TTCTTTCATGCTTCCGTTC	165
Z198	(TAA)*13	F：<M13>-GTAACTGGTGGATTTGTCG	R：CCATAGTCGGACCCTCTTT	201
Z199	(TAA)*13	F：<M13>-GTAACTGGTGGATTTGTCG	R：CCATAGTCGGACCCTCTTT	220
Z204	(TG)*9	F：<M13>-TGCTGGAAAGGAACGAAAC	R：CATTGCATGGCTGCTCATA	244
M13通用引物 <FAM>或<HEX>-TGTAAAACGACGGCCAGT
M13 universal primer

基因 Gene	序列号 ID	引物序列（5′-3′） Primer sequence		产物大小/bp Product size
SEP1	Cm158770.1	F：GCTTAGGCTTAGAGGAGTTG	R：GAATGAGCAGGTTGGTTG	244
SEP2	Cm240370.1	F：AGCGAACTCAGAGGAATC	R：ACCTAGTTGCTGCTCACC	263
SEP4	Cm146780.1	F：CCTGAGGAAGAAGTTGGAC	R：CGATAGAGGCTGAAAGAACC	142
SHP1	Cm078390.1	F：GGAATCGGCAGAGTCAGA	R：GCGTACTGGTGATTGGGT	260
SHP2	Cm164940.1	F：AATACTGGGTCTATCTGTGAA	R：TCTCGGCTATCTTTGCTC	289
SPL6	Cm158790.1	F：GACTCATCTGTTGACCCTT	R：CACTACCGACTTCACCATTA	211
SPL8	Cm162720.1	F：AGATGAAGCGACTCAAGAAT	R：GTGGGTGGGAGCGTAAGA	286
SPL9	Cm119150.1	F：TCTGATCGACTTCAGTGCATATC	R：AATTCCAGGACCAGAGAAACC	208
YABBY1A	Cm015380.1	F：ATCCCAAATCCATCTCCGAAC	R：TGAATCGGTTGTACGCAGAG	157
YABBY1B	Cm112520.1	F：ACTTCCCTCACATCCATTTCG	R：ACTACAACGAAACCCCAGC	192
YABBY4	Cm075530.1	F：CAAACATGGCTCACAAGGAAG	R：CTTCAAAGTTCCAAGCTCGC	132
YABBY5A	Cm147040.1	F：TGCGAACACCCACTAACAAG	R：AATGGATGTGAGGGAAGTGTG	198
CRC	Cm158810.1	F：CTCTCACCACTTCTCTCTTCTTC	R：AGAAGTTGCAGCGGACATAG	204
WOX1	Cm275910.1	F：ATTCTACTACGACTGCTATT	R：TCTCTCTTGTTTTGTTCACC	190
PRS	Cm223590.1	F：CCATAAAGCAAGAGAGAGGC	R：AGCTGGGTTAGAAAAGGAAC	190
LUG	Cm111030.1	F：CCCACCTCCTAAATGGCAATA	R：TAGCCGCATCATCCAAAGAA	150
AGL66	Cm105600.1	F：ATGTGAAGGAGAAATGCTGG	R：AATCTTTGGGTTGAGACTGG	388
AP2	Cm077350.1	F：AATCTAAAATAATTGCCCAA	R：AATAACTCAACTCACTCTGC	324
TIR1	Cm233920.1	F：GACGCAGCTCTACTCTCAGG	R：CTTTCTTGGTCCAGCAACAG	204
CsActin	Cs1g05000.1	F：CCAAGCAGCATGAAGATCAA	R：ATCTGCTGGAAGGTGCTGAG	101

基因 Gene	序列号 ID	引物序列（5′-3′） Primer sequence		产物大小/bp Product size
SEP1	Cm158770.1	F：GCTTAGGCTTAGAGGAGTTG	R：GAATGAGCAGGTTGGTTG	244
SEP2	Cm240370.1	F：AGCGAACTCAGAGGAATC	R：ACCTAGTTGCTGCTCACC	263
SEP4	Cm146780.1	F：CCTGAGGAAGAAGTTGGAC	R：CGATAGAGGCTGAAAGAACC	142
SHP1	Cm078390.1	F：GGAATCGGCAGAGTCAGA	R：GCGTACTGGTGATTGGGT	260
SHP2	Cm164940.1	F：AATACTGGGTCTATCTGTGAA	R：TCTCGGCTATCTTTGCTC	289
SPL6	Cm158790.1	F：GACTCATCTGTTGACCCTT	R：CACTACCGACTTCACCATTA	211
SPL8	Cm162720.1	F：AGATGAAGCGACTCAAGAAT	R：GTGGGTGGGAGCGTAAGA	286
SPL9	Cm119150.1	F：TCTGATCGACTTCAGTGCATATC	R：AATTCCAGGACCAGAGAAACC	208
YABBY1A	Cm015380.1	F：ATCCCAAATCCATCTCCGAAC	R：TGAATCGGTTGTACGCAGAG	157
YABBY1B	Cm112520.1	F：ACTTCCCTCACATCCATTTCG	R：ACTACAACGAAACCCCAGC	192
YABBY4	Cm075530.1	F：CAAACATGGCTCACAAGGAAG	R：CTTCAAAGTTCCAAGCTCGC	132
YABBY5A	Cm147040.1	F：TGCGAACACCCACTAACAAG	R：AATGGATGTGAGGGAAGTGTG	198
CRC	Cm158810.1	F：CTCTCACCACTTCTCTCTTCTTC	R：AGAAGTTGCAGCGGACATAG	204
WOX1	Cm275910.1	F：ATTCTACTACGACTGCTATT	R：TCTCTCTTGTTTTGTTCACC	190
PRS	Cm223590.1	F：CCATAAAGCAAGAGAGAGGC	R：AGCTGGGTTAGAAAAGGAAC	190
LUG	Cm111030.1	F：CCCACCTCCTAAATGGCAATA	R：TAGCCGCATCATCCAAAGAA	150
AGL66	Cm105600.1	F：ATGTGAAGGAGAAATGCTGG	R：AATCTTTGGGTTGAGACTGG	388
AP2	Cm077350.1	F：AATCTAAAATAATTGCCCAA	R：AATAACTCAACTCACTCTGC	324
TIR1	Cm233920.1	F：GACGCAGCTCTACTCTCAGG	R：CTTTCTTGGTCCAGCAACAG	204
CsActin	Cs1g05000.1	F：CCAAGCAGCATGAAGATCAA	R：ATCTGCTGGAAGGTGCTGAG	101

重复类型	重复次数 Number of repeat									总数	比例/%
Repeat type	5	6	7	8	9	10	11	12	>12	Total	Percentage
二核苷酸Dinucleotide	—	712	415	283	243	166	101	59	150	2 129	43.37
三核苷酸Trinucleotide	1 322	605	311	131	49	35	18	8	22	2 501	50.95
四核苷酸Tetranucleotide	115	36	7	6	2	0	0	0	0	166	3.38
五核苷酸Pentanucleotide	20	8	3	0	0	0	0	0	0	31	0.63
六核苷酸Hexanucleotide	59	13	7	0	1	2	0	0	0	82	1.67
总数Total	1 516	1 374	743	420	295	203	119	67	172	4 909
比例/% Percentage	30.88	27.99	15.14	8.56	6.01	4.14	2.42	1.36	3.50		100.00

香橼与佛手F₁杂种鉴定及果形差异分析

Identification of F₁ Hybrids and Analysis of Fruit Shape Difference Between Fingered Citron and Citron

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重复类型	重复基序	所占比例/%
Repeat type	Repeat motif	Percentage
二核苷酸	AG/CT	55.61
Dinucleotide	AC/GT	15.45
	AT/AT	21.18
	CG/CG	0.75
三核苷酸	AAG/CTT	24.32
Trinucleotide	AGG/CCT	4.37
	ACC/GGT	5.19
	AGC/CTG	11.69
	ATC/ATG	16.06
	CCG/CGG	3.30
	ACG/CGT	2.48
	AAC/GTT	7.20
	AAT/ATT	24.20
	ACT/AGT	1.18

引物序号 Primer code	N_a观测等位基因数 Observed number of alleles	N_e有效等位基因数 Effective number of alleles	H_o观测杂合度 Observed heterozygosity	H_e期望杂合度 Expected heterozygosity	P_i多态性信息量 Polymorphism information content
Z1	28	21	1.000	0.981	0.956
Z5	18	16	0.188	0.966	0.932
Z7	17	14	0.429	0.963	0.924
Z15	20	21	0.667	0.958	0.932
Z17	28	17	0.941	0.988	0.957
Z24	25	18	0.611	0.968	0.938
Z52	19	14	0.929	0.968	0.930
Z54	27	20	0.750	0.981	0.954
Z92	25	19	0.526	0.977	0.949
Z124	17	16	0.438	0.948	0.912
Z131	32	21	0.952	0.986	0.961
Z171	25	19	0.579	0.974	0.946
Z178	26	19	0.474	0.982	0.954
Z198	28	21	1.000	0.978	0.953
Z199	30	21	0.762	0.985	0.960
Z204	22	20	0.550	0.967	0.940
均值 Mean	24.188	18.563	0.675	0.973	0.944

基因 Gene	相关系数 Correlation coefficient		基因 Gene	相关系数 Correlation coefficient
基因 Gene	现瓣期 Petals emerging stage	即将开放期 Pre-anthesis stage	基因 Gene	现瓣期 Petals emerging stage	即将开放期 Pre-anthesis stage
SEP1	-0.358	0.086	YABBY4	0.501	0.258
SEP2	-0.645	-0.775^*	YABBY5A	0.645	0.430
SEP4	0.788^*	-0.430	CRC	0.358	0.775
SHP1	-0.072	-0.775^*	WOX1	0.788^*	-0.775^*
SHP2	-0.645	-0.775^*	PRS	0.620	0.430
SPL6	-0.358	0.775^*	LUG	-0.072	-0.775^*
SPL8	-0.501	-0.430	AGL66	-0.358	0.086
SPL9	0.501	0.258	AP2	-0.501	-0.602
YABBY1A	0.358	0.258	TIR1	0.501	0.775^*
YABBY1B	0.501	0.258

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