Genetic analysis on heterosis and important traits of ornamental pepper hybrids

LIN Zixiang; MA Xiaowei; WU Jianguo; ZHU Zhujun; ZHU Biao

doi:10.11833/j.issn.2095-0756.20200745

Volume 38 Issue 4

Aug. 2021

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LIN Zixiang, MA Xiaowei, WU Jianguo, ZHU Zhujun, ZHU Biao. Genetic analysis on heterosis and important traits of ornamental pepper hybrids[J]. Journal of Zhejiang A&F University, 2021, 38(4): 820-827. doi: 10.11833/j.issn.2095-0756.20200745

Citation:

LIN Zixiang, MA Xiaowei, WU Jianguo, ZHU Zhujun, ZHU Biao. Genetic analysis on heterosis and important traits of ornamental pepper hybrids[J]. Journal of Zhejiang A&F University, 2021, 38(4): 820-827. doi: 10.11833/j.issn.2095-0756.20200745

Genetic analysis on heterosis and important traits of ornamental pepper hybrids

doi: 10.11833/j.issn.2095-0756.20200745

Key Laboratory of Quality Improvement of Agricultural Products of Zhejiang Province, College of Horticulture Science, Zhejiang A&F University, Hangzhou 311300, Zhejiang, China

Received Date: 2020-11-29
Rev Recd Date: 2021-04-28

Available Online: 2021-08-09

Publish Date: 2021-08-20

Abstract

Objective This research aims to explore the heterosis of ornamental pepper and the genetic law of ornamental traits, to promote the cultivation of new varieties of ornamental pepper and the retention of excellent traits. Method The oval leaf inbred line (P15) of cluster pepper (Capsicum annuum ‘Salsa XP Red’) and the lanceolate leaf inbred line (P6) of single pepper (C. annuum ‘Salsa Deep Orange’) were used as parents, an orthogonal combination P6×P15 and an inverse combination P15×P6 were configured, and F₂ population was constructed to explore the inheritance of fruit bearing pattern, leaf shape and plant type of ornamental pepper. Result There were significant differences between P6 and P15 in leaf size, plant height and number of lateral branches (P＜0.05). All F₁ combinations of cross ornamental pepper showed high heterosis in plant height, plant width and leaf shape. In F₂ population of reciprocal cross, the descending order of agronomic characters with the largest variance was leaf length, fruit number per plant, petiole length, plant width and plant height, and there was a significant correlation between plant type and leaf type traits (P＜0.01). The Chi-square test showed that in the results of the segregation ratio of quality traits in the F₂ generations of reciprocal crosses, the fruit cluster traits were inherited in a single gene recessively for solitary traits and limited growth versus infinite growth. The developmental plant type versus the erect plant type and the lanceolate leaf versus the oval leaf were both controlled by two genes, and had recessive epistasis. Conclusion There are significant differences between ornamental pepper hybrid parents and F₁ generation in 5 traits, such as the number of fruit per plant and the number of branching grades. Among them, the coefficient of variation of plant type is the smallest, the coefficient of variation of fruit weight and fruit number per plant is larger, and the difference of leaf length and fruit number per plant is the most significant. The analysis of the inheritance law of the important traits of ornamental pepper reveals that the fruiting bearing pattern of clusters versus solitary and limited branch versus infinite branch are recessive genetic traits controlled by single gene.[Ch, 2 fig. 8 tab. 22 ref.]
- ornamental pepper,
- reciprocal combination,
- heterosis,
- analysis of variance,
- genetic analysis

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观赏辣椒为辣椒属Capsicum中一类主要作为观赏用途的特殊类型，通常指那些果形奇特、果色斑斓、株型紧凑、适合盆栽的辣椒品种，既可用于日常食用，又可作园艺用途、花坛布置与家居装饰，具有良好的开发应用前景^[1]。多年来，国外育种者开始有针对性地对观赏辣椒的表型性状进行遗传改良，培育出了很多具有新奇性状的新品种。ASMA等^[2]用杂交方式培育了营养价值高出普通辣椒12%且适合观赏和盆栽种植的观赏辣椒品种；METTUPALAYAM等^[3]在研究辣椒果实提取物对糖尿病治疗疗效的过程中，发现多个观赏辣椒品种果实提取物对糖尿病患者的疗效要优于普通辣椒果实；MANDARSARI等^[4]发现：观赏辣椒的遗传特性明显优于普通辣椒，对高温干旱环境的适应能力更强。中国学者对观赏辣椒也进行了专门研究，如郭咏梅等^[5]以3个紫色观赏辣椒品种的果实为试材，进行花青素的提取工艺优化以及pH示差法测定条件的确定，并进一步证实观赏辣椒果实花青素含量与色差值存在显著的相关性；王云梅等^[6]通过杂交方式育成了6个适合凉山地区种植，耐寒且极具观赏价值的观赏辣椒品种。杂交育种是当前观赏辣椒新品种培育中的常用方法。目前，针对普通辣椒果实品质与辣椒素含量遗传规律的研究较多，对于观赏辣椒观赏特性、营养特性和环境适应性方面的研究也见报道，但针对观赏辣椒果实着生方式、叶形与株型以及辣椒花器等性状的遗传规律研究还未见报道。果实着生方式、叶形与株型等都是决定观赏价值的重要特性，了解其遗传规律将极大地提高育种效率。本研究以直立型簇生椒与开展型单生椒为材料，配置正反交组合及构建F₂群体，测量和统计亲本、F₁和F₂的叶形、果实着生方式、株型性状，研究其遗传规律，为后续新品种选育提供理论基础。

1. 材料与方法

1.1. 试验材料

本研究选用从美国进口单生椒Capsicum annuum ‘Salsa Deep Orange’的高代自交系P6，簇生椒C. annuum ‘Salsa XP Red’的高代自交系P15，杂交获得F₁，F₁自交获得F₂。P6最显著的特点为开展型株型，果实着生方式单生，披针形叶片，青熟果颜色黄色；P15最显著的特点为直立型株型，果实着生方式簇生，卵圆形叶片，青熟果颜色绿色(表1)。

材料	青熟果色	果形	果实着生状态	叶形	分枝类型	株型
P6	乳黄	短指形	单生	披针形	无限生长	开展
P15	绿	长锥形	簇生	卵圆形	有限生长	直立
P6×P15	绿	长锥形	单生	长卵圆	无限生长	半直立
P15×P6	绿	长锥形	单生	长卵圆	无限生长	半直立

Table 1. Main characteristics of ornamental pepper parents and their reciprocal cross combinations

1.2. 试验方法

试验在浙江农林大学实验基地的塑料温室内进行。试验材料于2019年8月12日播种，9月13日田间定植，10−12月观察记录。亲本定植各20株，正反交F₁各定植20株，正反交F₂群体分别定植150株。株行距45 cm×50 cm，采取常规田间管理。材料进入果实成熟期后调查其表型性状。

1.3. 性状调查

依据《辣椒种质资源描述规范和数据标准》^[7]选定13个农艺性状进行统计分析，其中品质性状4个，分别为果实着生方式、分枝类型、叶形、株型；数量性状9个，分别为株高、株幅、侧枝数、分枝级数、单株结果数、叶长、叶宽、叶柄长及单节叶腋着生花数；数量性状使用美耐特游标卡尺测量，每个指标设置10次重复。使用Canon EOS30D单反相机拍摄记录性状。使用SPSS和Excel软件对测量所得数据进行统计分析、相关性分析、方差分析、性状变异频率和卡方检验分析。变异系数(V_C)计算方法参考段晓铨^[8]方法： V_C=D_S/N_M×100%，D_S为标准偏差，N_M是平均值；中亲优势=(F₁−M_P)/M_P×100%，超亲优势=(F₁−H_P)/H_P×100%，F₁为杂交组合F₁代表型，M_p与H_p为双亲表型性状的均值与高值亲本的表型。

3. 讨论

相较于张宇斌等^[9]、王述彬等^[10]通过境外引种，多代筛选辣椒优异品种的育种方式，国内的观赏辣椒自主育种起步较晚。很多育种者在辣椒的农艺性状表型上做了大量改良^[11-12]。多数研究认为：单株产量、单株结果数均有杂种优势，使用杂种优势选育适合国内推广的观赏辣椒品种是可行的。

通过对不同株型和叶形的观赏辣椒进行杂交组合，后代群体的表型性状优势指数与前人研究结果类似。陈蕊红^[13]选择了8个亲本辣椒，组配成16个不完全双列组合，发现：超中优势效果明显，单株产量杂种优势明显。这与本研究得出的单株结果数有显著杂种优势结论一致。果实数量和辣椒观赏性密切相关，对于辣椒观赏价值起决定性作用。杂种优势往往也表现在株高、冠幅、叶片等在常规辣椒育种中常被忽视的表型性状上。JEONG等^[14]在研究盆景辣椒杂交育种过程中就指出：观赏辣椒遗传特性主要表现为叶长变异性强，侧枝数量和株型性状变异不明显，在亲本选择的时候可以优先考虑选取侧枝表现较优的品种。尽管观赏辣椒属于辣椒属中的一个特异群体，但其在株高、株幅、叶形上较高的变异特性和杂种优势与常规辣椒基本一致。通过杂交育种可改善观赏辣椒生长势、分枝性等性状，更有利于观赏辣椒在全国范围内的推广。

辣椒诸多表型性状间存在错综复杂的关系^[15]。徐睿等^[16]分析了40份观赏辣椒种质资源的遗传性状，认为单株果实数量与株幅、株高存在正相关。陈倩倩等^[17]对19个辣椒品种的12个表型性状进行遗传分析，结果表明：不同辣椒品种之间性状差异显著，变异系数为8.49%~37.77%。叶长与叶宽、始花节位、对椒单果质量呈极显著正相关。可根据育种目标对辣椒性状的要求，有目的地筛选育种材料。本研究中辣椒正反交F₂代群体的植株性状与叶片性状间存在显著正相关，与前人研究结果基本一致。辣椒正反交组合F₁、F₂代群体中，单株结果数、分枝级数、单节叶腋着生花数和叶长都存在显著变异，不易通过杂交育种保留其叶形与开花性状，而株高和株幅变异较少，遗传稳定性较好。MAMEDOR等^[18]、王昆等^[19]也得出了相似的结果。

辣椒遗传规律受到基因的作用与控制，不同学者在辣椒各表型性状上进行了研究。隋益虎等^[20]使用紫色辣椒和绿色辣椒品种配制杂交组合，测定其叶片花青素相对含量，研究其遗传规律，发现辣椒叶片花青素含量遗传模式符合2对加性-显性-上位性主基因+加性-显性-上位性多基因模型，因此，对辣椒紫叶性状的遗传选择应在分离早期世代进行。本研究针对辣椒叶形的研究显示：长卵圆形叶片对披针形叶片为显性，且披针形叶片对卵圆形叶片具有隐性上位作用。此研究结果可为选育具有奇特叶型的辣椒种质材料提供理论依据。李全辉等^[21]以线辣椒C. annuum var. longunt黄色突变体为母本，野生型红色自交系辣椒为父本进行杂交，发现辣椒果实红色对黄色为显性，线辣椒成熟果色黄色性状受1对隐性基因控制。本研究针对辣椒果实着生方式进行研究，发现辣椒簇生对单生和有限分枝对无限分枝均为单基因控制隐性遗传性状。株型和叶形受2对等位基因控制，这与LÜ等^[22]的研究结果一致。株型、叶形、果实的着生方式是观赏性辣椒重要的表现性状，可为选育优良性状的观赏辣椒种质资源提供依据。

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[1]	杨莎, 陈文超, 张竹青, 等. 观赏盆栽辣椒品种及关键栽培技术[J]. 湖南农业科学, 2015(5): 93 − 95.	YANG Sha, CHEN Wenchao, ZHANG Zhuqing, et al. Key cultivation techniques of ornamental potted hot pepper varieties [J]. Hunan Agric Sci, 2015(5): 93 − 95.
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世代	株高/cm	株幅/cm	侧枝数/个	分枝级数/级	单株结果数/个	叶长/mm	叶宽/mm	叶柄长/mm	单节叶腋着生花数/朵
P6	21.64±1.61 c	41.27±2.35 cd	8.50±1.72 b	7.40±1.17 a	26.10±4.84 d	63.94±1.66 c	23.57±0.95 b	34.11±1.08 c	4.00±1.49 d
P15	24.90±1.79 b	38.93±3.12 d	3.30±0.82 d	2.40±0.52 e	37.30±4.00 c	87.94±1.13 a	34.53±1.24 a	48.45±0.88 a	9.20±1.32 a
P6×P15 F₁	25.33±2.91 b	53.27±5.02 a	8.10±1.60 b	5.50±1.08 b	71.10±5.63 a	54.72±5.04 d	21.05±3.45 b	22.69±3.89 d	7.00±1.63 bc
P15×P6 F₁	25.65±3.15 b	44.04±2.95 bc	11.90±2.73 a	4.70±1.06 cd	68.60±16.02 a	61.01±3.48 c	21.92±1.60 b	23.25±1.31 d	5.70±1.89 c
P6×P15 F₂	28.45±2.55 a	45.48±3.18 b	6.47±0.72 c	5.02±0.41 bc	55.77±8.10 b	69.61±7.39 b	35.30±3.28 a	37.14±3.77 b	6.91±0.99 bc
P15×P6 F₂	29.04±3.30 a	44.72±2.37 b	5.29±1.08 c	4.23±0.48 d	56.31±11.93 b	71.86±5.89 b	36.91±4.97 a	38.99±4.47 b	7.36±0.93 b
说明：数据为平均值±标准差，其中：P6和P15样本统计量为20，F₁样本统计量为20，F₂样本统计量为150；同列不同字母表示不同　　　　　世代间差异显著(P＜0.05)

指标	株高/cm	株幅/cm	侧枝数/个	分枝级数/级	单株结果数/个	叶长/mm	叶宽/mm	叶柄长/mm	单节叶腋着生花数/朵
平均值	28.45	45.48	6.47	5.02	55.77	69.61	35.30	37.14	6.91
最大值	43.56	84.91	11.00	9.00	110.00	132.17	53.67	65.51	13.00
最小值	14.45	27.59	2.00	2.00	19.00	26.83	14.70	19.96	1.67
变幅	29.11	57.32	9.00	7.00	91.00	105.34	38.98	45.55	11.33
方差	49.85	81.34	4.48	4.06	437.05	607.46	128.22	129.33	9.57
标准差	7.06	9.02	2.12	2.01	20.91	24.65	11.32	11.37	3.09
变异系数/%	25	20	33	40	37	35	32	31	45
说明：统计F₂样本数量为150

指标	株高/cm	株幅/cm	侧枝数/个	分枝级数/级	单株结果数/个	叶长/mm	叶宽/mm	叶柄长/mm	单节叶腋着生花数/朵
平均值	29.04	44.72	5.29	4.23	56.31	71.86	36.91	38.99	7.36
最大值	45.53	69.37	11.00	10.00	115.00	119.29	69.93	68.46	13.00
最小值	13.42	23.34	3.00	2.00	17.00	28.03	16.80	14.52	2.33
变幅	32.11	46.03	8.00	8.00	98.00	91.26	53.12	53.94	10.67
方差	63.75	105.71	3.33	2.59	541.05	652.22	174.25	181.43	9.37
标准差	7.98	10.28	1.83	1.61	23.26	25.54	13.20	13.47	3.06
变异系数/%	27	23	35	38	41	36	36	35	42
说明：统计F₂样本数量为150

项目	株高	株幅	侧枝数	分枝级数	单株结果数	叶长	叶宽	叶柄长	单节叶腋着生花数
株高	1
株幅	0.63**	1
侧枝数	0.42**	0.68**	1
分枝级数	−0.24**	0.27**	0.26**	1
单株结果数	0.58**	0.72**	0.73**	0.11	1
叶长	0.35**	−0.00	−0.16	−0.47**	0.02	1
叶宽	0.38**	−0.02	−0.17*	−0.51**	0.00	0.91**	1
叶柄长	0.21**	−0.21**	−0.30**	−0.55**	−0.12	0.80**	0.88**	1
单节叶腋着生花数	0.31**	−0.09	−0.25**	−0.55**	−0.04	0.88**	0.93**	0.93**	1
说明：*表示在0.01 水平(双侧)上显著相关。表示在0.05 水平(双侧)上显著相关

项目	株高	株幅	侧枝数	分枝级数	单株结果数	叶长	叶宽	叶柄长	单节叶腋着生花数
株高	1
株幅	0.69**	1
侧枝数	0.43**	0.32**	1
分枝级数	−0.01	0.32**	0.32**	1
单株结果数	0.75**	0.54**	0.62**	0.16	1
叶长	0.10	0.04	0.06	0.08	0.11	1
叶宽	0.06	0.07	−0.05	0.07	0.05	0.89**	1
叶柄长	0.07	0.06	−0.04	0.05	0.06	0.93**	0.96**	1
单节叶腋着生花数	0.05	0.09	−0.01	0.03	0.02	0.89**	0.88**	0.90**	1
表示：*表示在0.01 水平(双侧)上显著相关。表示在0.05 水平(双侧)上显著相关