甘草优良种质遴选指标与传粉特性研究
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摘要
对甘草进行优良品种选育是提高和稳定甘草药材产量和品质的重要途径。利用或创造变异,通过人工定向选择将有利的变异固定下来,可实现新品种的培育。其中,选择与评价是甘草育种工作的主要部分。选择不仅作为独立的育种手段,而且也是其他各种育种途径不可缺少的重要环节。同时,甘草的多用性决定其育种目标的多样性,所以如何依据不同育种目标制定适当的评价指标体系就显得十分必要。基于此,本论文以甘草品种选育基础工作为主线,通过4个不同种质(种源)2年4点栽培对比、10个类型甘草栽培比较及野外人工控制授粉等试验项目,采用数量遗传学分析方法并结合相关统计分析,对甘草优良种质的选择指标体系、不同育种目标的甘草种质评价指标体系以及传粉生物学特性三部分进行研究,其主要研究结果如下：
1、分析了甘草数量性状间遗传变异与相互关系,并建立目标性状间接选择指标体系
甘草栽培群体内存在较丰富的遗传变异,可为优良类型的选择提供材料。其中,主根长、侧根数和根鲜重变异系数在20%以上,而甘草苷、异甘草苷、甘草素变异系数则在30%以上。大部分表型性状与根鲜重呈显著正相关(p<0.05),成分性状间存在极显著正相关(p<0.01),并与主根长关系密切。D20、D芦头与根长、主根长分别对根鲜与成分性状正向直接作用最大。对于2年生甘草,其株高、主根长、甘草苷和异甘草苷不但有较高的遗传变异系数、遗传力,且相对遗传进展也较大,而对于3年生甘草,与成分性状相比,表型性状选择潜力较小。
借助部分表型性状及其组合,可实现对甘草目标性状的有效选择。其中,以根鲜重为目标性状时,株高、根长与D20、侧根数可分别作为2、3年生甘草的间接选择指标,且不同性状组合比直接对根鲜重选择效率最大可提高40%以上。以成分PC1为目标性状时,单性状的选择效果不明显,但以分蘖数、主根长和根长组成的方程,其选择相对效率是最高的,最大可提高选择效率148%。以甘草酸为目标性状时,主根长和尖削度组合可较直接选择效率提高16.8%。
2、种质和环境因素对甘草主要数量性状影响各异,且与气候因子关系密切
大部分性状在不同种质间和不同环境下差异达到显著水平(p<0.05)。种质与环境对所有性状影响显著(p<0.05),而茎粗、分蘖数和甘草酸在两年间均受种质和环境互作影响显著(p<0.05)。同时,种质对分蘖数、主根长和尖削度影响大于环境,而环境对D芦头、D20和侧根数影响大于种质,剩余大部分性状受两者共同影响。此外,甘草苷和异甘草苷受种质影响较大,甘草素和异甘草素受环境影响较大,而甘草酸受种质和环境共同影响；从根鲜重和成分性状的主成分双标图显示种质G2 (WNT-01)和G3(JX-01)为优良种质。同时,环境E4(酒泉)对成分积累有利,而E2(赤峰)和E3(杭锦旗)分别在2010年和2011年时对根鲜重和成分积累有利。
通过性状与地理气候因子间逐步回归分析结果表明,对数量性状影响较大的地理气候因子主要为日照时数,其次为降雨量、月均温与七月均温。其中,株高、茎粗等地上性状受日照条件影响较大,根长、根粗、侧根数及根鲜重等根部性状受日照、温度和降雨量三个因素的影响。而对于成分性状,甘草酸主要受日照条件的影响,甘草苷、甘草素和异甘草素则受日照和降雨量的影响。3、不同变异类型甘草生长及光合性能存在一定变异,且指标间相互关系密切
不同类型甘草生长特性有一定的差异,尤其在不同来源地间差异更明显。各类型株高进入始盛期、高峰期和盛末期的时间分别为出苗后的第28 d、47 d和65 d左右。茎粗进入高峰期和盛末期的时间分别为第16 d和50 d左右。其中,株高和茎粗进入高峰期时间最早和最晚相差10 d。不同类型株高生长期的差异主要来自类型的不同来源地之间的差异；不同类型甘草光合性能指标存在较大的变异,且相互关系密切。各指标间的相关分析表明,叶绿素含量和比叶重某种程度上能反映甘草光合能力。而株高、茎粗仅在生长前期与净光合速率(Pn)呈显著(p<0.05)正相关关系。
4、对不同类型甘草进行综合评价,并依据不同育种目标建立相应的评价指标体系
通过主成分分析可对甘草目标性状信息较好的提取,将不同育种目标的甘草多性状转换成2-3个独立的综合指标,能够解释不同目标的含义,并筛选出不同特点的种质材料。其中,作为高酸型育种材料,V3、V1和V6较好；无论作为饮片型还是浸膏型育种材料V8、V7和V3均较好；作为饮片型育种材料V1和V2得分分列4、5位,而作为浸膏型育种V9和V10则列4、5位。
以根条形(根长与尖削度)、产量(根鲜重)和成分含量(甘草酸、甘草苷、异甘草苷、甘草素、异甘草素、总黄酮和总皂苷)为评价指标,根据主成分提取的综合变量信息,初步对不同育种目标甘草评价指标体系进行构建,其中,高酸型甘草主要考虑甘草酸及药材产量；饮片型甘草主要考虑成分指标,并兼顾根条形；浸膏型甘草主要考虑成分指标,并兼顾药材产量。
5、对甘草传粉生物学特性研究,并对甘草品种选育与扩繁途径进行了探讨
通过P/O、OCI值间接估计以及人工控制授粉直接验证方法对甘草繁育系统类型进行研究,推测甘草繁育系统类型为自交、异交亲和,但以异交为主。根据其繁殖特点,可实现多种方式结合进行品种选育。采用选择育种时,对于混合选择和单株选择后代,要采取适当的隔离措施,以免花粉污染。同时,可对高世代选择材料可实施人工辅助自交,培育自交系。人工辅助授粉的时期选择在花蕾长约10～15cm,根据花粉活力和柱头可授性变化,针对于人工自花或异花授粉选择不同时期进行。
本文的研究特色和创新：
(1)借助部分表型性状及其组合,建立了甘草根鲜重与成分性状的间接选择指标体系；(2)从多育种目标角度,建立了高酸型、饮片型和浸膏型甘草的种质评价指标体系；(3)证实甘草自然群体中自交、异交同时存在,但通常以异交为主。
Establishing good-breeding for licorice(Glycyrrhiza uralensis Fisch.) is an important way to improve and control yield and quality. Using or creating variation, and fixing down the favorable genes through the artificial selection, may realize the breeding of the new varieties. Among them, the selection and evaluation are the main parts of the breeding. Selection is not only an independent breeding method itself, but also an indispensable segment of all other sorts of breeding approaches. Meanwhile, the multi-purpose of licorice determines its diversity of breeding objectives. Therefore, it is very necessary to make appropriate evaluation method according to multi-objective breeding of licorice. Consequently, this paper aimed at analysing three aspects, including selection index of elite germplasm, evaluation index of elite germplasm of licorice according to multi-objective breeding and pollination biology characteristics, which is to lay the foundation for the good-breeding of licorice seeds. For that purpose, the main quantitative traits of various cultivated licorice genotypes (provenance and variation types) were investigated at four locations in 2010 and 2011. The main results were as follows:
1. Genetic variation and relationship between the quantitative traits of cultivated licorice was analyzed, and selection index system for target traits was established It can provide the germplasm for the selection of elite types, for abundant genetic variation existed in cultivated licorice population. Among them, the coefficients of variation (CV) of taproot length (TRL), lateral root number (LRN) and root fresh weight (RFW) were above 20%, the CVs of liquiritin (LQ), Isoliquiritin (ILQ) and liquiritigenin (LQG) were above 30%. Persons correlation between traits showed that most of phenotypic traits had significant(p< 0.05) positive correlation with RFW, and among bioactive components traits had highly significant(p<0.01) positive correlation, and existed close relationship with TRL. Results of the path coefficient analyses showed that D20, root diameter (RD) and root length (RL) and TRL had the greatest positive direct effect on RFW and bioactive components traits respectively. For two-year licorice, its plant height (PH), TRL, LQ and ILQ not only had a higher genetic variation coefficient (GCV) and heritability, but also bigger relative genetic advance (RGA). However, the select potential of phenotypic traits was poor, compared with the bioactive component traits for the three-year licorice.
It can realize effective indirectly selection for objective traits of licorice by certain phenotypic trait and their combination. Among them, the highest relative value of correlative genetic advance (CGA) for RFW in 2a and 3a were predicated indirectly by combined PH with RL and D20 with LRN, respectively. And its relative value increased by 40%than that of RFW. For principal bioactive component traits (PC1), the highest relative value of combination with tillering number (TL), TRL and RL by selection index was increased by 148%; nevertheless, selection index of single phenotypic trait was not effective.
2. Germplasm (Genotype) and environment had different effects on the main quantitative traits of licorice, which also had close relations with climate factors
The differences among various genotypes and different environment for most evaluated traits were significantly (p<0.05) through Duncan's multiple range tests. The quantitative traits of licorice were influenced significant (p<0.05) by genotype (G) and environment (E), and stalk diameter (SD), TN and glycyrrhizin (GL) were influenced significantly by genotype and environment interreaction (GEI) in all both two years. Meanwhile, genotypes have lager effect on TN, TRL, taperingness (TR) than environment, and environment have larger effect on RD, D20 and LRN than genotype, respectively. And genotype and environment have similar effect on the other traits. In addition, LQ and ILQ were influenced more by genotype than by environment, in contrast LOQ and isoliquiritigenin (ILQG) were influenced more by environment than by gennotype, and GL were influenced by both genotype and environment. Biplot of the principal component analysis of RFW and bioactive component traits of licorice showed that, germplasm from G2 (WNT-01) and G3 (JX-01) were elite, and the environment in E4 (Jiuquan) was favorable for the accumulation of bioactive component. Moreover, E2 (Chifeng) and E3 (Hangjinqi) were favorable for the RFW and accumulation of bioactive component in 2010 and 2011, respectively.
Stepwise regression analysis (SRA) of 15 quantitative traits and nine geographical climate factors showed that sunshine duration had the greatest effect on quantitative traits, followed by rainfall, the average monthly temperature and mean temperature in July. Among them, the upground traits PH and SD were influenced most strongly by sunshine duration, and the root part traits RL, RD, and LRN were influenced by sunshine duration, temperature and rainfall simultaneously. And for bioactive component traits, GL was mainly influenced by sunshine duration, and LQ, LQG and ILQG were mainly influenced by sunshine duration and rainfall.
3. The growth and photosynthetic performance for different variation types of cultivated licorice existed certain variation, and the relationships between those parameters were close
There existed certain difference in growth characteristics for different variation types of cultivated licorice, and these differences were more significant especially among different locations of origin. For PH, the time of entering bloom prime stage, bloom luxuriant stage and bloom final stage was 28 d,47 d and 65 d after seedling, respectively. However, for SD, the time of entering bloom luxuriant stage and bloom final stage was 16 d and 50 d, respectively. Among these types, for SD and PH, the period between the earliest time and the latest time of entering the bloom luxuriant stage was 10d. The difference of PH in growth period mainly came from the difference among the origin location; Photosynthetic performance parameters existed great variation and there existed close relationship in different variation types of cultivated licorice. Person's correlation analysis among parameters showed that chlorophyll content (Chl) and specific leaf weight (SLW) can reflect photosynthetic performance of licorice to a certain extent. However, PH and SD only had significant(p<0.05) positive correlation with net photosynthetic rate (Pn) only in the early growth period.
4. Different variation type of cultivated licorice was evaluated comprehensively, and corresponding evaluation index system was established according to different breeding objectives
The information of target traits in licorice could be extracted preferably by means of principal components analysis (PCA), and then multi-characters of different objectives breeding of licorice could be converted into two to three independent comprehensive traits (PCs), which could explain different breeding objectives, and germplasm with various characteristics could be selected. Among them, V3, V1 and V6 performed better as "hiGL type", and V8, V7 and V3 performed better as both "yinpian type" and "extractum type". As "yinpian type", the comprehensive PCs score of V1 and V2 ranged the fourth and fifth as "yinpian type", respectively. The comprehensive PCs score of V9 and V10 ranged the fourth and fifth as "extractum type", respectively. Evaluation indexes system of licorice germplasm for different breeding objectives were constructed preliminarily, according to different traits extracted by PCs of different traits, including root strip (RL and TR), root yield (RFW) and bioactive component traits (LQ, LQG, GL, ILQ, ILQG, total flavonoids (TF) and total saponins (TS)). Among them, "extractum type" mainly considered comprehensive bioactive components, and also took the root yield into account; "hiGL type" mainly considered the total amount of GL (absolute amount); "yinpian type" mainly considered the root strip, and also gives consideration to the comprehensive bioactive components.
5. Pollination biology characteristics was explored, and breeding technology of licorice was discussed
The breeding system of licorice was presumed to be selfing and outcrossing compatible, but outcrossing took the leading role, according to pollen-ovule ratio (P/O), outcrossing index (OCI) and artificial pollination. Thereby, we may realize breeding by using combination of many ways, according to the breeding characteristics of licorice. For mass selection breeding offspring, quarantine measures should be taken to avoid the pollen pollution. In addition, selfing could be taken by artificial pollination for selected materials of high generation to foster inbred lines. When the length of bud is 10-15 cm approximately, it was time for artificial pollination. Artificial pollination origin selfing and outcrossing were carried out in different periods, based on pollen vitality and stigma receptivity.
In this paper, there are three characteristics and innovations:
(1) In this paper, indirect selection index system between RFW and bioactive component traits was established by means of certain phenotypic traits and their combination.
(2) The germplasm evaluation index system of "hiGL type","yinpian type"and" extractum type"was extablished from the point of multi-objective breeding.
(3) This paper proved that selfing and outcrossing exist simultaneously in licorice natural population, and outcrossing dominates in it.
1、分析了甘草数量性状间遗传变异与相互关系,并建立目标性状间接选择指标体系
甘草栽培群体内存在较丰富的遗传变异,可为优良类型的选择提供材料。其中,主根长、侧根数和根鲜重变异系数在20%以上,而甘草苷、异甘草苷、甘草素变异系数则在30%以上。大部分表型性状与根鲜重呈显著正相关(p<0.05),成分性状间存在极显著正相关(p<0.01),并与主根长关系密切。D20、D芦头与根长、主根长分别对根鲜与成分性状正向直接作用最大。对于2年生甘草,其株高、主根长、甘草苷和异甘草苷不但有较高的遗传变异系数、遗传力,且相对遗传进展也较大,而对于3年生甘草,与成分性状相比,表型性状选择潜力较小。
借助部分表型性状及其组合,可实现对甘草目标性状的有效选择。其中,以根鲜重为目标性状时,株高、根长与D20、侧根数可分别作为2、3年生甘草的间接选择指标,且不同性状组合比直接对根鲜重选择效率最大可提高40%以上。以成分PC1为目标性状时,单性状的选择效果不明显,但以分蘖数、主根长和根长组成的方程,其选择相对效率是最高的,最大可提高选择效率148%。以甘草酸为目标性状时,主根长和尖削度组合可较直接选择效率提高16.8%。
2、种质和环境因素对甘草主要数量性状影响各异,且与气候因子关系密切
大部分性状在不同种质间和不同环境下差异达到显著水平(p<0.05)。种质与环境对所有性状影响显著(p<0.05),而茎粗、分蘖数和甘草酸在两年间均受种质和环境互作影响显著(p<0.05)。同时,种质对分蘖数、主根长和尖削度影响大于环境,而环境对D芦头、D20和侧根数影响大于种质,剩余大部分性状受两者共同影响。此外,甘草苷和异甘草苷受种质影响较大,甘草素和异甘草素受环境影响较大,而甘草酸受种质和环境共同影响；从根鲜重和成分性状的主成分双标图显示种质G2 (WNT-01)和G3(JX-01)为优良种质。同时,环境E4(酒泉)对成分积累有利,而E2(赤峰)和E3(杭锦旗)分别在2010年和2011年时对根鲜重和成分积累有利。
通过性状与地理气候因子间逐步回归分析结果表明,对数量性状影响较大的地理气候因子主要为日照时数,其次为降雨量、月均温与七月均温。其中,株高、茎粗等地上性状受日照条件影响较大,根长、根粗、侧根数及根鲜重等根部性状受日照、温度和降雨量三个因素的影响。而对于成分性状,甘草酸主要受日照条件的影响,甘草苷、甘草素和异甘草素则受日照和降雨量的影响。3、不同变异类型甘草生长及光合性能存在一定变异,且指标间相互关系密切
不同类型甘草生长特性有一定的差异,尤其在不同来源地间差异更明显。各类型株高进入始盛期、高峰期和盛末期的时间分别为出苗后的第28 d、47 d和65 d左右。茎粗进入高峰期和盛末期的时间分别为第16 d和50 d左右。其中,株高和茎粗进入高峰期时间最早和最晚相差10 d。不同类型株高生长期的差异主要来自类型的不同来源地之间的差异；不同类型甘草光合性能指标存在较大的变异,且相互关系密切。各指标间的相关分析表明,叶绿素含量和比叶重某种程度上能反映甘草光合能力。而株高、茎粗仅在生长前期与净光合速率(Pn)呈显著(p<0.05)正相关关系。
4、对不同类型甘草进行综合评价,并依据不同育种目标建立相应的评价指标体系
通过主成分分析可对甘草目标性状信息较好的提取,将不同育种目标的甘草多性状转换成2-3个独立的综合指标,能够解释不同目标的含义,并筛选出不同特点的种质材料。其中,作为高酸型育种材料,V3、V1和V6较好；无论作为饮片型还是浸膏型育种材料V8、V7和V3均较好；作为饮片型育种材料V1和V2得分分列4、5位,而作为浸膏型育种V9和V10则列4、5位。
以根条形(根长与尖削度)、产量(根鲜重)和成分含量(甘草酸、甘草苷、异甘草苷、甘草素、异甘草素、总黄酮和总皂苷)为评价指标,根据主成分提取的综合变量信息,初步对不同育种目标甘草评价指标体系进行构建,其中,高酸型甘草主要考虑甘草酸及药材产量；饮片型甘草主要考虑成分指标,并兼顾根条形；浸膏型甘草主要考虑成分指标,并兼顾药材产量。
5、对甘草传粉生物学特性研究,并对甘草品种选育与扩繁途径进行了探讨
通过P/O、OCI值间接估计以及人工控制授粉直接验证方法对甘草繁育系统类型进行研究,推测甘草繁育系统类型为自交、异交亲和,但以异交为主。根据其繁殖特点,可实现多种方式结合进行品种选育。采用选择育种时,对于混合选择和单株选择后代,要采取适当的隔离措施,以免花粉污染。同时,可对高世代选择材料可实施人工辅助自交,培育自交系。人工辅助授粉的时期选择在花蕾长约10～15cm,根据花粉活力和柱头可授性变化,针对于人工自花或异花授粉选择不同时期进行。
本文的研究特色和创新：
(1)借助部分表型性状及其组合,建立了甘草根鲜重与成分性状的间接选择指标体系；(2)从多育种目标角度,建立了高酸型、饮片型和浸膏型甘草的种质评价指标体系；(3)证实甘草自然群体中自交、异交同时存在,但通常以异交为主。
Establishing good-breeding for licorice(Glycyrrhiza uralensis Fisch.) is an important way to improve and control yield and quality. Using or creating variation, and fixing down the favorable genes through the artificial selection, may realize the breeding of the new varieties. Among them, the selection and evaluation are the main parts of the breeding. Selection is not only an independent breeding method itself, but also an indispensable segment of all other sorts of breeding approaches. Meanwhile, the multi-purpose of licorice determines its diversity of breeding objectives. Therefore, it is very necessary to make appropriate evaluation method according to multi-objective breeding of licorice. Consequently, this paper aimed at analysing three aspects, including selection index of elite germplasm, evaluation index of elite germplasm of licorice according to multi-objective breeding and pollination biology characteristics, which is to lay the foundation for the good-breeding of licorice seeds. For that purpose, the main quantitative traits of various cultivated licorice genotypes (provenance and variation types) were investigated at four locations in 2010 and 2011. The main results were as follows:
1. Genetic variation and relationship between the quantitative traits of cultivated licorice was analyzed, and selection index system for target traits was established It can provide the germplasm for the selection of elite types, for abundant genetic variation existed in cultivated licorice population. Among them, the coefficients of variation (CV) of taproot length (TRL), lateral root number (LRN) and root fresh weight (RFW) were above 20%, the CVs of liquiritin (LQ), Isoliquiritin (ILQ) and liquiritigenin (LQG) were above 30%. Persons correlation between traits showed that most of phenotypic traits had significant(p< 0.05) positive correlation with RFW, and among bioactive components traits had highly significant(p<0.01) positive correlation, and existed close relationship with TRL. Results of the path coefficient analyses showed that D20, root diameter (RD) and root length (RL) and TRL had the greatest positive direct effect on RFW and bioactive components traits respectively. For two-year licorice, its plant height (PH), TRL, LQ and ILQ not only had a higher genetic variation coefficient (GCV) and heritability, but also bigger relative genetic advance (RGA). However, the select potential of phenotypic traits was poor, compared with the bioactive component traits for the three-year licorice.
It can realize effective indirectly selection for objective traits of licorice by certain phenotypic trait and their combination. Among them, the highest relative value of correlative genetic advance (CGA) for RFW in 2a and 3a were predicated indirectly by combined PH with RL and D20 with LRN, respectively. And its relative value increased by 40%than that of RFW. For principal bioactive component traits (PC1), the highest relative value of combination with tillering number (TL), TRL and RL by selection index was increased by 148%; nevertheless, selection index of single phenotypic trait was not effective.
2. Germplasm (Genotype) and environment had different effects on the main quantitative traits of licorice, which also had close relations with climate factors
The differences among various genotypes and different environment for most evaluated traits were significantly (p<0.05) through Duncan's multiple range tests. The quantitative traits of licorice were influenced significant (p<0.05) by genotype (G) and environment (E), and stalk diameter (SD), TN and glycyrrhizin (GL) were influenced significantly by genotype and environment interreaction (GEI) in all both two years. Meanwhile, genotypes have lager effect on TN, TRL, taperingness (TR) than environment, and environment have larger effect on RD, D20 and LRN than genotype, respectively. And genotype and environment have similar effect on the other traits. In addition, LQ and ILQ were influenced more by genotype than by environment, in contrast LOQ and isoliquiritigenin (ILQG) were influenced more by environment than by gennotype, and GL were influenced by both genotype and environment. Biplot of the principal component analysis of RFW and bioactive component traits of licorice showed that, germplasm from G2 (WNT-01) and G3 (JX-01) were elite, and the environment in E4 (Jiuquan) was favorable for the accumulation of bioactive component. Moreover, E2 (Chifeng) and E3 (Hangjinqi) were favorable for the RFW and accumulation of bioactive component in 2010 and 2011, respectively.
Stepwise regression analysis (SRA) of 15 quantitative traits and nine geographical climate factors showed that sunshine duration had the greatest effect on quantitative traits, followed by rainfall, the average monthly temperature and mean temperature in July. Among them, the upground traits PH and SD were influenced most strongly by sunshine duration, and the root part traits RL, RD, and LRN were influenced by sunshine duration, temperature and rainfall simultaneously. And for bioactive component traits, GL was mainly influenced by sunshine duration, and LQ, LQG and ILQG were mainly influenced by sunshine duration and rainfall.
3. The growth and photosynthetic performance for different variation types of cultivated licorice existed certain variation, and the relationships between those parameters were close
There existed certain difference in growth characteristics for different variation types of cultivated licorice, and these differences were more significant especially among different locations of origin. For PH, the time of entering bloom prime stage, bloom luxuriant stage and bloom final stage was 28 d,47 d and 65 d after seedling, respectively. However, for SD, the time of entering bloom luxuriant stage and bloom final stage was 16 d and 50 d, respectively. Among these types, for SD and PH, the period between the earliest time and the latest time of entering the bloom luxuriant stage was 10d. The difference of PH in growth period mainly came from the difference among the origin location; Photosynthetic performance parameters existed great variation and there existed close relationship in different variation types of cultivated licorice. Person's correlation analysis among parameters showed that chlorophyll content (Chl) and specific leaf weight (SLW) can reflect photosynthetic performance of licorice to a certain extent. However, PH and SD only had significant(p<0.05) positive correlation with net photosynthetic rate (Pn) only in the early growth period.
4. Different variation type of cultivated licorice was evaluated comprehensively, and corresponding evaluation index system was established according to different breeding objectives
The information of target traits in licorice could be extracted preferably by means of principal components analysis (PCA), and then multi-characters of different objectives breeding of licorice could be converted into two to three independent comprehensive traits (PCs), which could explain different breeding objectives, and germplasm with various characteristics could be selected. Among them, V3, V1 and V6 performed better as "hiGL type", and V8, V7 and V3 performed better as both "yinpian type" and "extractum type". As "yinpian type", the comprehensive PCs score of V1 and V2 ranged the fourth and fifth as "yinpian type", respectively. The comprehensive PCs score of V9 and V10 ranged the fourth and fifth as "extractum type", respectively. Evaluation indexes system of licorice germplasm for different breeding objectives were constructed preliminarily, according to different traits extracted by PCs of different traits, including root strip (RL and TR), root yield (RFW) and bioactive component traits (LQ, LQG, GL, ILQ, ILQG, total flavonoids (TF) and total saponins (TS)). Among them, "extractum type" mainly considered comprehensive bioactive components, and also took the root yield into account; "hiGL type" mainly considered the total amount of GL (absolute amount); "yinpian type" mainly considered the root strip, and also gives consideration to the comprehensive bioactive components.
5. Pollination biology characteristics was explored, and breeding technology of licorice was discussed
The breeding system of licorice was presumed to be selfing and outcrossing compatible, but outcrossing took the leading role, according to pollen-ovule ratio (P/O), outcrossing index (OCI) and artificial pollination. Thereby, we may realize breeding by using combination of many ways, according to the breeding characteristics of licorice. For mass selection breeding offspring, quarantine measures should be taken to avoid the pollen pollution. In addition, selfing could be taken by artificial pollination for selected materials of high generation to foster inbred lines. When the length of bud is 10-15 cm approximately, it was time for artificial pollination. Artificial pollination origin selfing and outcrossing were carried out in different periods, based on pollen vitality and stigma receptivity.
In this paper, there are three characteristics and innovations:
(1) In this paper, indirect selection index system between RFW and bioactive component traits was established by means of certain phenotypic traits and their combination.
(2) The germplasm evaluation index system of "hiGL type","yinpian type"and" extractum type"was extablished from the point of multi-objective breeding.
(3) This paper proved that selfing and outcrossing exist simultaneously in licorice natural population, and outcrossing dominates in it.
引文
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