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Article

Genetic Variability in a Population of Oilseed Rape DH Lines Developed from F1 Hybrids of a Cross between Black- and Yellow-Seeded DH Lines. II. Seed Quality

by
Laurencja Szała
1,*,
Zygmunt Kaczmarek
2,
Marcin Matuszczak
1 and
Teresa Cegielska-Taras
1
1
Department of Genetics and Breeding of Oilseed Crops, Plant Breeding and Acclimatization Institute—National Research Institute, Strzeszyńska 36, 60-479 Poznań, Poland
2
Department of Biometry and Bioinformatics, Institute of Plant Genetics–Polish Academy of Sciences, Strzeszyńska 34, 60-479 Poznań, Poland
*
Author to whom correspondence should be addressed.
Agriculture 2022, 12(3), 340; https://doi.org/10.3390/agriculture12030340
Submission received: 29 November 2021 / Revised: 4 February 2022 / Accepted: 22 February 2022 / Published: 27 February 2022
(This article belongs to the Section Crop Genetics, Genomics and Breeding)
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Abstract

:
One of the main reasons for oilseed rape breeding is to improve the quality and composition of seeds by increasing fat and protein content, and to reduce dietary fibre. We attempted to obtain such varieties by crossing two DH lines of winter oilseed rape of different origin, H2-26 and Z-114, which have black and yellow seeds, respectively. The cross was followed by in vitro androgenesis, resulting in a population of 44 DH lines that were studied in a field experiment over two seasons. The following characters of the resulting seeds were analysed: fat, protein, neutral detergent fibre, acid detergent fibre, glucosinolate content and seed colour. The main objective was to check the variability of the DH lines obtained from F1 hybrid black- and yellow-seeded oilseed rape. The range of variability of the DH lines significantly exceeded the range of variability of the parental lines for all traits. These experiments showed that by choosing the appropriate parental genotypes of oilseed rape it is possible to break the negative correlation between protein and fat content. The high level of heritability of traits related to seed quality raises the possibility of improving breeding lines through selection based on phenotype.

1. Introduction

Growing demand for oilseed rape (Brassica napus L. var. oleifera) exerts a permanent pressure on breeders to improve seed yield and quality. The main selection goals in oilseed rape breeding have remained unchanged for several years and include increasing productivity, obtaining oil with a different composition of fatty acids depending on the intended use of oil and increasing the quality of oilseed rape meal.
These goals can, in part, be achieved by improving the chemical composition of oilseed rape seeds, which consist mainly of lipids (37–50%) and proteins (15–26%) as well as carbohydrates, lignins, ash and secondary metabolites (e.g., phenolic compounds) [1,2].
Oilseed rape seeds are a source of high-quality oil, which is used, with modifications, for many food and industrial purposes. Moreover, an important by-product of canola oil extraction is oilseed rape meal—a very rich raw material that contains up to 50% protein on a dry weight basis [3]. It is also rich in calcium, phosphorus, magnesium and zinc. Oilseed rape meal has a similar content of vitamins E, B1, B2, B6 as soybean meal, but it contains much more folic acid, vitamin PP, biotin and choline. The proteins contained in the seeds of oilseed rape are characterized by a favourable composition of amino acids, especially a good content of the exogenous amino acids lysine, threonine and tryptophan. However, the nutritional value of both the expeller and the post-extraction meal of oilseed rape is limited by its high fibre content, which results in the low digestibility of the feed.
In general, dietary fibre is a collection of heterogeneous substances, mainly non-starch polysaccharides and lignins, which derive from plant cell walls [4]. These substances can be divided into four basic groups of compounds: pectin, hemicellulose, cellulose and lignin. Pectins, which are easily dissolved in water, are rapidly fermented and have good digestibility. The remaining three groups of compounds form various fractions of dietary fibre. So-called neutral detergent fibre (NDF) contains all three compounds, while acid detergent fibre (ADF) contains only cellulose and lignin. The third fraction–ADL–is made solely of lignin [5].
Much oilseed rape breeding research has focused on reducing levels of fibre and anti-nutritional substances such as glucosinolates (GSL) and phenolic compounds and increasing protein and fat content. The breeding of yellow-seeded oilseed rape is very promising, because the seeds of such varieties have a low fibre content due to their transparent coat [6,7,8]. Reducing the fibre content of oilseed rape cake or post-extraction meal increases the availability of protein and improves the overall digestibility of the feed.
Modern breeding programmes make extensive use of biotechnology [9,10]. In winter oilseed rape breeding, doubled haploid (DH) technology, which uses an in vitro androgenesis process, has already become a routine way of obtaining completely homozygous and fully genetically stable lines in one generation. The benefits of DH lines are primarily acceleration of the breeding process by reducing the time taken to obtain homozygous lines compared to inbreeding and an increase in the efficiency of selection in early generations [11]. In addition, for polygenic traits, DH technology requires significantly fewer genotypes compared to classical breeding [12,13].
The main objective of the study was to evaluate the DH line population of winter oilseed rape derived from F1 hybrids obtained by crossing black- and yellow-seeded DH lines for fat, protein, fibre and GSL content and to select the best genotypes.

2. Materials and Methods

2.1. Plant Material

This study used the population of DH lines of winter oilseed rape derived from F1 hybrids obtained by crossing the yellow-seeded DH line Z-114 and the black-seeded DH line H2-26. The cross was followed by in vitro androgenesis in isolated microspore culture [14,15]. The plant material thus comprised a set of 44 DH lines together with the 2 parental lines. The parental yellow-seeded line Z-114 was selected from the population of DH lines developed from yellow-seeded breeding materials obtained at the Plant Breeding and Acclimatization Institute–National Research Institute (PBAI-NRI) in Poznań, Poland, from a natural mutant with brighter seeds and the Canadian spring line [16]. The seeds of DH Z-114 line were characterized by excellent yellow colour and low fibre content. In contrast, the black-seeded parental line DH H2-26 obtained at PBAI-NRI by in vitro androgenesis from breeding line 10199 had relatively high protein content, good plant habit and high resistance to frost.

2.2. Experimental Conditions

The DH line population along with the parental lines were used in a field experiment conducted over 2 seasons, 2016/2017 and 2017/2018. The field experiment was carried out in a randomized design with 4 replications in the sandy loam soils at the Łagiewniki Experimental Station of the Smolice Plant Breeding company. Each genotype was grown in a plot 1.2 m × 5.0 m in size. Oilseed rape genotypes were sown on 28 August in 2016 and 31 August in 2017 and harvested 23 July 2017 and 20 July 2018, respectively, at the seed maturity stage using a plot harvester. In both seasons, the level of mineral fertilization was the same and was as follows: 205 N, 40 P2O5, 60 K2O, and 30 S (kg·ha−1). Full chemical protection of plants was applied, and other agronomic practices were similar to those used for commercial planting in the area. The weather conditions varied during 2 analysed growing seasons, especially during spring and early summer. The sums of precipitation were 365.1 mm from March to July 2017 and 303.8 mm from March to July 2018. Mean temperatures for these periods at the experimental site were 12.8 °C and 13.2 °C for the first and second year, respectively. The monthly temperatures and precipitations in each growing season, from the beginning of March till the end of July (harvest), are presented in Table 1.
Just before flowering, several plants in each plot were covered with bags to obtain self-pollinated seeds that were then analysed for GSL content, protein and two fractions of dietary fibre, NDF and ADF. In contrast, the fat content was measured in samples of seeds collected from open-pollinated plants because the instrument for fat content analysis in place required a sample of 60–80 g per analysis. Seed samples for chemical analyses were taken from each plot in every replication of the trial.
GSL content was analysed by gas chromatography [17]. To determine the content of protein and both fibre fractions, a near-infrared reflectance spectroscopy (NIRS) method was used. The fat content in dry seeds (about 5% moisture) was determined by pulsed nuclear magnetic resonance (NMR; MQA7005, Oxford Instruments, Abingdon, UK). Seed coat colour was determined with a Color Flex spectrophotometer on a scale from 0 to 5 (black to yellow; Figure 1).

2.3. Methods of Statistical Analysis

Experimental data were analysed using uni- and multivariate statistical methods [18,19]. Two-factor analysis of variance was performed for seed quality traits and seed colour. This enabled the determination of statistical characteristics for all genotypes for each individual year. Heritability in the broad sense was calculated using the following formula [20,21]:
H2 = σg2/σp2
where:
  • σg2—genotypic variance
  • σp2—phenotypic variance.
The phenotypic variance was calculated from the formula:
σp2 = σg2 + σge2/m + σe2/rm
where:
  • σge2—variance of genotype-environmental interaction
  • σe2—error variance
  • m—number of environments
  • r—number of replications
The assessment of the relationships between analysed traits was made on the basis of correlation coefficients for the appropriate pairs of variables (quantitative traits), determined on the basis of the mean values of DH lines for both years jointly. The results are presented in the form of a correlation matrix.
To identify transgressive lines with advantageous and significant effects for the studied traits, each DH line was compared with parent lines using the F-statistic. A given line was considered as transgressive if its value was significantly higher than the value of the better-scoring parent or significantly lower than the value of the lower-scoring parent. Positive transgressive effects were sought for fat and protein content, and negative transgressive effects were investigated for NDF, ADF, and GSL content. Moreover, each DH line with low fibre content was compared and tested by F-statistic with the Z-114 line to identify DH lines with significantly better effects than the yellow-seeded parental line in terms of other studied traits.
The studied genotypes were divided into homogeneous groups according to seed colour using the procedure of Gabriel [22]. The affiliation of genotypes to particular colour groups was described by Szała et al. [23].
To classify the studied DH lines in terms of six traits treated simultaneously, hierarchical clustering was performed using Ward’s method based on Mahanalobis distance, which was treated as a measure of multitrait phenotypic distance between genotypes.

3. Results

3.1. Statistical Characteristics of Studied Traits

A total of 44 DH lines and their parental lines H2-26 and Z-114 were studied in a field experiment over two seasons. The following characters were analysed: fat, protein, NDF, ADF, GSL content and seed colour. The results of two-factor analyses of variance for genotypes (DH lines and parental lines) and environments (years), carried out for all the investigated traits, are presented in Table 2. They indicate that both sources of variation tested in the trial, environments and genotypes, were significant (p < 0.01) for all monitored traits, apart from NDF content. With regard to NDF content, only genotype effects were significantly expressed. Furthermore, a significant genotype by environment interaction was found for fat, protein, ADF and GSL content as well as for seed colour.
Coefficients of variation for all traits were higher for the DH line population than for the parental lines (Table 3). Among the seed quality traits, the highest variation in the DH line population was observed for GSL content (CV = 17.3% and CV = 22.1%, in the first and second year of the experiment, respectively), and for ADF content (CV = 20.5% and CV = 18.1%, in the first and second year of the experiment, respectively). The lowest variability among the DH lines was obtained for fat content (CV = 3.3% and CV = 4.8%, in the first and second year, respectively). This latter trait also turned out to be the most constant, as its coefficients of variation for both parental lines and in both years ranged from 0.7% to 2.8%.
The broad-sense heritability coefficients were high for all six traits and ranged from 0.69 for GSL content to 0.97 for ADF content (Table 3).
Frequency distribution of seed biochemical traits and seed colour for two years were presented as histograms in Figure 2. The studied DH population showed typical quantitative variation for fat and protein contents. The distribution of NDF and GSL contents was close to quantitative, whereas ADF content and seed colour were qualitative. The GSL content was significantly lower in the first year of the study. The protein content tended to be lower in the first year as well, but the difference was very small (less than 1 percentage point on average). In addition, in the first year of the study, seeds were slightly darker than in the second year.
The profiles of main seed compounds in each individual DH line, based on the two-year means, were shown in Figure 3. Mean values for all monitored traits for individual genotypes and both years were presented in Supplementary Table S1.

3.2. Correlation between Fat, Protein, NDF, ADF and GSL Content and Seed Colour

The results of our two-year investigation revealed a weak but statistically significant negative correlation between seed colour and content of fat (r = −0.34 **), and a strong negative correlation between seed colour and NDF (r = −0.85 **), as well as between seed colour and ADF (r = −0.91 **), which means that seeds of a lighter colour have both a lower fat content and a lower fibre content (Table 4). A weak, but significant positive correlation was noticed between fat and NDF content (r = 0.35 **), and fat and ADF content (r = 0.32 **), while protein content was negatively correlated with NDF (r = −0.33 **) and with ADF (r = −0.27 **) content. However, a significant correlation between protein and fat content was not observed (r = −0.12). This last result suggests that it should be possible to develop genotypes with simultaneously increased protein and fat content in further breeding work.

3.3. Transgressive Effects

DH lines were also examined for positive transgressive effects for fat and protein content and negative transgressive effects for NDF, ADF, and GSL content (Table 5). Among 44 analysed genotypes, 10 DH lines revealed positive transgressive effects for fat content, and 2 DH lines showed such effects for protein content. We found three DH lines with negative transgression for NDF content and one DH line for ADF content. However, for GSL content, no transgressive lines were detected. Among the transgressive lines in terms of fat content, six lines showed black seed colour, one line was yellow-brown and three lines were yellow seeded. Transgressive effects for protein content occurred only in yellow-seeded lines. In the DH population, three lines with yellow seeds were particularly distinctive: line DH-20, which showed favourable transgressive effects for protein, NDF and ADF content; line DH-7, with favourable transgressive effects for protein and NDF content; and line DH-16, with significantly increased fat content and significantly reduced NDF content.

3.4. Selection of the Best Yellow-Seeded DH Lines

Each DH line with low fibre content, i.e., not significantly higher than the fibre content in seeds of parental line Z-114, was compared with this parental line for the studied traits (Table 6). Among 17 selected DH lines, five lines were notable: DH-16, DH-22, DH-30, DH-33 and DH-37. These lines were characterized by significantly higher fat content and protein content (p < 0.01) than the yellow-seeded parental line. Two lines, DH-16 and DH-22, were among five lines with the highest sum of fat and protein contents (70%) in the studied population (Figure 3). Moreover, line DH-16 had significantly reduced NDF content (p < 0.05) compared to parental line Z-114. These five lines also had a significantly higher GSL content than the Z-114 line, but the GSL level in seeds was still within the standard range. The lines DH-7 and DH-20 are also worth mentioning, as they had the highest protein content and significantly reduced fibre content.

3.5. Hierarchical Cluster Analysis

Based on the Mahalanobis distance of the DH lines, a dendrogram was created of similarities between DH lines and parental lines with respect to the quality traits and seed colour (Figure 4). The cluster analysis showed that the genotypes could be divided into two main groups: a group comprising 16 black-seeded lines and a group of 30 lines with yellow, yellow-brown and brown seed colour.

4. Discussion

The development of yellow-seeded cultivars of Brassica napus is especially desirable because of their improved seed meal quality [24]. Yellow-seeded cultivars have a thinner seed coat, higher protein and lower fibre content than black-seeded cultivars, resulting in a higher fat content in the seed [7,25]. Breeding of yellow-seeded oilseed rape is one way to reduce the fibre level in oilseed rape meal, and this helps to increase its digestibility and to improve energy intake from the feed [26,27,28].
The cross between the black-seeded H2-26 line and the yellow-seeded Z-114 line aimed to produce high-yielding genotypes [23] with low fibre content, the cake of which would be a valuable animal feed. Additionally, analyses of the protein, NDF, ADF and GSL contents, as well as the assessment of seed colour, allowed the variability in these traits and the relationships between them to be determined. Seeds of all genotypes collected in the first year tended to be darker than the seeds coming from the second year, with the exception of DH-1 and DH-7 lines. It was a reaction to heavy rainfall during the last stage of seed maturation, with lower sunlight and temperature. Seed colour is not a selective trait itself, but the light seed colour that results from a very thin, transparent seed coat may be a determinant and phenotypic marker of low fibre content. Relatively high values of coefficients of variation for the content of both fractions of dietary fibre, especially ADF, show high genetic variability for these traits among DH lines. In contrast, low coefficients of variation for parental lines and high coefficients of heritability for all studied genotypes indicate a high level of genetic variation for both these traits. The substantial efforts of breeders to reduce the fibre content in oilseed rape seeds aim to increase the digestibility of feeds obtained from oilseed cake or meal. According to many authors, high fibre content in oilseed rape meal can reduce energy and protein availability in monogastric animals [7,29,30,31]. In the DH population studied here, three lines were found with a negative transgression for NDF content. One of these lines also contained significantly less ADF and significantly more protein than the best parent. It is interesting that the low-fibre DH lines identified in this study tended to have a significantly increased protein content, with no corresponding effect on fat content. The same tendency was noticed by Wittkop [32] and Liu [33], who observed the relationship between ADL (acid detergent lignine) and protein contents. According to these authors, the tendency for a higher level of protein in lines with low ADL, without a corresponding increase in fat content suggests a direct biochemical relationship between the seed coat and the biosynthesis of storage proteins in the embryo. The consequence of the low fibre content is a reduction in the content of fibre-related tannins and polyphenols, as well as the yellow colour of seeds, or rather the colour of the cotyledons visible under the transparent seed coat. The correlation coefficients between the colour of seeds and the content of NDF and ADF confirm a very strong relationship between these features, which is explained by some authors to be due to their genetic control being shared in part [34].
The protein content in seeds was strongly influenced by the environment and genotype-environment interaction, but a genetic factor also had a significant influence on this trait. The variability among DH lines was low and slightly exceeded the variability of the parental lines in both years of the experiment, and the heritability rate was relatively high. It is worth noting that the yellow-seeded Z-114 parent line has a significantly lower protein content than the black-seeded H2-26 parental line, with similar fat content in both lines. However, transgressive effects for protein content occurred only in the yellow-seeded DH lines. Similar results were obtained by Wolko et al. [35], indicating that selection for a lighter colour could lead to higher protein content in seeds.
Cultivars with different protein content can be found among black-seeded oilseed rape, although the genetic variability within this trait is small. Among the 15 cultivars of winter oilseed rape assessed by Kowalska et al. [36], the protein content in seeds ranged from 17.7 to 20.1% in the first year, 18.8 to 23.1% in the second year, and 21.2 to 23.3% in the third year, respectively. Similar results were obtained by Marjanović-Jeromela et al. [37], who assessed 30 genotypes of different geographic origins. Against this background, the variability we observed within the DH lines seems high (18.1–26.4% in the first year and 20.0–26.6% in the second year of the study). Higher variability for seed protein content was observed by Chao et al. [38] and Wolko et al. [35] in mapping populations.
The lowest coefficient of variation was found for the fat content. In the studied population of DH lines, this was mainly due to genetic variation, which was confirmed by the high value of the heritability coefficient. A high degree of heritability in two very numerous populations of DH lines was also observed by Delourme et al. [39], who conducted research on the genetic determinants of fat content in black-seeded oilseed rape. However, the generated variability in the analysed population turned out to be sufficient to find 10 segregants with positive transgressive effects based on two years of experience. Furthermore, although fat content was positively correlated with dark seed colour, three transgressive lines had a light seed colour. The positive significant correlation between dark seed colour and seed fat content did not confirm many reports of increased fat content in yellow-seeded oilseed rapes [7,25,40]. Many reports also indicate the additive effect of genes as the main factor controlling the fat content in seeds [39,41,42,43,44]. According to Zhao et al. [45], crossing appropriately selected varieties and the derivation of DH lines allows the breeding of genotypes with a higher fat content than the parents by combining the positive alleles of genes derived from different gene pools. Nevertheless, Wang et al. [46] believe that the inheritance of seed fat content is explained by a more complex model, where additive and dominant effects play major roles, but there is also a small epistatic effect.
The correlation between protein and fat content in our sample turned out to be negative but insignificant, which contrasts with many reports of a very strong negative correlation between these features [38,41,47,48]. This indicates the possibility of finding desirable “correlation breakers” for these traits. Progress in breeding is largely dependent on breaking unfavorable linkage between genes by recombination through appropriate selection of parental components.
GSL content turned out to be the most variable of the biochemical traits. The coefficients of variation for genetically stable parental forms were high, and there were significant differences in the content of these compounds in both years of the study. Of all the quality traits, GSL content was characterized by the lowest heritability. Apart from the genetic factors, the content of GSL is determined by the availability of sulfur in soil and moisture conditions. Under conditions of rainfall deficiency, sulfur content increases significantly [49]. Jensen et al. [50] indicated that a higher increase of GSL content is due to the drought conditions in the period of vegetative development than during the pod development. Low rainfall in April in the second year of the study might result in a higher GSL content in the seeds. The critical importance of rainfall in April was pointed out by Wójtowicz [51], who found that the content of GSL increased under low rainfall conditions before and during the initial flowering period. For GSL content, no line with negative transgressive effects was found. This is not favorable, because alkene GSLs, especially progoitrin, are anti-nutritional substances and their levels in seeds need to be reduced to further improve the biological value of the meal. On the other hand, breeding for reduced seed GSL content could have a negative impact on seed yield and resistance to pests and diseases [52].
The study of this population was previously described in terms of yield and its components [23]. Four yellow-seeded lines gave significantly higher yields than the parental line Z-114, including the DH-16 and DH-30 lines, which were characterized by low fibre content and increased fat and protein content.
Breeding programmes for many crop species involve simultaneous improvement of a number of traits, usually taking into account both productivity and yield quality [24,53,54]. The plant materials are evaluated for multiple traits from the initial stages of breeding. Cluster analysis on the basis of a multidimensional evaluation of genotypes enables them to be comprehensively assessed for practical use in plant breeding. However, a multi-trait evaluation of our DH lines for seed quality and colour simultaneously did not identify the best genotypes. Seed colour had the most discriminating power to distinguish cluster groups, which means that the other traits had much less variation.

5. Conclusions

Crossing two DH lines of different origin, line H2-26 and line Z-114, followed by in vitro androgenesis, resulted in a population of DH lines characterized by a significant range of variability. Although the described population was relatively small, the range of variability of the obtained DH lines exceeded the range of variability of the parental forms. The calculated coefficients of variation for the traits studied in the DH population took into account the overall variability, while the coefficients of variation of the parental lines were indicative of non-heritable variability. Our data show that by choosing appropriate parental lines it is possible to break the negative correlation between protein content and fat content in seeds.
The high level of heritability of traits relating to seed quality suggests the possibility of improving breeding materials through effective selection based on phenotype. It seems that the selected DH-16 and DH-30 lines can be the starting material for breeding cultivars with reduced fibre content and increased fat and protein content.

Supplementary Materials

The following supporting information can be downloaded at: https://www.mdpi.com/article/10.3390/agriculture12030340/s1. Table S1: Mean values of monitored traits of 44 DH lines and parental lines for individual years and both years together.

Author Contributions

Conceptualization, L.S.; methodology, Z.K.; software, Z.K.; validation, L.S. and T.C.-T.; formal analysis, Z.K.; investigation, L.S.; resources, T.C.-T.; data curation, L.S. and M.M.; writing—original draft preparation, L.S.; writing—review and editing, L.S., Z.K., M.M. and T.C.-T.; visualization, M.M.; supervision, T.C.-T.; project administration, T.C.-T. All authors have read and agreed to the published version of the manuscript.

Funding

This research received no external funding.

Conflicts of Interest

The authors declare no conflict of interest.

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Agriculture 12 00340 g001 550
Figure 1. Seed colour on a scale from 0 (black) to 5 (yellow).
Figure 1. Seed colour on a scale from 0 (black) to 5 (yellow).
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Figure 2. Frequency distribution of oilseed rape seed biochemical traits and seed colour for two years in the population of DH lines (no parental lines included). The (A) fat, (B) protein, (C), neutral detergent fibre—NDF (D) acid detergent fibre—ADF, (E) glucosinolates—GSL content, and (F) scaled seed colour values were shown.
Figure 2. Frequency distribution of oilseed rape seed biochemical traits and seed colour for two years in the population of DH lines (no parental lines included). The (A) fat, (B) protein, (C), neutral detergent fibre—NDF (D) acid detergent fibre—ADF, (E) glucosinolates—GSL content, and (F) scaled seed colour values were shown.
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Figure 3. The profiles of main seed compounds in each individual DH line and two parental oilseed rape lines included in the study. The population mean was also shown.
Figure 3. The profiles of main seed compounds in each individual DH line and two parental oilseed rape lines included in the study. The population mean was also shown.
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Figure 4. Clustering dendrogram obtained on the basis of six studied traits.
Figure 4. Clustering dendrogram obtained on the basis of six studied traits.
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Table
Table 1. Meteorological data for seasons of experiments (mean value).
Table 1. Meteorological data for seasons of experiments (mean value).
MonthTemperature (°C)Precipitation (mm)
2017201820172018
March3.94.145.432.1
April11.29.459.219.2
May13.812.762.5138.5
June15.717.972.850.3
July19.522.0115.263.7
Mean temperature12.813.2--
Sum of precipitation--355.1303.8
Table
Table 2. Two-factor analysis of variance of genotypes (DH lines and parents) and environments (years) for six traits of interest.
Table 2. Two-factor analysis of variance of genotypes (DH lines and parents) and environments (years) for six traits of interest.
Source of VariationDegree of FreedomMean Squares for Content ofColour of Seeds
FatProteinNDFADFGSL
Genotypes451755.00 **8.76 **44.10 **49.22 **32.55 **13.93 **
Environments (years)118.81 **39.77 **5.41158.28 **2639.40 **39.78 **
Interaction G × E451.92 **1.82 **1.831.37 **0.49 **0.61 **
Error2761.201.191.460.492.810.12
** —significant at α = 0.01 level.
Table
Table 3. Characteristics of trait variability in the DH population and parental lines.
Table 3. Characteristics of trait variability in the DH population and parental lines.
TraitGroup2016/20172017/2018Broad-
Sense Heritability
RangeMeanCV (%)RangeMeanCV (%)
Fat content
(%)		DH41.0–48.544.53.338.5–51.845.04.80.89
H2-2643.7–44.544.20.944.3–47.145.42.8
Z-11443.9–44.544.10.743.6–44.843.91.3
Protein content
(% in dry matter)		DH17.5–26.422.57.418.9–26.623.16.00.79
H2-2622.4–26.024.15.423.0–24.723.74.1
Z-11418.8–22.120.46.121.1–22.221.72.1
NDF content
(% in dry matter)		DH17.6–27.422.211.517.6–27.821.911.00.93
H2-2622.0–25.724.75.122.9–24.623.83.5
Z-11418.9–21.120.14.219.9–20.520.21.2
ADF content
(% in dry matter)		DH9.1–18.713.120.58.4–16.611.718.10.97
H2-2615.4–16.916.13.313.6–14.714.13.5
Z-11410.3–11.510.84.09.9–10.310.11.8
GSL content (μmol g−1 of seeds)DH4.5–17.59.417.37.1–20.112.722.10.69
H2-266.6–9.18.013.612.3–13.212.83.7
Z-1144.5–7.36.313.77.3–8.98.48.6
Seed colourDH0.2–5.01.964.50.5–5.42.653.90.96
H2-260.4–0.60.410.60.7–0.80.86.6
Z-1144.0–5.04.49.14.6–4.74.61.1
Table
Table 4. Correlation coefficients for the six traits in winter oilseed rape DH lines.
Table 4. Correlation coefficients for the six traits in winter oilseed rape DH lines.
Trait123456
1. Fat content1
2. Protein content−0.121
3. NDF content0.35 **−0.33 **1
4. ADF content0.32 **−0.27 **0.91 **1
5. GLS content0.010.080.04−0.181
6. Colour of seeds−0.34 **0.38 **−0.85 **−0.91 **0.121
**—significant at α = 0.01 level.
Table
Table 5. DH lines with transgressive effects.
Table 5. DH lines with transgressive effects.
TraitDH LineMean
(%)		Effect of TransgressionColour of Seeds
EstimateF-Test
Fat contentDH-146.201.426.79 **0.60
DH-1548.513.7446.72 **0.53
DH-1646.511.7410.10 **3.93
DH-1745.891.114.14 *0.50
DH-2245.901.124.23 *2.65
DH-3046.491.719.81 **3.67
DH-3947.212.4419.87 **3.33
DH-4247.712.9428.86 **0.64
DH-4347.072.3017.69 **0.61
DH-4446.391.618.70 **0.67
Protein contentDH-725.351.7610.44 **3.78
DH-2025.201.618.76 **3.15
NDF contentDH-718.84−1.314.70 *3.78
DH-1618.89−1.264.38 *3.93
DH-2018.78−1.375.13 *3.15
ADF contentDH-209.57−0.876.13 *3.15
*—significant at α = 0.05 level; **—significant at α = 0.01 level.
Table
Table 6. Estimates and testing results of contrasts between chosen DH lines with low fibre content and yellow-seeded parental DH line Z-114.
Table 6. Estimates and testing results of contrasts between chosen DH lines with low fibre content and yellow-seeded parental DH line Z-114.
DH LineContent ofColour of Seeds
FatProteinNDFADFGLS
DH-2−0.111.93 **0.250.143.79 **−1.42 **
DH-7−2.17 **4.30 **−1.31 *−0.613.92 **−0.74 **
DH-8−2.22 **3.16 **−0.370.255.42 **−1.79 **
DH-9−0.761.20 **0.400.195.64 **−1.79 **
DH-162.47 **3.27 **−1.26 *−0.553.17 **−0.59 **
DH-191.16 *0.460.21−0.213.27 **−0.34 **
DH-20−0.764.15 **−1.37 *−0.87 *3.06 **−1.37 **
DH-221.86 **3.04 **−0.33−0.226.65 **−1.87 **
DH-251.021.76 **0.790.303.06 **−1.80 **
DH-260.612.67 **−1.18−0.512.91 **−0.19
DH-27−0.012.05 **−0.250.296.14 **−1.57 **
DH-281.052.18 **0.51−0.082.01 *−1.39 **
DH-302.45 **1.63 **0.330.031.70 *−0.85 **
DH-331.39 *1.68 **−0.010.143.44 **−1.42 **
DH-35−0.441.48 **1.070.72 *6.79 **−1.09 **
DH-371.55 **2.46 **−0.58−0.214.48 **−1.42 **
DH-401.011.58 **0.250.263.18 **−1.19 **
Z-11444.0421.0520.1510.447.334.52
*—significant at α = 0.05 level; **—significant at α = 0.01 level.
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Szała, L.; Kaczmarek, Z.; Matuszczak, M.; Cegielska-Taras, T. Genetic Variability in a Population of Oilseed Rape DH Lines Developed from F1 Hybrids of a Cross between Black- and Yellow-Seeded DH Lines. II. Seed Quality. Agriculture 2022, 12, 340. https://doi.org/10.3390/agriculture12030340

AMA Style

Szała L, Kaczmarek Z, Matuszczak M, Cegielska-Taras T. Genetic Variability in a Population of Oilseed Rape DH Lines Developed from F1 Hybrids of a Cross between Black- and Yellow-Seeded DH Lines. II. Seed Quality. Agriculture. 2022; 12(3):340. https://doi.org/10.3390/agriculture12030340

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Szała, Laurencja, Zygmunt Kaczmarek, Marcin Matuszczak, and Teresa Cegielska-Taras. 2022. "Genetic Variability in a Population of Oilseed Rape DH Lines Developed from F1 Hybrids of a Cross between Black- and Yellow-Seeded DH Lines. II. Seed Quality" Agriculture 12, no. 3: 340. https://doi.org/10.3390/agriculture12030340

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