Morphological characterization and genetic diversity analysis of yield and yield contributing parameters in brinjal (Solanum melongena L.) genotypes

doi:10.21203/rs.3.rs-3744782/v1

A field experiment was carried out to assess the significance of variance, heritability, genetic advance, correlation, path coefficient and genetic diversity for fifteen yield and yield contributing features in twenty brinjal genotypes. The study was conducted at Sher-e-Bangla Agricultural University in Dhaka from November 2021 to June 2022 using a Randomized Complete Block Design (RCBD) with three replications. In this experiment, the existing genotypes showed significant differences for all characters. The phenotypic co-efficient of variation was higher than the genotypic co-efficient of variation for all the characters. Highest PCV (68.60) was found in percent of BSFB infestation and GCV (59.50) was found in percent of BSFB infestation. High heritability associated with high genetic advance and high genetic advance in percentage of mean was found in plant height, individual fruit weight and percent of BSFB infestation, which indicated selection on the basis of phenotype would be effective. Assessments of character association indicating that fruit yield per plant had highest significant positive correlation with days of last harvesting (r_g = 0.495, r_p = 0.454), number of secondary branches per plant (r_g=0.356, r_p=0.315), leaf blade length (r_g = 0.714, r_p = 0.462), leaf blade width (r_g = 0.674, r_p = 0.498) and number of fruits per plant (r_g= 0.798, r_p= 0.787) in both genotypic and phenotypic level. Path analysis revealed that the number of flowers per plant (1.421) had the maximum positive and direct effect on yield per plant. According to PCA, D² and Cluster analysis, twenty brinjal genotypes were grouped into were grouped into five different clusters. Cluster I, II, III, IV and V composed of three, three, four, six and four genotypes respectively. The maximum intra cluster distance was exhibited by cluster III (1.918), whereas the highest value for inter cluster distance was recorded between cluster II and V (95.69). Mean performance of cluster revealed the highest number of fruits per plant (13.45) and yield per plant (0.71) value in cluster IV. Therefore, considering cluster distances, contribution of characters towards divergence, magnitude of cluster mean performance, genetic parameters and based on the objectives, the genotypes G10 (Choto Katali) and G20 (BARI Bt Begun 2) from cluster III, G11 (BARI Hybrid Begun 2) and G8 (BARI Begun 6) from cluster II, G5 (BARI Hybrid Begun 4) along with G18 (Local Black Long) from cluster IV might be used in future breeding program.

Brinjal

BSFB

Characterization

Diversity

Vegetables are one of the leading items of our daily diet. Brinjal is an important solanaceous vegetable crop of our country. Its scientific name is Solanum melongena L. having chromosome number 2n = 2x = 24. It is considered as a native to India (Taher et al., 2017, De Candolle, 1886). Brinjal is usually a self-pollinated crop, but the extent of cross-pollination has been reported as high as 48% and hence it is classified as often cross-pollinated crop (Agrawal, 1980). In 2020, the estimated world total production of brinjal was 56,618,843 metric tons, up by 2.2% from 55,376,521 metric tons in 2019. China ranked first (36,557,611 metric tons; 65% of world's total). Asia produce maximum (more than 90%) brinjal in terms of production (FAOSTAT, 2021).

In Bangladesh, it is cultivated in 53,642 ha area throughout the year with annual production 5,87,212 metric tons in the year 2020–2021 (BBS, 2022). Brinjal is a warm season vegetable crop mostly cultivated in tropical and subtropical regions of the world (Taher et al., 2017). It is one of the most common, popular and principal vegetable crops grown throughout the country except higher altitudes. It is a versatile crop adapted to different agro-climatic regions and can be grown throughout the year. A number of cultivars are grown in our country. The varieties show a wide range of fruit shapes and colors. Consumers preference being dependent upon fruit color, size and shape. In Bangladesh, to meet up the market demand, brinjal is widely cultivated during both the summer and winter seasons. During summer season, it is cultivated in 49,274 acres with 2,02,371 M tons and in winter season 83,277 acres with 3,84,841 M tons (BBS 2022).

BARI released 20 varieties of brinjal including two hybrids and four transgenic varieties for cultivation in Bangladesh. The fruit and shoot borer resistant, trangeneic varieties are ‘BARI Bt Bagun-1’, ‘BARI Bt Bagun-2’, ‘BARI Bt Bagun-3’ and ‘BARI Bt Bagun-4’ (Azad et al., 2017). Different seed companies released more than 60 varieties including hybrid (SCA, 2018). In addition, a lot of landraces are cultivated all over the country. Brinjal is considered as most popular vegetable with high nutritional value and lots of health benefits. Fresh brinjal has 92% water, 6% carbohydrates, 1% protein, a negligible amount of fat, and low amounts of essential nutrients (San José et al., 2014). It is highly beneficial for the regulation of blood sugar levels which helps to control the absorption of glucose (Saha et al., 2019) and its fruits are used to cure diabetes (Verma et al., 2018). Brinjal extracts have superb healing effects on different disorders like burns, warts, inflammatory infections, gastritis, stomatitis and arthritis (Im et al., 2016). Fiber contents present in eggplant helps in digestion by removing toxins and harmful materials from our stomach thus by reducing stomach and colon cancer (Fraikue, 2016). It is a rich source of magnesium, manganese, potassium, and copper that are important for health bones. The fruits are important in the treatment of various disorders like asthma, dysuria, dysentery, high blood pressure, and also to cure osteoporosis, arthritis, diabetes and bronchitis, heart diseases and strokes (Sekara et al., 2007., Seneff et al., 2011). The extracts of brinjal root and leaves can cure problems such as skin diseases, cough, toothache, piles, inflammation, throat problems, and stomach problems (Barik et al., 2020).

Now-a-days brinjal yield is much satisfactory but its production is greatly hampered due to different insect-pest, they play a central role for lowering the yield of brinjal by attacking them from the nursery stage to harvesting (Borkakati, 2019). More than 70 number insect species attack the brinjal (Subbarathnam, 1982). More than 80% yield loss occur due to BSFB (Brinjal Shoot and Fruit Borer) attack (Mishra, 2008; Jagginavar et al., 2009). It feeds internally on the fruit and excrete inside the fruit resulting worthless for human consumption (Baral et al., 2006). Brinjal production is also massively affected by several fungal and bacterial diseases (Singh et al., 2014). Most common diseases for brinjal includes: bacterial wilt, verticillium wilt, fusarium wilt, fruit rot, damping off, Phytophthora blight, Phomopsis blight and fruit rot, leaf spot, little leaf of brinjal, and mosaic (Taher et al., 2017). To diminish extensive yield loss, the farmers are applying many hazardous chemicals incessantly which will increase the production cost in addition to environmental contamination, human health hazard (Dadmal et al., 2004) and leads to insecticide resistance in insect-pests (Harish et al., 2011). To meet up the demand of faster growing population with limited availability of cultivated land it is immensely necessary to improve our cultivated brinjal varieties. It is also important to identify the natural mechanisms prevailing in the brinjal land races to utilize them in the improvement program. Brinjal improvement programs need to focus on developing higher yield potential varieties, better fruit quality, good shelf-life, resistance to major insect, pest and diseases with greater adaptation to different stress conditions (Daunay and Hazra, 2012). Brinjal improvement program largely depends upon the nature and magnitude of the genetic variability existing in breeding material as well as the parental selections in set hybridization programs (Taher et al., 2017). Higher the diversity of the parents, better the chances of improving economic traits under consideration in the offspring. To get suitable genotypes for higher yield or any other desirable characters, collection of germplasm and its genetic analysis plays a crucial role. So, it is fundamental need to pursue the diversity of local cultivars and improve the yield potential of existing genotypes through suitable breeding program. Therefore, this research was performed to characterize as well as to assess the genetic diversity among different existing genotypes, to screen different BSFB tolerant varieties under field conditions and to identify the best genotypes for the varietal improvement of brinjal in Bangladesh.

The experiment was conducted during the period from 03 November 2021 to 26 June 2022 (rabi season) at the Farm of Sher-e-Bangla Agricultural University, Dhaka-1207, Bangladesh. The soil of the research plot had the general shallow red-brown terrace soils. The texture the topsoil was sandy loam. The experiment was carried out in Randomized Complete Block Design (RCBD) with three replications.

The brinjal genotypes were distributed randomly in each block. Five plants were planted for each genotype in every single row. Each replication contains 100 plants. The spacing was 60 cm × 90 cm (plant to plant distance 60 cm and row to row 90 cm). Block to block distance was 1 m.

Twenty brinjal genotypes were used for this experiment. The genetically pure seeds of twenty genotypes were collected from Bangladesh Agricultural Research Institute (BARI), Department of Genetics and Plant Breeding and the local market of Dinajpur.

Brinjal seedlings were raised in seedbed. 19 days old healthy and uniform brinjal seedlings were uprooted separately from seedbed and transferred to polybags due to sudden rainfall and fungal contamination. 40 days old healthy and vigorous seedlings were transplanted in the main field. To provide vigorous growth and proper development of the brinjal plants fertilization and intercultural operations including weeding, irrigation, stalking, earthing up etc. were done as per requirement. Fruits were harvested on the basis of horticultural maturity, size, color, and age. Frequent picking was done throughout the harvesting period.

From every blocks, three plants were chosen randomly for each genotype and tagged properly for recording data. At different growth stages, data were recorded on days to first flowering, days to 50% flowering, days to first fruit harvest, days to last fruit harvest, fruit length, fruit diameter, number of primary branches per plant, number of secondary branches per plant, number of fruits per plant, leaf blade length, leaf blade width, plant height, individual fruit weight BSFB infestation percentage and yield per plant. The obtained data from twenty genotypes for aforementioned characters were statistically analyzed to observe the genotype of brinjal in relation to yield and yield contributing characters. The analysis of variance (ANOVA), the mean values of all the characters and the significant difference among the treatment means were calculated by using R package (variability, agricolae). The generalized distance method of Mahalanobis (1936) was used to determine genetic diversity (D²) using GENSTAT software.

Variability, heritability and genetic advance:

The results of analysis of variance (ANOVA) showed that the brinjal genotypes varied significantly for all the studied characters (Table 1), revealing that significant variations were seen

Table 1

Analysis of variance of fifteen characters of twenty genotypes of brinjal
Source of Variation	df	Mean sum of square
Source of Variation	df	DFF	DFPF	DFH	DLH	PH	NPBP	NSBP	LBL	LBW	FW	FL	FD	NFPP	BSFBI	YPP
Replication	2	71.47	32.85	3.35	5.60	220.37	1.07	56.11	12.40	2.04	41.92	0.02	0.12	6.22	101.68	0.02
Genotype	19	327**	243.93**	207.42**	539.87**	575.93**	5.14**	49.28*	13.76**	14.08**	1363.24**	51.86**	7.73**	70.77**	1232.14**	0.19**
Error	38	12.26	12.38	11.82	23.06	55.00	1.71	25.72	3.65	2.68	51.20	1.28	0.12	3.84	121.90	0.01
df = Degrees of freedom, ** = 1% level of significance, * = 5% level of significance

Table 2

Estimation of genotypic and phenotypic variation, genotypic and phenotypic co-efficient of variations, heritability, genetic advance and genetic advance percentage over mean for fifteen characters of brinjal
Characters	σ²_g	σ²_p	GCV	PCV	h²b	GA	GAPM
DFF	104.92	117.17	9.66	10.21	90	19.97	18.82
DFPF	77.19	89.56	7.81	8.42	86	16.80	14.94
DFH	65.20	77.02	6.14	6.67	85	15.30	11.64
DLH	172.27	195.33	6.63	7.06	88	25.39	12.82
PH	173.64	228.64	17.51	20.09	76	23.66	31.43
NPBP	1.15	2.85	12.76	20.13	40	1.40	16.65
NSBP	7.86	33.57	17.37	35.92	23	2.79	17.31
LBL	3.37	7.03	10.08	14.55	48	2.62	14.38
LBW	3.80	6.48	15.34	20.03	59	3.08	24.20
FW	437.35	488.54	31.49	33.28	90	40.76	61.37
FL	16.86	18.14	29.79	30.90	93	8.16	59.16
FD	2.54	2.65	37.69	38.54	96	3.21	75.94
NFPP	22.31	26.15	46.36	50.19	85	8.99	88.20
BFSBI	370.08	491.98	59.50	68.60	75	34.37	106.30
YPP	0.06	0.07	48.38	53.67	81	0.46	89.85

(σ²_g - Genotypic variation, σ²_p - Phenotypic variation, GCV- Genotypic co-efficient of variation, PCV- Phenotypic co-efficient of variation, h²b- Heritability percentage in broad sense, GA- Genetic advance, GAPM- Genetic advance percentage over mean), DFPF- Days to first flowering, DFPF- Days to 50% flowering, DFH- Days to first harvesting, DLH- Days to last harvesting, PH- Plant height (cm), NPBP- No. of primary branches per plant, NSBP- No. of secondary branches per plant, LBL- Leaf blade length (cm), LBW- Leaf blade width (cm), FL- Fruit length (cm), FD- Fruit diameter (cm), FW- Individual fruit weight (g), NFPP- No. of fruits per plant, BSFB- Percent of BSFB infestation, YPP- Yield

among all genotypes for all investigated parameters. The significant differences among the genotypes indicated the presence of genetic variability in the material used and provide a good opportunity for improvement. For all of the studied attributes, the existence of genetic variability was investigated using heritability, genetic advance, genetic advance in percentage of mean, genotypic co-efficient of variation, and phenotypic co-efficient of variation (Table 2).

For all the characters under study, the phenotypic variance was higher than the corresponding genotypic variance, indicating a greater influence of the environment on these characters' expression. Characters like, individual fruit weight, fruit length, fruit diameter, number of fruits per plant, percent of BSFB infestation and yield per plant exhibited high genotypic and phenotypic co-efficient of variation. Plant height, number of primary branches per plant and number of secondary branches per plant exhibited high phenotypic co-efficient of variation. The phenotypic co-efficient of variation was higher than the genotypic co-efficient of variation for all the characters. Maximum difference between phenotypic and genotypic coefficient of variation were 35.92 and 17.37, respectively which indicated that the number of secondary branches per plant was mostly depended on the environmental condition. Highest phenotypic co-efficient of variation (68.60) was found in percent of BSFB infestation and genotypic co-efficient of variation (59.50) was found in percent of BSFB infestation. High heritability associated with high genetic advance and high genetic advance in percentage of mean was found in plant height, individual fruit weight and percent of BSFB infestation which indicated that additive gene expression on this character and selection methods can be accomplished for the improvement of these traits. Fruit length, fruit diameter, number of fruits per plant and yield per plant showed high heritability with low genetic advance and high genetic advance in percentage of mean that might be presence of additive and non-additive gene expression. High heritability with high genetic advance and low genetic advance in percentage of mean were found in days to last harvesting. High heritability with low genetic advance and low genetic advance in percentage of mean were found in days to first flowering, days to 50% flowering and days to first harvesting. Moderate heritability with low genetic advance and low genetic advance in percentage of mean were found in number of primary branches per plant, leaf blade length and leaf blade width. Number of secondary branches per plant showed low heritability with low genetic advance and low genetic advance in percentage of mean.

Table 3

Mean performance, range, standard error and coefficient of variation in respect of fifteen characters of twenty brinjal genotypes
Genotypes	DFF	DFPF	DFH	DLH	PH	NPBP	NSBP	LBL	LBW	FW	FL	FD	NFPP	BFSBI	YPP
G1	105.34fg	111ef	128ef	176.33fg	71.23d-f	8.55a-d	12.77c-e	16.9c-e	10ef	50.44ij	14.07de	3.2e	10.22d-g	10.26e-h	0.29f-h
G2	109.67d-f	113de	130d-f	183f	68.46ef	9a-c	17.1a-d	18.56bc	13.37a-c	61.22g-i	14.49d	3.85d	8.88e-g	43.83bc	0.34e-g
G3	109.67d-f	113.67de	129ef	196de	61.25fg	7.44b-d	15.88a-e	18.67a-c	14.03a-c	61.45g-i	14.06de	3.82d	12.78cd	12.31d-h	0.65c
G4	96.34h	106.67f	126f	197cde	71.81d-f	9.11ab	9.88de	18.63a-c	12.1c-e	70.03e-g	9.45g	6.07bc	8.55fg	49.19b	0.43d-g
G5	106.33e-g	111.33ef	128ef	207.67ab	85.68b	9.abc	24.22a	20.76ab	15.27ab	75.58de	10.07g	5.95bc	22.33a	29.06cd	1.10a
G6	106e-g	117.33b-d	128ef	204bc	62.58fg	9.22ab	13.21c-e	16.93c-e	10.17d-f	79.67de	9.62g	6.92a	7.33gh	22.6def	0.44d-g
G7	103.34g	114.67de	129.33d-f	204bc	78.68b-e	8.55a-d	15.49b-e	18.7a-c	13.23a-c	63.07f-h	12.7d-f	3.92d	13.05b-d	20.49d-g	0.53c-e
G8	117.33ab	121.33ab	140b	203b-d	70.15d-f	9.a-c	19.61a-c	17.63b-d	13.7a-c	117.3a	10.57g	6.78a	2.39j	29.96cd	0.38e-g
G9	119a	120.67bc	139b	196de	81.21b-d	6.94cd	12.33c-e	18.43bc	13.57a-c	93.11bc	10.5g	6.07bc	3.83ij	28.33c-e	0.32fg
G10	70.67i	82h	105g	168.33h	39.76h	7.55b-d	17a-d	14.06ef	8.73f	29.42l	9.35g	3.08e	11.33c-f	7.48f-h	0.42d-g
G11	113b-d	115c-e	146a	204bc	99.75a	9.a-c	23.5ab	20.2ab	14.57a-c	101.33b	12.6ef	3.92d	4.78h-j	45.23bc	0.43d-g
G12	108.33d-g	114de	132c-e	204.33bc	84.7bc	9.a-c	19.66a-c	21.73a	14.83ab	74.1d-f	11.09fg	5.52c	13.88bc	48.69b	0.87b
G13	108.67d-g	113.67de	139b	210.33ab	78.75b-e	8.5a-d	15.88a-e	20.53ab	13.9a-c	55.92h-j	19.12bc	2.78e	8.16fg	61.46ab	0.40d-g
G14	105.67e-g	112d-f	128ef	206.67ab	72.90c-f	8.33a-d	15.88a-e	19.6a-c	14.2a-c	84.67cd	10.58g	6.38ab	14.11bc	28.92cd	1.06ab
G15	106e-g	111.67d-f	135b-d	207.67ab	87.05b	9.77a	19.66a-c	19.13a-c	15.67a	53.8h-j	18.52bc	2.78e	16.05b	52.93b	0.66c
G16	111.33c-e	115.67b-e	139b	209ab	80.4b-e	8.78a-d	16.33a-e	18.3bc	13.06a-c	56.11h-j	20.01ab	2.08f	7g-i	72.92a	0.26gh
G17	106.33e-g	114.67de	132cde	213a	87.65ab	9.22ab	15.1c-e	17.96bc	12.8b-d	61.31g-i	19.93ab	2.75e	11.9c-e	50.85b	0.59cd
G18	109.66d-f	114de	132.33c-e	207.67ab	89ab	9.89a	18.1a-d	19.3a-c	13a-c	56.39h-j	17.27c	2.87e	11.11c-f	28.97cd	0.47c-f
G19	92.34h	99.67g	127ef	193e	52.49g	4.11e	12.88c-e	14.8d-f	9.47ef	46.11jk	21.72a	2.88e	13.11b-d	3.2gh	0.39e-g
G20	116.33a-c	127a	137.33bc	170gh	81.7b-d	6.83d	8.11e	13.56f	8.53f	37.33kl	9.98g	2.93e	3j	0h	0.12h
MAX	119.00	127.00	146	213	99.75	9.89	24.22	21.73	15.67	117.3	21.72	6.92	22.33	72.92	1.10
MIN	70.67	82.00	105	168.33	39.76	4.11	8.11	13.567	8.53	29.42	9.35	2.08	2.39	0.00	0.12
MEAN	108.48	112.45	131.50	198.05	75.26	8.39	16.13	18.22	12.71	66.42	13.78	4.23	10.19	32.33	0.51
CV%	3.30	3.06	2.67	2.42	9.85	15.54	31.43	10.48	12.88	10.77	8.18	7.84	19.23	34.15	19.70
SE	2.86	2.87	2.81	3.92	6.06	1.07	4.14	1.56	1.34	5.84	0.92	0.28	1.60	9.01	0.10
LSD	5.79	5.81	5.68	7.94	12.26	2.16	8.38	3.16	2.71	11.83	1.87	0.56	3.24	18.25	0.20

SE- Standard error, CV- Co-efficient of variation, LSD- Least Significant Differences, DFPF- Days to first flowering, DFPF- Days to 50% flowering, DFH- Days to first harvesting, DLH- Days to last harvesting, PH- Plant height (cm), NPBP- No. of primary branches per plant, NSBP- No. of secondary branches per plant, LBL- Leaf blade length (cm), LBW- Leaf blade width (cm), FL- Fruit length (cm), FD- Fruit diameter (cm), FW- Individual fruit weight (g), NFPP- No. of fruits per plant, BSFB- Percent of BSFB infestation, YPP- Yield per plant (kg

According to the mean performance, G10 (70.67 DAS) had the shortest length needed for first flowering, whereas G9 (119 DAS) had the longest. The maximum time was observed in G20 (127 DAS) and the shortest duration needed for 50% flowering was identified in G10 (82 DAS). G10 had the 105 DAS minimum period needed for first fruit harvesting while G11 had the 146 DAS maximum length. The longest period needed for the last fruit harvesting has been found in G17 (213 DAS), and the shortest period was reported in G10 with 168.33 DAS. The G11 plant measured 99.75 cm in height at its maximum, whereas the G10 plant measured 39.76 cm at its shortest. The genotype G18 scored the most primary branches (9.89), while the genotype G19 recorded the fewest (4.11). G5 was found to have the most secondary branches (24.22), whereas the genotype G20 had the fewest (8.11). The genotype G12 had the longest leaf blades (21.73 cm), while genotype G20 had the shortest (13.56 cm). The genotype G15 had the widest leaf blades (15.67 cm), whereas genotype G20 had the narrowest (8.53 cm). The largest fruit weight in G8 was 117.3 g, and the smallest fruit weight in G10 was 29.42 g. The genotype G19 had the longest fruit length at 21.72 cm, while genotype G10 had the shortest fruit length (9.35 cm). The genotype G6 had the largest fruit diameter (6.92 cm), whereas G16 had the smallest (2.08 cm). Genotype G5 produced the most fruits per plant (22.33) and the genotype G8 produced the fewest (2.39). G16 had the highest brinjal shoot and fruit borer infestation (72.92%), whereas the genotype G20 had the lowest infestation (0%). The largest fruit yield per plant 1.10 kg emerged in G5, and the lowest 0.12 kg was found in G20.

Correlation co-efficient

Yield is a complex attribute that is impacted by a number of quantitative factors. Some of these characteristics have a strong relationship to yield. For the purpose of developing selection criteria, it is crucial to analyze how these qualities interact and how they relate to yield. Tables 4 and 5 display the genotypic and phenotypic correlation co-efficient between yield and yield-contributing features of twenty brinjal genotypes. Correlation co-efficient values vary from − 1.0 to 1.0. While a correlation of 1.0 displays a perfect positive connection, one of -1.0 displays a perfect negative correlation. There is no correlation between the movements of the two variables when it is 0. Assessments of character association (Tables 4 & 5) indicating that fruit yield per plant had highest significant positive correlation with days of last harvesting (r_g = 0.495, r_p = 0.454), number of secondary branches per plant (r_g=0.356, r_p=0.315), leaf blade length (r_g = 0.714, r_p = 0.462), leaf blade width (r_g = 0.674, r_p = 0.498) and number of fruits per plant (r_g= 0.798, r_p= 0.787) in both genotypic and phenotypic level indicating the importance of these trait in selection for increasing yield and were identified as yield attributing characters. Thus, selection can be effective upon these characters for the genetic improvement of brinjal yield.

Table 4

Genotypic correlation co-efficient among different pairs of yield and yield contributing characters for twenty genotypes of brinjal
Characters	DFF	DFPF	DFH	DLH	PH	NPBP	NSBP	LBL	LBW	FW	FL	FD	NFPP	BFSBI
DFPF	0.984**
DFH	0.907**	0.868**
DLH	0.389^NS	0.354 ^NS	0.505*
PH	0.736**	0.712**	0.800**	0.580**
NPBP	0.296^NS	0.328 ^NS	0.236 ^NS	0.536*	0.673**
NSBP	0.093^NS	-0.175 ^NS	0.237 ^NS	0.651**	0.480*	0.654**
LBL	0.509*	0.375 ^NS	0.492*	0.877**	0.721**	0.832**	0.884**
LBW	0.561*	0.399 ^NS	0.566**	0.845**	0.694**	0.656**	0.974**	0.956**
FW	0.572**	0.478*	0.555*	0.518*	0.385 ^NS	0.349 ^NS	0.533*	0.593**	0.607**
FL	0.031^NS	-0.053 ^NS	0.260 ^NS	0.354 ^NS	0.139 ^NS	-0.164 ^NS	0.061 ^NS	0.041 ^NS	0.096 ^NS	-0.388 ^NS
FD	0.189^NS	0.220 ^NS	-0.003 ^NS	0.197 ^NS	-0.054 ^NS	0.203 ^NS	0.046 ^NS	0.329 ^NS	0.246 ^NS	0.751**	-0.748**
NFPP	-0.370^NS	-0.407^NS	-0.458*	0.262 ^NS	-0.046 ^NS	0.143 ^NS	0.637**	0.384NS	0.349 ^NS	-0.290 ^NS	0.155 ^NS	-0.044 ^NS
BFSBI	0.278^NS	0.204 ^NS	0.483*	0.709**	0.590**	0.777**	0.514*	0.829**	0.746**	0.280 ^NS	0.378 ^NS	-0.068 ^NS	-0.010 ^NS
YPP	-0.048^NS	-0.103 ^NS	-0.182 ^NS	0.495*	0.170 ^NS	0.366 ^NS	0.885**	0.714**	0.674**	0.249 ^NS	-0.182 ^NS	0.416 ^NS	0.798**	0.147 ^NS
(5% level of significance) * (1% level of significance) NS (Non- significant)
DFPF- Days to first flowering, DFPF- Days to 50% flowering, DFH- Days to first harvesting, DLH- Days to last harvesting, PH- Plant height (cm), NPBP- No. of primary branches per plant, NSBP- No. of secondary branches per plant, LBL- Leaf blade length (cm), LBW- Leaf blade width (cm), FL- Fruit length (cm), FD- Fruit diameter (cm), FW- Individual fruit weight (g), NFPP- No. of fruits per plant, BSFB- Percent of BSFB infestation, YPP- Yield per plant (kg)

Path co-efficient analysis

Path co-efficient analysis is the most commonly used statistical method for determining the direct or indirect effects of yield contributing attributes on yield per plant and measuring the relative significance of each component on yield per plant. Table 6 shows the partitioning of genotypic correlations into direct and indirect effects of significant features using path co-efficient analysis of brinjal. Path analysis showed that highest positive direct effect was number of fruits per plant (1.421) and the lowest positive direct effect was fruit length (0.083). Days to 50% flowering, days to first harvesting, number of primary branches per plant, number of secondary branches per plant, leaf blade width, individual fruit weight, fruit length, fruit diameter, number of fruits per plant and percent of BSFB infestation showed positive direct effect on fruit yield per plant indicating that direct selection based on these traits may be effective in evolving high yielding varieties of brinjal. On the other hand, negative direct effect was found by days to first flowering, days to last harvesting, plant height and leaf blade length. The residual effect (R) of path co-efficient analysis was 0.1033 which reported that the traits under study contributed 89.67% of the fruit yield per plant. It can be said that there were some other traits those contributed 10.33% to the yield per plant that were not included in the present study.

DFPF- Days to first flowering, DFPF- Days to 50% flowering, DFH- Days to first harvesting, DLH- Days to last harvesting, PH- Plant height (cm), NPBP- No. of primary branches per plant, NSBP- No. of secondary branches per plant, LBL- Leaf blade length (cm), LBW- Leaf blade width (cm), FL- Fruit length (cm), FD- Fruit diameter (cm), FW- Individual fruit weight (g), NFPP- No. of fruits per plant, BSFB- Percent of BSFB infestation, YPP- Yield per plant (kg)

Table 5

Phenotypic correlation co-efficient among different pairs of yield and yield contributing characters for twenty genotypes of brinjal
Characters	DFF	DFPF	DFH	DLH	PH	NPBP	NSBP	LBL	LBW	FW	FL	FD	NFPP	BFSBI
DFPF	0.913**
DFH	0.854**	0.748**
DLH	0.345**	0.288*	0.413**
PH	0.611**	0.555**	0.683**	0.496**
NPBP	0.210 ^NS	0.172 ^NS	0.128 ^NS	0.359**	0.358**
NSBP	0.060 ^NS	-0.023 ^NS	0.091 ^NS	0.312*	0.267*	0.328*
LBL	0.285*	0.176 ^NS	0.309*	0.565**	0.504**	0.376**	0.434**
LBW	0.344**	0.227 ^NS	0.353**	0.571**	0.496**	0.395**	0.490**	0.853**
FW	0.489**	0.447**	0.462**	0.444**	0.312*	0.243 ^NS	0.288*	0.389**	0.462**
FL	0.03 ^NS	-0.041 ^NS	0.216 ^NS	0.319*	0.130 ^NS	-0.077 ^NS	0.017 ^NS	0.041 ^NS	0.097 ^NS	-0.325*
FD	0.179 ^NS	0.204 ^NS	-0.002 ^NS	0.180 ^NS	-0.042 ^NS	0.116 ^NS	0.107 ^NS	0.205 ^NS	0.194 ^NS	0.720**	-0.696**
NFPP	-0.373**	-0.377**	-0.421**	0.242 ^NS	0.017 ^NS	0.041 ^NS	0.240 ^NS	0.283*	0.283*	-0.240 ^NS	0.135 ^NS	-0.042 ^NS
BFSBI	0.288*	0.196 ^NS	0.403**	0.553**	0.436**	0.404**	0.227 ^NS	0.462**	0.485**	0.194 ^NS	0.322*	-0.088 ^NS	-0.039 ^NS
YPP	-0.121 ^NS	-0.135 ^NS	-0.205 ^NS	0.454**	0.163 ^NS	0.134 ^NS	0.291*	0.462**	0.479**	0.238 ^NS	-0.166 ^NS	0.370**	0.787**	0.058 ^NS
(5% level of significance) * (1% level of significance) NS (Non- significant)

DFPF- Days to first flowering, DFPF- Days to 50% flowering, DFH- Days to first harvesting, DLH- Days to last harvesting, PH- Plant height (cm), NPBP- No. of primary branches per plant, NSBP- No. of secondary branches per plant, LBL- Leaf blade length (cm), LBW- Leaf blade width (cm), FL- Fruit length (cm), FD- Fruit diameter (cm), FW- Individual fruit weight (g), NFPP- No. of fruits per plant, BSFB- Percent of BSFB infestation, YPP- Yield per plant (kg)

Table 6

Path coefficient analysis showing direct and indirect effects of different characters on yield per plant of brinjal
Characters	DFF	DFPF	DFH	DLH	PH	NPBP	NSBP	LBL	LBW	FW	FL	FD	NFPP	BFSBI	r
DFF	-1.209	0.900	0.952	-0.160	-0.375	0.165	0.010	-0.469	0.321	0.237	0.003	0.069	-0.526	0.035	-0.048^NS
DFPF	-1.190	0.915	0.910	-0.145	-0.362	0.183	-0.018	-0.345	0.228	0.198	-0.004	0.081	-0.578	0.026	-0.103 ^NS
DFH	-1.097	0.794	1.049	-0.207	-0.407	0.131	0.024	-0.453	0.323	0.230	0.022	-0.001	-0.651	0.061	-0.182 ^NS
DLH	-0.471	0.324	0.529	-0.411	-0.295	0.299	0.066	-0.808	0.483	0.215	0.030	0.072	0.372	0.090	0.495*
PH	-0.890	0.651	0.839	-0.238	-0.509	0.375	0.049	-0.664	0.397	0.160	0.012	-0.020	-0.066	0.075	0.170 ^NS
NPBP	-0.358	0.300	0.247	-0.220	-0.343	0.557	0.067	-0.766	0.375	0.145	-0.014	0.074	0.203	0.098	0.366 ^NS
NSBP	-0.113	-0.160	0.248	-0.267	-0.244	0.364	0.102	-0.814	0.557	0.221	0.005	0.017	0.906	0.065	0.885**
LBL	-0.615	0.343	0.516	-0.360	-0.367	0.463	0.090	-0.921	0.546	0.246	0.003	0.121	0.546	0.105	0.714**
LBW	-0.679	0.365	0.594	-0.347	-0.353	0.365	0.100	-0.881	0.571	0.252	0.008	0.090	0.495	0.094	0.674**
FW	-0.692	0.437	0.582	-0.213	-0.196	0.194	0.054	-0.547	0.347	0.414	-0.032	0.276	-0.412	0.035	0.249 ^NS
FL	-0.038	-0.048	0.273	-0.146	-0.071	-0.091	0.006	-0.038	0.055	-0.161	0.083	-0.275	0.220	0.048	-0.182 ^NS
FD	-0.229	0.202	-0.003	-0.081	0.028	0.113	0.005	-0.303	0.141	0.311	-0.062	0.367	-0.063	-0.009	0.416^NS
NFPP	0.448	-0.372	-0.480	-0.108	0.024	0.080	0.065	-0.354	0.199	-0.120	0.013	-0.016	1.421	-0.001	0.798**
BFSBI	-0.336	0.186	0.507	-0.291	-0.301	0.433	0.053	-0.764	0.426	0.116	0.031	-0.025	-0.014	0.127	0.147 ^NS
*Residual: 0.1033, r indicates genotypic correlation coefficient with NS (Non- significant), 5% () and 1% () level of significance

DFPF- Days to first flowering, DFPF- Days to 50% flowering, DFH- Days to first harvesting, DLH- Days to last harvesting, PH- Plant height (cm), NPBP- No. of primary branches per plant, NSBP- No. of secondary branches per plant, LBL- Leaf blade length (cm), LBW- Leaf blade width (cm), FL- Fruit length (cm), FD- Fruit diameter (cm), FW- Individual fruit weight (g), NFPP- No. of fruits per plant, BSFB- Percent of BSFB infestation, YPP- Yield per plant (kg)

Genetic diversity of twenty brinjal genotypes

A two-dimensional scatter diagram utilizing component score 1 as the X-axis and component score 2 as the Y-axis was constructed based on the values of principal component scores 2 and 1 obtained from the principal component analysis. This diagram is shown in Fig. 1. There was clearly a five-group distribution of genotypes positions in the scatter diagram, indicating that there was a high degree of genotypic diversity.

Following components with latent roots greater than the unity contribution of the various morphological characters towards divergence were discussed from the latent vectors of the first two principal components, which were computed using the correlation matrix and genotype scores obtained from the first components. With the accounting for the variation among the genotypes, the principal component analysis produced Eigen values of each Principle component axes. Six of these Eigen values above unity accounted for 93.89% in Table 7 of the total variation and the last nine principal axes accounted for 6.11% of the total variation.

Table 7

Eigen values and percentage of variation in respect of fifteen characters in brinjal
Principal Component Axis	Eigen values	Percent variation	Cumulative percent variation
Days to first flowering	6.15	40.97	40.97
Days to 50% flowering	3.20	21.38	62.35
Days to first harvesting	2.34	15.63	77.98
Days to last harvesting	0.98	6.56	84.54
Plant height (cm)	0.78	5.19	89.73
No. of primary branches per plant	0.62	4.16	93.89
No. of secondary branches per plant	0.31	2.10	95.99
Leaf blade length (cm)	0.23	1.59	97.58
Leaf blade width (cm)	0.12	0.79	98.37
Individual fruit weight (g)	0.08	0.53	98.90
Fruit length (cm)	0.07	0.49	99.39
Fruit diameter (cm)	0.05	0.35	99.74
Number of fruits per plant	0.02	0.15	99.89
Percent of BSFB infestation	0.01	0.08	99.97
Yield per plant (kg)	0.005	0.03	100

According to PCA, D² and Cluster analysis, the genotypes were grouped into five different clusters. Cluster I, II, III, IV and V composed of three, three, four, six and four genotypes respectively (Table 8).

Table 8

Distribution of twenty brinjal genotypes in five different clusters
Clusters	No. of genotypes	Name of genotypes
I	3	G2, G4, G12
II	3	G8, G9, G11
III	4	G1, G10, G19, G20
IV	6	G3, G5, G6, G7, G14, G18
V	4	G13, G15, G16, G17

Plate 1. Leaves from twenty genotypes of brinjal

Plate 2. Flowers from twentygenotypes of brinjal

Plate 4. BSFB infected fruits from sixteen genotypes of brinjal

Table 9

Cluster mean for fifteen characters of twenty genotypes of brinjal
Characters	Clusters
Characters	I	II	III	IV	V
Days to first flowering	104.78	116.44	96.16	106.77	108.08
Days to 50% flowering	111.22	119.00	104.91	113.83	113.91
Days to first harvesting	129.33	141.67	124.33	129.28	136.25
Days to last harvesting	194.78	201.00	176.92	204.33	210.00
Plant height (cm)	74.99	83.71	61.30	75.02	83.46
No. of primary branches per plant	9.03	8.31	6.76	8.74	9.07
No. of secondary branches per plant	15.55	18.48	12.69	17.13	16.75
Leaf blade length (cm)	19.64	18.76	14.83	18.99	18.98
Leaf blade width (cm)	13.43	13.94	9.18	13.32	13.86
Individual fruit weight (g)	68.45	103.91	40.82	70.13	56.78
Fruit length (cm)	11.68	11.22	13.78	12.38	19.40
Fruit diameter (cm)	5.15	5.59	3.02	4.97	2.60
Number of fruits per plant	10.44	3.67	9.41	13.45	10.78
Percent of BSFB infestation	47.23	34.50	5.23	23.72	59.54
Yield per plant (kg)	0.55	0.38	0.30	0.71	0.48

From the clustering mean values, it was observed that cluster I produced the highest mean values for leaf blade length (19.64 cm) in comparison with other five clusters (Table 9). These group possessed genotypes with the second highest cluster mean for no. of primary branches per plant (9.03), fruit diameter (5.15 cm), number of fruits per plant (10.44), percent of BSFB infestation (47.23) and yield per plant (0.55 kg). On the other hand, cluster II produced the highest mean values for days to first flowering (116.44 days), days to 50% flowering (119 days), days to first harvesting (141.67 days), plant height (83.71 cm), no. of secondary branches per plant (18.48), leaf blade width (13.94 cm), individual fruit weight (103.91 g) and fruit diameter (5.59 cm) (Table 9). Cluster III produced the second highest mean value for fruit length (13.78 cm) and the lowest mean value for days to first flowering (96.16 days), days to 50% flowering (104.91 days), days to first harvesting (124.33 days), days to last harvesting (176.92 days), plant height (61.30 cm), no. of primary branches per plant (6.76), no. of secondary branches per plant (12.69), leaf blade length (14.83 cm), leaf blade width (9.18 cm), individual fruit weight (40.82 g), percent of BSFB infestation (5.23) and yield per plant (0.30 kg).

Cluster IV had the highest mean values for number of fruits per plant (13.45) and yield per plant (0.71 kg). These genotypes had the second highest cluster mean for days to last harvesting (204.33 days), secondary branches per plant (17.13), leaf blade length (18.99 cm), and individual fruit weight (70.13 g). Cluster V had the highest mean values for the number of primary branches per plant (9.07), fruit length (19.4 cm), and percentage of BSFB infestation (59.54). Cluster V contains the second highest mean value for days to first flowering (108.08 days), days to 50% flowering (113.91 days), days to first harvesting (136.25 days), plant height (83.46 cm), leaf blade width (13.86 cm) and fruit diameter (2.6 cm) (Table 9).

Table 10

Average intra (bold) and inter cluster distances (D²) for twenty brinjal genotypes
Cluster	I	II	III	IV	V
I	0.819
II	71.42	0.855
III	14.44	72.24	1.918
IV	7.95	64.79	11.94	0.957
V	24.37	95.69	30.1	31.72	0.604

Results showed the maximum inter cluster distance was recorded between cluster II and V (95.69), followed by between II and III (72.24), and II and I (71.42). So, genotypes from these clusters if involved in hybridization might produce a wide range of segregating populations, as genetic variation was very distinct among these groups. The lowest inter cluster distance was found between the cluster I and IV (7.95) followed by III and IV (11.94), I and IV (14.44) representing a close relationship among these clusters. The intra cluster divergence varied from 1.918 from cluster III which comprised of four cultivars of diverse origin to 0.604 from cluster V which comprised four genotypes. Higher inter cluster distances indicated greater genetic variation among the genotypes of different clusters. This study's clustering pattern showed that genotypes gathered from the same locations did not group together into a single cluster (Table 10).

The relative contribution of fifteen quantitative traits to genetic divergence among the twenty germplasms of brinjal is presented in Fig. 2. Among the yield contributing characters, the maximum contribution towards divergence was made by fruit diameter (36.4%) followed by fruit length (16%), days to first flowering (7.5%), days to last harvesting (7.1%), individual fruit weight (6.8%), number to fruits per plant (4.5%), days to 50% flowering (4.4%), days to first harvesting (4.1%), leaf blade width (2.7%), percent of BSFB infestation (2.7%), plant height (1.8%), yield per plant (1.8%), number of primary branches per plant (1.6%), leaf blade length (1.6%), number of secondary branches per plant (1.2%) (Fig. 2).

Significant variations were observed among all examined parameters of brinjal genotypes. for each character. Among all the genotypes, maximum (1.10 kg) yield per plant was found in G5 due to better performances in important yield contributing characters such as number of secondary branches and number of fruits per plant. Phenotypic variance was considerably higher than the genotypic variance for all the examined characters. Characters like, individual fruit weight, fruit length, fruit diameter, number of fruits per plant, percent of BSFB infestation and yield per plant exhibited high genotypic and phenotypic co-efficient of variation. The phenotypic co-efficient of variation was higher than the genotypic co-efficient of variation for all the characters. Maximum differences between the phenotypic and genotypic co-efficient of variation were found in number of secondary branches per plant indicating the more environmental effect to control the characters. High heritability associated with high genetic advance and high genetic advance in percentage of mean was found in plant height, individual fruit weight and percent of BSFB infestation which indicated that additive gene expression on this character, suggesting the high possibility of selecting the genotypes for this traits. Path analysis showed that direct positive effect on fruit yield per plant was observed by days to 50% flowering, days of first harvesting, number of primary branches per plant, number of secondary branches per plant, leaf blade width, fruit length, fruit diameter, number of fruits per plant, individual fruit weight and percent of BSFB infestation. Considering cluster distances, genetic parameters and based on the objectives, the genotypes such as G10 (from cluster III) for early flowering and fruit setting, G11 (from cluster II) for plant height, G5 (from cluster IV) for highest yield per plant, maximum number of fruits per plant and maximum number of the secondary branches per plant, G18 (from cluster IV) for maximum number of primary branches, G8 (from cluster II) for highest individual fruit weight and G20 (from cluster III) for lowest BSFB infestation. Therefore, it may be decided that the genotypes G10, G11, G5, G18, G8 and G20 might be used in future breeding program.

Author Contribution

M.A.Z. carried out the experiments, analyzed the data and wrote the first draft of the manuscript; S.P., M.A.R. and M.H.R. supervised the student, developed the concepts and designed experiments, and edited the manuscript; M.A.I. analyzed the data, and prepared figures, K.M.H. edited the manuscript and M.R.S. helped to carry out the experiment and data collection. All authors reviewed the results and approved the final version of the manuscript.

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Plates are available in the Supplementary Files section.

No competing interests reported.

Plates.docx

Morphological characterization and genetic diversity analysis of yield and yield contributing parameters in brinjal (Solanum melongena L.) genotypes

Status:

Version 1

Abstract

Figures

Introduction

Materials and methods

Results and discussion

Variability, heritability and genetic advance:

Correlation co-efficient

Path co-efficient analysis

Genetic diversity of twenty brinjal genotypes

Conclusion

Declarations

Author Contribution

References

Plates

Additional Declarations

Supplementary Files

Status:

Version 1