Current Plant Biology 16 (2018) 27–31

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Current Plant Biology

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Heterosis and inbreeding depression for fruit yield attributing traits in T

eggplant
⁎
Chintan R. Mistrya, , Keshubhai B. Kathiriaa, Srikanth Sabolua, Sushil Kumarb
a
Department of Plant Breeding & Genetics, Anand Agricultural University, Anand 388 110, India
b
Department of Agricultural Biotechnology, Anand Agricultural University, Anand 388 110, India

A R T I C LE I N FO A B S T R A C T

Keywords: An experiments was performed to study heterobeltiosis and relative heterosis in F1 and inbreeding depression in
Eggplant F2 generation in eggplants. A set of four F1 hybrids were generated by crossing six parents. In all crosses (Doli
Hybrid vigor 5 × GBL1, Doli 5 × KS 331, Pusa Uttam × KS 331, AB-07-02 × GOB 1), fruit length, fruit volume and fruit yield
Heterosis per plant exhibited significantly positive heterobeltiosis which indicates that the hybrid vigor can be utilized on
Inbreeding depression
commercial scale for these traits. The higher magnitude of heterobeltiosis for all the morphological traits sug-
Yield
gested the presence of over-dominance. In F2 generation, negatively significant estimates of inbreeding de-
pression was observed for plant height and days to first picking in cross Doli 5 × KS 331; fruit per plant, fruit
weight and fruit yield per plant in cross Pusa Uttam × KS 331; primary branches per plant, fruit volume and fruit
weight in cross Doli 5 × GBL1 and fruit weight in cross (AB-07-02 × GOB 1) indicated the possibilities to get the
desired segregants to improve the traits. The cross with least inbreeding depression and high heterosis should be
further exploited at commercial scales.

1. Introduction develop a variety suitable for different environment and consumer

Eggplant (Solanum melongena L.), a member of family Solanaceae, is
an important Solanaceous vegetable crop after potato, tomato and
pepper [1]. Eggplant, grown throughout the year, is a common and
popular vegetable crop in the subtropics and tropics regions [2].
Globally, India ranks second in production (27% of world's total) of
eggplant after China (57% of world's total). This warm-weather crop
was probably originated in India. Though it is a self-pollinated crop, but
cross-pollination is reported upto the extent of 30–40% [3]. In Asian
and the Mediterranean countries, eggplant also called brinjal, ranks
among the top five major vegetable crops [1]. Immature fruit of culti-
vated species of eggplant are commonly used as a summer vegetable.
This low-cost vegetable is used as sautéing, grilling, baking, frying and
even barbecuing by both poor and rich people. With low caloric value
and high oxygen radical absorbance capacity, this healthy vegetable is a
good source of dietary minerals, vitamins, chlorogenic acid, antho-
cyanins and antioxidants [4].
Despite the economic and nutritional importance, breeding efforts
in eggplant are limited, consequently eggplant production is low than
other solanaceous crops. In general, eggplant breeding program's ob-
jective is to develop high-yielding varieties, mostly F1 hybrids, with
good stress tolerance potential [5]. However, it is not possible to
preferences. Therefore, development of locally preferred hybrids with
certain fruit characters along with high yield and adaptation is essen-
tially achieved through heterosis breeding [6]. Heterosis (or hybrid
vigor) is a natural phenomenon whereby hybrid offspring of genetically
diverse individuals display superior (or inferior) performance relative
to the mid-parent value (average heterosis), or to the superior parent
(heterobeltiosis), or over check cultivar (standard heterosis) [7]. Many
decades back, researchers suggested the biochemical and physiological
concept of heterosis [8]. But the recent advancements in molecular
genetics have confirmed that the cause of heterosis is purely genetical
[9]. Heterosis breeding involves evaluation of elite-parents and first
filial generations to detect heterotic hybrids and appropriate parents
[10].
Exploitation of heterosis/hybrid vigor in crop plants for significant
enhancement in yield and in other quantitative characters is a time-
tested approach in crop improvement. The estimation of heterosis for
yield and its component would be useful to judge the best hybrid
combination for exploitation of superior hybrids. Nagai and Kida [11]
were the first to observe hybrid vigor in eggplant. The commercial
exploitation of heterosis in eggplant is possible due to the low cost of F1
seed production and the low seed requirement per unit area.
Heterosis and inbreeding depression are results of the process of

⁎
Corresponding author.
E-mail address: chintanmistery123@gmail.com (C.R. Mistry).

https://doi.org/10.1016/j.cpb.2018.10.004
Received 8 May 2018; Received in revised form 16 October 2018; Accepted 18 October 2018
2214-6628/ © 2018 The Authors. Published by Elsevier B.V. This is an open access article under the CC BY-NC-ND license
(http://creativecommons.org/licenses/BY-NC-ND/4.0/).
C.R. Mistry et al.
changing individual genetic diversity in two reverse (increase and de-
crease) directions [12]. Inbreeding depression (ID) i.e. the decline in the
vigor of inbred caused by inbreeding is an opposite phenomenon of
heterosis and the amount of documented inbreeding depression varies
for different species [13]. Both heterosis and inbreeding depression are
due to non-additive gene action [14]. Despite poor results of in-
breeding, it is a valuable tool in crop breeding and is essentially re-
quired to develop superior genotype. The information on nature and
magnitude of ID is helpful in determining the effectiveness of selection
[15]. Hence, the present study was undertaken with an objective to
study the extent of heterosis in different crosses to develop superior F1
hybrids and to examine the magnitude of inbreeding depression in F2
generation and then utilization in future crop improvement programs.
2. Materials and methods
2.1. Field trial
The experiment was performed at Main Vegetable Research Station
(MVRS), Anand Agricultural University (AAU), Anand during July 2011
(latitude/longitude 22°–35′ North latitude and 72°–55′ East longitude).
This location has inceptisol soils of sandy loam texture and pH of 7.2.
The experimental material was comprised of six parents. Crosses viz.,
Doli-5 × GBL-1 (cross 1), Doli-5 × KS-331 (cross 2), Pusa Uttam × KS-
331 (cross 3) and AB 07-02 × GOB 1 (cross 4) were made between
parents by manual emasculation and pollen transfer. Salient features of
parental lines used in the study are given in Sabolu et al. [2]. F1 plants
were selfed to obtain seeds for the F2 generation. Parents, F1 and F2
generations were planted in separate experimental plots each in a
compact family block design with three replications. All agronomic
practices and preventive measures were taken to raise a healthy crop.
Each plot had one row each of two parents (P1 and P2) and F1 and four
rows of F2. Each row consisted of fifteen plants and inter and intra-row
spacing were 90 and 60 cm, respectively. The number of plants eval-
uated varied depending on the treatment and was larger for the seg-
regation generations such as the F2 (with sixty plants per repetition)
than for treatments non-segregating generations such as the P1 and P2
parents and the F1 generation (with fifteen plants per repetition each).
Plantlets were thinned to one plant per planting hole when the seed-
lings reached a height of 10 cm.
For evaluation of twelve yield associated traits, marketable and
fresh fruits were harvested from 10 plants. The observations were re-
corded for days to first flowering, days to first picking, plant height,
primary branches per plant, secondary branches per plant, fruit length,
fruit girth, fruits per plant, fruit yield per plant, pedicel length, fruit
volume and fruit weight. To avoid border effect, plants at the end of
each row were not considered [16].
2.2. Statistical analysis
Heterosis for each trait was worked out by using the overall mean of
each hybrid over replications for each trait.
Relative Heterosis was estimated as per cent deviation of hybrid
value from its mid-parental value. The formula used for estimating
relative heterosis was as follows
F1 − MP
Relative Heterosis (di) = × 100
MP
Where,
di = Heterosis over mid parental value (relative heterosis),
F1 = Mean hybrid performance, and MP = Mid parental value i.e.,
the arithmetic mean of two parents involved in the respective cross
combination
Heterobeltiosis was calculated at the superiority of hybrid from the
better parent as follows.
28
Current Plant Biology 16 (2018) 27–31
F1 − BP
Heterobeltiosis (dii) = × 100
BP
where,
dii = Heterobeltiosis i.e. heterosis over better parent,
BP = Average performance of better parent in the respected cross
combination
Inbreeding Depression: It was calculated by using the following
formula:
F1 − F2
ID (%) = × 100
F2
Where,
ID (%) = Inbreeding depression,
F1 = Mean hybrid performance, and
F2 = Mean of F2 population for a trait
Significance of the estimates of heterosis, heterobeltiosis and in-
breeding depression was tested using Student t-test by comparing the
calculated value of ‘t’ with the tabulated value of ‘t’ at 0.05 and 0.01
levels of probability of error.
3. Results and discussion
The degree of heterosis is measured as superiority of F1 hybrids over
the best parent (heterobeltiosis). The possibility of exploitation of hy-
brid vigor depends on feasibility of hybrid seed production at com-
mercial scale. Heterobeltiosis is an indicator of level of transgressive
segregants as superiority of hybrids helps in identification of promising
cross combinations having potential to produce the highest level of
transgressive segregants in conventional crop improvement program. In
the present study, heterosis is reported over mid parent (relative het-
erosis) and over better parent (heterobeltiosis) (Tables 1 and 2).
3.1. Days to first flowering
For this trait, heterosis in negative direction is desirable as it im-
parts early flowering. Relative heterosis was non-significant in all four
crosses for days to first flowering which is undesirable for this trait.
Similarly, heterobeltiosis was also non-significant in all crosses except
in cross 3 (Pusa Uttam × KS 331) which expressed highly significant
and positive heterobeltiosis (13.20%). In preview of above results the
cross 3 depicted significant heterobeltiosis towards positive direction,
which is not desirable for early flowering. The results of positive het-
erobeltiosis for days to first flowering in present study were in agree-
ment with Chadha and Sidhu [17], Chadha et al. [18] and Kapadia
[19]. With prolong vegetative growth, flowering time deviates in F1 but
reason for such phenomenon is still not clear. Genetic and epigenetic
reprograming of genes in particular hybrids due to blending of two
genomic make up may be one of the reason for positive heterobeltiosis
for delayed flowering [20]
3.2. Plant height
Except cross 4 (AB-07-02 × GOB 1), mid-parent heterosis was sig-
nificant in all crosses. Though, shorter than parents, relative heterosis
was negative (−13.04%) in cross 2 (Doli 5 × KS 331) which is not
desirable. Significantly positive heterobeltiosis was detected for cross 1
(Doli 5 × GBL 1 = 19.12%), cross 3 (Pusa Uttam × KS 331 = 18.95%)
and cross 4 (AB-07-02 × GOB 1 = 10.6%). Similar to relative heterosis,
heterobeltiosis was significantly negative (−13.90%) in cross 2 (Doli
5 × KS 331). The significant heterobeltiosis towards positive direction
showed its desirability for this trait. These findings are in congruence
with the results of Choudhary and Mishra [21], where negative to po-
sitive range of heterobeltiosis for plant height was reported. Only po-
sitive values of heterobeltiosis were calculated by Singh and Kumar
[22] Singh et al. [23], Patil and Shinde [24] and Sidhu and Chadha [25]
C.R. Mistry et al. Current Plant Biology 16 (2018) 27–31

Table 1
Heterosis (MP%), heterobeltiosis (BP%) and inbreeding depression (ID%) for various characters in four crosses in eggplant.
Crossa Percent heterosis over ID (%) Percent heterosis over ID (%)

MP (%) BP (%) MP (%) BP (%)

Days to first flowering Days to first picking

I −0.50 0.53 9.63** 1.63 −1.53 4.06**
II −2.37 −1.48 −7.38 1.76 −1.10 −12.96**
III 4.58 13.20** 7.08** 0.44 2.00 6.15**
IV 1.83 4.81 8.93** −0.56 2.81 3.33

Plant height Primary branches per plant

I 12.8** 19.12** 6.69** 2.91 −5.12** −18.24**
II −13.4** −13.90** −14.78** 13.82 7.69** 4.76**
III 12.8* 18.95** 16.30** −1.17 −2.56** −7.23**
IV 7.08 10.06* 7.47* 9.2 7.89** 0.60

Secondary branches per plant Fruit length

I 120.3** 100.00** 13.66** 15.08* 13.52* 1.19
II 112.6 107.50** 7.41** 22.72** 6.12** 8.80*
III 60.9** 47.17** 21.15** 18.28** 17.30** 6.31
IV 17.09** 11.86 21.67** 47.08** 40.49** 8.28

Fruit girth Fruits per plant

I 8.30 3.91 3.29 29.70** 26.68** 9.54**
II 12.67** 5.84 6.46* 6.26 5.75 7.00*
III 18.62** 17.47** 12.26** 7.67 1.69 −16.80**
IV 19.15** 14.66* 14.90** 30.48** 28.50** 3.82*

Fruit yield per plant Pedicel length

I 21.47** 21.00** −5.73 31.54** 27.55** −7.80
II 29.53** 24.47** 1.52 35.64** 29.77** 11.28**
III 16.88 13.50* −35.69** 14.45 8.05 13.04
IV 29.18** 20.55** 0.59 7.67 −8.27 −12.37

Fruit volume Fruit weight

I −26.05** 2.41** −35.28** 6.53** 3.77 −21.62**
II 95.52** 88.48** 22.39 19.80 15.23 −8.85
III 45.10** 11.71** 48.94 9.51** 5.72 −26.08**
IV 33.71** 7.52** −4.55 11.29 9.93 −24.72**

a
I = Doli-5 × GBL-1; II = Doli-5 × KS-331; III = Pusa Uttam × KS-331; IV = AB 07-02 × GOB 1.
* p < 0.01.
** p < 0.05.

Table 2 3.4. Secondary branches per plant

Cross-wise significant and desirable heterobeltiosis for different traits in four
crosses of eggplant.
Cross Traits
Doli 5 × GBL 1 Plant height, Secondary branches per plant, Fruits
length, Fruit per plant, Fruit yield per plant
Doli 5 × KS 331 Plant height, Primary branches per plant, Secondary
branches per plant, Fruit length, Fruit yield per plant
Pusa Uttam × KS 331 Plant height, Secondary branches per plant, Fruit length,
Fruit girth, Fruit yield per plant
AB-07-02 × GOB 1 Plant height, Fruit length, Fruit girth, Fruit per plant,
Fruit yield per plant, Primary branches per plant
For this trait, three crosses i.e., Doli 5 × GBL 1 (cross 1), Doli 5 ×
KS 331 (cross 2), Pusa Uttam × KS 331 (cross 3) showed highly sig-
nificant heterobeltiosis and relative heterosis as well as highly sig-
nificant and positive magnitude of inbreeding depression was also ob-
served. This result indicates dominance gene effects for obtaining
desirable segregants in yield improvement. Whereas, cross 4 (AB-07-02
× GOB 1) showed highly significant and positive estimate of relative
heterosis (17.09%) which is desirable for this trait while heterobeltiosis
was non-significant and positive. However, inbreeding depression was
found to be significant and positive.

for this trait. 3.5. Fruit length

In all four crosses, heterobeltiosis was found highly significant and

3.3. Primary branches per plant
The trait, primary branches per plant revealed highly significant and
positive heterobeltiosis, indicating presence of dominance gene effect in
cross 2, (Doli 5 × KS 331 = 7.69%) and cross 4 (AB-07-02 × GOB
1 = 7.89). Whereas, the cross 1 (Doli5 × GBL 1 = −5.12%) and cross
3 (Pusa Uttam × KS 331 = −2.56%) showed negative heterobeltiosis
and inbreeding depression. In this trait negative ID is desirable to in-
crease the yield indicating undesirable segregants. Negative to positive
range of heterobeltiosis for this trait was reported by Choudhary and
Mishra [21], though; Singh et al. [23], Chadha and Sidhu [17], Patil
and Shinde [24] and Sidhu and Chadha [25] reported it in positive
direction for this trait.
positive i.e., the cross 1 (13.52%), heterosis over mid (22.72%) and
better parent (6.12%) in the cross 2 (Doli 5 × KS 331), the cross 3 (Pusa
Uttam × KS 331) expressed highly significant heterosis over mid and
better parent, as well as in the cross4 (AB-07-02 × GOB 1) highly
significant and positive value of heterobeltiosis (47.08%) and relative
heterosis (40.49%) was found for this trait.
The positive heterobeltiosis in all four crosses suggested that the
fruit length may remain constant or may get reduced in F2 generation in
comparison to F1 hybrids due to less dominant gene action in F2. The
results of positive heterobeltiosis for fruit length in present study were
in agreement with Chadha and Sidhu [17], Patil and Shinde [24], Sidhu
and Chadha [25] and Dixit and Gautam [26]. Positive magnitude of
inbreeding depression was exhibited in most of the crosses for fruit

29
C.R. Mistry et al.
length suggesting fewer chances for beneficial segregants in F2 popu-
lations.
3.6. Fruit girth
In this trait, two crosses viz.,cross 3 (Pusa Uttam× KS 331) and
cross 4 (AB-07-02 × GOB 1) showed significant and positive magni-
tudes of relative heterosis (18.62%, 19.15%) and heterobeltiosis
(17.47%, 14.66%).The magnitude of inbreeding depression for this trait
was also found significantly positive and which is not desirable and to
improve the trait one should avoid the self-pollination. Whereas, two
crosses i.e., cross 1 (Doli 5 × GBL 1) and cross 2 (Doli 5 × KS 331)
showed non-significant values of both heterobeltiosis as well as relative
heterosis. Though, the value of inbreeding depression was non-sig-
nificant for cross 1 (Doli 5 × GBL 1) and significant (6.46%) for cross 2
(Doli 5 × KS 331).
3.7. Fruits per plant
The trait, fruits per plant manifested highly significant hetero-
beltiosis as well as relative heterosis in positive direction suggesting
higher potentiality of this hybrid for producing more number of fruits
per plant in the cross 1 (Doli 5 × GBL 1) and cross 4 (AB-07-02 × GOB
1) in F1generation. These findings were supported by the results of
Vijay and Nath [27], Bhutani et al. [28], Patel [29], Kalloo et al. [30],
Kapadia [19], Ingale and Patil [31], Kumar et al. [32], Jha [33] and
Patel [34], as they reported negative to positive range of heterobeltiosis
for this trait. Whereas, cross 3 (Pusa Uttam × KS 331) expressed sig-
nificant and negative inbreeding depression indicating more chances
for beneficial segregants in F2 population. Kapadia [19] reported
moderate heritability for this trait.
3.8. Fruit yield per plant
Yield is a complex trait and is the end product of several basic yield
components. The higher magnitude of heterobeltiosis as well as relative
heterosis in almost all four crosses indicated the presence of over-
dominance, which suggested that exploitation of heterosis through
heterosis breeding may prove to be effective for improvement at a
greater extent of the fruit yield per plant in F1population. These find-
ings are in congruence with the results of Singh et al. [23], Singh [35],
and Jha [33]. For fruit yield per plant, the inbreeding depression (ID)
was recorded for crosses 2 and 4. Only cross 3 (Pusa Uttam × KS 331)
depicted significant inbreeding depression in negative direction for this
trait showing significant increase in F2 over F1, which is more preferred
to obtain beneficial segregants. The higher estimates of various het-
erotic effects in positive direction and presence of undesirable in-
breeding depression suggested that heterosis breeding would be of
immense value for improvement of fruit yield in brinjal. These findings
are in conformity with the results of Singh [35], Gopinath and Mada-
lageri [36] and Jha [33] who reported positive estimates of inbreeding
depression for this trait.
3.9. Pedicel length
In this trait, the two crosses viz., cross 1 (Doli 5 × GBL 1) and cross
2 (Doli 5 × KS 331) revealed highly significant and positive values of
both relative heterosis and heterobeltiosis which are helpful for making
effective selection in succeeding generations. The cross 4 (AB-07-02 ×
GOB 1) expressed negative and non-significant values of both hetero-
beltiosis (−8.27%) and inbreeding depression (−12.37%), which
suggested undesirable effect of heterosis and desirable effect of IB, re-
spectively, on this trait.
30
Current Plant Biology 16 (2018) 27–31
3.10. Fruit volume
All the four crosses viz., Doli 5 × GBL 1, Doli 5 × KS 331, Pusa
Uttam × KS 331 and AB-07-02 × GOB 1 exhibited higher magnitude of
heterobeltiosis in positive direction, while negative value of inbreeding
depression in the cross cross1 (−35.28%) and cross 4 (−4.55%) in-
dicated possibilities for obtaining desirable segregants that can be im-
proved in succeeding generations.
3.11. Fruit weight
Non-significant values for heterobeltiosis in all crosses were found
for the trait fruit weight. However, relative heterosis in two crosses i.e.
1 and 3 was significantly positive (p < 0.01). The crosses 1 (Doli 5 ×
GBL 1), 3 (Pusa Uttam × KS 331), and 4 (AB-07-02 × GOB 1) had
significantly negative values (p < 0.01; F2 mean higher than F1 mean)
of inbreeding depression. Although, ID for cross 2 was non-significant
however it also showed negative value. The magnitude of inbreeding
depression was found negative and highly significant in two crosses viz.,
cross 3 (−26.08%) and cross 4 (−24.72%) indicated existence of
chances for the improvement of this trait. The result also suggested that
negative ID would increase the fruit weight with selfing in further
generations.
3.12. Inbreeding depression
The inbreeding depression is an important criterion for crop
breeding program. Due to minimal genetic load self-pollinated crop
species like eggplant, show little inbreeding depression [37]. Natural
selection and/or plant breeding would be expected to eliminate dele-
terious gene mutations with large effects. Therefore, the inbreeding
depression observed in present study is assumed not to be due to ex-
pression of homozygous deleterious alleles as a case in cross-pollinated
crops [38]. The character and cross wise results of inbreeding depres-
sion are presented in Table 1. The magnitude of inbreeding depression
varied from cross to cross indicating influence of genetic constitution of
crosses. In the present study, either low or moderate amount of in-
breeding as well as outbreeding depression was detected. It is desirable
to have highly significant and positive heterosis with low inbreeding
depression. Positive and significant inbreeding depression was observed
in cross 1 (Doli 5 × GBL 1), cross 3 (Pusa Uttam × KS 331) and cross 4
(AB-07-02 × GOB 1) except cross 2 (Doli 5 × KS 331) for days to first
flowering, indicating possibilities to get the desirable segregants in the
segregating generations. Likewise, significantly positive ID was also
observed for days to first picking in crosses 1 and 3. Kapadia [19] re-
ported positive and significant inbreeding depression for both the traits.
In case of plant height (in all crosses) and primary branches per plant
(in three crosses), significant inbreeding depression in both negative
and positive directions was observed.
In case of fruit traits, significant and positive magnitude of in-
breeding depression was observed in the cross 2 (Doli 5 × KS 331), and
cross 4 (AB-07-02 × GOB 1) for fruit girth and fruits per plant. A sig-
nificantly positive ID for fruit length was observed only in cross 2 (Doli
5 × KS 331) indicating less chances for beneficial segregants in F2 po-
pulation. Sao and Mehta [39] reported the high degree of inbreeding
depression for fruit yield per plant and its related traits in eggplant.
To avoid inbreeding depression, in F2 and onward generations,
desired segregants should be crossed in bi-parental mating fashion to
get desirable plant types in progenies. The higher estimates of various
heterotic effects in positive direction and presence of undesirable in-
breeding depression suggested that heterosis breeding would be of
immense value for improvement of fruit yield in eggplant. These find-
ings are in conformity with the results of Singh [35], Gopinath and
Madalageri [36] and Jha [33], who reported positive estimates of in-
breeding depression for these traits.
C.R. Mistry et al. Current Plant Biology 16 (2018) 27–31

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