preliminary studies on heterosis in snakegourd...
TRANSCRIPT
2011 vol. 74, 25-38
DOI: 10.2478/v10032-011-0002-x ________________________________________________________________________________________
Corresponding author:
e-mail: [email protected] © Copyright by RIVC
PRELIMINARY STUDIES ON HETEROSIS IN SNAKEGOURD
(TRICONSANTHES CUCUMERINA)
F.N. AHSAN, A.K.M.A. ISLAM, M.G. RASUL, M.A.K. MIAN, M.M. HOSSAIN
Department of Genetics and Plant Breeding
Department of Horticulture
Bangabandhu Sheikh Mujibur Rahman Agricultural University
Gazipur 1706, Bangladesh
Received: October 19, 2010; Accepted: May 8, 2011
Summary
The analysis of variance showed significant difference among the parents
and hybrids of snakegourd for most of the characters except days to first female
flower, days to first fruit setting and node number of first fruit setting. The hy-
brid SG-18 × SG-01 produced the highest number of fruits per plant and fruit
yield per plant followed by SG-04 × SG-26. The hybrid SG-04 × SG-26 took
minimum 81-83 days to produce female flower. Both positive and negative mid
parent and better parent heterosis was obtained for different characters of hy-
brids of which few hybrids showed desirable and significant values. The highest
mid parent heterosis in negative direction (-9.66%) was found for days to first
male flower in hybrid SG-06 × SG-18 and for days to first female flower open-
ing in hybrid SG-04 × SG-26 (-9.30%). The hybrids SG-01 × SG-18, SG-04 ×
SG-26, SG-06 × SG-18, SG-06 × SG-25, SG-18 × SG-01 and SG-18 × SG-25
showed significant desirable mid parent heterosis for number of fruits per plant,
and SG-04× SG-26, SG-06 × SG-18, SG-06 × SG-25, and SG-18 × SG-25
showed better parent heterosis. Four hybrids (SG-04 × SG-26, SG-06 × SG-18,
SG-06 × SG-25 and SG-18 × SG-25) exhibited significant and desirable level of
both mid and better parent heterosis for number of fruits per plant. The hybrids
SG-04 × SG-26, SG-06 × SG-18, SG-06 × SG-25 and SG-18 × SG-25 could be
identified as promising combination for commercial cultivation. Further inves-
tigation should be carried out to confirm the heterosis exhibited by the hybrids.
key words: Snakegourd (Triconsanthes cucumerina), hybrid, underutilized crop,
heterobeltiosis, summer vegetables
INTRODUCTION
Snakegourd (Triconsanthes cu-
curminata) belongs to the family Cu-
curbitaceae having chromosome
number, 2n=22 (Chakarbati 1982,
Jeffery 1980). Snakegourd grows
more or less in every part of Bangla-
desh. Snakegourd is a day neutral type
of vegetable which can be grown in
26 VEGETABLE CROPS RESEARCH BULLETIN 74
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any time of the year except acute win-
ter. Usually it is grown well from
March to October both in the field and
homestead garden. As a result, it can
meet up the vegetable demand during
early kharif season (mid March to mid
June) when there exists an acute
shortage of vegetable in Bangladesh.
In addition it has got tremendous ex-
port potentiality because of its excel-
lent keeping quality. The yield of
snakegourd is very low and its pro-
duction is also restricted to only to 3-4
months in Bangladesh (BBS 2007).
There are number of cultivars with
wide range of variability in size,
shape and color of fruits available in
this country (Rashid 1993) and for
this we can easily fulfill the gap by
producing more snakegourd in the
lean period by hybrid variety.
Heterosis or hybrid vigor can
play a vital role in increasing the yield
quality of vegetable crops. It refers to
the phenomenon in which F1 hybrid
obtained by crossing of two genetical-
ly dissimilar inbred lines or genotypes
which shows increased or decreased
vigor over the better parent or mid
parent value (Poehlman 1979). In
cucurbits, heterosis was first noted by
Hays and Jones (1961). At present, in
most advanced and developing coun-
tries of the world like USA, Japan,
Netherlands, Taiwan, Thailand etc.
almost all common varieties of vege-
table crops are being replaced by F1
hybrids. There is limited information
available about the magnitude and
nature of heterosis for yield and yield
contributing characters in snakegourd
as it is an underutilized crop. Due to
monoecism, much scope for exploita-
tion of heterosis in snakegourd exists
virtually obligatory out crossing sys-
tem of snakegourd open the scope of
hybrid variety. In spite of that, a few
numbers of superior hybrid varieties
have been developed for cultivation in
different countries like India, Thai-
land etc. However, very little attention
has been given to develop high yield-
ing hybrid variety of snakegourd be-
ing monoecious. Therefore there is a
bright scope to study the heterosis,
which is the prerequisite for develop-
ing high yielding snakegourd varieties
or hybrid variety in Bangladesh. Con-
sidering the above idea in mind the
present investigation was undertaken
with the objectives (i) to determine
the magnitude of heterosis as well as
hybrid performance for some desira-
ble yield contributing characters and
(ii) to identify high heterotic parental
combination in order to develop hy-
brid varieties.
MATERIALS AND METHODS
The experiment was conducted
at the experimental farm of Banga-
bandhu Sheikh Mujibur Rahman Ag-
ricultural University (BSMRAU),
Salna, Gazipur, Bangladesh during the
months from March to June, 2007.
Seven parents (P1=SG-01, P2=SG-04,
P3=SG-06, P4=SG-10, P5=SG-18,
P6=SG-25 and P7=SG-26) and their
twelve F1s were used as an experi-
mental materials. The experiment was
laid out in a Randomized Complete
Block Design (RCBD) with three
replications. Seeds of the seven par-
ents and their F1s were first allowed to
soak water for 24 hours. The soaked
seeds were then sown in poly bag
(size 15 × 15 cm), containing a mix-
ture of soil and well decomposed cow
dung (1:1). Thirty five days old seed-
F.N. AHSAN et al. – PRELIMINARY STUDIES OF HETEROSIS … 27
_____________________________________________________________________________________________________
lings were transplanted in main field
and four pits for every genotype for
each replication were maintained. The
spacing between rows and between
plants was 2.0 m and 1.5 m, respec-
tively. Recommended doses of ma-
nure and fertilizers were applied in the
experimental field according to BARI
(1991). Data were collected on each
of the seven parents and their F1s for
the characters such as days to first
male flower, days to first female
flower, days to first fruit setting, node
number of first fruit setting, number
of primary branches per plant, length
of vine in each plant (cm), total num-
ber of fruits plant, fruit weight (g),
fruit diameter (cm), fruit length (cm),
fruit yield/plant (g). Heterosis was
recorded as mid parent heterosis (Hm)
and better parent heterosis (Hb) ac-
cording to the formula adopted by
Falconer & Mackay (1996) and was
estimated following Mather & Jinks
(1971). The significant test (t-test)
was employed following given formu-
la whether F1 hybrid mean were statis-
tically different from mid parent and
better parent values (Singh &
Choudhary 2006).
RESULTS
The analysis of variance showed
significant difference among the par-
ents and hybrids for most of the char-
acters except days to first female
flower, days to first fruit setting and
node number of first fruit setting (Ta-
ble 1 & 2). The estimates of percent
mid parent heterosis and better parent
heterosis observed in the F1 genera-
tion are presented in Table 3 and 4.
The heterosis analysis in the present
study was done to identify the best
combinations of parents giving high
degree of heterosis.
Days to first male flower Mid parent heterosis for most of the
crosses were earlier than their mid
parent except SG-06 × SG-25 for first
male flower opening. Mid parent het-
erosis was ranged from 2.52 to -9.66
percent. The highest mid parent heter-
osis in negative direction (-9.66%)
was found for this trait in hybrid SG-
06 × SG-18 (Table 3). Thus the cross
combination SG-06 × SG-18 may be
selected for best cross combination
for getting early male flower. The
male flower bloomed in hybrids were
earlier than their parents (Table 1).
Better parent heterosis ranged from
-0.41 to -12.60 percent. The highest
negative better parent heterosis
(-12.60%) was found for this charac-
ter in hybrid combination SG-06 ×
SG-18 and SG-10 × SG-18 (Table 4).
The hybrid SG-06 × SG-18 exhibited
significant negative mid parent heter-
osis.
Days to first female flower
The crosses SG-01 × SG-26 (-8.50%),
SG-04 × SG-26 (-9.30%), SG-06 ×
SG-10 (-5.23%) and SG-18 × SG-25
(-4.54%) showed significant negative
heterosis. Heterosis for earliness in
female flower ranged from -0.60 to -
9.30%. The highest significant nega-
tive heterosis was noticed for this trait
in hybrid SG-04 × SG-26 (Table 3)
and heterobeltiosis for earliness in
female flower ranged from -0.80% to
-10.00%. The maximum significant
negative heterobeltiosis was observed
in the hybrid SG-04 × SG-26 followed
by SG-01 × SG-26 (Table 4).
28 VEGETABLE CROPS RESEARCH BULLETIN 74
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Table 1. Performance of seven parental genotypes for yield and yield related characters
in snakegourd
Parents/crosses
Days to
first
male
flower
Days to
first
female
flower
Days to
first
fruit
setting
Node
no. of
first
fruit
setting
No. of
primary
branches
per plant
Length
of vine
(m)
Parents
SG-01 81.33 83.00 89.66 19.33 8.33 4.83
SG-04 76.33 83.0 85.00 21.66 10.33 4.97
SG-06 76.66 82.33 87.00 23.33 11.00 5.59
SG-10 75.00 83.00 88.00 13.66 10.00 4.64
SG-18 82.00 83.33 89.00 16.00 10.66 6.17
SG-25 82.00 84.33 89.00 18.33 11.66 5.54
SG-26 74.66 81.667 83.66 21.33 9.33 4.63
F test * NS NS * ** **
X 78.28 82.95 87.33 19.190 10.190 5.19
CD (0.05) 5.28 3.80 4.54 5.57 0.95 0.16
Hybrids
SG-01×SG-10 76.33 86.00 88.33 17.50 10.66 4.92
SG-01×SG-18 77.00 84.00 87.00 21.50 10.33 5.15
SG-01×SG-26 73.33 75.33 78.66 23.00 7.66 4.79
SG-04×SG-26 72.33 74.667 79.00 20.33 10.66 6.36
SG-06×SG-10 72.00 82.66 86.00 18.00 10.66 5.91
SG-06×SG-18 71.66 82.33 85.00 20.00 7.33 4.49
SG-06×SG-25 81.33 84.33 87.66 20.00 11.33 6.33
SG-10×SG-06 72.66 78.33 84.66 18.66 7.33 5.85
SG-10×SG-18 71.66 79.00 83.00 26.00 9.66 5.02
SG-10×SG-25 72.33 82.66 84.66 14.66 11.00 5.07
SG-18×SG-01 81.66 82.66 84.66 17.66 9.00 4.74
SG-18×SG-25 76.00 80.00 82.33 17.33 11.33 5.78
F test ** ** ** NS ** **
X 74.86 81 84.250 19.417 9.750 5.37
CD (0.05) 3.02 3.69 3.46 5.82 0.91 0.23
Note: ** significant at 1% level of significance
F.N. AHSAN et al. – PRELIMINARY STUDIES OF HETEROSIS … 29
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Table 2. Performance of twelve hybrids for yield and yield related characters in
snakegourd
Parents/crosses
Total no.
of fruits
per plant
Fruit
weight (g)
Fruit diam-
eter (mm)
Fruit
length
(cm)
Fruit yield
per plant
(kg)
Parents
SG-01 14.33 111.82 34.45 27.31 1.60
SG-04 32.33 144.37 37.61 22.14 4.66
SG-06 20.33 101.89 29.89 24.12 2.07
SG-10 30.00 158.82 44.13 21.28 4.76
SG-18 28.00 151.55 50.85 17.73 4.24
SG-25 16.00 136.80 41.05 20.50 2.18
SG-26 13.00 159.45 26.18 52.79 2.07
F test ** ** ** ** **
X 22.00 137.81 37.74 26.55 3.08
CD (0.05) 1.02 1.54 0.99 0.88 4.03
Hybrids
SG-01×SG-10 14.66 154.58 39.94 25.91 2.26
SG-01×SG-18 28.00 124.13 40.49 24.17 3.47
SG-01×SG-26 9.00 145.16 42.82 31.58 1.30
SG-04×SG-26 42.00 210.32 46.42 33.75 8.83
SG-06×SG-10 23.33 137.85 35.71 24.53 3.21
SG-06×SG-18 35.33 131.17 36.10 28. 00 4.63
SG-06×SG-25 22.33 133.95 31.30 30.87 2.99
SG-10×SG-06 25.33 150.65 34.22 27.77 3.81
SG-10×SG-18 18.66 140.85 49.66 21.85 2.62
SG-10×SG-25 32.33 110.26 35.14 25.60 3.56
SG-18×SG-01 51.66 189.24 50.58 28.25 9.77
SG-18×SG-25 31.66 152.81 44.69 24.89 4.83
F test ** ** ** ** **
X 27.86 148.41 40.59 27.26 4.27
CD (0.05) 1.07 1.19 0.81 0.65 2.07
Note: see Table 1
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F.N. AHSAN et al. – PRELIMINARY STUDIES OF HETEROSIS … 31
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32 VEGETABLE CROPS RESEARCH BULLETIN 74
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Days to first fruit setting
Mid parent heterosis for days to first
fruit setting were ranged from -0.38 to
-9.23 percent (Table 3). The highest
mid parent heterosis in negative direc-
tion (-9.23%) was observed in the
hybrid combination SG-01×SG-26.
Better parent heterosis ranged from
-1.48 to -12.26 percent and the highest
negative better parent heterosis
(-12.26%) was found in hybrid com-
bination SG-01×SG-26 (Table 4).
Node number for first fruit setting
Mid parent heterosis varied from
-1.70 to -8.31 percent. The highest
(-8.31%) mid parental heterosis for
node number of first fruit setting was
observed in the cross combination
SG-10×SG-25 followed by SG-10×
SG-25 (-5.39%) and SG-06×SG-25
(-3.98%) (Table 3). Better parent het-
erosis ranged from 11.22 to -22.80
percent. The highest (-22.80%) better
parent heterosis for this trait was
found in the cross combination SG-06
× SG-10 (Table 4).
Number of primary branches per
plant
The cross combination SG-06×SG-25
showed no heterosis (0). The highest
(16.37%) positive heterosis was found
in SG-01×SG-10 (Table 3), could be
used as a desirable combination for
remaining the no of primary branches.
The better parent heterosis was ranged
from -9.38 to -34.30 percent (Table 4).
Length of vine in each plant
Heterosis over mid parent ranged
from -0.39 to 32.5 percent (Table 3).
The hybrid combination SG-04×SG-
26 (32.5%) showed the highest mid
parent heterosis value for length of
vine followed by SG-06×SG-10
(15.65%), SG-10×SG-06 (14.48%)
and SG-06×SG-25 (13.84). The high-
est better parent heterosis was ob-
served in the hybrid combination SG-
04×SG-26 (27.96%). The hybrid SG-
06×SG-10 exhibited significant posi-
tive mid parent heterosis and het-
erobeltiosis (Table 4).
Total number of fruits per plant
Seven hybrids out of twelve showed
significant mid parent heterosis for
total number of fruits per plants. The
highest significant positive heterosis
was found in the cross combination
SG-18×SG-01 (144.08%) (Table 3).
In case of better parent heterosis, six
cross combinations showed signifi-
cant positive heterosis for number of
fruits per plant. The range of better
parent heterosis was -51.13% to
84.5%. The highest (84.50%) signifi-
cant positive heterosis was observed
in the cross combination SG-18×SG-
01 (Table 4).
Average fruit weight
The highest mid parent value for this
trait was found in the cross combina-
tion SG-18×SG-01 (43.71%) followed
by SG-04×SG-26 (38.5%). Better
parent heterosis ranged from -2.08 to
31.90 (Table 4). Only three cross
combinations (SG-04 × SG-26, SG-18
×SG-01 and SG-18×SG-01) showed
positive and significant better parent
heterosis.
Fruit diameter
Mid parent heterosis for average fruit
diameter were ranged from -2.74 to
45.56 percent. The highest significant
positive mid parent heterosis was
found by the cross combination SG-
04×SG-26 (45.56%) (Table 3). Better
parent heterosis for this trait ranged
from -0.53 to 24.29 percent. Two
hybrids SG-01×SG-26 (24.29%) and
F.N. AHSAN et al. – PRELIMINARY STUDIES OF HETEROSIS … 33
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SG-04×SG-26 (23.42%) exhibited
significant and positive heterosis.
Fruit length
Heterosis over mid parent for average
fruit length ranged from -9.9 to 38.36
percent. The highest mid parent heter-
osis effect was observed in hybrid
SG-06×SG-25 (38.36%) followed by
SG-06×SG-18 (33.81%) and SG-18×
SG-25 (30.24%) (Table 3). Better
parent heterosis ranged from -5.12 to
27.96 percent.
Fruit yield per plant
Mid parent heterosis ranged from -
41.77 to 234.58 percent for fruit yield
per plant. The cross combinations,
SG-18×SG-01 (234.58%), SG-04×
SG-26 (162.79%), SG-18×SG-25
(50.46%), SG-06×SG-18 (46.98%)
and SG-18×SG-01 (41.03%) showed
positive significant mid parent hetero-
sis (Table 3). Better parent heterosis
ranged from -130.48% to 89.48% for
fruit yield per plant (Table 4).
DISCUSSION
Days to first flowering is an im-
portant character as it reflects earli-
ness or lateness of parents and their
hybrids which has great importance in
breeding for earliness in any crop
species. Among the twelve crosses, all
the crosses were earlier than their
better parent heterosis. The highest
negative better parent heterosis was
found in hybrid combination SG-06×
SG-18 and SG-10×SG-18 which indi-
cates these are the best performing
heterotic hybrid combination over the
better parents. Significant negative
mid parent heterosis and heterobelti-
osis for hybrid SG-06×SG-18 re-
vealed the over dominance type of
gene action for this cross. The magni-
tude of heterosis over better parent
was high for days to first male flower
in ridge gourd (Shaha & Kale 2003).
Hybrid exhibited a negative heterosis
for the trait days to first female flower
opening in snakegourd, indicating the
earliness in crop growth (Kumaresan
et al. 2006). The hybrid SG-04×SG-
26 could be used as the best cross
combination for days to first female
flower as this hybrid showed signifi-
cant negative heterosis. Significant
highest negative heterosis for female
flowering was also reported in
snakegourd (Rahman 2004). The hy-
brid SG-01×SG-26 exhibited signifi-
cant negative mid parent heterosis and
heterobeltiosis that revealed the over
dominance type of gene action for this
cross. The highest better parent heter-
osis was found in the cross combina-
tion SG-06×SG-10 for node number
of first fruit setting. Similar results
were found by Banik (2003) and
Rahman (2004) in snakegourd. Singh
& Joshi (1979) also found these re-
sults in bitter gourd. The highest
standard heterosis for early fruit yield
per plant was observed in bottle gourd
(Liu et al. 2003). The cross combina-
tion SG-04×SG-26 can be used as a
desirable combination for getting the
longest vine. The hybrid SG-06×SG-
10 exhibited significant positive mid
parent heterosis and heterobeltiosis
which revealed the over dominance
type of gene action for length of the
vine. Significant positive mid parent
and better parent heterosis for total
number of fruits per plants was found
in seven and six hybrids, respectively
and could be used as the best combi-
nation. Solanki et al. (1982) recorded
maximum heterosis (42.12%) for
number fruits per plant in cucumber
34 VEGETABLE CROPS RESEARCH BULLETIN 74
_____________________________________________________________________________________________________
hybrid. Podder (2006) also reported
similar results in snake gourd and
Mohanty and Mishra (1999) in pump-
kin. Three cross combinations showed
positive and significant better parent
heterosis. For getting larger size fruit
these combinations could be used for
hybrid breeding. Banik (2003) also
found both positive and negative het-
erosis over better parent for fruit
weight. Significant positive heterosis
for fruit characters and fruit yield was
also observed by Hedau & Sirohi
(2004) in ridge gourd. Four hybrids
showed significant positive mid par-
ent heterosis for average fruit diame-
ter, which can be used as a desirable
combination for large fruit diameter.
Out of twelve cross combinations,
only two hybrids exhibited significant
and positive heterosis. The cross
combination SG-06×SG-25 exhibited
the highest better parent heterosis and
can be used as best cross combination
for getting long fruit.
The aim of heterosis in this study
was to identifying the best heterotic
combinations and its exploitation for
commercial purpose. In the present
investigation heterosis of the hybrids
for fruit yield and yield components
were studied over mid parent and in
addition better parent (Table 5). None
of the hybrid studied had recorded
maximum heterosis for all the traits,
but significant and desirable level of
heterosis over mid parent and better
parent were notes in several hybrids
for the different traits (Table 5). Sig-
nificant and desirable level of mid
parent heterosis was observed in sev-
en hybrids for fruit yield per plant and
better parent heterosis in four hybrids.
The heterosis exhibited for fruit char-
acters and fruit yield was mostly posi-
tive in bitter gourd (Pal et al. 2005).
The maximum heterosis for yield was
recorded in ash gourd by Sudhakar et
al. (2005). Similar results were also
found by Ahmed (1998), Banik
(2003) and Rahman (2004) in
snakegourd. The mid and better parent
heterosis was observed to be as high
as ~165% for yield per vine in ash
gourd (Verma & Behera 2007). Six
hybrids in both case showed signifi-
cant and desirable heterosis for num-
ber of fruits per plant. Only four hy-
brids exhibited significant and desira-
ble levels of heterosis over both mid
and better parent (Table 6, Photo 1).
The cross combinations SG-04×SG-
26, SG-06×SG-18, SG-06×SG-25,
SG-18 ×SG-25 showed significant
positive mid parent heterosis and het-
erobeltiosis (Photo 1) revealed the
over dominance type of gene action
operating these crosses. These cross
combinations could be identified as
desirable hybrids for commercial cul-
tivation. So these four hybrid viz SG-
04×SG-26, SG-06×SG-18, SG-
06×SG-25, and SG-18×SG-25 were
identified as promising for commer-
cial cultivation. Performance of these
hybrids needs to be evaluated in mul-
tilocation and on farm trial prior to
commercial use.
F.N. AHSAN et al. – PRELIMINARY STUDIES OF HETEROSIS … 35
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36 VEGETABLE CROPS RESEARCH BULLETIN 74
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F.N. AHSAN et al. – PRELIMINARY STUDIES OF HETEROSIS … 37
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WSTĘPNE BADANIA EFEKTU HETEROZJI U GURDLINY WĘŻOWEJ
(TRICHONSANTHES CUCUMERINA L.)
Streszczenie
Analiza wariancji wykazała istotne różnice wśród form rodzicielskich i mieszań-
ców gurdliny wężowej pod względem większości cech z wyjątkiem: liczby dni do
pierwszego kwiatu żeńskiego, liczby dni do pierwszego zawiązania owoców oraz liczby
węzłów w pierwszym zawiązaniu owoców. Mieszaniec SG-18 × SG-01 wytworzył
największą liczbę owoców na roślinę i plon owoców na roślinę, a za nim SG-04 × SG-
26. Mieszańcowi SG-04 × SG-26 zajęło minimum 81-83 dni żeby wytworzyć kwiat
żeński. Zarówno pozytywny jak i negatywny efekt heterozji względem średniej cechy
obojga rodziców (mid-parent heterosis) oraz względem lepszego z rodziców (better-
parent heterosis) uzyskano dla różnych cech mieszańców, pod względem których nie-
wiele mieszańców wykazywało pożądane i istotne wartości. Najwyższą heterozję mid-
parent w kierunku ujemnym (-9.66%) stwierdzono dla liczby dni do pierwszego kwiatu
męskiego u mieszańca SG-06 × SG-18 oraz dla liczby dni do otwarcia pierwszego
kwiatu żeńskiego u mieszańca SG-04 × SG-26 (-9.30%). Mieszańce SG-01 × SG-18,
SG-04 × SG-26, SG-06 × SG-18, SG-06 × SG-25, SG-18 × SG-01 oraz SG-18 × SG-25
wykazały istotny pożądany efekt heterozji mid-parent dla liczby owoców na roślinę, a
mieszańce SG-04× SG-26, SG-06 × SG-18, SG-06 × SG-25, oraz SG-18 × SG-25 efekt
heterozji better-parent. Cztery mieszańce (SG-04 × SG-26, SG-06 × SG-18, SG-06 ×
SG-25 oraz SG-18 × SG-25) wykazały istotny i pożądany poziom heterozji zarówno
mid-parent jak i better-parent pod względem liczby owoców na roślinę. Mieszańce SG-
04 × SG-26, SG-06 × SG-18, SG-06 × SG-25 oraz SG-18 × SG-25 można określić jako
obiecujące skrzyżowania do hodowli dla celów handlowych. Należy przeprowadzić
dalsze badania w celu potwierdzenia heterozji wykazywanej przez te mieszańce.