Chapter 5
Chapter 5
DISCUSSION
97
In the opinion of Joshi and Dhawan (1966), desirable improvement could be
possible by creation and exploitation of genetic variability with the help of
characterization of genetic variance. In the light of early reports, it seems that further
improvement in yield potential of barley can be brought firstly, by suitably altering the
genetic makeup of existing varieties through the process of recombination and secondly
by employing the appropriate selection method for exploiting the different diverse
populations.
Therefore, present investigation was undertaken to obtain and compare the nature
and magnitude of gene effects, heterosis and selection parameters of yield and yield
contributing characters in barley in normal and saline sodic soils.
98
Heritability estimates provide information on transmission of characters from
parents to the progeny. Such estimate facilitates evaluation of hereditary and
environmental effects on phenotypic variation and thus aid in selection. Heritability
estimates are used to predict expected genetic advance under selection so that breeders
are able to anticipate improvement from different selection intensity. Johnson et al.
(1955) have suggested that heritability estimates in association with genetic advance are
much useful for selection than heritability alone.
The results obtained on various aspects of the present investigation are discussed
in the light of available literature in this chapter under following sections:
5.1 Genetic variability
5.1.1 Coefficient of variation
5.1.2 Heritability and genetic advance
5.2 Combining ability
5.2.1 General combining ability and its effects
5.2.2 Specific combining ability and its effect
5.3 Proportional contribution of lines, testers and line x tester
5.4 Nature and magnitude of heterosis for yield and its components
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for these traits are in close agreement with the results of Mishra et al (2007) Singh et al
(2008), Al-Tabbal et al. (2012) and Drikvand et al. (2011) in barley.
The characters which exhibited moderate estimates (10-20%) of PCV and GCV
were recorded for Grains/ Spike, Spikelets/ Spike and Grain Yield/ Plant under both
(E1&E2) and 1000- grains weight and harvest index saline-sodic soil (E2). Number of
effective tiller/plant, and biological yield has moderate PCV and low GCV under both
normal (E1) and saline-sodic condition (E2), while harvest index in normal soil (E 1)
remaining traits viz., plant height, days to 50% flowering and days to maturity showed
low estimates (< 10%) of PCV and GCV under both normal (E 1) and saline-sodic
condition (E2), while1000- grains weight in normal (E1).
5.1.2 Heritability and genetic advance:
Heritability in broad sense gives an idea of the magnitude of environmental
influence on the phenotypic expression of a character. Heritability in narrow sense, which
gives the precise idea of response to selection, broad sense heritability can also be
considered as the upper limit of the transmissibility of the character.
In the present study, high estimates of broad sense heritability (> 75%) were
recorded for grains/ spike (94.20%), spikelets/ spike (93.53%), Biological yield
(91.49%), plant height (90.00%), grain yield/plant (89.14%), Harvest Index (85.25%),
1000-grain weight (84.69%), and days to 50% flowering (84.54%) under normal
condition (E1) and under saline-sodic soil (E2) the high estimates of broad sense
heritability (> 75%) were recorded for plant height (98.78%), grains/ spike (97.11%),
1000-grain weight (94.49%), Harvest Index (91.04%), Biological yield (83.62%), days to
50% flowering (83.26%), spikelets/ spike (82.60%), grain yield/plant (81.87%) and days
to maturity (77.65%), confirming the earlier observations that these are less influenced by
environmental factors. Observations on higher estimates of heritability for these traits are
in close agreement with the results of Yassin et al. 2015, Porumb et al. 2016 and Hailu et
al. 2016.
100
reported had high estimates of heritability was recorded for protein content, husk content
of F1 & F2 generation in barley.
Since heritability in broad sense contains non-additive components of genotypic
variance in addition to additive genetic variance. Genetic advance calculated with the
help of this heritability is usually an over-estimate. However, Panse (1957) has reported
that high heritability in broad sense coupled with higher genetic advance gives an
indication of the predominance of additive effects in controlling the character.
Heritability estimates together with genetic advance are generally regarded to be more
useful in predicting the genetic gain through selection (Johnson et al. 1955). In the
present investigation, high genetic advance in per cent of mean (>20%) was recorded for
number of grains/ spike (31.91%), spikelets/ spike (29.86%) and grain yield/plant
(27.95%), whereas Biological yield (19.38%), harvest index (18.44%), number of
effective tillers/plant (15.79%), plant height (13.52%) and 1000-grain weight (12.76%)
showed moderate estimates of genetic advance in per cent of mean (10-20%) and other
showed low estimates of genetic advance in per cent of mean (<10%) in normal soil
(E1).Under saline-sodic soil (E2), high i.e. >20% genetic advance in per cent of mean was
recorded for number of grains/ spike (30.98%), 1000-grain weight (24.42%), spikelets/
spike (23.60%), harvest index (23.52%) and grain yield/plant (20.82) whereas, plant
height (19.02%), biological yield (17.61%) and number of effective tillers/plant (11.45%)
showed moderate estimates of genetic advance in per cent of mean (10-20%) and other
showed low estimates of genetic advance in per cent of mean (<10%).
High heritability coupled with high genetic advance in per cent of mean was
recorded for number of grains/ spike ear, grain yield/plant and number of spikelets/ spike
in normal soil (E1), while, high heritability coupled with moderate genetic advance in per
cent of mean was recorded for biological yield, harvest index, plant height and 1000-
grain weight. High heritability coupled with low genetic advance in per cent of mean was
recorded for days to 50% flowering and low heritability coupled with high genetic
advance in per cent of mean was recorded for number of effective tillers/plant while; low
heritability coupled with low genetic advance in per cent of mean was recorded for days
to maturity. In saline-sodic soil (E2), high heritability coupled with high genetic advance
in per cent of mean was recorded for number of grains/ spike, 1000-grain weight, harvest
index, number of spikelets/ spike and grain yield/plant while, high heritability coupled
with moderate genetic advance in per cent of mean was recorded for plant height, and
biological yield.
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High heritability coupled with low genetic advance in per cent of mean was
recorded for days to 50% flowering and days to maturity while, low heritability coupled
with moderate genetic advance in per cent of mean was recorded for number of effective
tillers/plant. In earlier studies, high genetic advance in per cent of mean coupled with
high heritability have also been reported for grain yield per plant by Yassin et al. 2015,
Porumb et al. 2016, Ahmadi et al. 2016, Hailu et al. 2016 and Pesaraklu et al. 2016.
The significant and positive gca effects for seed yield per plant were exhibited by
three lines and two tester which in order of merit were NDB 1245, DWRUB 64 and
Karan 741 among lines and NDB 943 and BHS 352 among the testers in normal soil (E1)
while, two lines and two tester in saline sodic soil (E2) which in order of merit were HUB
113 and NDB 1173 among lines and NDB 943 and BHS 352 among testers.
On the basis of gca effects and mean performance, parent Karan 741was found
good combiner for grain yield per plant along with days to 50% flowering, number of
spikelets per spike, number of grains per spike and harvest index for normal soil (E1) and
number of grains per spike and biological yield per plant for saline sodic soil (E2) and
DWRUB 64 for grain yield per plant in addition to days to maturity, 1000 grains weight
and harvest index in normal soil (E1) and reduce to plant height in saline sodic soil (E2)
and NDB 1245 for grain yield per plant in addition to days to maturity, plant height and
harvest index in normal soil (E1) and reduce to plant height, days to 50% flowering and
days to maturity and in addition to1000 grains weight in saline sodic soil (E2). Parent
NDB 1173 for days to 50% flowering, number of spikelets per spike, number of grains
per spike, days to maturity and harvest index in normal soil (E1) and for grain yield per
plant with biological yield per plant in saline sodic soil (E2) and HUB 113 for grain yield
per plant with days to 50% flowering, plant height, number of spikelets per spike, number
of grains per spike, biological yield per plant and harvest index in saline sodic soil (E2).
NDB 943 was found good general combiner for grain yield per plant in addition
to number of effective tillers per plant, number of spikelets per spike, number of grains
per spike and harvest index in normal soil (E1) and grain yield per plant in addition to
days to 50% flowering , number of spikelets per spike, number of grains per spike,
harvest index and reduce plant height for saline sodic soil (E2) while, BHS 352 was found
good general combiner for grain yield per plant in addition to plant height and biological
yield per plant in normal soil (E1) and grain yield per plant in addition to days to
maturity, number of effective tillers per plant, harvest index and reduce days to 50%
flowering for saline sodic soil (E2) .
The available literature also indicates significant and positive gca effects for seed
yield and yield components in barley (Sayed et al. 2008, Bornare et al. 2013, Zhang et al.
103
2015 and Patial et al. 2016). The above maintained lines and testers may be
recommended for exploitation in hybridization programme aimed at improving the yield
components for which they emerged as good general combiners.
The above five parents in E1 and four parents in E2 showing positive and
significant gca effects for seed yield and other important traits as mentioned in above
paragraphs may serve as valuable parents for hybridization programme or multiple
crossing programme for obtaining high yielding variety or transgressive segregants for
developing varieties of barley.
Some other lines identified as good general combiners in desirable direction for
characters other than seed yield per plant are listed in Table 5.1 and 5.2, respectively.
The promising F1’s exhibiting significant sca effects in desirable direction may be
incorporated in future barley improvement programme. The general combining ability
and specific combining ability is associated with interaction effects, which may be due to
dominance and epistatic components of genetic variation that are non-fixable in nature.
The maximum merits of significant and positive sca effects in desirable direction are
presented in Table 5.2(a) & 5.2(b) for normal and saline sodic soil conditions
respectively.
105
5.2.(a). A perusal of Table 5.2.(a) revealed that the good specific combiners involved
parents of low x low, average x high, average x low, low x high and high x high general
combining ability effects. From these findings, it is obvious that best cross combinations
are not always resulted from high x high general combiners, but may also be occurred in
other type of parental combinations. However, in majority of cases, the crosses exhibiting
high sca effects were found to have both or one of the parents as good general combiner
for the characters under study. The results are in agreement with the findings of Singh et
al. (1996), Sharma et al. (2003), Singh et al. (2005), Saad et al. (2005) Sayed et al.
(2008), Verma et al. (2009), Zhang et al. (2015) and Patial et al. (2016).
106
Prakash et al. (2006), Singh et al. (2007), Verma et al. (2009), Zhang et al. (2015) and
Patial et al. (2016).
5.4 Heterosis:
The desirable sca effects may not be of practical utility until and unless per se
performance of the combinations is compared to that of respective better parent and with
standard varieties (SV). In pursuance to this objective, estimates of heterobeltiotic
responses as well as response relative to standard variety SV were computed for all the
characters in different cross combinations.
In the present study, high manifestation of heterosis for seed yield under normal
soil (E1) was observed as evident by the superiority of best crosses over BP ranging from
NDB 1245 X NDB 943 (10.82%) to Azad X BHS 352 (12.22%) and for best crosses
over standard variety, which ranged Azad X DWR 28 (11.74%) from to NDB 1245 X
NDB 943 (53.44%) per cent over SV and under saline sodic soil (E 2) high manifestation
of heterosis for seed yield was observed as evident by the superiority of best crosses over
107
BP ranging from Karan 741 X NDB 943 (11.73%) to HUB 113XNDB 943 (35.61%) and
for best crosses over standard variety, ranged BH 902 X NDB 943 (10.20%) from to
NDB 1465 X NDB 943 (47.45%) per cent over SV,respectively.
A perusal of Table 5.3 and 5.4 revealed that heterosis in grain yield was
proportional to the heterosis observed for yield components. In majority of cases
heterosis in most of the components registered heterosis for grain yield. The top four
crosses showing significant heterobeltiosis for grain yield were also found to register
significant positive heterobeltiosis under normal soil (E1). However, top ten crosses,
which showed positive heterosis over SV for grain yield, were also having positive and
significant standard heterosis for almost characters while, in case of saline soil (E2)
significant heterobeltiosis for grain yield were found to register significant positive
heterobeltiosis. However, top ten crosses, which showed positive heterosis over SV for
grain yield, were also having positive and significant standard heterosis for almost
characters.
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Table 5.1: Summary of general combining ability affects of parents for 10 characters in Barley (Environment-I)
S.No. Line
Days to Days to Effective Spikelets/ Grains/ 1000 Biological Grain Yield/
Plant height Harvest
50% maturity Tillers/ Spike Spike grains yield /plant Plant
Index
flowering Plant weight
1 NDB 1245 0 - - 0 0 0 - 0 + +
2 NDB 1592 + + + - + + 0 + - -
3 NDB 1465 + - + 0 0 0 - + - 0
4 NDB 1173 + + 0 - + + - - + -
5 Karan 741 + 0 0 - + + 0 - + +
6 DWRUB -
- - - - - - + + +
64
7 RD 2794 0 0 - - 0 0 - - 0 -
8 HUB 113 - - 0 - - - - 0 - -
9 BH 902 - - - 0 - - - - - -
10 Azad - - + + - - + + - 0
Testers
1 NDB 943 - - - + + + - - + +
2 BHS 352 - 0 - - - - - + 0 +
3 DWR 28 0 - + - - - - - - -
Where,
+ = Good combiner (Significant and Positive) - = Poor combiner (Significant/Non Significant and Negative)
O = Average combiner (Positive but not significant)
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Table 5.1: Summary of general combining ability affects of parents for 10 characters in Barley (Environment-II)
S.No. Line Days to Days to Plant Effective Spikelets/ Grains/ 1000 Biological Harvest Grain
50% maturity height Tillers/ Spike Spike grains yield Index Yield/
flowering Plant weight /plant Plant
1 NDB 1245 + + + 0 - - + 0 - -
2 NDB 1592 + - + - - - + - - -
3 NDB 1465 - - + 0 0 + + - + 0
4 NDB 1173 - - - - 0 - - + - +
5 Karan 741 0 0 - 0 0 + - + - -
6 DWRUB 64 - - + - - - - - 0 -
7 RD 2794 0 - - - + + - - + -
8 HUB 113 + 0 - 0 + + - + + +
9 BH 902 - - + 0 0 0 - - + -
10 Azad 0 0 - 0 - - + + - -
Testers
1 NDB 943 - 0 + 0 + + + - + +
2 BHS 352 + - - + - - - 0 + +
3 DWR 28 + 0 - - - - - - - -
Where,
+ = Good combiner (Significant and Positive) - = Poor combiner (Significant/Non Significant and Negative)
O = Average combiner (Positive but not significant)
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Table 5.3: Relationship of positive heterobeltiosis for seed yield with heterobeltiosis of other characters (Environment-I)
Characters Grain Plant Effective Spikelets Grains/ Days to Days to 1000 Biological Harvest
Yield/ height Tillers/ / Spike Spike maturity 50% grains yield Index
S.No. Plant Plant flowering weight /plant
Crosses
1. NDB 1245 X NDB 943 + 0 0 + + 0 0 + 0 +
2. + - 0 0 + - 0 0 + -
HUB 113 X BHS 352
3. NDB 1592 X NDB 943 + - 0 0 0 - - + + 0
4. Azad X BHS 352 + - + - - - - + + 0
5. NDB 1592 X BHS 352 0 - + 0 0 - 0 0 0 0
6. Azad X NDB 943 0 - 0 0 0 - - 0 0 0
7. NDB 1173 X NDB 943 0 - 0 + + - - 0 0 0
8. DWRUB 64 X NDB 943 0 - + + + 0 0 + - 0
9. RD 2794 X NDB 943 - - + + + - - 0 - +
10. Karan 741 X DWR 28 - - 0 + + - - + 0 0
Where,
+ = Good combiner (Significant and Positive) - = Poor combiner (Significant/Non Significant and Negative)
O = Average combiner (Positive but not significant)
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Table 5.3: Relationship of positive heterobeltiosis for seed yield with heterobeltiosis of other characters (Environment-II)
Characters Grain Plant Effective Spikelets/ Grains/ Days to Days to 1000 Biological Harvest
S.No. yield per height Tillers/ Spike Spike maturity 50% grains yield Index
plant (g) Plant flowering weight /plant
Crosses
1. HUB 113XNDB 943 0
+ - + 0 + - 0 0 +
2. NDB 1173XNDB +
+ - 0 0 + - - + +
943
3. RD 2794XBHS 352 -
+ - 0 + + + - - +
4. Karan 741XBHS 352 0
+ - 0 0 + 0 0 + 0
5. NDB 1173XBHS -
352 + - 0 0 + + - + +
6. NDB 1465XNDB +
943 + - + 0 + - - - +
7. HUB 113XDWR 28 -
+ + 0 0 + 0 - + 0
8. Karan 741XDWR 28 -
+ + 0 0 + 0 0 0
9. NDB 1592XNDB +
+ - + - - - - + +
943
10. HUB 113XBHS 352 +
+ - + - 0 0 - 0 +
Where,
+ = Good combiner (Significant and Positive) - = Poor combiner (Significant/Non Significant and Negative)
O = Average combiner (Positive but not significant)
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Table 5.4: Relationship of positive heterobeltiosis SV (NDB943) for seed yield with standard heterosis of other characters
(Environment-I)
Where,
+ = Good combiner (Significant and Positive) - = Poor combiner (Significant/Non Significant and Negative)
O = Average combiner (Positive but not significant)
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Table 5.4: Relationship of positive heterobeltiosis SV (NDB 943) for seed yield with standard heterosis of other characters
(Environment-II)
Characters Grain Effective Spikelets/ Grains/ Days to Days to 50% 1000 Biological
S. yield per Tillers/ Spike Spike maturity flowering grains yield /plant
Plant Harvest
plant (g) Plant weight
No. height Index
Crosses
1. NDB 1465XNDB 943
+ - + 0 + - - + 0 +
Where,
+ = Good combiner (Significant and Positive) - = Poor combiner (Significant/Non Significant and Negative)
O = Average combiner (Positive but not significant)
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Table 5.2: Best specific combiners in relation to mean performance of the cross and general combining ability (Environment-I)
S.
Characters Cross with significant effects Mean performance of crosses gca effects of parents
No.
NDB 1245 X BHS 352 78.57 A×A
Karan 741 X NDB 943 78.70 L×A
1 Days to 50% flowering Karan 741 X DWR 28 80.53 L×A
NDB 1465 X DWR 28 81.27 L×A
RD 2794 X BHS 352 82.83 A×A
RD 2794 X BHS 352 114.13 A×A
NDB 1173 X NDB 943 114.33 L×A
2 Days to maturity NDB 1245 X BHS 352 115.00 H×A
NDB 1173 X DWR 28 115.40 L×A
NDB 1465 X DWR 28 115.90 A×A
NDB 1592 X DWR 28 71.77 L×L
HUB 113 X DWR 28 73.13 A×L
3 Plant height NDB 1465 X DWR 28 73.93 L×L
Karan 741 X DWR 28 74.37 A×L
NDB 1173 X NDB 943 75.00 A×H
Azad X BHS 352 7.13 H×A
NDB 1465 X NDB 943 7.10 A×H
Number of effective
4 RD 2794 X NDB 943 7.07 A×H
tillers/plant
Karan 741 X BHS 352 7.03 A×A
BH 902 X DWR 28 6.97 A×A
NDB 1245 X NDB 943 25.50 A×H
NDB 1592 X NDB 943 25.33. H×H
5 Spikelets/ Spike NDB 1465 X NDB 943 25.27 A×H
HUB 113 X DWR 28 25.17 L×L
NDB 1592 X BHS 352 24.83 A×L
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NDB 1245 X NDB 943 63.83 A×H
NDB 1465 X NDB 943 63.70 A×H
6 Grains/ Spike NDB 1592 X NDB 943 63.47 H×H
Karan 741 X DWR 28 62.27 H×L
NDB 1592 X BHS 352 62.17 H×L
NDB 1592 X NDB 943 42.37 A×A
DWRUB 64 X BHS 352 41.50 H×A
7 1000 grains weight Azad X BHS 352 40.40 H×A
Karan 741 X DWR 28 40.27 A×A
NDB 1245 X NDB 943 40.27 A×A
HUB 113 X BHS 352 36.23 A×H
Azad X BHS 352 35.43 H×H
8 Biological yield /plant NDB 1465 X DWR 28 33.33 H×A
NDB 1592 X DWR 28 32.37 H×A
NDB 1592 X NDB 943 31.93 H×L
NDB 1245 X NDB 943 41.00 H×H
DWRUB 64 X NDB 943 39.73 H×H
9 Harvest index RD 2794 X NDB 943 38.77 A×H
HUB 113 X NDB 943 38.53 A×H
NDB 1173 X BHS 352 38.27 H×A
NDB 1245 X NDB 943 12.63 H×H
DWRUB 64 X NDB 943 12.40 H×H
10 Grain Yield/ Plant HUB 113 X BHS 352 11.93 A×H
NDB 1465 X DWR 28 11.73 A×L
NDB 1173 X NDB 943 11.67 A×H
H = High (significant and positive), L= Low (significant and negative), A = Average (non-significant)
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Table 5.2: Best specific combiners in relation to mean performance of the cross and general combining ability (Environment-II)
S.
Characters Cross with significant effects Mean performance of crosses gca effects of parents
No.
HUB 113 X BHS 352 72.10 L×L
HUB 113 X DWR 28 75.47 L×L
1 Days to 50% flowering NDB 1245 X DWR 28 77.80 L×L
NDB 1245 X BHS 352 78.27 L×L
NDB 1592 X DWR 28 78.30 L×L
NDB 1465 X NDB 943 114.40 A×A
Azad X NDB 943 114.57 A×A
2 Days to maturity NDB 1245 X BHS 352 115.33 L×H
NDB 1592 X BHS 352 115.57 A×H
NDB 1245 X NDB 943 115.80 L×A
NDB 1245 X BHS 352 69.00 L×A
BH 902 X NDB 943 69.93 L×L
3 Plant height NDB 1245 X NDB 943 71.33 L×L
NDB 1465 X DWR 28 71.43 L×A
BH 902 X DWR 28 71.53 L×A
NDB 1245 X NDB 943 6.17 A×A
NDB 1465 X NDB 943 5.87 A×A
Number of effective
4 BH 902 X BHS 352 5.80 A×H
tillers/plant
DWRUB 64 X NDB 943 5.73 A×A
HUB 113 X BHS 352 5.70 A×H
NDB 1245 X NDB 943 18.80 L×H
BH 902 X NDB 943 17.97 A×H
5 Spikelets/ Spike HUB 113 X DWR 28 17.90 H×L
RD 2794 X BHS 352 17.50 H×L
Karan 741 X NDB 943 17.37 A×H
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NDB 1245 X NDB 943 49.73 A×H
RD 2794 X BHS 352 49.13 H×H
6 Grains/ Spike HUB 113 X DWR 28 47.70 H×A
RD 2794 X DWR 28 47.43 H×A
NDB 1465 X DWR 28 45.97 H×A
NDB 1592 X NDB 943 37.07 H×H
A×H
NDB 1173 X NDB 943 36.90
H×H
7 1000 grains weight NDB 1245 X NDB 943 35.57
L×L
HUB 113 X BHS 352 35.33
H×H
NDB 1465 X NDB 943 34.37
Karan 741 X BHS 352 26.67 H×A
NDB 1173 X BHS 352 24.97 H×A
8 Biological yield /plant Azad X DWR 28 24.40 H×A
HUB 113 X DWR 28 24.20 H×A
NDB 1173 X NDB 943 24.00 H×A
NDB 1465 X NDB 943 47.30 H×H
BH 902 X BHS 352 45.27 H×H
9 Harvest index RD 2794 X BHS 352 42.43 H×H
DWRUB 64 X BHS 352 42.33 A×H
HUB 113 X NDB 943 41.40 H×H
NDB 1465 X NDB 943 9.63 A×H
HUB 113 X NDB 943 9.27 H×H
10 Grain Yield/ Plant NDB 1245 X NDB 943 9.07 A×H
NDB 1173 X NDB 943 9.00 H×H
BH 902 X BHS 352 8.90 A×H
H = High (significant and positive), L= Low (significant and negative), A = Average (non-significant)
118