6. Arida (Genetic) Revisi Daf...


BIOTROPIA Vol. 20 No. 2, 2013: 122 - 133

GENETIC VARIATION, HERITABILITY AND
CORRELATION BETWEEN RESIN PRODUCTION

CHARACTER OF HIGH RESIN
YIELDER (Hry)

Pinus merkusii

ARIDA SUSILOWATI *, ISKANDAR Z. SIREGAR , SUPRIYANTO ,
IMAM WAHYUDI and CORRYANTI

(Kopertis Wilayah I Sumut-NAD)

Recipient of BIOTROP Research Grant 2010/Accepted 18 April 2013

Tree improvement programs for high resin yielder were started in 2006 through
a series of survey and morphological identification of candidate trees with high resin
production. Specific information about genetic parameter of resin yielder candidates in
Cijambu Seedling Seed Orchard is still not determined yet, although based on resin distribution
trend this SSO has the highest mean of resin production. In this research, individual and family
heritability, coefficient genetic variation and genetic-phenotypic correlation were estimated for
resin production and growth data from 15 open polinated families of planted in
1982 (set 1) - 1983 (set 2) in Cijambu SSO. The results showed high value of coefficient genetic
variation (CVG: 14.5-28.43%), individual narrow sense heritability values for resin production
character (0.58-0.77) which resemble with previous researches. This indicates that genetic
factor was dominant for resin production and selection activities were effective to get high
yielder superior candidate. Phenotypic and genotypic correlation found that bark thickness,
crown length and stem diameter character was positively significant correlated to resin
production, whereas severity attack level of pests-diseases and number of branches were
negatively significant correlated to resin production.

Selection, high resin yielder, , heritability, phenotypic

1 2 2

3 4

1

2

3

4

Coordinatior of Private Higher Education Region I North Sumatera-Aceh, Indonesia

Silviculture Department, Faculty of Forestry, Bogor Agricultural University, Bogor, Indonesia

Forest Product Department, Faculty of Forestry Bogor Agricultural University, Bogor, Indonesia

Research and Development Center of Perum Perhutani, Bogor, Indonesia

P. merkusii

Pinus merkusii

P. merkusii, variation

ABSTRACT

Key words:

* Corresponding author : arida_iswanto@yahoo.co.id

DOI: 10.11598/btb.2013.20.2.1

122



INTRODUCTION

Pinus merkusii

et al

P. merkusii

P. merkusii

Pinus merkusii

is known as an important industrial species for pulp and paper and
sawn wood, gum rosin production, as well as considerable species for reforestation
and land rehabilitation in Indonesia (Suhardi . 1994). One valuable product of
resin and very demanded by international market is gondorukem (gum rosin).
Gondorukem is a potential product, grouped under pine chemical products and it
plays an important role as non timber forest product in Indonesia because it provides
high national income about US$ 50 million/year as well as more job opportunities
(Fachrodji 2010).

Problems faced in gondorukem export was lower productivity so that Indonesia
still ranks as third position after China and Brazil as gum rosin producer (Cunningham
2006). In order to sustain Indonesian gondorukem export, several activities have been
undertaken to increase resin production through tree breeding activities, application
of improved silvicultural techniques, improvement of tapping techniques and
management (Fahrodji 2010). Breeding activities focusing on resin production is the
most prospective way to be developed since early studies conducted in 2006, have
resulted to several plus trees producing high yield of resin (high resin yielder). High
resin yielder is the term given for pine genotype producing over 50 g/tree/3 days
(Fakultas Kehutanan UGM 2006) higher than the current production (21 g/tree/3
days).

breeding activities in Indonesia were started since 1976 through a series
of progenies test and descendants of plus tree selection which was focused on
stem straightness character (Soeseno 1988), while resin production is only a side
product. In considering high value of resin products, pine breeding activities were
recently conducted to obtain plus trees for resin production (Soeseno 2001). For
breeding activities concentrating on resin yielder, it is important to collect information
on genetic parameters such as coefficient of genetic variation, heritability and
association of characters related to resin production for directing efficient selection.
Several researches on conducted in early establishment of Seedling Seed
Orchard (SSO) in Java and other pine species showed that resin production character
has high heritability value, but specific information of Cijambu SSO is not determined
yet. Therefore, research on "Genetic variation, heritability and correlation between
resin production characters of High Resin Yielder (HRY) in Cijambu
Seedling Seed Orchard (SSO)" is needed. The result will be used to provide best
possible trees for further genetic improvement as well as to establish plantations using
improved genetic materials.

Experiment was carried out in Cijambu SSO, Sumedang. Data for this research
were obtained from progenies test plantation established in 1982 (Set 1) and 1983
(set 2). At early plantation the research focused on stem character, where 200 families

MATERIALS AND METHODS

123

Genetic variation, heritability and correlation between resin production character Arida Susilowati– et al.



were planted/year designed in Completely Randomized Block Design, 5-tree line
plot with ten blocks as replication at a spacing of 3 x 3 m. Together with selective
thinning leaving only 2-3 trees per plot. Second selection focused on resin production
conducted in 2006 found 96 families with 110 plus trees in progenies trial planted in
1978-1983. Planting year 1982 involved 20 open pollinated families and in 1983
involved 25 open pollinated families of resin yielder candidates spread in 2 different
blocks. For CVG and heritability estimation, 15 selected open polinated families were
used. In this research two assumption and 2 types of estimation were used, those
were: 1) Resin production was not different at the age 29 and 30 years, for these reason
we combined 2 set of families (1982 and 1983) into one estimation and 2). Resin
production was different at the age of 29 and 30 years so we used separate estimation
using set 1 and set 2.

Characters studied in this research were resin production and other quantitative
characters which assume to be related to resin production those were: total height
(TH), clear bole height (CBH), diameter, crown length (CL), crown width (CW)
number of branches (NB), first branch angle (FBA), bark thickness (BT), severity
attack level from pest and disease (SL), clear bole volume (VBC) and total volume
(Vtot). All characters were measured based on previous methods for forest trees
developed by Bacilieri (1996); Cantini (1999); Kremer (2002); Ginwal

(2004); Weber and Montes (2005); Baliuckas (2005) and Devagiri (2007).
Resin production was measured by calculating resin tapping weight for 3 days.

Analyses of variance (ANOVA) were carried out on Single Tree Plot design
according to Isik (2008) using individual tree basis for each progeny test with following
linear model:

Y : μ+ B + F +e

where: Y : individual tree observation; μ: the overall mean; B : the effect of i

block; F :the effect of j family; e : within plot error. Only 2 replications were included in
the analyses. Variance and covariance component were calculated by equating the
means square or mean cross product to their expectation. As progeny
trial assumed to be half-sib, heritabilities and standar error (S.E) were estimated by
using single site analyses formula:

heritability of family means:

h f/ + /nb+ ), with S.E h f: (S.E f)/ + /nb+ ),

individual tree heritability:

h : 4 + ), with S.E. h : (4S.E f)/ ( + )

where: h f: family mean heritability; h :individual tree heritability f: the
component due to family means : the component of block : the component of

variance within plot. The standard error of family component variance (S.E f) was
calculated by Anderson & Bancroft (1952) Hardiyanto (1996):

S.E f: [2/k (MS ) / (df +2)]

,

et al. et al. et al.
et al. et al. et al.

( (

in

Data Analysis

ijk i j ijk

ijk i

j ijk

f b e f b e

f b+ e f b+ e

b e

i i i

th

th

2 2 2 2 2 2 2 2 2 2

2 2 2 2 2 2 2 2 2 2

2 2 2

2 2

2

2 2 2 0.5

f : σ σ σ σ σ σ σ σ

f/ (σ σ σ σ σ σ

; σ
; σ ; σ

σ

σ

δ δ

BIOTROPIA Vol. 20 No. 2, 2013

124



i:

i.

where: k is the coefficient of family means square; MS the i means square used to

estimate n component; df:the number of degree of freedom for MS Criteria for

heritability f) was calculated according to Cotteril and Dean (1990).

Estimates of genetic variation coefficient performed using Cornelius (1994)
formula as follows:

Genetic and phenotypic correlation between resin production character and
growth character estimated using statistical formula as follows:

Estimates of variances and allied statistics for contributing sources of variability
are presented for each trait in Table 1. Based on Cornelius (1994) classification, genetic
variation in Cijambu SSO showed variation value with total character contribution
ranging from 4.05% (total height character) to 28.70% (severity attack level from pest
and disease). This result indicated that not all of the characters observed in Cijambu
SSO were influenced by genetic factors. Resin production and severity attack level from
pest-disease character have high value of coefficient genetic variations. Diameter,
crown, bark thickness, number of branches, clear bole volume, total volume, have
moderate value of coefficient genetic variations, while total height, clear bole height
and branching angle in set 2 have lower value of coefficient genetic variations.

High CVG value represents resin production character (14.5-28.43%) and severity
attack level from pest-disease (24.0- 28.70%). It indicates that genetic factors will have
a big impact on the appearance of characters. High value for resin production
character was obtained compared to previous research conducted by Roberds
(2003) in SSO (CVG:13.7%). High value for severity attack level from pest-
disease was also compared to Bastein & Alia (2000) in (CVG:26.8%). In
order to tree improvement focused on resin production in Cijambu SSO, these
characters are important to be studied in detail because of the influence of genetic
factors. CVG value for total height, clear bole height, crown width, crown length and
number of branches character has lower value. It indicated that these characters
belong to vegetative characters which is influenced by environmental factors.

th

th

2
(h

et al.
P.taeda

P. sylvestris

Coefficient Genetic Variation (CVG)

RESULTS AND DISCUSSION

Genetic variation, heritability and correlation between resin production character Arida Susilowati– et al.

125



Heritability

Based on Cotteril and Dean (1990) classification, heritability value for resin
production character at Cijambu SSO was high (h2f:0.700.09-0.820.08 and
h2:0.580.08-0.770.08), severity attack level from pest-disease also has high heritability
value (h2f: 0.640.02-0.800.16 and h2:0.640.13-0.690.04). Stem diameter, bark
thickness, crown length, number of branches and volume have lower to moderate
heritability value (Table 1).

Characters σ2f σ2b σ2e
CVG
(%) h

2f SE h2 SE

29 and 30 years old
Resin production days
(g/3)

581.8 338.1 86.1 21.6 0.70 0.09 0.58 0.08

Total height (m) 1.94 2.58 11.07 6.0 0.14 0.05 0.12 0.05
Clear bole height (m) 3.40 7.0 3.39 6.6 0.33 0.23 0.25 0.17
Diameter (m) 0.002 0.007 0.002 10.1 0.29 0.72 0.20 0.52
Bark thickness (cm) 0.36 0.89 0.26 8.8 0.34 0..83 0.24 0.58
Number of branches 55.92 69.15 33.17 12.8 0.45 0.10 0.35 0.08
Crown length (m) 1.507 3.122 1.74 10.6 0.31 0.30 0.24 0.23
Crown width (m) 1.272 2.96 1.55 12.5 0.30 0.30 0.22 0.21
First branch angle 105.97 34.32 58.12 11.2 0.58 0.10 0.53 0.09
Clear bole volume (m3) 0.002 0.009 0.04 13.9 0.35 0.22 0.33 0.20
Total volume (m3) 0.007 0.34 0.14 9.4 0.18 0.60 0.12 0.42
Severity attack level
from pest and disease

139.91 15.75 49.86 28.2 0.71 0.13 0.68 0.12

Set 1 (29 years old)

Resin production days
(g/3)

200.70 53.94 46.82 14.5 0.72 0.13 0.67 0.19

Total height (m) 0.67 3.5 21.83 3.3 0.04 0.01 0.03 0.03
Clear bole height (m) 1.69 3.5 6.08 4.89 0.21 0.11 0.15 0.13
Diameter (m) 0.001 0.0005 0.002 7.9 0.47 0.15 0.36 0.04
Bark thickness (cm) 0.27 0.36 0.12 6.5 0.39 0.15 0.36 0.54
Number of branches 15.73 77.16 40.13 13.2 0.14 0.03 0.12 0.09
Crown length (m) 2.42 3.02 1.13 13.0 0.40 0.48 0.37 0.18
Crown width (m) 1.11 1.56 3.26 10.3 0.26 0.18 0.19 0.05
First branch angle 6.4 36.15 106.2 2.8 0.07 0.01 0.04 0.02
Clear bole volume (m3) 0.02 0.002 0.07 13.1 0.31 0.16 0.19 0.06
Total volume (m3) 0.06 0.07 0.23 13.8 0.25 0.15 0.17 0.37
Severity attack level
from pest and disease

95.59 5.29 37.83 24.0 0.80 0.16 0.69 0.04

Table 1. Family f), block variance b), within plot error e), coefficient
of Genetic variation (CVG), family heritability (h f) and individual tree heritability for
resin production and growth characters

variance (σ (σ variance (σ
2 2 2

2

BIOTROPIA Vol. 20 No. 2, 2013

126



P. merkusii
et al. P. eliotii et al.

P. pinaster et al. P. taeda

et al. et al. et al.

et al. et al.

et al. et al.

Heritability for resin production shows high value (0.580.08-0.770.08). It seems to
be slightly different than previous research results (0.69) conducted by Leksono (1990)
at Cijambu and Sempolan on 12 years old progenies test plantation. This
result was higher than reported by Zhang (2010) on (0.37), Tadesse
(2001) on (0.5) and Roberds (2003) on (0.44 to 0.59), and similar
value was also found at stem diameter and branching quality. Higher value in this
research compared to previous researches at early SSO establishment suggested
that the materials used for this research originated from second selection focused for
resin production. For high resin production heritability values, Wenger (1984);
Burczyk (1998); Kassuth (1984); Mergen 1955, Gill (1998) explained that
resin production character is controlled by gene. This indicates that improvement
program for resin production character through genetic selection would provide
higher genetic gain not affected by other interaction factors.

Heritability value for height and stem diameter were slightly different from
previous researches conducted by Hardiyanto (1996) who focused on stem
straightness, and Leksono (1996) for resin yield at 12 years old progenies test (h2f: 0.40
dan h2f: 0.43). Heritability value for branching trait character is lowe in set 1 (0.12
and 0.07) but in set 2 the heritability value is high (0.44 dan 0.71). The changes of
heritability value in long rotation crops such as a tree is not surprising since genes
involved in growth may change with age (Namkoong 1980; Monteuis 2011),
and these changes also may be related to different growth phases (Franklin 1979).
Change of heritability value for diameter, tree height and bark thickness character at
different age probably is influenced by silvicultural practices such as thinning and
other management practices (Gwaze 1997 & Lopez-Upton 1999).

Result from CVG and heritability estimation in Cijambu SSO showed high value
for resin production, it indicated that more dominant genetic factors determine this
character. Based on this value, tree improvement program for high resin yielder can be
initially conducted by mass selection of individual trees with high resin production.

Characters σ2f σ2b σ2e
CVG
(%) h

2f SE h2 SE

Set 2 (30 years old)

Resin production (g/3
days)

1012.5 151.2
5

151.3 28.43 0.82 0.084 0.77 0.08

Total height (m) 1.15 1.02 0.86 4.64 0.46 0.06 0.38 0.52
Clear bole height (m) 3.08 6.75 0.33 4.91 0.45 0.09 0.30 0.06
Diameter (m) 0.002 0.01 0.001 8.49 0.19 0.07 0.12 0.411
Bark thickness (cm) 0.09 0.19 0.38 10.93 0.16 0.01 0.14 0.29
Number of branches 66.50 14.09 5.61 8.12 0.84 0.05 0.70 0.41
Crown length (m) 0.82 0.05 1.007 7.81 0.44 0.06 0.44 0.55
Crown width (m) 0.50 0 0.48 7.84 0.51 0.09 0.51 0.01
First branch angle 201.95 18.75 75 5.26 0.71 0.10 0.68 0.10
Clear bole volume (m3) 0.0090.04 0.03 12.04 0.62 0.09 0.50 0.06
Total volume (m3) 0.14 0.33 0.009 13.06 0.38 0.08 0.26 0.07

Table 1. Continued

Genetic variation, heritability and correlation between resin production character Arida Susilowati– et al.

127



BIOTROPIA Vol. 20 No. 2, 2013

This condition agrees with Tadesse . (2001) who stated that when a population
has high heritability value for a character, mass selection method would be more
efficient for improving the character. Furthermore, White . (2007) stated that mass
selection was appropriate to be implemented in early selection and for high heritability
value characters because the phenotypes of individual trees describe its genetic ability.

Phenotypic and genotypic correlation (Table 2) showed low to moderate
coefficient value, indicated that not overall phenotypic appearance describing genetic
expression because interaction between environment and genetic factor also
influencing phenotypic expression of trees. Phenotypic correlation showed resin
production character was positively significant correlated with stem diameter, bark
thickness and crown length. Furthermore, number of branches and severity attack
level from pest -disease character was negatively significant correlated with resin
production. Positive correlation between resin productions with some characters,
indicated that resin production will increase equally with the increase of character
value. On the other hand, a negative correlation between resin production with some
characters indicates that resin production will decrease with high value of character
components.

Genetic correlation between resin production and stem diameter, bark thickness,
and crown length indicated that resin production will increasedequally with the
increase of these characters. On the contrary resin production will decrease equally
with the increase of branching number and severity attack level of pest-disease.
Different results were obtained in set 2 (30 years old progenies trial), in set 2 we found
negative correlation between resin production and tree height, it indicates that resin
production will decrase with the increase of tree height.

Correlation between resin production with stem diameter and crown length are in
accordance with previous research results conducted by Pswaray (1996); Coppen

(1984) on Westbork (2011) on and Tadesse (2001) on
. Panshin & De Zeeuw (1984) also explained that a good tree is

characterized by large diameter and large crown size because trees need more wider
light absorbance for photosynthesis process. Correlation between resin production
and stem diameter have been reported also by Coppen 1984 who stated that wider
diameter trees have wider annual ring and giving great chance to have more resin ducts
and produce higher resin yield than smaller diameter trees.

Number of branches and severity attack level from pest and disease have
negatively significant effect to resin production meaning that resin production
decreased with the increasing number of branches and severity attack level from pest.
Papajiannopoulos (2002) in also found that tree with canopy openes (lower
branch number) have higher resin yield compared with higher branch number which
assume to be related to photosyntetic process, resin viscocity and accumulation.
Correlation between resin production and severity attack level from pest-disease have
been deeply studied in other pine species and conifers. Previous researches conducted

et al

et al

et al.
et al. P.elliottii; P.taeda et al.
P.pinaster naval store

et al.

P.halepensis

Phenotypic and Genetic Correlation Between Growth Character and Resin
Production

128



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Genetic variation, heritability and correlation between resin production character Arida Susilowati– et al.

129



BIOTROPIA Vol. 20 No. 2, 2013

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130



in temperate regions by Kleinhentz . (1998); Blada (2000); Kim . (2003); Rafael
. (2005) concluded that pest and disease caused significant decrease of resin

production quantities. Furthermore, Raffa & Berryman (1982) in also found
severity attack level which caused loss of resin yield quantities. However, specific
studies about correlation between resin production and severity attack level in
Cijambu SSO have not been conducted yet. From this research we found that both
resin production and severity attack level in Cijambu SSO have high CVG value, it
indicated that selection activities focused for resin production also can escorted
together with resistance for pest and disease.

Although some characters have a correlation to resin production, further research
still need to be conducted because phenotypic observation is influenced by growth
phase and environment, so it could not differentiate recessive genotype with resemble
morphological appearance such as secondary metabolites (Finkeldey 2005). To
overcome this problem molecular marker such as RFLP, RAPD, AFLP and
microsatellite can be used.

Results from genetic variation and heritability estimation of resin yielder
candidates character in Cijambu SSO showed high coefficient of genetic variation
value (CVG: 14.5-28.43%) and heritability value (h2:0.580.08-0.770.08) for resin
production character. It indicated that genetic factor strongly affected resin
production character.

Genetic and phenotypic correlation found that stem diameter, bark thickness and
crown length character were positively significant correlated to the resin production,
whereas severity attack level from pest-disease and number of branches was negatively
significant correlated to resin production. It indicated t genetically improvement on
stem diameter, bark thickness and crown length increasing resin production.
Whereas, higher severity attack level from pest-disease and number of branches
decreased resin production.

This research was part of Arida Susilowati's studies toward doctorate degree from
Bogor Agricultural University (IPB). I would like to express my sincerest thank to
SEAMEO-BIOTROP for supporting my research with

and Development Center of the Perum Perhutani for the
access to their experimental plot in Cijambu SSO and in providing samples as well as
for technical assistance during the fieldwork.

et al et al
et al

P.taeda

PhD research grant. My sincerest
appreciation also goes to Research

CONCLUSIONS

ACKNOWLEDGMENTS

Genetic variation, heritability and correlation between resin production character Arida Susilowati– et al.

131



BIOTROPIA Vol. 20 No. 2, 2013

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