Oil Palm CHP Aik Chin
Oil Palm CHP Aik Chin
Oil Palm CHP Aik Chin
Oil Palm
Aik Chin Soh, Choo Kien Wong, Yuk Wah Ho, and Chieh Wean Choong
11.1 Introduction
The oil palm is the world’s most important oil crop producing 24.9% of total
vegetable oils and fats surpassing soybean at 23.9% (Mielke, http://www.oil
world.bz, 31 March 2007). It produces two types of oil from its fruits, meso-
carp oil and kernel oil known as crude palm oil (CPO) and palm kernel oil
(PKO), respectively, in international trade. Total world production of CPO
stands at about 38 million tons worth around US$ 20 billion. The oils are
produced from some 13 million ha of plantations in the humid tropical
countries of Asia, Africa and Latin America: Indonesia (5.3 million ha),
Malaysia (4.2 million ha), Papua New Guinea, Colombia, Ivory Coast,
Nigeria and Thailand with the first two countries having the bulk of the
plantings. Palm oil is the largest internationally traded vegetable oil with its
main markets in China, European Union, Pakistan, India, Japan and Bangla-
desh. Palm oil is mainly used in food (80%), e.g. as cooking oil, margarine,
vanaspati or vegetable ghee and shortenings, and the remaining 20% are used
as oleochemicals replacing mineral oil to feed the detergents, cosmetics,
pharmaceutical/nutraceutical, plastics and lubricants industries. With the
recent high rise in petroleum prices and that the deadline for meeting the
requirements of the Cartagena Protocol on Biosafety in terms of ‘green’ or
renewable energy substitution is approaching, there has been a tremendous
demand for palm oil as a source of biofuel (biodiesel). Also, responding to
consumer health and environmental concerns, secondary and by-products
from the palm oil industry have spawned new industries, e.g. vitamins A and
E and other antioxidant health supplements from the oil, animal feed and
organic fertilizers from the kernel, and sludge cakes and wastes from oil
extraction mills have served as value additions.
The oil palm has been postulated to originate in Gondwanaland which disap-
peared when the American and African continents drifted apart in prehistoric
times (Zeven 1965) giving rise to the evolution of African oil palm (Elaeis
guineensis Jacq.) and American oil palm (E. oleifera or E. melanoccoca pre-
viously). The African oil palm is endemic to the equatorial belt of Africa
stretching eastwards from Guinea at the Atlantic coast to Madagascar Island
in the Indian Ocean and from Senegal in the sub-Sahara to the Angola-Namibia
border in the south. However, the centres of origin and diversity appear to be
concentrated in the tropical forests of the west (Ivory Coast, Nigeria, Ghana,
Cameroon) and central (Congo, Zaire, Angola) African countries where they
occur as semi-wild forest groves fringing rivers in the lowlands usually close to
settlements, although they can thrive in drier and higher areas (Hartley 1988;
Latiff 2000). The American counterpart is endemic to the tropical countries of
Central and South America, from Mexico in the north to the Amazon in the
south and from the Pacific to the Atlantic coasts. Its distribution appeared to be
more discontinuous, it is also found in open grasslands and river banks and
associated with native Indian migratory trails (Santos et al. 1986). The oil palm
fruit forms an important part of both the native West African and South
American diet as a source of fat, carbohydrate, protein and vitamins.
Interest in oil palm as an industrial crop arose from the need to substitute
animal fat in the production of candle wax, soap and margarine. The European
colonists started oil palm plantations in Indonesia and Malaysia to ensure a
steady supply of oil. Four seedlings planted in Bogor Botanic Gardens in
Indonesia derived apparently from the same fruit bunch in West Africa and
obtained via Amsterdam and Mauritius/Reunion, gave rise to the current
industry. Hybridizations and selections among the progenies of these four
progenitors, which had thick shell fruits or Dura (D) fruit form, were distrib-
uted to the plantations in Deli province in Sumatra and thence to Malaysia
(Rosenquist 1986). These Deli Ds were the commercial planting material for the
rapidly developing plantation industry in Malaysia and Indonesia from 1911 till
the early 1960s. Beirnart and Vanderweyen (1941) elucidated the monogenic
inheritance of the shell gene: the cross between the thick shell D parent palm and
the shell-less (usually female sterile) pisifera (P) parent would give rise to 100%
thin-shell tenera (T) palms (Fig. 11.1) exhibiting incomplete dominance of the
shell gene; the T T, the T P and D T crosses would give rise to segregating
progenies in the classical Mendelian ratios of 1D:2T:1P for the first (Fig. 11.2),
1T:1P for the second and 1D:1T for the last cross, respectively. Teneras from
West Africa have been brought in by breeders and with this revelation the
switch to the T hybrid as the commercial material was very rapid (Hartley
1988). This process was spurred on by private plantation companies becoming
interested in commercial hybrid seed production and investing in oil palm
breeding. Mixed T or D P hybrids have been the dominant commercial
11 Oil Palm 335
planting materials till today. Recently, oil palm clones from tissue culture have
become commercially available although still in limited quantities as compared
to the total demand for oil palm seeds (Soh et al. 2006).
In oil palm, there are no varieties in the strict sense as the commercial materials
are mixed hybrids from non-fully inbred and sometimes out-crossed parents
(Soh 1999). They can therefore be considered as inter-population hybrids. The
parent populations e.g. Deli, were usually derived from very few progenitors
336 A.C. Soh et al.
Costa Rica and Thailand. This BPRO confers smaller fruits but with high oil
content to their progenies.
Calabar. The genetic base of the Nigerian Institute for Oil Palm Research is
much broader from collections made in Aba, Calabar, Ufuma and Umuaibi.
Progenies from its Calabar selections, particularly palm NF 32.3005 have been
distributed to Ghana, Costa Rica, Indonesia and Malaysia.
Derived and recombinant BPROs. A number programmes have initiated the
recombinant phase of breeding having interbred or introgressed parents from
various traditional BPROs to form new BPROs with mixed lineages or recom-
binant BPROs, e.g. URT (Ulu Remis teneras), Dumpy.AVROS, Dumpy.
Yangambi.AVROS, La Me x Yangambi, La Me x Dumpy.AVROS (Soh
et al. 2006).
Recognizing the very narrow genetic base (Deli, AVROS, Yangambi, La Me)
of the existing oil palm breeding programmes and considering that Malaysia
had a very high stake in the rapidly expanding industry, MARDI (Malaysian
Agricultural Research and Development Institute) and subsequently PORIM
(Palm Oil Research Institute of Malaysia), currently known as MPOB
(Malaysian Palm Oil Board), launched a series of expeditions to Africa and
Latin America to prospect for new E. guineensis and E. oleifera germplasm for
genetic base broadening, breeding and conservation in its ex situ genebank
(Ooi et al. 1973; Rajanaidu et al. 1979; Obasola et al. 1983; Rajanaidu and
Rao 1988). The first prospection was systematically done in Nigeria in colla-
boration with NIFOR and the then International Board of Genetic Resources
with the objectives of genetic conservation and studying its population genetic
structure. Genetic studies done on the prospected progenies revealed that
genetic variability was higher within families than between families and popu-
lations. Subsequent collections in Angola, Cameroon, Gambia, Guinea,
Madagascar, Sierra Leone, Tanzania and Zaire were guided by this finding
and also targeted towards accessions with prospective economic and agro-
nomic traits. Prospections for E. oleifera in Latin America were carried out in
Brazil, Colombia, Costa Rica, Ecuador, Honduras, Panama, Peru and Sur-
iname and included also other oil bearing palms (Bactris gossipaes or Pejibaye,
Jessenia-Oenocarpus, Orbignya martiana or Babassu) with unusual fatty acids
and other uses. The accessions have been planted and maintained as living
collections in Malaysia with a sample retained by the host country (Rajanaidu
1990; Rajanaidu and Jalani 1994). These prospections proved to be scientifi-
cally successful in that a number of extremely desirable traits and traits absent
in existing breeding populations have been discovered (Table 11.1) and intro-
gressed into advanced breeding populations or developed into new popula-
tions (Sharma 1999; Rajanaidu et al. 2000).
338 A.C. Soh et al.
Table 11.1 Oil palm collections from various countries and their special attributes
Accessions Useful traits
A. E. guineensis
Nigeria High in unsaturated fatty acids, dwarfness, high stearic acid, low lipase
Angola Large fruits, high carotene content
Zaire Tolerance to Ganoderma disease
Cameroon Tolerance to Ganoderma disease
Tanzania Thin shell
B. E. oleifera
Colombia Very high unsaturated fatty acids content
Brazil Thicker oil bearing mesocarp
Costa Rica Compact stature, good oil yielding
Ecuador Thicker oil bearing mesocarp, very small palms
Suriname Very compact palm tree stature
Davidson (1993) attributed 70% of the oil palm yield improvement in Malaysia
for the previous 50 years to breeding improvement and 30% to improved
agronomic practices. Undeniably, the mere switch-over from the thick shell,
thinner oil-bearing mesocarp (ca. 60% mesocarp to fruit content) D to the thin
shell, thicker mesocarp T (ca. 80%) variety would account for at least 30% of
the yield increase, notwithstanding improved FFB yield. There have been at
least two generations (ca. 20 years) of improved T materials since and oil yields
have improved from about 5 t/ha/yr to about 10 t/ha/yr based on trial results
then. Hardon et al. (1987) estimated that there had been an average yield
improvement of about 15% per generation over two generations of breeding
in the D, but they did not translate this into T hybrid improvement. Subse-
quently, Lee et al. (1990) and Rajanaidu et al. (1990) estimated a resultant T
improvement of only 6–7% when they progeny tested the same Ps on two
successive generations of selected Ds. Lee and Yeow (1985) reported that
selecting the best P (top 15%) from the progeny-test of seven Ps would give
12% improvement. In commercial seed production, at least 3–4 progeny tested
Ps (top 30–50% selection) need to be used reducing the improvement to about
5%. Reconciling the estimated improvements in the two parental populations,
the estimated improvement of 10–15% per generation for the T hybrids was not
unreasonable (Soh et al. 2003a). The grossness of the estimate is inevitable as
there were no common standard crosses linking the different trials of different
generation materials and the difficulty of identifying a standard (single hybrid,
sample of mixed hybrids) treatment to represent a particular generation of
commercial mixed hybrids. Breeding programmes based on the MRRS (mod-
ified reciprocal recurrent selection) system (Soh et al. 1999) were better placed
11 Oil Palm 339
The projected 30% yield increase by cloning the best individual palms from
commercial mixed hybrids provided the impetus in the development of the
tissue culture clonal propagation technique of the oil palm (Jones 1974; Hardon
et al. 1987; Meunier et al. 1988). Soh (1986), however, contended that the likely
increase would be about 13% with the first cloning, based on general theory
because of the low heritability for yield in advanced commercial hybrids. The
results from the CIRAD group comparing the improvements made by their
clones and improved hybrids from the MRRS programme were in general
agreement with Soh’s estimate (Nouy et al. 2006). However, the mean yield
advantage of clones over commercial hybrids summarized from five trials
testing 68 clones by AAR (Applied Agricultural Resources) was about 18%.
There were clones exceeding hybrids by 30% which could be recloned, but at the
same time hybrid improvement would have caught up by 10–15% (Soh et al.
2003a,b, 2006). Yields of up to 11–12 t/ha/yr oil have been reported in trial and
commercial clonal plantings (Mohd Isa et al. 2005; Soh et al. 2006).
Besides breeding for yield per se, there are programmes which also emphasize
other desirable agronomic/economic traits, e.g. dwarfness or improved oil
quality. The semi-dwarf Dumpy.AVROS variety which was about 20%
shorter than the popular but tall AVROS variety was available since the
early 1980s and represented about 10–20% of Malaysia’s annual oil palm
plantings till recently (Soh et al. 2006; Mohd Isa et al. 2005). An improved
version of the Dumpy.AVROS variety, the Dumpy.Yangambi.AVROS
(Fig. 11.3) with better physiological traits and thus higher yield potential
has been developed (Soh et al. 2006). The development of dwarf palms high
in unsaturated fatty acids derived from their Nigerian E. guineensis accessions
by MPOB and E. oleifera E. guineensis hybrid-derived compact statured
clones by ASD have been announced (Rajanaidu et al. 2000; Escobar and
Alvarado 2003). Few groups have also stated to produce limited numbers of
340 A.C. Soh et al.
biclonal (clonal parents on both sides) and semi-clonal (clonal parent on one
side, usually the dura) hybrid seeds from proven parents. The impact from the
commercial planting of these newly developed special materials remains to
be seen.
Owing to the versatility of the crop in terms of the myriad uses of its oil, many
suggestions for various desirable traits to be improved in the palm and its oil
have been made. To reconcile opinions and lobbies, MPOB in 2003 organized
a workshop comprising breeders, agronomists, biotechnologists, oleochemists
and technologists, palm oil end-users and traders for prioritizing traits for
improvement in the oil palm, as it is inefficient if not impossible for breeders
to consider all the useful traits in a breeding program. The four top priority
traits in ranking order were: high oil yield, dwarf stature, resistance to Gano-
derma disease and high oleic acid oil (Table 11.2). The first three are agronomic
traits related to yield. This is not surprising as palm oil is still essentially
a commodity crop mainly used as food where high yield ensures lower cost of
production and competitiveness against other vegetable oils, e.g. soybean oil.
Except for high oleic acid content, other oil quality components and value
addition traits such as stearic acid, carotene or tocotrienols were given lower
11 Oil Palm 341
Table 11.2 Priority list (top four traits) of desirable traits for genetic improvement of oil palm
in Malaysia
Priority Trait Rationale
1 High palm oil Commodity crop, mainly used as food. Lower cost of
yield production. Oil already versatile in its uses.
2 Dwarf stature Current palm varieties grow too tall too fast. Inefficient
harvesting. Scarcity and high cost of workers.
3 Ganoderma Ganoderma basal stem rot is becoming a serious problem in
resistance young second cycle plantings, decimating the stand. Cultural
and chemical controls are ineffective.
4 High oleic acid Healthy monounsaturated oil. Liquid oil in temperate countries
for use in salad dressing and cooking. More suitable for
production of oleochemicals and biodiesel.
High palm oil yield is still the prime goal of most if not all breeding pro-
grammes. However, there are many facets in terms of achieving a high yield.
Potential yield. This is the maximum yield achieved by a variety when grown
under stress-free conditions to which it is adapted (Evans and Fischer 1999).
Modern high-yielding varieties, resulting from a plant ideotype breeding
approach are typically smaller statured plants that can be planted at higher
density, are consequently capable of high biomass production and possess a
high harvest index or better conversion of dry matter directed to yield rather
than to vegetative growth. The combination of high biomass production and
high harvest index results in a super high yield (Soh 2005). Such varieties are
single cross hybrids maximizing heterosis and stand uniformity. Early oil palm
breeding efforts were biased toward high early individual palm yield resulting in
aggressive plant types, and their use in mixed hybrid planting would not fully
exploit the potential yield of the crop. The ideotype approach in oil palm
breeding has been advocated since the early 1980s and some early results from
these efforts are available (Breure and Corley 1983; Squire 1984; Breure 1986;
Henson 1998; Soh et al. 2006).
Harvestable/recoverable/realizeable yield. Harvesting oil palm fruit bunches
particularly from older and taller (5–12 m) palms is still a tough and tedious
manual operation (man with sickle on long pole) with no imminent prospective
342 A.C. Soh et al.
As palm oil’s principal use is in food production, its dietary quality is under
close scrutiny by the current health conscious consumers. Palm oil is often
regarded mistakenly as a saturated tropical fat with the connotation that its
consumption will lead to elevated levels of blood cholesterol and risk of cardi-
ovascular heart disease (CVD). Although it contains about 42% saturated fatty
acids, it is in the form of palmitic acid which is not known to be cholestrolemic;
unsaturated fatty acids such as the monounsaturated oleic acid (40%) and
polyunsaturates (linoleic and linolenic acids, ca. 15%) mainly comprise the
rest. Oleic acid has been identified as a top priority trait for improvement
because it confers a healthy liquid oil similar to olive oil that is marketed in
temperate countries for salad dressing and as cooking oil. It also serves as a very
useful feed stock for other oleochemical and biofuel industries. Palm oil also
contains other useful organic components such as carotene (vitamin A), toco-
pherols and tocotrienols (vitamin E) as well as other antioxidants which not
only confer health (anti-CVD, anti-cancer) properties to palm oil, but can also
be isolated and spun-off in the health-care and cosmetics industries (Yusoff
2000; Khosla and Sundram 1996; Sundram and Chandrasekharan 2000).
The oil palm is a cross-pollinated perennial tree crop. Not surprisingly, it has
adapted breeding methodologies developed in maize, e.g. recurrent selection,
and animal breeding, e.g. sire (pisifera) testing, index and BLUP selection, and a
close temporal and genetic correspondence of commercial hybrid production
with each cycle of breeding (Soh 1999).
The major oil palm breeding programmes adopt either the modified recur-
rent selection method (MRS; Soh 1999) or the modified reciprocal recurrent
method (MRRS; Meunier and Gascon 1972). The former is practiced by
programmes particularly in the Far East linked to or influenced by the pro-
grammes of the Unilever plantations group, while the latter is adopted by
countries in West Africa and Indonesia advised by CIRAD.
In the modified recurrent selection scheme (Fig. 11.4), selection of Deli D
parents for further breeding and for mother palms in commercial hybrid seed
production is based on family and individual palm performances, hence Rosenquist
(1990) chose to name the method as FIPS (family and individual selection).
Tenera parent selection for further P breeding is also based on FIPS.
Being female sterile, P is selected as the male parent for DP hybrid seed
production based initially on its T sib performance in the TT family followed
by a DP progeny test by crossing it with a sample of usually 3–5 of the selected
344 A.C. Soh et al.
D female parents in a nested mating design (NCM 1), i.e. top-cross or sire
testing. The resultant commercial mixed hybrid is akin to a synthetic in the
plant breeding literature (Simmonds 1979; Allard 1960) except that its sub-
sequent progeny seeds should not be saved for commercial planting. The
variability within the commercial mixed hybrids varies with the genetic diver-
sity and inbreeding status of the parents used. The main advantages of this
scheme are that more recombinant crosses and genotypes can be turned over
within shorter time and smaller space without the need for extensive progeny
tests, and that the genetic variability of the commercial hybrids would ensure a
good adaptability and reduced risk of genetic vulnerability. This breeding
scheme exploits only general combining ability (GCA), but not specific com-
bining ability (SCA). Also, the assumption that the additive or GCA effects
expressed within the parental DD and TT crosses will be reflected in the
DT/P inter-population hybrid yield performance may be untenable, espe-
cially when parents become more restricted and inbred (Soh 1999; Soh and
Hor 2000).
In modified reciprocal recurrent selection (Fig. 11.5), parents selected for
further breeding and for commercial hybrid seed production have been progeny
tested. Instead of DP progeny testing, a DT progeny test of the selected D
and T parents from the DD and TT families is done. To save time, selfs and
sibs of the parents undergoing progeny testing are made and planted simulta-
neously as the progeny test crosses. The best hybrid crosses are identified from
the DT progeny test. The best hybrid is then readily reproduced as commer-
cial DP hybrid using the Ds from the selfs of the D parent and the Ps from the
selfs of the T parent. This scheme exploits both GCA and SCA, and the
11 Oil Palm 345
Breeding system. The oil palm is monecious bearing male and female inflor-
escences on the same palm occurring in different but overlapping cycles. Stress
conditions such as drought, malnutrition or plant competition favour male
inflorescence production (Corley and Tinker 2003). Pisifera palms do produce
female inflorescences especially under good growing conditions, but they
usually are aborted. Cross pollination in the oil palm is effected efficiently by
the pollinating weevil, Elaidobius kcameroonicus, in its natural home in Africa.
Prior to the weevil’s introduction to the Far East, pollination occurred by wind
and thrips (Thrips hawaiiensis) which was less efficient. Stringent controlled-
pollination procedures had to be adopted after discovering that serious illegi-
timate pollination had occurred in the commercial hybrid seeds resulting in high
frequency of D contaminants in young commercial T fields soon after the
introduction of the weevils to the plantations in the Far East. In male and
female inflorescences, anthesis and receptivity occur about 2 months after
emergence or appearance. The pollen viability and receptivity periods of the
male and female inflorescences are about 3–5 days. The set fruit ripens, chan-
ging colour from dark purple or black to reddish orange, in about 5 months.
Controlled pollination. The formalin surface-cleaned male inflorescence is
isolated with a permeable terelene/canvas/paper bag about 7–10 days prior to
anthesis. The anthesized male inflorescence is harvested and air-dried for about
3 h in an oven, chamber (38–398C) or air-conditioned chamber; the pollen is
shaken out, sieved and further dried in filter paper envelopes at 38–398C in an
oven or desiccator till 6% moisture content and stored in test or specimen tubes
in a freezer. Properly processed pollen can retain its viability for 6 months up to
a year; freeze-drying is recommended for longer storage. Isolation of the female
inflorescence is done in exactly the same manner. Controlled pollination is
carried out when the female inflorescence is observed to be receptive, by puffing
a 1:10–1:20 pollen: talcum powder mixture through a hole made in one or more
plastic windows sown onto the isolation bag (Fig. 11.6). The bag is removed
after a month and the fertilized bunch allowed to ripen in about 5 months. As a
stringent quality control measure against illegitimate pollination, the inflores-
cence is discarded if there is any hole or tear in the bag or the presence of weevil
spotted inside the bag.
Seed germination. The oil palm seed is recalcitrant and requires heat treatment
at 37–388C and 17–19% moisture content for D seed (20–21% for tenera seed)
for 40–60 days for germination at ambient conditions after rehydrating to
21–23% moisture for D (27–28% for T seeds). Germination of very thin-shelled
T and shell-less fertile P seeds is erratic (Arasu 1970) and in vitro germination
(embryo rescue) is advisable.
348 A.C. Soh et al.
needed. Augmented designs (Federer et al. 2001) would also be useful in such
situations. Incomplete block designs, e.g. the balanced lattice (Cochran and
Cox 1957) to circumvent soil heterogeneity effects within large blocks asso-
ciated with large sets of crosses have been attempted in oil palm but were found
to be cumbersome and did not improve trial efficiency much (Soh et al. 1990).
Plot sizes of 10–20 palms (without border trees) planted in 3–6 replicates are
commonly used with the large plots preferred if between-cross differences
especially in growth habits are suspected. When the trial is replicated over
locations, 2–3 replicates per location would suffice. Soh et al. (1989, 1990)
found that a trial size of 6 replicates of 16 palm plots or 5 replicates of 20
palm plots is suitable to detect treatment differences of 15% with a trial
coefficient of variation of 10% under Malaysian conditions. Single palm plots
have been found to be not efficient unless highly replicated to about 100 times,
and they are also not experimenter-friendly for field visual appraisal. Single
palm plots in completely randomized design (CRD) would be useful in field
screening for disease resistance and for estimating between palm within progeny
variability in genetic studies.
Data collection. The following data are collected on an individual palm basis:
For vegetative growth measurements (Corley et al. 1971; Corley and Breure
1981), height, girth, leaf production, leaf area and weight measurements are
made annually. For fresh fruit bunch yield (FFB), number and weight of ripe
bunches are recorded in each 10 day harvesting round. Yield data is collected
from start of harvesting at about 3 years after field planting till about 9 years.
For bunch analysis (Blaak et al. 1963), bunches are sampled over the yield
recording period to determine bunch and fruit quality and oil content. Fruit
sub-samplings are made to estimate the ratio of fresh fruit weight in the bunch
(F/B%), the ratio of mesocarp weight in the fruit (M/F%), the ratio of kernel
weight in the fruit (K/F%) and the ratio of oil weight in the mesocarp (O/M%).
The product F/B M/F O/M gives O/B% (oil to bunch) and F/B K/F
gives K/B (kernel to bunch). For the determination of a progeny mean for O/
B% about 160 bunches from about 40 palms are sampled over the yield
recording period. For individual palm means usually more than 5 analyses are
needed. The following trait data can be generated from the above
measurements:
Vegetative growth: girth, height and height increment, total leaf production,
leaf area index, leaf area ratio, vegetative dry matter production (VDM).
Yield: bunch dry matter yield, BDM = FFB dry matter/fresh matter.
Bunch analysis: oil yield, OY = FFB O/B; kernel yield, KY = FFB K/B;
kernel oil yield, KOY = FFB K/B 0.5. These are reflective of the
commercial oil extraction, kernel extraction and kernel oil extraction rates
achieved in the mills. Total dry matter production, TDM = BDM +
VDM; bunch index, BI = BDM/TDM; harvest index, HI = BI 0.4.
All these primary yield, yield component and morpho-physiological traits,
either formally based on measured data or visually/intuitively scored, are taken
350 A.C. Soh et al.
in consideration in most oil palm breeding programs, besides oil quality and
stress resistance in others.
Fig. 11.7 Somaclonal variation in oil palm: mantled parthenocarpic fruits (left), bunch
abortion and sterility (right)
Fig. 11.8 Oil palm tissue culture system (gel and liquid systems)
Fig. 11.9 Commercial oil palm tissue culture laboratory with two million plantlet production
capacity (Applied Agricultural Resources, S.B., Malaysia)
11 Oil Palm 353
Explant sampling. The explants are the young leaf pinnae of the youngest
non-emerged spear leaves of the selected palm. The harvested spear is surface-
sterilized before the leaves are unraveled in the laminar flow chamber and
1,000–2,000 explants of 1 cm2 pinnal segments are inoculated onto agar med-
ium in test tubes.
Callogenesis. Callus growth is induced on the explant using MS (Murashige
and Skoog 1962) medium which may be supplemented with additional nutri-
ents, variable levels and proportions of phytohormones, typically 2,4-D and/or
NAA and with or without the addition of charcoal. Some laboratories also
apply explant pretreatments. The explant cultures are incubated at ambient
conditions (26–288C, 90% relative humidity) in the dark or in lighted rooms.
Calli begin to appear on the leaf explants at about 3 months and continue
proliferation up to a year or more.
Embryogenesis. Callus differentiates into somatic embryos or embryoids
when nodular callus on explants is sub-cultured on medium with reduced levels
of hormones to stimulate callus differentiation into embryoids or somatic
embryos in air-conditioned culture rooms maintained at 26–288C and
50–60% relative humidity.
Embryoid proliferation and germination. Good embryoids are transferred
onto embryoid proliferation medium and sub-cultured usually at bimonthly
intervals. In the case of liquid suspension culture, embryogenic calli are trans-
ferred to liquid medium in a conical flask on an orbital shaker for proliferation.
At each monthly sub-culture, embryoids of the desired size are sieved out and
transferred to fresh medium.
Plantlet regeneration. In the gel system, shoots begin to germinate at about
the 5th sub-culture on the polyembryogenic mass consisting of secondary
embryoids, primary embryoids and callus. The shoots are harvested from the
polyembryogenic mass and put onto a liquid medium in a test tube for shoot
development and root induction. The polyembryogenic mass is put back onto
fresh medium for further proliferation. In the liquid suspension system, mature
embryoids are inoculated onto plated medium for germination. The shoots are
subsequently transferred individually into tubes for growth and root induction
as for the gel system.
Acclimatization. Rooted plantlets of about 8–12 cm height are selected and
planted out in trays containing inert potting mixture, e.g. sand, peat fibre,
vermiculite in a plastic incubation chamber in the conditioning nursery with
75% shade, 90% relative humidity and 27–308C temperature. Shade and
humidity are gradually reduced by opening the flaps of the chamber for increas-
ing lengths of time to condition the plantlets. After 3 weeks the trays of plantlets
are taken out from the plastic chamber and put under 50% shade to be
hardened and foliar fertilized. Hardened 3–4 leaved plantlets are harvested,
cleaned of its potting medium and treated with a fungicide before dispatching as
‘bare root seedlings’ contained in moistened plastic bags packed in carton boxes
to the plantation nurseries, where they are raised as for normal seedlings.
354 A.C. Soh et al.
The commercial planting of oil palm clones, totaling less than 20,000 ha out of
the world’s total oil palm planting area of 14 million ha, can be considered in its
infancy or at only a larger scale pilot commercial testing state (Fig. 11.10). The
clones are produced by a handful of commercial tissue culture laboratories with
mainly used to saturate the majority of oil palm genetic maps (Mayes et al. 1997;
Billotte et al. 2005; Ting et al. 2005).
Some of these polymorphic markers are used for oil palm DNA fingerprint-
ing, particularly useful for genotype identification and genetic diversity assess-
ment (Cheah and Wooi 1995; Cheah et al. 1999). As concerns about breeder’s
rights on planting materials increase, Malaysia has gazetted the Plant Variety
Protection Act in 2004. The Malaysian Department of Agriculture aided by
MPOB and industry members is in the process of drafting test guidelines for the
conduct of tests for distinctness, uniformity and stability in compliance with
UPOV (Union for the Protection of New Varieties of Plants). DNA fingerprint-
ing for genotype identification is identified as one of the elements to comple-
ment morphological markers. Of particular interest are elite clones and inbred
parents for hybrid seed production that need protection from piracy and sub-
sequently ensure profit return to the rightful breeder. Discriminating commer-
cial ‘varieties’ of oil palm is envisaged to be more difficult as they are mixed
hybrids from parents with very similar genetic backgrounds.
In advanced breeding programmes, it is becoming increasingly difficult to
select for extreme outliers because of the highly selected nature of the genetic
materials. Genetic diversity studies with the estimation of genetic distance or
relatedness between oil palms can help identify suitable divergent parents for
base broadening breeding or for crossing to enhance hybrid vigour in the
progeny or seed produced. The same can be applied to inbreeding programmes
to assess homozygosity or inbreeding level of the resulting materials and in
backcrossing programmes to recover the recurrent genotype. Genetic diversity
studies can assist in hastening the prediction of parents with good combining
ability in that a pre-selection of materials can be included in a breeding trial to
expedite creation of homozygous inbred parents.
The ability of molecular markers to assess relatedness also introduces the
possibility of assessing illegitimacy of a cross or a commercial hybrid (Corley
2005). With increasing efforts and expenditure invested in oil palm breeding
programmes and commercial field planting of hybrids, illegitimacy in the
breeding and commercial hybrid materials cannot be tolerated. With molecular
marker technology, crossing programmes can be quality-checked and breeding
as well as commercial hybrid seed production will become more efficient.
With the construction of genetic maps, it is possible to identify the genomic
location of specific traits by using genetic markers. Marker based selection can
be used as an early screening technique for specific traits of interest such as
desirable fatty acids, amenability to tissue culture, fruit colour, shell thickness,
stem height and mature frond length, even before the palms mature or before
the trait is expressed. Marker assisted selection can help reduce the breeding
trial size thus increasing the chances of selecting desirable extreme outliers for a
given trait and thereby expedite the improvement process. Single genes control
fruit colour and shell thickness, therefore single markers can be developed for
their selection (Mayes et al. 1996; Moretzsohn et al. 2000; Billotte et al. 2005).
The issue at hand is whether the marker is close enough to the target gene to
11 Oil Palm 357
prevent recombination and thus is reliable. For a marker for virescent fruit it
appeared to be so, but not for the shell gene. Other traits mentioned above are
likely to be controlled by multiple genes or quantitative traits and several
markers are needed for their selection. Markers controlling quantitative traits
can be found in the same or across different linkage groups. The location of
these quantitative trait loci (QTL) can be identified using fine mapping techni-
ques, such as the bulked segregant analysis (BSA) as described in Weising et al.
(2005). Currently, the interest is to analyze single nucleotide polymorphisms
(SNPs) to further saturate the genetic map so that more precise markers may be
located (Rajinder and Cheah 2005).
Fluorescent in situ hybridisation (FISH) was used to detect collectively the
amount of E. oleifera’s DNA in interspecific crosses of E. oleifera (O) with E.
guineensis (G). The whole genome of O can be labelled, hybridised to the O G
genome and detected with fluorescent dye. Chromosomes that contain genome
from O will show fluoresce under fluorescent microscope revealing the compo-
sition of genetic material from both parents. The inheritance of genetic material
in the F1 hybrid is 50% from each parent. However, as reintroduction of
desirable G characteristics by backcrossing with G continues, percentages of
genetic content from O would gradually reduce from 50 to 0%. Monitoring
with FISH therefore enables targeting of progeny with the desirable genetic
composition and reduces the number of backcrosses needed (Maria et al.
1998a,b; Cheah et al. 1999) prior to screening for other desirable traits (e.g.
oil quality), which is essential in long life cycle crops.
As alluded to earlier, the current thought on the cause of the floral abnorm-
ality somaclonal variant is that of methylation (Jaligot et al. 2005) and alteration
in the expression of MADS box gene (Van der Linden et al. 2005). In order to
detect methylation in the MADS box gene, use of methylation sensitive MADS-
box directed profiling is currently under investigation (Auyong et al. 2005).
There has been also an increasing interest in the expression marker system
for the detection of complicated biological events such as embryogenesis and
floral abnormality. These markers should detect expression level changes coin-
ciding with a biological event, making it possible to detect an event of interest.
Genes involved in embryogenesis have been isolated from cDNA libraries
(Ong-Abdullah et al. 2005). Currently, more genes are to be analysed and
identified using the microarray technology (Low et al. 2005), which can screen
through thousands of expressed genes in one single analysis. Several genes
involved in floral abnormality and embryogenesis have been identified and
are undergoing expression studies and functional analysis.
Genetic transformation of oil palm via biolistics is now a reality with
BASTA-resistant oil palm produced (Ghulam Kadir et al. 2005) and used as
a selection means for transformation. Of particular interest is the creation of
oil palms that produce desirable fatty acids. This can be done by enhancing
and/or suppressing key genes that produce certain fatty acids. Some of these
key enzymes are: b-ketoacyl-ACP synthase II (KASII) which catalyses con-
version of palmitic acid to stearic acid, stearoyl-ACP desaturase which
358 A.C. Soh et al.
The various steps of oil palm seed processing are described according to
Periasamy et al. (2002):
Depericarping. The ripe bunch from controlled pollination harvested from
the D mother palm is chopped to separate the fruit spikelets which are then
allowed to ret for days in a basket or gunny/raffia sack to detach and soften the
fruit. The mesocarp of the fruits is then stripped off using a depericarping
machine to give the fresh seeds. Fresh seeds are then further cleaned of the
adhering fibres, washed with a detergent and treated with a fungicide.
Fresh seed quality checking. A sample of the seeds is taken from each bunch,
the shells cracked and the kernels and embryos scored for quality. The kernels
must be present, fresh looking and not diseased or rotting. The embryos must be
well-formed and not shrivelled, cylindrical in shape and white in colour with a
pale yellow green tinge. Bunches with more than 80% good quality seeds are
acceptable for further processing into commercial seeds.
Seed moisture adjustment and storage. A sample of seeds from each bunch is
checked for its moisture content. The seeds from the bunch are then soaked or
surface air-dried to achieve 18–19% moisture content for storage in an air-
conditioned room at 218C.
Heat treatment. Stored fresh seeds (at about 18–19% moisture) when
required to be germinated are sent to the germinator (hot room) to be heat
treated at 37–398C for about 50 days. The heat treated seed can be directly
processed for seed germination or cold stored again as ‘preheated’ seed.
Resoaking and germination. Preheated seeds need to be remoistened to
21–22% moisture before setting them to germination at ambient conditions.
Germination begins after a week with weekly flushes for about 6 weeks. With
good quality seeds, more than 90% germination rates would be achieved by the
third flush.
Seed selection and dispatch. Germinated seeds/seedlings with pearly white
plump and balanced plumules and radicles of <5 cm length are selected and
11 Oil Palm 359
collected in plastic bags with 200–250 seeds in each. These are then packed in
polystyrene-lined and bead buffered carton boxes for dispatch. With current
ease of air and surface transport systems, commercial oil palm seeds are now
exclusively dispatched as germinated seeds all over the world instead of pre-
heated seeds.
The world trade in oil palm seed is about 250–300 million seeds worth US$ 100
million at around 50 US cents per seed. The main supply and demand is in
Indonesia with about 117 million seeds, Malaysia with 85 million seeds and all
others with 85 million seeds as well. The break-down of the oil palm seed
suppliers, many of which are plantation-based companies and their estimated
capacities are given in Table 11.3.
The high crude palm oil prices boosted by oil demands for biofuel have
prompted the opening of more land in Indonesia and other countries for oil
palm cultivation resulting in increased demand for oil palm seeds, and conse-
quently many seed companies have stepped-up production. The supply of oil
palm clonal planting material from about 6–8 tissue culture laboratories is still
Table 11.3 Oil palm seed production companies and their estimated annual seed supply
Estimated supply
Country Company (in million seeds)
Indonesia Indonesian Oil Palm Research Institute (IOPRI) 35
PT. Socfm Indonesia 20
PT. London Sumatera Indo 20
PT. Bina Sawit Makmur 20
PT. Tunggal Yunus Estate 6
PT. Dami Mas Sejahtera 10
PT. Tania Selatan 6
Malaysia Felda Agricultural Services S.B. 27
Guthrie Research Chemara S.B. 20
Golden Hope Research S.B. 10
United Plantations Bhd. 8
Applied Agricultural Resources S.B. 6
Ebor Research, Sime Darby Plantations Bhd. 3
EPA 3
Sawit Kinabalu 3
Others 8
Costa Rica ASD 20
Papua New Guinea Dami 20
Africa Ivory Coast, Zaire, Nigeria 17
Thailand Univanvic 5
South America Colombia, Brazil, Ecuador 14
360 A.C. Soh et al.
very small constituting less than 1% (ca. 2 million) of plants and may rise to
5–10 million in 5 years time; the high price and limited supply of cloned plantlets
is attributed to the still inefficient commercial tissue culture process.
Acknowledgements We wish to express our sincere thanks to our company, Applied Agri-
cultural Resources Sdn. Bhd. and its principals, Boustead Plantations Bhd. and Kuala
Kepong Bhd. for permission to publish this paper.
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