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Designing the oil palm of the future

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Journal of Oil Palm Research Vol. 29 (4) December 2017 p. 440 – 455
JOURNAL OF OIL PALM RESEARCH 29 (4) (DECEMBER 2017)
DOI: https://doi.org/10.21894/jopr.2017.00015

DESIGNING THE OIL PALM OF THE FUTURE

ZULKIFLI, Y*; NORZIHA, A*; NAQIUDDIN, M H*; FADILA, A M*; NOR AZWANI, A B*; SUZANA, M*;
SAMSUL, K R*; ONG-ABDULLAH, M*; SINGH, R*; GHULAM KADIR AHMAD PARVEEZ* and KUSHAIRI, A*

ABSTRACT
In order to allow for future uninterrupted growth of the oil palm industry in Malaysia, there is a need
to increase yield, value-addition of the oil and reduce production costs. The collection and exploitation
of germplasm material is considered a major factor in contributing towards a tremendous impact on the
Malaysian oil palm industry. Improved oil palm varieties with higher yields, good oil quality, compact
in architecture, better adaptation to climate change and higher tolerance to diseases have been prioritised
to meet future demands of the oil palm industry. It is only natural that biotechnology and bioinformatics
have been identified as the core areas to expedite improvements to achieve these objectives. The use of a
combination of conventional and molecular breeding approaches is useful as they open up new avenues for
crop improvement and have the potential for producing novel oil palm varieties, in terms of sustainability
and its contribution to human well-being.

Keywords: oil palm, breeding, planting materials, biotechnology, genomic tools.

Date received: 17 October 2017; Sent for revision: 21 October 2017; Received in final form: 24 Novembar 2017; Accepted: 30 November 2017.

INTRODUCTION worldwide. The oil palm has many other uses in the
non-food sector, with palm kernel oil mainly used in
Originally a minor crop in the 1960s with a yield the oleochemical and animal feed industries. Many
of 94 000 t (Basiron, 2000), palm oil production has biomass-derived products such as plywood and
steadily increased to become the world’s major fibreboard, are also produced from oil palm biomass
vegetable oil. The United States Department of (Suhaily et al., 2012). Oil palm industries produce
Agriculture (USDA) estimates that global palm oil an enormous quantity of lignocellulosic biomass, a
production in 2017 will be 62.88 million tonnes, huge source of fibre and cellulosic materials which
which is an increase of about 4.08 million tonnes have potential for further commercialisation. It is
(6.94%) from 2016 (USDA, 2017). Collectively, projected that conversion of cellulosic fibre into
Indonesia and Malaysia produce about 57 million liquid biofuel makes the oil palm a future source of
tonnes of palm oil, making up over 85% of global renewable energy which will enhance sustainable
palm oil production. The momentum of high development (Basiron, 2007).
productivity at a lower cost than other vegetable The oil palm belongs to the genus Elaeis, which
oils, has increased the demand of palm oil especially consists of two species, Elaeis guineensis and Elaeis
in India and China (Murphy, 2007). Palm oil is oleifera (Schultes, 1990). The two species are closely
mainly used as an edible product and has good related and can be easily inter-crossed to produce
resistance to heat and oxidation, making palm oil an interspecific hybrid (Hardon and Tan, 1969).
an ideal ingredient in frying and dietary component The major commercially important oil palm is the
E. guineensis species. A comprehensive description
of oil palm biology is provided by Hartley (1988).
* Malaysian Palm Oil Board, Essentially, oil palm has a solitary unbranched stem
6 Persiaran Institusi, Bandar Baru Bangi,
43000 Kajang, Selangor, Malaysia. that is topped by 35-60 pinnate fronds and its height
E-mail: [email protected] can reach over 20 m. The palm reaches maturity

440
 DESIGNING THE OIL PALM OF THE FUTURE

when it is about three years old. The crop bears flourishes in Malaysia and elsewhere outside its
fruits throughout the year and can live up to 200 home continent, owing to the favourable tropical
years, but the productive life of each tree is between climate with good sunshine and rainfall throughout
25–30 years (Cheah, 1997). The average annual the year. However, weather patterns might change,
yield of oil palm plantations is currently around 4 t and to ignore climate change, the oil palm industry
ha-1. In addition, planting materials producing over would fall into the trap of its own complacency.
8-10 t ha-1 oil have been obtained through selected Until recently, agricultural scientists, including
breeding trials and clonal propagation. oil palm breeders and agronomists, have strictly
With the projection of population growth used conventional and hybrid plant breeding
touching 9 billion, agricultural products need to to continually improve performance of oil
increase by 70% and net exports of oilseeds and palm productivity. Although results have been
vegetable oils are expected to triple by 2050. Despite outstanding, additional gains in agricultural
oil palm’s inherent high productivity, its narrow productivity are demanded at an even faster pace
genetic base is a barrier to the industry. The mother due to continuing changes in agricultural practices
palms used were and still is mainly the Deli duras and consumer preferences. Modern genomics tools
which are direct descendants of the four seedlings allow substantial knowledge to be attained on the
brought to Indonesia in 1848 (Rajanaidu et al., 2000). biological traits of a species which can be exploited
The Malaysian oil palm industry had realised the in improving productivity and obtain better product
risk in relying on a limited genetic base and saw quality. The whole spectrum of breeding approaches
the need for an adequate genetic base for effective and biotechnology is being deployed to meet these
selection in breeding programmes. Furthermore, objectives. In this review, we focus on the exploration
selection progress was hampered by the narrow of oil palm germplasm, traits improvement and how
genetic base. A lingering concern was also on the modern genomics tools can contribute to design this
susceptibility of palms to pests and diseases due remarkable crop.
to its narrow genetic base. The need for genetic
variability for future improvements triggered a
PLANTING MATERIALS
systematic exploration of oil palm germplasm in
Africa and South America spearheaded by MPOB Germplasm: Elaeis guineensis
back in the early 1970s. This led to MPOB having
now in possession of the world’s largest ex situ Oil palm germplasm that exhibits economically
germplasm collection, comprising collections from important traits is a valuable resource for oil palm
11 African countries for Elaeis guineensis whilst Elaeis improvement. Recognising this, E. guineensis
oleifera collections were obtained from eight South germplasm was collected from Africa to broaden
American sites (Rajanaidu et al., 2013). the genetic base of oil palm breeding materials
The exploitation of germplasm material is in Malaysia at that time. The first attempt was
beginning to bear its rewards and shows a lot accomplished in Nigeria in 1973 and evaluated at the
more genetic diversity. Initial collections have MPOB Research Station at Kluang, Johor, Malaysia
provided to date a source of planting materials for (Rajanaidu, 1986). Besides Nigeria, prospection
important traits such as dwarfness and oil quality was made from other countries in Africa such
(Kushairi et al., 1999a, b). Meanwhile, parental lines as Angola, Cameroon, Zaire (now known as the
of advanced breeding materials are continuously Democratic Republic of Congo), Ghana, Guinea,
being improved through extensive breeding Gambia, Madagascar, Sierra Leone, Senegal and
programmes. Coupled with improving breeding Tanzania (Rajanaidu and Rao, 1987; Rajanaidu et al.,
and mass propagation methods to increase yield, 2013). Extensive screenings should be continuously
the oil palm sector should also focus on higher carried out for those materials collected in order to
quality oils and high value niche products for its exploit their genetic potential in oil palm breeding
long-term growth (Murphy, 2007). Stimulating R&D programmes.
to generate resistance planting materials to major Population diversity studies among MPOB
pests and diseases such as Ganoderma, and improved oil palm germplasm collection are very crucial
palm architecture will also have significant impact for a foundation set of germplasm with minimal
to the development of new oil palm varieties in repetitive and maximal genetic diversity (Zulkifli,
the coming decades. In addition, there is also an 2012). This core germplasm collection could be
increasing demand to accelerate oil palm breeding established by field and molecular evaluation. Field
for better adaptation to drought, heat, and other evaluation includes analysing bunch yield, yield
abiotic stresses (Suthashinikisan, 2015). More than components, vegetative measurement and fatty acid
a decade ago, the tagline Palm Oil: Nature’s Gift composition (FAC) for each germplasm either using
to Malaysia and Malaysian Gift to the World which Cluster or Principal Component Analysis which
describes the blessings of palm oil to Malaysia, was systematically reduce the dimensionality of the
coined. Though originated from Africa, oil palm data. This method combined with molecular studies

441
JOURNAL OF OIL PALM RESEARCH 29 (4) (DECEMBER 2017)

would be very useful to detect genetic relationships more than 20% crude protein in the kernel (PS14)
among accessions and subsequently determine the were identified in germplasm collections from
overall degree of polymorphism within germplasm Ghana, Cameroon, Gambia, Senegal, Tanzania and
collections. It could be a complete guide to develop Zaire (Noh et al., 2014). The introduction of these
strategies to conserve unique genetic sources as best materials into commercial populations can improve
as possible and avoid any redundancy which will the protein content in the palm kernel.
not bring informative values to the next generation. Parental lines of advanced breeding materials
MPOB regularly transfers new technologies are continuously being improved through MPOB’s
(breeding and planting materials) as Planting extensive breeding programmes. The breeding
Series (PS) palms to the industry and to date 14 PS scheme primarily involves a reciprocal recurrent
have been transferred. Evaluation of the Nigerian selection scheme (RRS) which has been adapted by
germplasm indicated some interesting attributes a number of oil palm breeders. In this scheme, the
such as dwarf (PS1), high iodine value (PS2) and dura and pisifera genotypes are kept as distinctly
high kernel (PS3) (Rajanaidu et al., 2000). High separate base populations. In the form of hybrids
oleic acid breeding populations (PS12) were also (tenera), the heterosis effect is obtained by crossing
obtained from the Nigerian germplasm (Isa et al., origins with complementary characteristics. The
2006). The Angola palms exhibited large fruit dura performance of inter-origin crosses is attributed to
(PS6) (Kushairi et al., 2003a) which is similar to Deli the additive effect of favourable genes combination
duras. Individual Angola palms were also used for a from the parents. It was reported that the RRS
long stalk programme (PS10) to improve efficiency scheme has increased oil yield by almost 18% per
in harvesting (Noh et al., 2005). Besides that, Angola cycle compared to the base population (Rajanaidu
germplasm exhibited wide variability in FAC in et al., 2000). On the other hand, the way to broaden
their oils compared to those in DxP materials at that their genetic diversity is through introgression
time (Noh et al., 2002). Some palms from Nigeria of current materials with selected materials from
and Madagascar also exhibited iodine value (IV) of the germplasm developed by MPOB. In order to
more than 60, which is relatively higher than current achieve the target, various crossings of oil palms
levels (<50) (Kushairi et al., 2003b). with different economically desirable characters
A study by Marhalil et al. (2015) reported that were conducted through breeding programmes.
MPOB germplasm materials assembled from Through new combinations, the best materials
Senegal, Nigeria, Cameroon, Tanzania, Madagascar would be further evaluated and later exploited for
and Angola had experienced extreme low rainfall. future commercial seed production. Rajanaidu et al.
Palms from these collections are potentially useful (2011) reported that MPOB’s latest Nigerian-based
for the development of drought and heat tolerant planting material has the qualities to develop an
planting materials. Some palms from Tanzania are ideal palm with balanced bunch number and bunch
known for thin-shelled tenera (PS5) (Kushairi et weight of 15 x 15 kg. The individual fruit weight is
al., 2003c) and high bunch index (PS7) (Junaidah close to 15 g. Musa and Gurmit (2008) introgressed
et al., 2004). Recently, Fadila et al. (2016) reported high IV value materials into the present breeding
that Tanzania, Cameroon, Nigeria and Angola populations have resulted in increased IV (57 – 58)
germplasms possessed high bunch index (BI) which which is relatively higher than current levels (<50).
is an important trait to consider for the selection of The oil palm industry in Malaysia should rapidly
materials towards increasing yields. exploit the genetic potential of germplasm materials
Selected palms from the Angola, Cameroon, to achieve improved productivity and remain
Nigeria, Tanzania and Zaire germplasm collections competitive and sustainable in the future.
produced oils with high vitamin E (PS8) which
is measured based on the levels of tocopherols Germplasm: Elaeis oleifera, Interspecific Hybrids
and tocotrienols in palm oil (Kushairi et al., 2004). and Backcrosses
The mean carotene content (PS11) of E. guineensis
collected from the Tanzanian germplasm ranged Another oil palm species is the Elaeis oleifera,
between 2059 and 2280 ppm (Mohd Din et al., which originates from Central-South America. Elaeis
2006). These palms can be exploited to improve the oleifera has brought its own impact to the oil palm
carotene contents of current planting materials with industry especially in South-Central America for its
a reported content of 500 - 700 ppm. Low lipase contribution in curbing diseases (Corley and Tinker,
genotypes (PS13) are also available in MPOB oil palm 2016). A prospection made by MPOB for E. oleifera
germplasm materials collected from Cameroon, species covered eight countries viz. Honduras,
Guinea and Tanzania (Maizura et al., 2008). Ebongue Brazil, Panama, Colombia, Costa Rica, Suriname,
et al. (2008) reported that low lipase activity among Ecuador and Peru (Rajanaidu, 1994; Rajanaidu and
oil palm ripe fruits is desirable to delay free fatty Rao, 1987). Even though it is relatively low in oil
acid (FFA) accumulation which could reduce the yield, its superior unsaturation in oil composition
quality of palm oil. Genetic materials possessing and slow height increment traits to E. guineensis are

442
 DESIGNING THE OIL PALM OF THE FUTURE

the major areas of interest to oil palm breeders. Apart In terms of clonal performance, Khaw and
from that, a high carotene value trait (PS4) was also Ng (1997) reported that clonal plantlets derived
sourced from this oil palm species (Mohd Din et al., from selected ortets were significantly superior
2002). In spite of the economically beneficial traits of to commercial DxP seedlings. The clones yielded
E. oleifera compared to current DxP material, these at least 20% more than DxP standards (Rohani et
materials show a low germination rate (below 30%) al., 2000; Corley and Tinker, 2003; Tan et al., 2003).
which results from a strong dormancy. In fact, this Based on the good attributes that tissue culture has
species also tends to have poor fruit sets due to low to offer, the interest to go ahead with large scale
pollen viability (Norziha et al., 2015). propagation of oil palm clones is flourishing. This
The present and absent properties of both is proven by the growing number of tissue culture
species has led to the hybridisation of E. oleifera laboratories that applied for licenses from MPOB for
and E. guineensis species, aimed at incorporating commercial production. To date, there are currently
desirable qualities from E. oleifera (such as oil 14 licensed laboratories producing almost 5 million
quality and palm height) and E. guineensis (such ramets annually for local supply. Application of
as bunch yield) (Corley and Tinker, 2016). Most tissue culture is not only restricted to multiplying
research carried out on interspecific hybrids high yielding palms, but also to produce oil palm
generally revealed intermediate values for oil yield, elite lines that feature other desirable characteristics
IV and carotene content (Hardon, 1969; Mohd such as high BI, low height increment, compact,
Din, 2000; Zulkifli et al., 2014). Morphologically, improved tolerance to biotic and abiotic stresses,
the hybrids tend to have thin shell with a lack virescens fruit sets, etc.
of fibre rings (Corley and Tinker, 2016). For the
purpose of increasing homozygosity to the desired
characteristics, backcrosses to E. guineensis have to TRAITS FOR IMPROVEMENT
be continuously conducted. Being a perennial (at
10 years per generation), it may take quite a long Yield
period of time to achieve the final product which
would require several generations of backcrosses Global demand for vegetable oils may reach
to the recurrent parent to fix the genes. However, 240 million tonnes by the year 2050 (Barcelos et al.,
through genomics tools such as quantative trait 2015). Due to expanding populations especially in
loci mapping, selection process for the desired India and China which make up 37% of the world
product can be accelerated. population, as well as rising income levels in
many developing countries in the 1990s, demand
Clonal Palms for vegetable oil is steadily increasing (Murphy,
2007). There are four major oilseed crops in the
High yielding materials have been propagated world, i.e. oil palm, soyabean, sunflower and canola
through the traditional oil palm breeding technique. (rapeseed). Harvested area of these crops covered a
However, the problem of segregation in seed derived total of 196 470 ha with oil palm occupying the least
progenies still persists and this technique is tedious land area of only 9% of the total land area (Oil World
and time-consuming. Therefore, oil palm needs to Annual, 2016). Surprisingly, oil palm has the highest
be propagated vegetatively through tissue culture to productivity, recording 29% of the total of 203.91
maximise the potential of selected genotypes. The million tonnes oil in 2016, followed by soyabean
tissue culture technique has emerged as a feasible (25%), rapeseed (12%) and sunflower (8%) (MPOB,
and cost-effective tool for developing elite oil palm 2017a).
materials (Sakhanokho and Kelley, 2009). Oil palm The improvement of oil palm yield in Malaysia
tissue culture, which was introduced in 1970s for the previous half century was attributed to
(Jones, 1974), has been widely exercised to fast track breeding improvement and improved agronomic
breeding programmes. Central to producing elite practices (Davidson, 1993). Corley (2006) estimated
clones is the availability of high quality ortets. The that potential oil yields of about 18 t ha-1 with
selection of ortets undergoes stringent criteria such possibilities of oil to bunch (O/B) at 50%, fruit to
as data collection of up to four years successively bunch (F/B) at 80%, mesocarp to fruit (M/F) at 95%
for fresh fruit bunch, oil to bunch and a set of data and oil to wet mesocarp (O/WM) at 65%. However,
from a minimum of 30 palms per trait. This exercise the average oil yield in Malaysia has been stagnant
contributed to a 25% increase in yield from clonal for more than 20 years. In 2016, the Malaysian palm
planting (a) (b)
materials in comparison (c)
to commercial (d)
oil industry produced 17.32 million tonnes of crude
DxP (Rohani et al., 2000; Tan et al., 2003). In fact, the palm oil (CPO), 15.91 t ha-1 fresh fruit bunch (FFB)
ortet selection criteria, MS 2099:2008 (SIRIM, 2008), yield with an oil extraction rate (OER) of 20.18%
was enacted by Department of Standards, Malaysia (MPOB, 2017a). Exclusive usage of Deli dura x
as a standard to ensure only superior ramets are AVROS pisifera (DxP) crosses by most commercial
produced from high quality ortets. seed producers in this country is starting to limit

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JOURNAL OF OIL PALM RESEARCH 29 (4) (DECEMBER 2017)

further improvement of commercial planting levels in the air are currently at their highest in
material (Corley and Tinker, 2016). 650 000 years, which is more than 406 ppm or an
Expansion of current plantation area to increase increase of 6% in the last 10 years (NASA, 2017).
oil palm production by economies of scale is no Global temperature has increased sharply since
longer a viable option. The only alternative to boost 2009 with increases between 0.64°C and 0.99°C per
productivity is by increasing yield and profitability year (NASA, 2017). Due to global climate change,
per hectare on existing land. The introduction of some effects on the environment had already been
exotic germplasm by MPOB to increase genetic observed. As global demand for vegetable oils is
variability was an important step in the oil palm increasing, there is a need to address the effects of
breeding programme. Deli dura x MPOB-Nigeria climate change to oil crops especially in relations to
pisifera crosses (PS1.1) showed a 45% improvement yield or productivity.
over the national average in terms of FFB yield A few studies were conducted to recognise
(Table 1). Together with dwarf character, these palms the impact of climate change on oil palm growth
should be incorporated as planting material in and yield as well as to strategise research and
future replanting programmes. In addition to seed- development in breeding and agronomy for climate
derived palms, planting clonal materials is also change. Development of models to simulate the
promising for yield improvement. Clone P456 and change in oil palm growth and yield in Malaysia due
P325 are both high oil yield (OY) candidates, which to climate change have been reported (Christopher,
have performed outstandingly in a few trials (Table 2015; Nizam et al., 2015; Mohamad Faizal, 2015).
1). These clones also displayed high success rates in These studies were essential to understand oil
tissue culturing and very low mantling rates in the palm’s response to abiotic stresses such as increased
field. Another potential future clone, P126, features levels of CO2 and salinity in water supply.
a distinctive characteristic, which is short rachis Some effects of climate change on crops
length (RL) of approximately 4.4 m as compared including oil palms and approaches to deal with
to 5-6 m of DxP and other clones. The short RL of it in oil palm plantations have been reported
this clone makes it suitable for high density (HD) by Heslop (2015) and Nuzul (2015). Changes in
planting (198 palms ha-1 for HD as compared to 148 genetics of crops are needed besides applying
palms ha-1 for normal density). P126 also produces adaptation strategies in agricultural practices.
smaller bunches, which is favourable for harvesting. Meanwhile, screening of oil palm germplasm for
Another clone, P325, showed outstanding O/B and breeding programmes to develop planting material
OY rates. with potentially sustainable yields under extreme
weather was initiated by the Malaysian Palm Oil
TABLE 1. PERFORMANCE OF MPOB SUPERIOR Board (MPOB) and Felda Global Ventures Sdn
PLANTING MATERIALS Bhd (FGV) (Marhalil et al., 2015; Suthashinikisan,
2015). Senegal, Nigeria, Cameroon, Tanzania,
Material FFB O/B OY
(t ha-1 yr-1) (%) ( t ha-1 yr-1)
Madagascar and Angola germplasms with high
bunch number characteristic were identified for
PS 1.1 28.6 28.0 >8.0 the purpose.
Clone P456 33.0 29.0 8.0 – 10.0
Clone P325 29.0 37.5 10.4
Compact Architecture
Clone P126 (HD) 32.3 28.0 9.0
National average* 18.2 20.3 3.7
One practical approach to increase oil palm yield
Note: *Average of five years data, from 2012 to 2016. is by increasing the number of productive oil palms
FFB - fresh fruit bunch. per hectare. Oil palms are commercially planted
O/B - oil to bunch. with a density of 120–150 palms ha-1 (Woittiez et al.,
OY - oil yield.
2017). Jusoh et al. (2003) stated that the maximum
income could be obtained from a planting density of
On the other hand, efficient management system 148 palms ha-1, contrary to the conventional practice
which includes better harvesting methods, less crop of 136 - 148 palms ha-1. However, the planting distance
losses due to pests and diseases, less spoilage during is normally denser on peat soils, at 160 palms ha-1.
transport and storage, and more efficient processing Palat et al. (2012) reported that an effective strategy
in mills, also play a role in increasing the yields of oil for yield maximisation is high-density planting
palm (Murphy, 2007). followed by selective thinning at 8 - 9 years after
planting. Some early signs of depressive effect on
Adaptation to Climate Change yields were found for high planting densities (180
and 205 palms ha-1). Such a negative impact was not
Some data on climate change revealed by the severe enough to counteract the effects of the higher
National Aeronautics and Space Administration number of palms per hectare (Bonneau et al., 2014).
(NASA) were very alarming. Carbon dioxide (CO2) Recently, Breure (2017) reported that optimal density

444
 DESIGNING THE OIL PALM OF THE FUTURE

can be derived from the leaf area index (LAI). The In terms of oil quality, E. oleifera has higher
LAI value at mature crown size or at about 10 years levels of oleic and linoleic acids and its IV can
after planting would give the most realistic estimate reach up to more than 90 (Rajanaidu et al., 2000),
of optimal planting density. For maximum yield per thus this species should be examined and assessed
hectare, the optimal LAI was assumed to lie between effectively. In E. guineensis, several promising lines
5.5 and 6 (Breure, 2010). Besides planting density, of germplasm with 50%-60% oleic acid have already
oil palms with lower height increment (HTi) found been identified and show great promise to produce
in some Nigerian-derived progenies are amenable a combination of altered fatty acid profile and high
to easier harvesting and mechanisation (Junaidah OY with existing commercial lines (Isa et al., 2006).
et al., 2008; Isa et al., 2008). Based on the extensive Meanwhile, several varieties of oil palm germplasm
screening of MPOB germplasm, breeding for shorter from both oil palm species have been identified with
RL and lower HTi will open up the possibility of high carotene content which has stimulated great
developing compact palms suitable for higher interest in the potential of palm oil in the lucrative
density planting and subsequently increase oil palm health foods market (Rajanaidu et al., 1999; Mohd
yields. Numerous studies on the development of Din et al., 2006). In recent years, a few molecular-
compact palms have been reported. Guzman and based studies have identified quantitative trait
Peralta (2010) reported that six compact clones with loci linked to FAC in oil palm (Singh et al., 2009;
excellent agronomic characteristics such as short Montoya et al., 2013; Ting et al., 2016). This is in view
leaves and stems, precocious and high yielding, to understand the molecular mechanisms associated
have been selected for commercial production. with the inheritance of higher unsaturated oil.
Standard E. guineensis lines such as Deli, Ghana and Breeders and researchers should take multi-
Nigeria crossed with COMPACT palms produced pronged approaches to address this challenge using
leaves of 6.6 - 6.9 m long and can be planted at conventional and molecular breeding strategies to
a density of 170 palms ha-1 which is still higher achieve the prospects for higher oil quality.
than the industry standard of 138-143 palms ha-1
(Corley and Tinker, 2003). Current research in Tolerance to Diseases
MPOB reported that P126 palms exhibited short
RL of about 4.4 m on average which were possible Basal Stem Rot (BSR), caused by a white rot
for use in high density plantings (Zamzuri, 2011). fungus known as Ganoderma boninense, is a major
Recently, Norziha et al. (2017) reported that MPOB- threat to the oil palm industry (Cooper et al.,
Tanzania germplasm showed good potential as 2011), especially in South-east Asia. This currently
breeding material for compactness with reasonable incurable disease has caused major economic losses
yields and would be introgressed into advanced to Malaysia and Indonesia, the two largest palm
breeding populations to generate future value- oil producing countries. Malaysia alone recorded
added planting materials in Malaysia. approximately USD 500 million of yearly losses
(Assis et al., 2016). Far from solving this very
Oil Quality problem, experts predicted that the infected area
would expand to 443 430 ha in 2020 (Roslan and
The quality of oil is a reflection of its FAC and Idris, 2012) or approximately 8% of the current total
this determines its functionality. Palm oil is about oil palm plantation area in Malaysia. Application
50% saturated fat and equal in monounsaturated of chemical and biological agents to overcome the
fats. This profile restricts the potential uses of palm disease has been implemented but the results are less
oil beyond a relatively basic commodity function than satisfactory (Shamala, 2013). To address this
for either edible or oleochemical markets (Murphy, crucial issue, MPOB together with industry members
2007). It is envisaged that modifying or changing the had ranked Ganoderma resistance as one of the top
FAC of palm oil could improve or extend its usage. priorities for oil palm improvement in Malaysia
Particularly of interest is increasing the proportion through breeding programmes (Basri, 2003). Idris
of monounsaturated fatty acids (particularly oleic et al. (2004), who conducted seedling inoculation
acid) and reducing the content of saturated fatty screenings from multi-parental oil palm populations
acids (e.g. palmitic acid). Such oil has the potential in search for genetic resistance against Ganoderma,
to open up markets for oil palm in the liquid oil found that DxP Zaire x Cameroon progenies
sector and would compete more effectively with displayed partial resistance. This line of study
other oil crops such as soyabean, rapeseed and provides useful information on the genetic diversity
sunflower. Apart from the FAC, improving the IV, of multi-parental populations and is important for a
which is a measure of oil unsaturation, and carotene rational use of genetic resources. Moving forward,
content are also sought after. Oil high in carotene is plant breeders need to include analysis of genetic
desirable as there is a growing demand for natural variation of potential and elite planting materials
sources of carotene for food colouring (Corley and for a more holistic understanding. It will also help
Stratford, 1998). in monitoring germplasms as well as predicting the

445
JOURNAL OF OIL PALM RESEARCH 29 (4) (DECEMBER 2017)

potential of Ganoderma tolerance trait. In addition, forms at the nursery stage, prior to field planting.
to increase precision in selection of tolerant planting It beneficially serves as quality control tool to
material, a combination of molecular marker eliminate dura contamination in commercial planting
technologies with advanced quantative trait loci materials. Another assay, the SureSawit VIR kit,
mapping technique may provide a powerful tool allows planters to select the more easily identified
to unravel candidate genes controlling qualitative mature virescens (Vir) palms instead of planting
disease tolerance, not only towards Ganoderma, but nigrescens (Nig), thus helping to reduce loose fruits
also for other oil palm associated diseases including and optimal oil content could be harvested. Oil
Fusarium wilt, Coconut Cadang-Cadang Viroid palm production can then be increased by close
(CCCVd), bud rot, etc. to one-third without the burden of expanding oil
palm land area. With the available oil palm genome
sequence, it is foreseen to help oil palm researchers
TECHNOLOGIES FOR IMPROVEMENT across the world in creating sustainable oil palms
for the future.
Genome Sequencing and Molecular Markers Linkage mapping has been a foundation in the
genetic analyses of quantitative trait loci in plants.
Conventional oil palm breeding is influenced Quantative trait loci mapping in oil palm was
by environmental factors, phenotyping accuracy constructed using various marker technologies to
and requires long generation cycles (10-12 years). identify quantative trait loci associated with desired
Efforts to increase palm oil yield and other traits of traits such as yield (Rance et al., 2001), height
interests have been initiated via various research (Pootakham et al., 2015), oil composition (Singh
strategies involving biotechnological techniques. et al., 2009; Montoya et al., 2013; Ting et al., 2016)
In 2013, MPOB with international collaborators and also for callogenesis and embryogenesis in oil
reported on the 1.8-gigabase (Gb) genome sequence palm tissue culture (Low et al., 2008). Searching for
of Elaeis guineensis (Singh et al., 2013a). The oil more important complicated traits such as yield
palm genome sequence greatly helped in mining and Ganoderma tolerance using quantative trait loci
genes of interest controlling important traits that mapping is now of greater concern for MPOB to
were previously targeted conventionally by oil ensure the sustainability of the Malaysian palm oil
palm breeders. Meanwhile, molecular markers are industry.
rapidly evolving and have been improved by far Linkage disequilibrium (LD) mapping or
with new technologies and knowledge in the oil association mapping (AM) has been used to find
palm genomics field. The ability to link specific linkages among independent populations (Soto-
markers with useful traits via molecular method Cerda and Cloutier, 2012). The mapping resolution
enables marker-assisted-selection (MAS) which will be higher with unlimited number of alleles,
allows selection based on DNA information is thus marker trait-association could be discovered
more reliable in speeding up genetic improvement more efficiently (Flint-Garcia et al., 2013). Recently,
compared to conventional breeding. In oil palm SNP markers with significant linkages to oil palm
research, different types of markers were developed yield, fruit to bunch (FTB) and oil to dry mesocarp
and used in different areas of studies such as genetic (OTDM) have been identified (Chuenporm and
diversity (Zulkifli et al., 2012; Zaki et al., 2012; Okoye Volakert, 2017). AM is suitable for the analysis of
et al., 2016), DNA fingerprinting (Lim and Rao, the MPOB germplasm collection of E. guineensis and
2005), quantitative trait loci mapping (Mayes et al., E. oleifera previously reported to possess high genetic
1997; Singh et al., 2009) and association mapping variability (Ong et al., 2015). Thousands of materials
(AM) (Chuenporm and Volakert, 2017). A modern from this collection could be extensively screened
molecular marker, single nucleotide polymorphism and outstanding palms with interesting traits such
(SNP), offers several advantages over classical as low height, thin shell, long stalk, high BI, vitamin
markers in terms of its abundance, genetic stability, E and carotene can be potentially identified.
and it is also amenable for high-throughput Two new approaches, namely genomic wide
automated analysis (Vignal et al., 2002). It is proposed association studies (GWAS) and genomic selection
that this marker should be extensively used for oil (GS), have emerged as more effective selection
palm improvement in future. The complete oil palm tools. In oil palm, GWAS has been implemented
genome sequence and high density SNP array (92 to increase selection gain per unit time and cost
459 SNP) are now available at MPOB. with small populations (Wong and Bernardo,
The SHELL gene, which has been successfully 2008), predict GS accuracy (Cros et al., 2015), select
mapped and identified, controls a monogenic trait higher oil content (Bai et al., 2017) and OTDM
responsible for the dura, pisifera and tenera fruit (Teh et al., 2016). GS is a method broaden from
forms in E. guineensis (Singh et al., 2013b). The MAS but uses a wide coverage of markers in the
SureSawit SHELL Gel Kit was then developed and genome so that breeding values of complex traits
commercialised for early screening of different fruit can be identified precisely (Goddard and Hayes,

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 DESIGNING THE OIL PALM OF THE FUTURE

2007) without needing accurate knowledge of their of pisifera descendants by ensuring that the
specific location. It allows the early selection of elite selected palms from existing pisifera descendants
materials and maximises genetic gain per unit time. are legitimate (Siti Hawa et al., 2015). Using DNA
In oil palm and other perennial crops, increasing fingerprinting, Rajinder et al. (2007) reported SSR
genetic gain requires adequate GS accuracy in markers can be used for clonal identification,
selecting candidates to shorten breeding cycles and monitoring line uniformity between and within
increase selection intensity (Cros et al., 2015). GS is clones and detecting culture mix-up.
not yet widely implemented by oil palm genome
researchers, hence the information on the reliability Phenomics and Phenotyping
of GS for oil palm is still limited. GWAS and GS have
now been seriously initiated and will hopefully Phenomics is defined as systematic high
accelerate genetic improvement of current materials throughput phenotype studies on a genome wide-
via genomics-guided breeding. scale (Bilder et al., 2009). It is a new experimental
approach developed to relate genomics features
Plant Variety Protection (PVP) and DNA to phenotype with the aim to link to phenotypic
Fingerprinting traits (Hancock, 2014). This technology accelerates
progress in understanding gene function and
Breeding of new plant varieties requires environmental responses. Plant phenomics study
investment of time, labour and money. It may take on plant growth, performance and composition
many years of research and development to release for valuable traits will enable breeders to develop
a new variety for commercial planting. Once a new new oil palm varieties to support future production.
variety is released, there is a high risk of it being Phenomics also makes it possible to trace traits and
reproduced by others, thus depriving its breeder/ facilitate the introgression of novel variation from
developer the opportunity to acquire his rightful wild germplasm (Tester and Langridge, 2010).
profit. The New Plant Variety Protection Act 2004 Phenotyping work is widely recognised as the
by the Department of Agriculture (DOA), Malaysia most laborious and technically challenging part.
was enforced with the objectives to provide rights, For a germplasm to be screened for a valuable
acknowledgement and protection to those bringing agricultural trait, replicated trials are necessary
in new plant varieties. Further investment and across multiple environments. Furthermore, this
development of more new plant varieties may also phenotyping work needs to be done precisely
be motivated due to this Act (Laws of Malaysia, especially when the phenotypic data will be used in
2004). Based on this Act, Oil Palm Test Guidelines molecular research. The phenotyping bottleneck can
(OPTG) for distinctness, uniformity and stability of now be addressed by combining novel technologies
new palm varieties has been drawn up (DOA, 2010). such as noninvasive imaging, spectroscopy, image
These test guidelines can be applied to all varieties analysis, robotics and high-performance computing
of E. guineensis Jacq. and its derivatives including (Furbank and Tester, 2011). As a result, field
interspecific hybrids (E. guineensis x E. oleifera), their evaluation of palms performance is much faster,
reciprocal crosses and backcrosses. Phenotypic traits more dynamic and reliable.
such as RL, petiole cross-sectional area, bunch shape,
fruit type, fruit shape and mean fruit weight are Bioinformatics and Database
recognised descriptors to differentiate each variety.
DNA-based fingerprinting could be the best option Bioinformatics has been described as new
to distinguish at a molecular level the varieties for discipline that involves the biologist, mathematicians
patent protection as well as registration under the and computer scientist in efforts to interpret a large
New Plant Variety Protection. scale of data generated by genome sequencing and
Legitimacy confirmation is important in other ‘omic technologies (Kanehisa and Bork, 2013;
oil palm breeding programmes as controlled Murphy, 2014). In the past decades, bioinformatics
pollination, seed germination, nursery and field has become critical part of genomic research and
planting are labour intensive and tedious which functional genomic technologies. The advancement
may lead to pollination contamination by stray of bioinformatics tools has brought about a
pollen or seedling mix-ups, hence some or all deeper understanding of genome architecture and
of the progeny would be illegitimate. With the accelerated biotechnological research innovations
deployment of DNA fingerprinting techniques, and huge data interpretation (MPOB, 2017b). The
quality control testing has been accelerated to as recent leap of sequencing projects has led to a huge
early as the seed stage which has been implemented amount of DNA that needs to be analysed and
by Sime Darby Plantation (Arutchelvam et al., assembled. Genetic maps, genotypes or expression
2017). FGV has a well-developed oil palm DNA information must be analysed to obtain relevant
fingerprinting method and their current focus is biological information. In oil palm, the Genomsawit
to assist breeders in improving the next generation web portal (Rosli et al., 2014) and MYPalmViewer

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JOURNAL OF OIL PALM RESEARCH 29 (4) (DECEMBER 2017)

genome browser (Low et al., 2015) were developed SNP arrays could enable reverse breeding of oil
by MPOB as resources to identify relevant palms (Ting et al., 2014). In future, haploid research
information of the genome data. Efforts will also be will have an enormous impact on conventional oil
focused on improving annotation and identification palm breeding because it can significantly reduce
of marker-trait association to support molecular- the time taken to develop a new planting material
based research. compared with traditional breeding techniques.
The process of collecting accurate data in field Furthermore, it can also be used to create palms
works is tedious and time-consuming. To meet with novel characteristics.
research requirements, data must be presented in
the most structured, systematic and scientific way. Epigenetics
In MPOB, a system called the MPOB Breeding
Information System (MPOB-BIS™) has been In addition to genomics research, epigenetic
developed as a significant tool in accumulating information may also provide opportunity for
field data (Mohd Din et al., 2012). This user-friendly traits enhancement. However, this area of research
conceptual system has enabled all information is relatively less explored compared to genomic
related to research data to be managed and studies. Epigenetic occurrences such as transgenic
implemented with the latest use of information silencing, paramutation, imprinting and DNA
technology in a more systematic manner. The system methylation are quite common phenomena in
also allows data to be interpreted and statistically plants and potentially offer molecular resources for
analysed more effectively. This module has also targeted crop improvement (Springer and Schmitz,
significantly reduced labour and material costs by 2017). A recent breakthrough by exploiting epigenetic
reducing the time needed for data checking and information has successfully unravelled the cause
analysis. of floral abnormality or known as ‘mantling’ of
clonal oil palm planting materials. Changes in DNA
Haploid Technology methylation of the ‘Karma’ retro-element, located
within an intron of EgDEF1, was found to be
Current oil palm seeds used for commercial involved in defining the mantled phenotype (Ong-
plantings are produced from individual parents Abdullah et al., 2015). Subsequently, a KARMA
selected from genetically heterogeneous populations screening assay was commercially developed to
of non-homozygous palms. Thus, seeds produced screen nursery plantlets (Ong-Abdullah et al., 2016).
from oil palm crosses are not genetically uniform. This would allow assessment of the mantling risk
This genetic variation impedes the oil palm industry and early culling of plants to circumvent economic
from selecting specific genotypes for high yield and time losses involved in the event of planting
or other desirable traits. With doubled haploid mantled palms. Therefore, incorporating epigenome
technology, homozygosity is achieved in one profiling and engineering into oil palm research may
generation. Thus, the oil palm breeder can eliminate complete the missing piece of the puzzle towards
the numerous cycles of inbreeding which is usually designing the oil palm of the future.
necessary to achieve practical levels of homozygosity
by conventional methods. Kumar and Sparjanbabu Genome Editing
(2013) reported various developments in haploid
breeding with respect to economically important Genetically modified (GM) crop plants through
palm species. Haploids exist at low frequency and conventional breeding have long been utilised for
can be obtained naturally or spontaneously. The first the objectives of improving oil yield and quality.
report of doubled haploid production in oil palm via Over the years, a new approach called genome
haploid tissue culture was published by Sumatra editing (GE) has emerged that enables researchers
Bioscience (Iswandar et al., 2010). A breakthrough in to precisely make improvements by manipulating
oil palm research has been the detection of naturally targeted genes in a diverse range of cell types and
occurring haploids and doubled haploids which organisms (Sattar et al., 2017). GE enables DNA to be
can be exploited immediately in F1 hybrid breeding either inserted, deleted or replaced in the genome
(Dunwell et al., 2010). The current research in MPOB using engineered nucleases, or ‘molecular scissors’.
involves screening of culled oil palm germinated There are few types of these nucleases with their
seeds obtained from germplasms as well as advanced respective pros and cons compared to each other. The
breeding populations. The germinated seeds were latest contemporary clustered regularly interspaced
then characterised based on their abnormalities, short palindromic repeats (CRISPR) along with
prior to planting and followed by flow cytometry CRISPR-associated protein 9 (Cas9) provides several
analysis. High throughput flow cytometry is a useful benefits over the like zinc finger nucleases (ZNF)
method to screen naturally occurring haploids in and transcription activator-like effector nucleases
abnormal oil palm germinated seeds (Wan Ibrahim (TALEN). Using GE technology, the scope of traits
et al., 2011). Genotyping doubled-haploids using that could be modified will be expanded and the

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 DESIGNING THE OIL PALM OF THE FUTURE

identification of key genes that regulate complex ACKNOWLEDGEMENT

traits by genomics and omic technologies will
be much easier. There is an optimistic prediction The authors would like to thank the Director-General
that GM crops will capture people’s attention and of MPOB for permission to publish this article.
commercial GM might be released before the end
of 2020s (Soh et al., 2017). GE studies in oil palm is
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