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Oil Palm and Coconut

Article · November 2013

DOI: 10.1007/978-1-4614-9572-7_11

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Chapter 11
Oil Palm and Coconut

S.A.C.N. Perera

Abstract Coconut and oil palm are the two important plantation crops grown in the
wet tropics. Both the palms are monocots and produce oil from their fruits. Oil palm
currently occupies the topmost position in the international vegetable oil market,
while coconut is a palm with diverse uses in addition to being an oil crop. Out of the
two palms, oil palm has shown a spectacular boom in the world oil trade during the
last 60 years. Systematic genetic improvement programmes of both these palms
have been carried out since the first quarter of the last century due to the importance
of oil palm and coconut. Oil palm and coconut share much similar challenges in
their genetic improvement for better traits owing both of these being perennial
monocot palms of massive stature, cross-pollinating heterogeneous populations and
certain other genetic implications. Despite these inherent constraints, considerable
gains have been achieved in the breeding of both oil palm and coconut through the
transfer of desirable genes from diverse sources into the cultivated material. African
oil palm populations have been most important in transfer of genes from popula-
tions of different origins. In addition, interspecific hybridization between African
and American oil palms has also been carried out with success. Similarly, in coco-
nut, much advancement has been made with respect to yield components, precocity
and shorter stature using intervarietal and interpopulation gene transfer methods.
These gene transfer methods have helped to bring oil palm to the topmost position
in the world oil trade and coconut to provide vegetable oil and livelihoods to
millions of people in the coconut-growing countries.

Keywords Coconut • Intervarietal hybridization • Interspecies hybridization • Oil

palm • Wide crosses

S.A.C.N. Perera (*)

Genetics and Plant Breeding Division, Coconut Research Institute of Sri Lanka,
Lunuwila 61150, Sri Lanka
e-mail: chandrikaperera2003@yahoo.com

A. Pratap and J. Kumar (eds.), Alien Gene Transfer in Crop Plants, Volume 2: 231
Achievements and Impacts, DOI 10.1007/978-1-4614-9572-7_11,
© Springer Science+Business Media, LLC 2014
232 S.A.C.N. Perera

11.1 Introduction

Oil palm and coconut are two of the most important tropical oil crops. Both these
plants are monocots belonging to the family Arecaceae and produce vegetable oil
from their fruits. Oil palm has been the most important oil crop in the world surpassing
soybean since 2006. Coconut oil, although produced in much less quantities, contin-
ues to be the main cooking oil in many of the coconut-growing countries in the
world. Programmes for the genetic improvement of the two palms have been in place
during the last several decades due to the importance of these crops in many tropical
countries. Oil palm and coconut share much similar challenges in their genetic
improvement for better traits owing to both of these being perennial monocot palms
of huge stature, cross-pollinating heterogeneous populations and a number of other
genetic implications. Despite these inherent constraints, considerable gains have
been achieved in breeding of both, oil palm and coconut, through the transfer of
desirable genes from diverse sources into the cultivated material. As a result, both
oil palm and coconut have managed to maintain their respective positions in the
world trade of oil and livelihoods of people depending on these palms.

11.2 Oil Palm

Oil palm cultivation first began in Africa where it still makes an important contribu-
tion in the diets of people and as an industrial raw material. However, oil palm made
an economic revolution in the Southeast Asia, especially in Malaysia and Indonesia,
with Thailand and Papua New Guinea also accommodating significant plantations.
Total cultivated area in Malaysia increased from 300,000 ha in 1970 to over 4.0
million ha in 2008. Total area under oil palm cultivation in Indonesia was over 5.0
million ha in 2008 giving evidence for the economic importance of this crop in these
countries and indicating the increase in the world demand for palm oil. Meanwhile the
production in the African countries has also begun to increase, and plantations have
been started in the South American countries also (Corley and Tinker 2003).
Among all the oil-producing plants in the world, oil palm produces the highest oil
content per hectare per year recording 12 t/ha in some experimental plots and averag-
ing between 3 and 7 t/ha in commercial cultivations. With this high productivity, palm
oil contributes to about one-fourth of the total world production of fats and oils
(Oil World Annual Report 2010). Two types of oils are extracted from the fruits of oil
palm. Palm oil or internationally known as crude palm oil (CPO) is extracted from the
mesocarp of the fruit, while palm kernel oil (PKO) is extracted from the kernel of the
fruit. CPO is the predominant oil out of the two, with a production of 95 % of the total
palm oil production. Palm oil is used in several different ways, mainly as a cooking
oil and margarine, vegetable ghee and shortening production, while about 10 % of
the palm oil produced is used for industrial purposes as oleo chemicals, cosmetics
and more recently as biofuels also. Biodegradable plastics, pharmaceuticals and
nutraceuticals are the recent products demanding palm oil.
11 Oil Palm and Coconut 233

11.2.1 Origin

The oil palm has been hypothesized to originate in Gondwanaland which disappeared
in the drift of American and African continents in prehistoric era (Zeven 1965)
resulting in the separate evolution of African oil palm (Elaeis guineensis Jacq.) and
American oil palm (E. oleifera, previously E. melanoccoca).
There are many historical (Opsomer 1956; Surre and Ziller 1963; Zeven 1965)
and fossil (Raymond 1961; Zeven 1964; Ergo 1997; Sowunmi 1999) evidences for
the evolution of the African oil palm along the gulf of Guinea. There are wild and
semiwild oil palm groves along the coastal belt starting from northernmost Senegal
through Sierra Leone, Liberia, Ivory Coast, Ghana, Togo, Benin, Nigeria, Cameroon
and Angola up to the southernmost Democratic Republic of Congo (Ngando-
Ebongue et al. 2010). Much of the diversity of African oil palm is believed to be
centred in the tropical forests of Nigeria, Cameroon, Congo and Angola. The oil
palm cultivation in Southeast Asia was started in 1848 with four seedlings derived
from the same fruit bunch of a palm in West Africa and planted in Bogor Botanic
Gardens in Java, Indonesia. The entire Southeast Asian palm groves are believed to
descend from these four palms. The American oil palm is endemic to the tropical
region of Central and South America, starting from Mexico in the north to the
Atlantic coast with a discontinuous distribution.

11.2.2 Biology/Botany

Oil palm belongs to the family Arecaceae and the genus Elaeis. The genus Elaeis
includes two species, viz., Elaeis guineensis (Jacq) (African oil palm) and Elaeis
oleifera (previously E. melanoccoca) (American oil palm). Oil palm is a diploid with
a chromosome number of 2n = 32 (Maria et al. 1995) with a genome of 1C = 980 Mbp
(Bennett and Smith 1991; Bennett and Leitch 2005), as discovered from the studies
on African oil palm. The plants are monoecious producing distinct male and female
inflorescences in different cycles. It is mainly outcrossing with the varying lengths of
male and female cycles largely depending on the genotype and the environment
(Corley and Donough 1995).

11.2.2.1 Elaeis guineensis

The African oil palm E. guineensis is the most widely planted commercial species.
This perennial palm displays an indeterminate growth of its cylindrical pseudostem
with a large crown consisting of 30–45 green leaves. After reaching the reproduc-
tive stage around 2–3 years after field planting, an inflorescence is initiated in the
axil of every leaf as compact and ovoid masses having many spines. Palms usually
produce 1–2 inflorescences per month, and these can be female or male depending
on whether the palm is in male or the female phase (Adam et al. 2005). The female
234 S.A.C.N. Perera

inflorescence, upon pollination, develops into a fruit bunch in about 22–26 weeks
after pollination (Ngando-Ebongue et al. 2010). The weight of fruit bunches varies
from about 10–50 kg having a total of about 500–4,000 fruits/bunch averaging
about 1,500. The oil palm fruit is a sessile drupe of generally ovoid shape weighing
about 3–30 g. The fruit turns red to brown at maturity and consists of the pulp which is
the mesocarp, shell and the kernel. The mesocarp is about 60–90 % of the fruit weight
and 35–55 % of the bunch weight and produces edible, orange-reddish-coloured oil
which is referred to as palm oil. The kernel or the endosperm produces a clear yellowish
colour oil, which is known as palm kernel oil, and this oil is similar to coconut oil.
In African oil palm thickness of the shell is a main characteristic, because it is the
criterion which distinguishes the three types of palms within this species. Shell thick-
ness is a trait, which shows monogenic inheritance of the shell gene (Sh). The three
types of oil palm populations are named Dura, Pisifera and Tenera out of which
Dura is the predominant type with a frequency of about 97 % in the wild palm
groves (Ngando-Ebongue et al. 2010). Dura palm is characterized by a thick shell
of about 2–8 mm in thickness and a mesocarp to fruit percentage of 35–75 %.
The shell is not present in the fruit of Pisifera which is the second type of
E. guineensis palm. This type contains a high mesocarp to a fruit ratio of 90–99 %.
Pisifera palms are rare in the wild, and it is characterized by a higher degree of
female sterility due to the common characteristic of dying of female inflorescences
before reaching maturity.
The third palm type is termed Tenera, and it contains a thin shell in the range of
0.5–2 mm and a thick fibrous mesocarp. The mesocarp to fruit ratio varies between 55
and 96 %. Tenera form resulted from a cross between Dura and Pisifera palms. Tenera
hybrids, produced by artificial pollination, have been widely used in commercial
plantations worldwide.

11.2.2.2 Elaeis oleifera

American oil palm E. oleifera has been evolved in the Central and South America
(Rajanaidu 1986). E. oleifera is characterized by a slow-growing shorter stem reaching
to about one-fifth of the height of E. guineensis. This palm also often becomes procum-
bent with an erect crown when it reaches maturity (Hartley 1988). The leaflets of
E. oleifera form only in one plane along the leaf petiole similar to coconut, while
in E. guineensis, more numbers of leaflets originate from several planes. E. oleifera
also produces distinct male and female fruit bunches, while the pollen is of foul smell.
The fruit bunches are of conical shape, spiked with shorter spines and they are partly
covered by spathes even at maturity. They bear smaller fruit bunches than the African
oil palm with an average weight of 8–12 kg, and the fruits are also smaller with a
shell thickness of 1–3 mm. The mesocarp to fruit ratio is between 29 and 50 % for
normal fruit amounting up to 80 % in parthenocarpic fruit (Corley and Tinker 2003).
The fruits, at maturity, turn orange in colour. The palm oil extracted from the mesocarp
of E. oleifera fruits contains a high unsaturated fatty acid content (59–90 %) as
compared with E. guineensis. E. oleifera, the American oil palm, is inferior with
respect to bunch yield and oil extraction rate when compared to E. guineensis.
11 Oil Palm and Coconut 235

Table 11.1 Genetic resources of oil palm and their important characteristics
Species Country Important characteristics
E. guineensis Nigeria Higher unsaturated fatty acid Content, short stature
E. guineensis Zaire and Cameroon Tolerance to fungal disease Ganoderma
E. guineensis Angola Larger fruits with high carotene content
E. oleifera Ecuador Thick mesocarp, small stature
E. oleifera Costa Rica High oil yield
E. oleifera Suriname Compact stature
E. oleifera Colombia High unsaturated fatty acids percentage

11.2.3 Goals in Oil Palm Breeding

Yield increment is the most important and the commonest goal in breeding any crop;
this holds true for oil palm also. Yield in oil palm consists of two components: bunch
yield (number of bunches and bunch weight) and oil content per bunch (number of
fruits/bunch and ratios of mesocarp/fruit and oil/mesocarp). In addition to yield, there
are several other oil palm improvement objectives which may vary from country to
country depending on their relative importance. For example, dwarf stature is an impor-
tant objective in countries like Malaysia where labour is a limitation for harvesting
tall palms. In addition to these, high oleic acid content, resistance to diseases such
as Ganoderma (mainly in Southeast Asia), Fusarium wilt resistance (in Africa), bud
rot and fatal yellowing (in South and Latin America) and drought tolerance (mainly
in Africa) are among the major goals in oil palm improvement programmes.
Certain genetic resources of oil palm have been reported to carry genes which are
essential to achieve the above objectives (Soh et al. 2010). Table 11.1 summarizes
some of the traits reported in different oil palm populations.

11.2.4 Genetics and Breeding of Oil Palm

There are two factors dominating oil palm breeding research, namely, shell thickness
of the fruit and the long selection cycles. Shell (endocarp) thickness is controlled by a
major gene Sh (Beinaret and Vanderweyen 1941). Dura palms are homozygous for the
thick shell, while the Pisifera palms are the other homozygotes producing no shell.
The heterozygote, Tenera, is thin shelled. The lack of a shell in Pisifera is proposed to
be caused by a mutation in the Sh gene resulting in failure of lignification of the region
where the shell would form (Sparnaagi 1969; Bhasker and Mohankumar 2001). In
Dura, wild gene homozygous alleles replace 30 % of the mesocarp with shell thereby
reducing the oil yield derived from the mesocarp. Pisifera, although lacking in a shell,
does not give increased yields because in most of the Pisifera germ plasm, fruit
bunches abort during development resulting in negligible yield. Certain female fertile
Pisifera germ plasm has also been identified, suggesting a close linkage of the shell
thickness Sh gene with a fertility gene.
236 S.A.C.N. Perera

Selection cycles in oil palm are long. For Dura female plants, it is about 10–12
years, while for Pisifera male parents, the selection cycles are of about 16 years.
This indicates a limited number of selection cycles over the oil palm improvement
programmes compared to many of the shorter duration plants.
Several breeding methods have been adopted in oil palm with due considerations
to the cross-pollinating breeding behaviour of oil palm. Out of these, reciprocal recur-
rent selection and the family and individual selection are the two most widely used
breeding methods. In addition to these two breeding methods, backcross breeding
has been used for the introgression of desirable characters from unimproved geno-
types to a commercially viable genetic background. Furthermore, breeding methods
for clonal propagation and index and BLUP selection (Soh 1994) have also been
attempted in oil palm breeding over the years.

11.2.5 Alien Gene Transfer in Oil Palm

Many of the above-mentioned breeding methods in oil palm are concerned with
transferring a trait of importance from one genetic background to another. This trans-
fer of novel genes to a different genetic background is called the alien gene transfer
in the sense that the transferred gene, or in most cases the allele, has previously been
lacking in the recipient genotypes. Alien gene transfer in oil palm in most occasions
has been between populations of different origins. However, more distant gene
transfers, i.e. interspecific hybridizations, have also been successfully achieved in
this perennial oil crop.
Transfer of genes essentially requires genetic materials which differ for the trait
that is transferred. As discussed previously, there are two species in the oil palm
genus Elaeis. However, the majority of the gene transfer in oil palm has been between
the populations of different origins. In oil palm genetic resources, there are no variet-
ies as per, the exact meaning of the word variety (Soh 1999). However, Dura, Pisifera
and Tenera are the three main types of oil palm which have formed the base material
for the genetic improvement of this palm. In addition to these palm types, popula-
tions of different origins of these genetic resources have been observed to have
considerable differences among them. These populations have widened the genetic
base of the oil palm breeding material over the last few decades.

11.2.6 Genetic Resources of Oil Palm for Alien Gene Transfer

Oil palm was introduced to Malaysia and Indonesia by the European colonists. Four
Dura seedlings known to have obtained from the same fruit bunch of a palm in West
Africa and planted in Bogor Botanic Gardens in Indonesia formed the base material
for the plantations in these two countries. The progenies resulted from selections
and hybridizations among these thick-shelled Dura plants were planted in Deli
11 Oil Palm and Coconut 237

Province in Sumatra and from there were taken to Malaysia (Rosenquist 1986).
This population was named as “Deli Dura” and became a commercial cultivar in
both Malaysia and Indonesia from 1911 to the 1960s. Such populations which are
derived from a few progenitors resulting in a narrow genetic base are referred to as
Breeding Populations of Restricted Origins (BPRO’s) (Rosenquist 1986).
Deli Dura BPRO has given rise to several subpopulations by distribution to
different localities and subsequent selections. La Me Deli, Ulu Remis Deli, Serdang
Deli and Elmina Deli are examples for such Deli populations. The two populations
Dumpy and Gunung Melayu are dwarf type mutants of Deli BPRO.
AVROS BPRO is a Pisifera population. It has been derived from a high-performing
single plant named Djongo located in Zaire and was developed in Sumatra. AVROS
BPRO has been widely used in oil palm breeding programmes in Malaysia,
Indonesia and Papua New Guinea, Colombia, Costa Rica and Thailand. Characteristic
features of AVROS BPRO are vigorous trunk growth and large fruits with thick
mesocarp producing high yield.
Yangambi BPRO was produced in Congo at Yangambi Research Station from
open pollinated progenies of the Djongo palm and other Tenera palms from
Yawenda, Isangi and N’gazi. Main characteristics of Yangambi BPRO are similar to
those of AVROS BPRO. However, a short-stemmed Yangambi variant also has been
developed in this BPRO.
• La Me BPRO was developed in Ivory Coast by CIRAD (French Agricultural
Research Centre for International Development), which was then known as IRHO.
The ancestral seeds of this Tenera population have been collected from the wild oil
palm groves in the Ivory Coast. Smaller stature, high number of smaller bunches
bearing smaller fruits and the tolerance to less favourable growing conditions are
the characteristics of this BPRO. La Me BPRO is a base material in West Africa
and Indonesia oil palm breeding programmes guided by CIRAD.
• Ekona BPRO has been developed from the wild palms in the Ekona area in
Cameroon by the Unilever plantation group. Certain progenies of this BPRO are
high yielding and show resistance to Fusarium wilt disease. This BPRO generally
bears smaller fruits with high oil content. Ekona BPRO has been distributed to
Malaysia, Costa Rica and Thailand through Unilever group.
• Calabar is a BPRO developed in Nigeria by the Nigerian Institute for Oil Palm
Research (NIFOR). Progenies of NIFOR’s Calabar selections have been distributed
to Ghana, Costa Rica, Indonesia and Malaysia also (Soh et al. 2010).

11.2.7 Derived and Recombinant BPROs

BPROs are interbred or introgressed to develop new BPROs from the traditional
BPROs. The main objective is to transfer the desirable genes from traditional BPROs
to new recombinants to have mixed favourable genes in the resultant progenies. Ulu
Remis teneras, Dumpy AVROS, Dumpy.Yangambi.AVROS, La Me × Dumpy AVROS
are some examples to derived and recombinant BPROs (Soh et al. 2006).
238 S.A.C.N. Perera

11.2.8 Achievements and Impacts of Alien Gene

Transfer in Oil Palm

Oil palm yields have recorded a considerable increase over the last 5–6 decades.
Out of the total increase, 70 % is reported to be due to genetic improvement and
30 % to the improvements in agronomic practices (Davidson 1993). A major portion
of the yield increment caused by breeding is due to the transfer of genes from
Pisifera parents into Dura parents to produce Tenera hybrids.

11.2.9 Tenera Hybrid Improvement

Dura (D) type Deli populations were the only commercial planting material in
Malaysia and Indonesia until the 1960s. With the elucidation of the monogenic
inheritance of the shell gene (Sh), the cross between the thick-shelled (D) parents
with the shell-less Pisifera (P) parents was observed to result in 100 % thin-shelled
Tenera (T) palm, indicating incomplete dominance of the shell gene. D palms were
used as the female parent while P palms brought to Southeast Asian region from
Africa always served as the male parent in T development owing to the common
female sterility of the P palms. The transfer of the wild allele of the gene Sh from P
populations to the cultivated type D is the first reported transfer of alien gene
between the populations of oil palm.
The evaluation of early Tenera hybrids revealed the presence of important differ-
ences with respect to vegetative characters, bunch yield components and the ratio of
oil to mesocarp between the palms from different origins. Based on these results,
two groups of E. guineensis were identified. The palm in group A is characterized
by a small number of large bunches (Southeast Asian Deli palms), while group B
palms produce a large number of smaller bunches, and these are generally from
Africa. The early studies further revealed the significantly higher yielding capacity
of crosses between Asian Deli and African populations. This inter origin gene transfer
led to the success of Tenera hybrids (Gascon and de Berchoux 1964). The findings
also led to the choice of parents from between the different origins; group A Deli as
female parents and group B, African Pisifera palms, as male parents.
Upon the introduction of T hybrid, it soon became the most favoured planting
material for commercial plantations of oil palm in a very short period of time
(Hartley 1988). The switchover of planting material from the thick-shelled thinner
mesocarp (60 % mesocarp to fruit content) producing D palms to thin-shelled,
thicker mesocarp producing T (80 % mesocarp to fruit content) would account for a
minimum of 30 % mesocarp oil yield increase in addition to the increase in the
yields of FFB (Soh et al. 2009). During a period of two decades, about two genera-
tions of improved T materials have been planted, reporting an increase of oil yield
from 5 to about 10 tons/ha/year in research trials. Lee et al. (1990) and Rajanaidu
et al. (1990) estimated a T improvement of 6–7 % by progeny testing the same Ps on
11 Oil Palm and Coconut 239

Fig. 11.1 A Tenera hybrid oil palm

two successive generations of selected Ds. Selecting the best 15 % of P in hybridization

resulted in a 12 % yield increase (Lee and Yeow 1985). However, in commercial
seed production, about 30–50 % of the top P parents are used resulting in a yield
increment of only 10–15 % per generation for the T hybrids, which is reasonable
with the selection of both the parental populations (Soh et al. 2003). Therefore, the
average yield increment of 15 % per generation over two successive generations
estimated by Hardon et al. (1987) is due to the T hybrid improvement with selection
of both the parental populations (see Fig. 11.1).

11.2.10 Interspecific Hybrids in Oil Palm

Much of the breeding and genetic improvement efforts in oil palm were centred on
the species E. guineensis, the African oil palm. The American oil palm species
E. oleifera received much less attention up until recently, due to its low bunch yield
and poor oil extraction rate compared to its African relative. However, E. oleifera
possesses its own desirable traits which have been left unexploited until recently in
commercial plantations. Oil extracted from the mesocarp of the fruits of E. oleifera
palms are characterized by a high saturated fatty acid content of 59–91 % as against
25–72 % for that of E. guineensis (Ngando-Ebongue et al. 2010). In addition, shorter
stem, compact growth habit and, most importantly, the resistance to fatal yellowing
are the economically important characteristics of E. oleifera.
240 S.A.C.N. Perera

Transfer of genes for fatal yellowing, shorter stature and changes in fatty acid
composition from E. oleifera to E. guineensis planting material has been given pri-
ority in the breeding programmes in several countries. To achieve this gene transfer,
the interspecific hybrid, E. oleifera × E. guineensis, has been attempted. Despite
being of two different species and geographical isolation, the two species were
found to be cross compatible, and fertile hybrids could easily be obtained (Hardon
1969; Hardon and Tan 1969; Amblard et al. 1995). However, the major constraint in
this effort was the poor fertility of the hybrids resulting sometimes in seed abortion
(Viegas and Muller 2000). Also on certain occasions, fruit set in interspecific hybrid
was found to be poor (Hardon 1969; Meunier and Boutin 1975). These problems
however were not common in all the attempts at producing and planting of this
interspecific hybrid. Countries plagued with fatal yellowing do not have the option
of planting E. guineensis, and therefore, moving forward with the interspecific
hybrid was essential. The fertility problem of this hybrid was observed to be worse
with the E. oleifera material from Central American origin. E. oleifera germplasm
has been categorized based on geographical location as Central American, Brazilian
and Surinamese. These different germplasm have been observed to produce different
hybrids with E. guineensis, although remarkable differences have also been observed
between progenies from the same origin (Meunier et al. 1976).
The oleifera material used in the hybridization programme has not undergone
much selection process compared to the stringent selection and improvement
programme of the E. guineensis material. Moreover, the germplasm of E. oleifera
has neither been conserved nor been characterized to the extent of E. guineensis
(Rajanaidu et al. 1989). Consequently, prospects for the E .oleifera × E. guineensis
hybrid are not gloomy if efforts are made to properly collect, conserve, characterize
and utilize the E. oleifera germplasm followed by selection of parents prior to
hybridization.
E. oleifera × E. guineensis interspecific hybrid, although is not competitive with
commercial E. guineensis with respect to crude palm oil production, provides the only
option of planting material in the fatal yellowing-affected areas such as in the
American region. The primary objective of the oil palm industry in these areas is to
supply the local market with refined palm oil, and for this purpose, the interspecific
hybrid is sufficiently competitive mainly due to its observed resistance/tolerance.
There are other uses of this hybrid in addition to the oil which is rich in unsaturated
fatty acids (Rajanaidu et al. 1989). The height increment of E. oleifera × E. guineensis
hybrid is much less than the commercially preferred cultivars, increasing the eco-
nomic lifespan up to about three times, depending on the combination of parents.
Artificial pollination may be a remedy for the low natural fruit set resulting in bunch
failure. The extent of the problem associated with E. oleifera × E. guineensis hybrid
is highly variable between progenies (Arnaud 1980; Schwendiman et al. 1982,
1983). In contrast, strikingly high natural fertility has been observed in interspecific
hybrids of Suriname descent (Closen 1987; Rao et al. 1989).
Consequent upon the recognition of the importance of E. oleifera × E. guineensis
interspecific hybrid, in vitro embryo rescue methods have been tested with success
(Alves et al. 2011a, b).
11 Oil Palm and Coconut 241

11.2.11 Transfer of Alien Genes in Oil Palm via

Backcross Breeding

Backcross breeding is an important method for the introgression of desirable genes

from a relatively unimproved genotype into a recurrent host genotype. A series of
backcross breeding programmes are in progress to introgress the desirable traits
(oil quality, dwarfness and disease resistance) of E. oleifera into the best known
combinations of E. guineensis genotypes (Obasola et al. 1977; Tam et al. 1977;
Sterling et al. 1988; Sharma and Tan 1990; Le Guen et al. 1993; Chin 1993; Din and
Rajanaidu 2000; Soh 1999; Sharma 2000). This method of alien gene transfer
although takes a long time, has merits in terms of maintaining the advantageous
genetic background of the recurrent parent.
In addition, backcross breeding method has been used in the development of
AVROS and the Dumpy.AVROS in developing the AVROS and to introgress the
dwarf trait in the Dumpy Deli to AVROS population, respectively. However, oil
palm, being a perennial, backcrossing in the strict sense to the original recurrent
genotype is not possible, rather it is done to the population or the progeny of the
recurrent parent.

11.2.12 Prospects of Alien Gene Transfer in Oil

Palm via Transformation

Gene introgression via genetic transformation is the recent and the most advanced
method of alien gene transfer. It offers the ultimate potential for the introgression of
genes from species that are not only distant but also quite far apart members in
the plant kingdom. The perfected protocols of tissue culture methods have provided
the basis for research into genetic transformation of oil palm. The first report on the
transient expression in oil palm was by using the biolistics approach (Alves et al.
2011a). Studies on genetic transformation in oil palm have made significant progress
since then. Te-chato et al. (2002), Zubaidah and Siti Nor (2003), Rohani et al. (2003),
Adam et al. (2005) and Lee et al. (2006) reported work on establishing conditions
for transformation. The genes needed to achieve genetic transformation for oil
synthesis was reported by Shah and Cha (2000) and Asemota and Shah (2004), for
carotenoids by Khemvong and Suvachittanont (2005) and for kernel expression by
Cha and Shah (2001).
The transformation methods, direct gene transfer and Agrobacterium-mediated
approaches have been used to transfer useful genes such as cowpea trypsin inhibitor
(CpT1) (Abdullah et al. 2003) and Bacillus thuringiensis (Bt) crystal insecticidal
protein genes as a solution for the problems caused by insect pests (Sharma 2000;
Sharma et al. 2002) and also chitinase to address the problems related to basal
stem rot. The synthetic gene Cry1A(b) has been successfully transformed using
particle bombardment method and the expression tested (Lee et al. 2006).
242 S.A.C.N. Perera

Cry1A(b) synthetic gene is intended against the damages caused by Lepidopteran

pests (Masson et al. 1999; Reardon et al. 2004). However, despite much advancement
made in the genetic transformation in oil palm, this method is still not in practice for
the production of planting material. Most Southeast Asian countries, where the oil
palm is planted in massive scale, are still in the process of developing biosafety
mechanisms for the planting of genetically modified crops. Therefore, it will take
more time for the beneficial effects of alien gene transfer via genetic transformation to
come into effect in oil palm.

11.3 Coconut

Beginning from the early twentieth century to about the 1960s, coconut oil occupied
the topmost position as a vegetable oil in the tropics. Lately certain health concerns,
raised due to coconut being saturated oil, reduced its value as a cooking oil in the
international trade. While these concerns are currently been reinvestigated, the
value of coconut in many of the coconut-growing countries remains undiminished
as the term “tree of life” that had been tagged onto the coconut palm remains intact
to date. Coconut palm is identified as a plant having a potential for alleviating poverty
in areas where coconut is grown due to its multitude of uses in addition to being an
oil crop. Coconut can be grown both in plantation scale as well as a home garden
crop making it possible to be used in small scale or cottage industries for its diverse
uses. Copra and oil, desiccated coconut, coir and fibre products, charcoal and other
shell products are the important products from a coconut tree, while tender nut
beverage coconuts, various uses of coconut leaves or fronds and timber industry are
some other examples for the diverse uses of coconut.

11.3.1 Origin

Several conflicting theories regarding the origin and domestication of coconut have
been put forward. A New World origin of coconut with subsequent dispersal to Asia
and Polynesia has been proposed by several groups (Guppy 1906; Cook 1910;
Ridley 1930; Bruman 1944). In these, the centre of origin of Cocoid palms, the closest
relative of coconut, is North-Western and South America. Evidence from fossils and
archaeological studies indicate a South West Pacific origin of coconut (Child 1974;
Purseglove 1965), while Indian fossils and the Madagascar forest coconut support
an Indian Ocean origin (Harries 1995). Even at present the issue of origin of coco-
nut is still not fully resolved. However, coconut is found extensively distributed
throughout the tropics including Central and South America, East and West Africa,
South and Southeast Asia and the Pacific islands. From its putative centre of origin,
coconut has been disseminated to both east and west by floating in the sea and also
by human dissemination (Ohler 1984).
11 Oil Palm and Coconut 243

11.3.2 Biology/Botany

Coconut (Cocos nucifera L.) is a monocot belonging to the family Arecaceae and to
the subfamily Cocoidae. Subfamily Cocoidae includes 27 genera and 600 species,
and coconut is currently the sole species of the genus Cocos. Coconut possesses a
diploid genome with 16 pairs (2n = 2x = 32) of chromosomes. Similar to oil palm,
coconut is a monoecious plant bearing both male and female flowers on the same
plant. However, unlike oil palm, coconut bears both male and female flowers on the
same inflorescence which are produced in a continuous succession on the adult
palms. A coconut palm upon reaching the reproductive stage produces about 12–15
inflorescences per year throughout its economic lifespan. Coconut inflorescence
consists of many flower-bearing spikelets situated on a central axis or a peduncle.
The female flowers are located at the base of the inflorescence while a numerous
number of male flowers occur along the length of the spikelets. Tall varieties of
coconut are predominantly outbreeding due to nonoverlapping of the male and
female phases of the inflorescence. The dwarf or short varieties of coconut are naturally
self-pollinating due to the overlapping of the two phases. The important phenomena
of the inflorescences with respect to the genetic improvement of the palm are the
lack of incompatibilities and the ability to artificial cross-pollination as well as the
artificial self-pollination.

11.3.3 Varietal Groups/Populations of Coconut

A variety is defined as “a general term to denote a single strain or a group of strains

which distinctly differ in structural or functional characters from one another or a
group of the same species which can be depended upon the ability to reproduce itself
true-to-type” (Menon and Pandalai 1958). Many reported types of coconuts in the
world may not qualify as distinct varieties according to the discrete considerations in
the definition. However, there are many different varieties, forms and types of coconut
recorded in the world, although the terminology may not be exact or strictly correct in
the different coconut-growing countries or regions within a country.
Although different classifications of coconut exist, there is one common feature
in all of them, the grouping of coconut as talls and dwarfs. Talls have their specific
characteristics and have gained popularity as commercial planting materials among
the unimproved planting material of coconut throughout the world. Dwarf was later
identified as a variety with a high potential to be used as a parent to transfer favour-
able genes to commercially grown tall coconuts. In certain countries a few different
intermediate coconut varieties, which display characteristics in between the talls
and the dwarfs, have also been reported. King coconut in Sri Lanka (Liyanage
1958), Niu Leka Dwarf in Fiji (Bourdeix et al. 2005a) and Gangabondom in India
(Menon and Pandalai 1958) are few of the examples for this semi-tall/semi-dwarf or
intermediate groups of coconut. A comparison of contrasting features of the different
types of coconuts is presented in Table 11.2.
244 S.A.C.N. Perera

Table 11.2 Comparison of different varieties of coconuts

Trait Tall Dwarf Intermediate
Palm height Tall (20–30 m) Short (10–15 m) Vary from short to tall
Trunk circumference Enlarged with a Thin and no bole Usually enlarged with
bulbous base formation a root bole
Lifespan 60–100 years 40–50 years 40–60 years
Time taken for flower 5–8 years 2.5–4 years Vary from 4 to 6 years
initiation
Mode of pollination Highly crossed Highly selfed Highly selfed
Bearing nature Continuous Seasonal Seasonal to continuous
Nuts/palm/year Average 40–60 Average 80–100 Average 50–100
Whole fruit size Very small to large Very small to medium Medium to large
Copra amount and quality 200 g/nut; good 80–100 g/nut; inferior 100–150; Mainly
inferior
Leaf and bunch attachment Strong Fragile Usually fragile
Pigmentation of nuts Mixture of green, Pure green, brown, Green, red
brown and yellow yellow or red

The main groups of coconut include palms of different morphologies despite

their basic similarity in stature and the pollination behaviour. For example, the talls
include morphotypes which show differences in nut colour and size, shell thickness,
mesocarp or endocarp differences. Liyanage (1958) referred to these as forms
within varieties and Bourdeix et al. (2005a,b) referred to them as phenotypically
distinct variants.

11.3.4 Goals in Coconut Breeding

Similar to other crops high yield is the primary objective of the genetic improvement
of coconut. Coconut has diverse uses but the main economically important yield
parameter is the weight of copra or kernel. Fruit and nut components, number of nuts
and the weight of kernel per nut are the determining factors of per palm coconut
yield. However, there is a negative correlation between the number of nuts and the
size of the nuts in a bunch. Consequently, improvements towards increasing the
rate of inflorescence (resulting in bunch) emission will be one way of overcoming
this problem. Efforts to maintain a balance between nut number and the nut size will
be another approach to increase the kernel/copra yield.
Precocity or the shortening of the vegetative phase is one major objective in
coconut breeding. Coconut is a perennial plant with a long vegetative phase as well.
As a result of the long vegetative phase, which lasts for about 6–7 years, the growers’
waiting period for returns to their investments also becomes long reducing the
appeal of coconut as an economic venture. The genetic potential of the dwarf coco-
nuts for early flowering and bearing provides the required genetic material for
breeding for precocity.
11 Oil Palm and Coconut 245

With the global scenario of climate change and the occurrence of prolonged
droughts, the attention of the coconut breeders has shifted on breeding coconuts for
tolerance to abiotic stresses, water stress being the most important. Coconut is a
monocot with a fibrous root system which does not grow into deeper layers in the
soil resulting in adverse effects from moisture stress than the dicot perennials having
a tap root. Hence, many countries are now focusing on breeding for tolerance to
moisture stress in their coconut breeding programmes to expand the cultivation of
coconut to newer areas and also for higher productivity of the existing plantations.
Breeding for tolerance to biotic stresses (pests and diseases) is a highly beneficial
way of managing and controlling them. So far the most devastating groups of patho-
gens of coconut are the phytoplasmas and viroids which are intracellular, causing
incurable deadly or debilitating diseases. The phytoplasma group is recorded to be
the causal organism for diseases such as lethal yellowing in Africa, Kalimantan wilt
in Indonesia and Kerala Root wilt in India, while Cadang-cadang in Malaysia is
reported to be caused by viroids. The countries plagued by phytoplasma diseases
include tolerance breeding for such diseases as a primary objective in their coconut
breeding programmes (Nair et al. 2006).
Development of short stature of the palms is another objective of coconut breeding.
Most of the old coconut stands are very tall and the scarcity of labour for picking
has become a major problem at present even in large plantations in many of the
coconut-growing countries. Furthermore, short stature is an important character in
cultivars for home gardens also.

11.3.5 Genetics and Breeding of Coconut

Coconut is a difficult to breed perennial plant. Long generation interval, cross-

pollination breeding behaviour of tall coconuts resulting in highly heterogeneous
populations, lack of a viable vegetative propagation method, low number of seeds
produced per palm and the massive stature of the palm are the most important con-
straints in coconut breeding. Despite these limitations coconut breeding programmes
have been started from the first quarter of the twentieth century. In these programmes
two breeding methods, mass selection and hybridization, have been widely used for
the genetic improvement of the coconut palm.
The majority of the coconut plantations in the world are derived from mass selec-
tion. Mass selection was scientifically recognized as the most fundamental method
for coconut breeding (Liyanage 1955). The common selection criteria was the yield
of copra per palm or one of its components such as number of fruits produced or per
nut copra content. In many of selection programmes, seednuts were selected mainly
based on the size, weight and shape of the nuts. The resulting coconut populations
ware subjected to successive selection for fruit characteristics. Around the mid-
twentieth century, the system of selection of the best trees within the best plots
was applied, and coconut breeders throughout the world adopted this system of
mass selection. Three methods of mass selection have been practiced depending on
246 S.A.C.N. Perera

the reproduction system used: mass selection using open pollination, selfing or
intercrossing between parents.
Intervarietal hybridization has been widely practiced in coconut to extract
heterosis from genetically diverse material. Much progress has been achieved with
this method world over mainly with respect to nut yields, precocity and copra
productivity.

11.3.6 Alien Gene Transfer in Coconut

Coconut is the sole species of the genus Cocos. Because of this the alien gene transfer
at interspecies level is not possible in coconut. However, hybridization between
varieties and different populations has been extensively practiced with success to
transmit the characters of interest to cultivated material.
The main coconut varieties tall and dwarf are the genetic material much used in
hybridization to mix important traits. Out of these two varieties, tall has been the
economically important commercial planting material. The tall coconuts are hardy
and tolerate the harsh environments better than the dwarf varieties. In contrast the
dwarfs display a certain amount of seasonality in bunch emission and nut setting
upon pollination. The talls are usually regular and continuous bunch bearers,
although a reduction in nut number is observed during prolonged drought periods.
The nuts of the talls also are bigger and produce more copra per nut (Table 11.2).
In comparison, dwarfs produce a larger number of nuts per year. However, one of
the most troublesome inferiorities in the commercially preferred tall coconut variet-
ies is the long vegetative phase which lasts for about 5–8 years. In comparison, the
dwarfs are precocious coming into bearing in about 2–4 years. Another important
advantage of dwarf coconuts is their short stature compared to tall coconuts.
Coconut hybridization programmes world over have focused on transmitting the
economically important traits precocity, high nut number and shorter stature from
the dwarf varieties to the commercial tall cultivars through hybridization between
the tall and dwarf varieties.
Tall coconuts in the world are reported to be of two types. The first is the popula-
tions of talls in the Southeast Asian and the Pacific origin. The nuts of these tall
coconuts are generally larger and round in shape. The second type is the tall coco-
nuts present in South Asia and the African region. The nuts of these tall coconut
populations are generally smaller and elongated in nut shape than the Pacific talls.
On the other hand the dwarf coconuts are believed to have evolved from the tall
coconuts of the Southeast Asian and the Pacific region (Perera et al. 2000). Even
within the two tall coconut types and the dwarf coconut type, there are many differ-
ent populations and/or subpopulations which are mainly geographically isolated
and distinguished. As such the accepted international naming most of the times
includes the country or the region where a particular coconut type, variety or a
population, has been grown. Sri Lanka Tall, San Ramon Tall, West African Tall and
Fiji Tall are some examples for such naming in tall coconuts, while Malayan yellow
11 Oil Palm and Coconut 247

Dwarf, Sri Lanka Yellow Dwarf and Madang Brown Dwarf are some examples
for dwarf coconuts. These different populations show morphological and genetic
variations, and they do provide the necessary raw material for transfer of important
genes in coconut.

11.3.7 Achievements and Impacts of Alien Gene

Transfer in Coconut

Different types of intervarietal hybridizations have been carried out to transfer

the economically important traits to commercially cultivated varieties of coconut.
The most common intervarietal coconut hybridization has been between the tall and
the dwarf varieties (D × T crosses). Hybridization between the tall and the dwarf is
considered a wide cross in coconut because wider crosses, such as interspecies
hybridizations, are not feasible due to coconut being the sole species of the genus
Cocos. Considerable achievements with respect to the goals of coconut breeding
have been made even with the intervarietal hybridizations between the talls and the
dwarfs in many coconut-growing countries.
The coconut hybrids Mawa (PB 121—cross between Malayan Yellow Dwarf
and West African Tall), Camren (PB 113—cross between Cameroon Red Dwarf
and Rennell Island Tall) produced in Ivory Coast, Matag (PCA 15-2 cross
between Malayan Red Dwarf and Tagnanan Tall) produced in the Philippines,
CRIC 65 (cross between Sri Lanka Green Dwarf and Sri Lanka Tall) produced in
Sri Lanka and Maren produced in the Solomon Islands are a few examples of
widely planted tall and dwarf hybrids. In all these intervarietal hybrids, the char-
acters, viz., high nut number, precocity and shorter stature, have been transmitted
to a certain extent to the tall coconut varieties (see Fig. 11.2). These improved
cultivars have been mass produced and widely cultivated recording yield incre-
ments varying from about 50 to 150 % than the tall cultivars. In addition, the tall
and dwarf hybridization has also resulted in shortening the vegetative phase by
about 30–50 % depending on the parents used. Also these hybrids are shorter in
height compared to their tall parents increasing their appeal to growers (Bourdeix
et al. 2005a, b).
Tall × tall hybridizations between populations of different origins have also been
carried out in coconut in several countries. The coconut hybrid Waren or PB 213 is
produced by crossing the two different tall varieties, West African Tall and Rennell
Island Tall. This cultivar is tall in habit and flowers earlier than its two tall parents
in addition to producing larger nuts. The improved cultivar Wavan or PB 214 is yet
another tall × tall hybrid produced by crossing two different tall populations West
African Tall and Vanuatu Tall. This hybrid has also brought precocity to a certain
degree to tall cultivars in addition to its comparable yields. CRISL 98 is yet another
tall × tall hybrid produced in Sri Lanka. The parents, Sri Lanka tall and San Ramon
tall, are two populations of tall from South Asian and Southeast Asian origins and
are therefore considered a wider cross than two tall populations of the same group.
248 S.A.C.N. Perera

Fig. 11.2 A dwarf × tall coconut hybrid in Sri Lanka

With transmittance of genes for higher copra content and high nut weights from
San Ramon, by now CRISL 98 is the highest per nut copra producer in Sri Lanka
(Perera et al. 2010).

11.4 Conclusion

As has been discussed above, remarkable achievements have been made in increasing
the quantity and quality of palm oil, tolerance to biotic stresses and reducing the
stature of oil palm cultivars through gene transfer between populations of different
origins and gene transfer through wider crosses and interspecies hybridizations in
oil palm. Similarly, in coconut also much advancement has been made with respect
to yield components, precocity and shorter stature using intervarietal and interpopula-
tion gene transfer methods. These gene transfer methods have helped to bring oil palm
to the topmost position in the world oil trade and coconut to provide vegetable oil and
livelihoods to millions of people in the coconut-growing countries. Nonetheless,
keeping in view that avenues for interspecific hybridizations and consequently alien
gene transfers are very limited, particularly in coconut, efforts need to be initiated
towards development of transgenics. Molecular tools also need to be integrated with
conventional breeding in these two most important oil crops in order to hasten the
breeding process as well as develop cultivars with specific traits. In general, as
compared to other commercial crops, genomic resources are a limitation in oil palm
and coconut, and this issue needs to be addressed on priority to initiate a target-ori-
ented and ameliorated breeding for improvement of oil palm and coconut.
11 Oil Palm and Coconut 249

Acknowledgements The author wishes to thank Miss H. D. M. A. C. Dissanayaka, Mrs. I.

Jayawardana and Miss W. G. R. Subhathma for their assistance in numerous ways in the prepara-
tion of the manuscript.

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