CHAPTER II

REVIEW OF LITERATURE

Coconut is one of the most important perennial palms, which

sustains lively -hoods of resource poor farm families in the coastal agro-

ecosystems in the tropics. The coconut palm is a versatile tree; its value is

innumerable to the society. Coconut has a distribution mainly in coastal

region at 20° on either side of the equator. In India, coconut is cultivated in

17 states and 3 union territories. Genetic improvement of coconut has been

aiding the farmers in obtaining high productivity and tolerance to biotic

and abiotic stress. Coconut breeding is constrained by many inherent

problems like lack of vegetative propagation, long juvenile phase, large

area needed for experimentation and seed production and high variability.

Methods of improvement employed in coconut are gene pool enrichment

and introduction, selection and varietal cross hybrids. Several workers

reported variation in morphological and reproductive characters in many

hybrids. Understanding the genetic architecture of the crop is an important

pre-requisite for any breeding program. Genetic architecture in coconut

palm is poorly understood. There are very few reports on inheritance of

important traits of growth, development and yield. A brief account of

biometrical studies, identification of hybrids using molecular markers and

sub cellular organell activities and influence of weather parameters on

yield is presented here.

2.1, Qualitative characters

Quantitative characters of coconut mainly includes colour of plant

parts,nature of inflorescence ,nature of endosperm etc.

Colour of the petiole has been an important character used to

identify hybrid seedlings. Two genes (R-G- Brown, rrG- Green, R-gg

Orange and rrgg yellow) control the colour of petiole (Bourdeix, 1999) in

coconut.

Spicata is a mutant in coconut with unbranched inflorescence with

greater female flower production. Open pollinated progenies of spicata

palm segregated in 1: 1 ratio of spicata and normal palms. Thus, earlier

workers assumed spicata trait to be in (Ss) heterozygous state and

dominant over normal (ss) inflorescence. Spicata palm also had shown

many meiotic irregularities and high degree (12 %) of pollen sterility

(Ninan and Satyabalan, 1963). Androgena palms also had shown meiotic

abnormalities such as aneuploidy (2n-l) in a few cells (Ninan et al., 1960).

Macapuno is a mutant coconut form, which has jelly like

endosperm inside the fruit. This has been exploited in confectionery

industry due to soft texture of endosperm. This trait is controlled by single

recessive gene (mm), which is associated with lethality (Cedo et al.,

1984). The embryos of this genotype can’t germinate normally in nature

and thus tissue culture techniques are utilized to germinate this rare

genotype.

9
2.2. Quantitative characters

2.2.1. Variation in vegetative and floral characters

Several workers reported variability for morphological and

reproductive characters among coconut across the coconut growing

countries.

Luntungun and Liyanage (1978) reported that D x T hybrids

produced significantly higher number of leaves than the tall parents from a

variability study. They also reported significant variation in number of

female flowers and nuts per bunch among the parents and hybrids.

The inflorescence of coconut is a spadix, which develops in the

axil of each leaf. Thus, normally, number of leaves produced annually also

determines the number of spadices developed (Patel, 1938; Child, 1974).

The inflorescence, from the primordial initiation stage, takes about 26

months to emerge out of the leaf axil (Patel, 1938). Initiation of the spadix

begins about thirty three months before it’s opening. The spadix consists

of a main branch with 20-65 branches or spikelets bearing flowers.

Pistillate (female) flowers are located at the base of the spikes, the rest of

the spike being fully covered by staminate (male) flowers. Each spikelet

may have one or more female flowers and generally about two to three

hundred male flowers (Ohler, 1999). Spadix initiation and production are

10
greater during March to September when average day length is more

(Wickramasurya, 1968).

A wide range of phenotypic variations for number of female

flowers, setting per cent, number of spathes per year was reported by

Louis (1981) through a variability analysis over 25 varieties and two

hybrids.

Ninan (1983) compared tall, dwarf green, dwarf orange and their

hybrids for variability. Variability was reported for mean number of

spathes, mean number of female flowers and yield of nut.

Nunez and De Paz (1990) studied the flowering and yield

characteristic of Macapuno- bearing dwarf x tall coconut hybrids and

reported variation in number of female flowers, pollen shedding, stigmatic

receptivity and intraspadix overlapping of inflorescence.

Balakrishnan et al. (1991) reported significant variation among the

16 coconut hybrid combinations for number of characters with high

estimates of phenotypic and genotypic variance for nuts per palm. The

number of leaves per palm was recorded with low phenotypic and

genotypic variance.

Ramanathan et al. (1992) reported significant differences for

reproductive and nut characters while comparing the performance of

coconut cultivars and hybrids. Among cultivars Andaman ordinary

recorded higher yield per palm per year while among the hybrids, West

11
Coast Tall x Malayan Dwarf Yellow and the reciprocal cross performed

better.

Luke Rathna Kumar et al. (1993) studied 18 growth, reproductive

and yield traits and reported that number of leaves per crown, petiole

length, leaflets per leaf, kernel weight, kernel thickness and number of

nuts per palms were the traits for selection of superior palms for high

yield.

2.2.2. Variation in Yield and yield components

2.2.2.1. Nut yield

Yield of nuts/palm/year is a very important character and, thus it

attracted the attention of many workers. The yield and its components are

influenced by climate of the region where the crop is growing (Ong et al,

1985; Santos et al, 1986).

Annual nut production per palm showed variation among the

coconut cultivars and hybrids (Kasturi Bai, 1993). Narayanan Kutty and

Gopalakrishnan (1991) found a correlation between chlorophyll content in

the leaves, the rate of apparent photosynthesis and annual yield of nuts.

The performance of coconut varieties may differ under different

ecological conditions. In a varietal trial, conducted in the Philippines at

three different locations with different climates and soils, Santos et al

(1986) found differences in copra production per palm at each location for

12
most of the varieties. There are indications that the yield potential of the

palms could be gauged from their initial yields and also that the height of

the palm and the number of functioning leaves on the crown are

significantly correlated with yield (Satyabalan et al., 1972). Jacob Mathew

et al. (1991) proposed prediction equations to estimate the annual yield of

coconuts based on one time total count of nuts present in the crown at

different stages in palms of local tall cultivars.

2.2.22. Nut components

The coconut palm is a large seeded monocotyledonous tree crop.

The fruit is a fibrous drupe developed from a tricarpellate ovary. Nut

development from the opening of spathe to fruit maturity requires about

one year of time. It has a thick pericarp and fibrous mesocarp together

constituting the husk. Inside the husk is the hard endocarp (shell) lined by

solid endosperm. The endosperm of fruits when dried to copra is hard and

rigid. Nut composition significantly varies between cultivars and hybrids,

which reflects in the variations in partitioning of nut diy matter towards its

components viz., husk (43-58%), shell (20- 27%) and copra (27-34%)

(Kasturi Bai et al.,1996). Importance of increased partitioning of total dry

matter towards copra at the expense of other nut components for yield

improvement has been emphasized by many workers (Green and Foale,

1961; Corley, 1983). For obtaining maximum number of hybrid seedlings

13
(Dwarf x Tall), Satyabalan and Rajagopal (1987) suggested to use

pistillate parents which yield nuts having low shell content and high copra

content.

Growing conditions have an influence on fruit development and

proportion of different components in fruit. Studies in Nigeria showed that

the number of dry days and sunshine hours are the factors responsible for

the variation in percentage of copra to nut and oil to copra (Akpan and

Obisesan, 1984). The copra content of fruits is maximum during summer

followed by winter and minimal in the post-monsoon period of October-

November (Nambiar and Rao, 1991). It was observed that the second

phase of nut development was influenced by surface air temperature

(Prasada Rao, 1991). Satyabalan (1997) studied the impact of weather

variables on different components of coconut fruit. Failure in rainfall not

only affects the nut production but the amount of copra per nut also gets

reduced (Murray, 1977). Significant correlation was observed between (1)

weight of fruit (unhusked) and weight of husked nut, (2) weight of fruit

and that of the husk, (3) weight of husked nut and fresh weight of kernel,

(4) weight of husked nut and weight of shell, (5) weight of husked nut and

weight of copra, (6) fresh weight of kernel and weight of copra and (7)

weight of shell and weight of copra in the nuts harvested during different

months of the year (Satyabalan and Mathew, 1984). Some slight

14
differences in the indices studied were attributed to seasonal effect during

fruit development.

Nut composition is influenced by soil type also. Partitioning

towards copra was higher in COD x WCT grown on laterite soils than on

sandy loam soils (Anonymous, 1988). Differential response of WCT and

WCT x COD palms to available soil moisture regimes was observed in

terms of water relation components and nut yield (Rajagopal et al, 1986).

2.2.3. Heritability studies

Heritability estimates represent the degree of inheritance of a

character from parent to progeny. Heritability estimates of different

characters would give information about the contribution by these

characters towards yield. Characters having high heritability could be

improved through selection since they are less affected by environment

(Lush, 1940).

High heritability along with high genetic gain could be considered

to determine the amount of heritable variation with accuracy (Johnson et

al., 1955)

Works done by earlier workers on heritability and genetic advance

for different traits in coconut are presented below

15
2.2.3.1. Yield and yield components

Studies conducted by Meunier et al. (1984) on yield attributing

characters in T x D and D x T hybrids revealed high heritability estimates

for copra yield per nut, oil yield per nut and number of nuts per palm. The

heritability estimates did not vary between Dwarf and Tall types. Muluk

(1987) reported high heritability estimates for plant height and bunch

number.

Nut production pattern as the palm advances in age seems to have

a role of maternal inheritance because, Bourdeix et al. (1992) observed

reciprocal differences based on eight hybrids involving Tall and Dwarf

cultivars. Whenever a Dwarf cultivar was used as mother parent the

progenies had initial high yields (-8 years). Nambiar et al. (1970a) found

the heritability of nut yield as 0.30 based on progenies. Meunier et al,

(1984) found the character to be of additive nature. But high yield

obtained in hybrids indicated heterosis and thereby the dominance of the

trait.

Nampoothiri et al. (1999) found the cumulative nut yield of

coconut to be controlled by both additive and non-additive genes. Based

on the diallel experiment they also indicated role of over dominance and

environmental influence on this trait. Seasonal variation in nut yield and

sex expression has already been reported by Nambiar et al. (1970b).

Heritability values of number of bunch were found to be very low (<0,23)

16
by earlier workers (Nambiar et al. 1970a, Meunier et al. 1984). There is

always negative correlation between number of nuts and copra/nut. Thus,

it is better to select varieties/hybrids with high copra yield/unit area than

high nut yield/palm.

2.2.3.2. Nut components

Coconut palm is a large seeded monocotyledonous tree. The fruit

is a fibrous drupe developed from a tricarpellary ovary. Nut development

from opening of spathe to fruit maturity requires about one year time. It

has a thick pericarp and fibrous mesocarp together constituting the husk.

Inside husk is the hard endocarp (shell) lined by solid endosperm. The

endosperm of fruits when dried to copra is hard and rigid. Nut

composition significantly varied between the cultivars and hybrids, which

reflected in variations in partitioning of nut dry matter towards its

components viz., husk (43-58%) shell (20- 27%) and copra (27-34%)

(Kasturi Bai et al, 1996). Importance of increased partitioning of total dry

matter towards copra at the expense of other nut components for yield

improvement was emphasized (Green and Foale, 1961; Corley, 1983). For

obtaining maximum number of hybrid seedlings (Dwarf x Tall),

Satyabalan and Rajagopal (1987) suggested to use pistillate parents, which

yield nuts having low shell content and high copra content.

17
 Growing conditions have an influence on fruit development and

proportion of different components in fruit. Studies in Nigeria showed that

the number of dry days and sunshine hours are the factors responsible for

the variation in percentage of copra to nut and oil to copra (Akpan and

Obisesan, 1984). The copra content of fruit is the maximum during

summer followed by winter and minimal in the post-monsoon period of

October-November (Nambiar and Rao, 1991). It was observed that the

second phase of nut development was influenced by surface air

temperature (Prasada Rao, 1991). Satyabalan (1997) studied the impact of

weather variables on different components of coconut fruit. Failure in

rainfall not only affects the nut production but the amount of copra per nut

also gets reduced (Murray, 1977). Significant correlation was observed

between (1) weight of fruit (unhusked) and weight of husked nut, (2)

weight of fruit and that of the husk, (3) weight of husked nut and fresh

weight of kernel, (4) weight of husked nut and weight of shell, (5) weight

of husked nut and weight of copra, (6) fresh weight of kernel and weight

of copra and (7) weight of shell and weight of copra in the nuts harvested

during different months of the year (Satyabalan and Mathew, 1984). Some

slight differences in the indices studied were attributed to seasonal effect

during fruit development.

18
 Nut composition is influenced by soil type also. Partitioning

towards copra was higher in COD x WCT grown on laterite soils than on

sandy loam soils (Anonymous, 1988). Differential response of WCT and

WCT x COD palms to available soil moisture regimes was observed in

terms of water relation components and nut yield (Rajagopal et al, 1986).

2.23.3. Oil content and Fatty acid profile

Coconut oil is the most important product of the coconut palm. The

oil content in coconut copra and oil yield per hectare varies among the

cultivars (Louis and Ramachandran, 1981; Naresh Kumar et al, 2000b).

Analysis of oil content of five cultivars and their hybrids indicated

maximum and minimum oil content in the cultivars Laccadive Ordinary

and Chowghat Orange Dwarf respectively (Nambiar and Rao, 1991). They

also observed variation in oil content with the seasons.

Oo and Stumpf (1979) studied the fatty acid biosynthesis in the

developing endosperm of coconut. The fatty acid and tri acyl glyceryl

(TAG) composition of oils of nine coconut hybrids revealed that the lauric

acid content among hybrids varied from 47.3 to 50.5 % (Rodriguez et al,,

1998). Coconut oil mainly contains saturated fatty acids like caproic (6C),

caprylic (8C), capric (10C), lauric (12C), myristic (14C), palmitic (16C),

stearic (18C) and arachadic (20C). It also contains imsaturated fatty acids

such as oleic (18 C: 1), palmitoleic (16 C:2), and linoleic (18 C:3) in lesser

19
quantities (Oo and Stumpf, 1979; Padua Resurreccion and Banzon, 1979;

Naresh Kumar et al, 2000b). Among these fatty acids, lauric acid is

present in maximum concentration. Variability among the coconut

cultivars for fatty acid composition and most suitable cultivars for

different uses has been reported (Naresh Kumar et al, 2000b; Naresh

Kumar and Rajagopal, 2000). The long chain fatty acids C18:3 and C22:0

have been found at all stages of fruit growth in nut water (Jayalekshmy et

al., 1988) but not in the kernel (Padua Resurreccion and Banzon, 1979;

Balachandran, et al., 1985). Traces of linolenic acid have also been

reported to be present in kernel (Udayasekhara Rao and Srinivasa Rao,

1981). Variations exist in the proportions and properties of fats at different

locations of the developing coconut kernel (Kartha, 1963; Lim and

Banzon, 1983). Coconut oil contains both neutral lipids and polar lipids

(Krishnamurthy and Chandrasekhara, 1983). Lipid fractionation by silicic

acid column chromatography indicated that neutral lipids formed major

fraction (about 94%) followed by glycolipids (3.5%) and phospho lipids

(2.5%) (Naresh Kumar et al, 2000b).

2.2.3.4. Heterosis for oil content

Oil content of hybrids was always the intermediate of parents.

Nallathambi et al (1986) observed heterosis and heterobeltiosis to the

tune of 12-13% for oil content in hybrids between East Coast Tall (ECT)

20
and Gangabondam or Kanyakumari Yellow Dwarf based on seven parents

and six direct and reciprocal crosses. The differences in opinions are due

to the differences in genetic make up of populations studied. Seasonal

variation was also noticed for the trait i.e. it was maximum in the rainy

season and minimum in summer (Nambiar and Rao, 1991).

Heterosis is also modified by the interactions between genotype

and environment in cultivation. Hybrid sorghum can show heterosis for

yield; but this effect varies widely between trials conducted at sites

differing in seasonal water supply (Chapman et al., 2000), so that it is

more meaningful to characterize a particular hybrid line as showing

heterosis for yield at a specific locality or under certain environmental

conditions

Heterosis for size, vigour or yield is most evident in out breeding

crops. The classic case is maize; in which heterosis has long been

exploited in the production of uniformly high-yielding Fl seed in

commercial quantities.

2.2.4. Inheritance of fruit component traits

Asmono et al. (1993) while working on heritability (broad sense)

values of fruit characters based on five tall and two dwarf populations

found high heritability (>0.80) for fruit length in all the coconut

21
populations studied. Broad base heritability values for husked nut weight

was found to be as high as 0.95 by Liyanage and Sakai (1960). Additive

effects were known to govern the volume of husked nut, weight of shell

and weight of kernel (Patil et al., 1993).

Renuga (1999) studied heritability and genetic advance for a

number of vegetative, reproductive and nut component characters and

reported high heritability coupled with high genetic advance weight of

dehusked nut, whole nut weight, length of petiole, number of nuts per

bunch, number of female flowers and number of nuts per bunch, number

of female flowers and number of internodes for one meter. It was

suggested that high heritability with genetic advance indicated additive

gene action and hence these characters could be used in selection

programmes. The characters stem girth, number of inflorescence per palm

and thickness of shell showed low heritability with low genetic advance.

2.3, Association of traits

2.3.1. Correlation studies

Several workers have worked out correlation co-efficients among,

morphological, reproductive and nut component characters of coconut.

Patel (1937) and Menon and Pandalai (1958) observed that number

of leaves on crown and girth of trunk were positively correlated with

number of nuts per year in Tall.

 Thampan reported a significant positive correlation between

number of female flowers produced and number of nuts per palm.

Satyabalan (1972) observed a positive correlation between nut yield per

palm and stem height of adult palm.

Nampoothiri et al (1975) showed that the number of spathes

produced and number of female flowers produced exhibited a positive

correlation with pre-flowering period. They also observed that the pre­

flowering period had a negative association with number of nuts produced

per year.

Abeywardena (1976) reported positive association between

number of spathes produced and number of nuts per palm. He also

reported a positive correlation between trunk girth and number of leaves

per palm. Balingasa and Carpio (1983) showed that the number of female

flowers produced and number of nuts produced were positively correlated.

Iyer (1980) concluded that the increase in plant height with

simultaneous increase in number of leaves would lead to increased nut

production. Sukumaran et al. (1981) observed that number of leaflets and

length of leaf was positively associated with number of nuts per year.

They also reported a positive correlation between number of bunches per

year and number of nuts per year. The number of nuts per year and outturn

of copra had significant positive association.

23
 Louis (1983) showed that the number of leaflets and number of

nuts per year were positively correlated. A negative association between

the number of female flowers produced and setting per cent was observed.

Ramanathan (1984) reported a positive correlation of nut yield per

palm with stem height of the adult palm. Prabhakaran et al, (1991)

reported that number of leaves was the major contributor, which correlated

with yield.

Kalathiya and Sen (1993) studied the correlation among floral and

yield characteristics. Nut yield was found to be significantly and positively

correlated with number of spadices and length of female phase where as

number of leaves produced and length of spadix exhibited significant

positive correlation with number of spadices. They suggested that the

number of spadices and length of female phase should be considered as

selection criteria for nut yield improvement in the variety Dwarf Green.

Narayanan Kutty and Gopalakrishnan (1993) reported association

among biometric, mineral and organic constituents with yield of coconuts.

They observed significant positive correlation for number of leaves and

length of leaves with yield. Among the mineral nutrients, nitrogen and

potassium content in leaves had significant positive correlation with yield.

Sindhumole and Ibrahim (2001) studied the relationship among

various vegetative, reproductive and economic characters and reported

lower phenotypic correlation than genotypic correlation co-efficients. The

24
genotypic correlation was mostly negative where vegetative characters

were involved, but positive for other pairs. Yield was positively correlated

with both vegetative and reproductive characters.

Bavappa and Sukumaran (1976), Balakrishnan and Vijayakumar

(1988), Gopimony (1988) and Patil et al. (1993) have reported association

among nut component characters.

Renuga (1999) worked out genotypic and phenotypic correlation

co- efficients and found that genotypic correlation co-efficients were

higher than phenotypic correlation co-efficients. Number of nuts per

bunch, number of bunches per palm and setting percentage had positive

and significant genotypic and phenotypic correlation with yield.

2.4. Path coefficient analysis

Study on path co- efficient analysis, developed by Wright (1921),

provides an effective means of partitioning the total correlation into direct

and indirect effects of various characters on dependentant variable. The

available works on path coefficient analysis in coconut is reviewed here.

Sukumaran et al. (1981) made the path analysis in coconut for

annual and cumulative nut yield. Average number of female flowers at

21-24 years, number of functional leaves, inter-nodal length, total leaf

production up to

First three years of sowing and time taken for flowering had direct:

and largest influence on cumulative and average yield. Bhagavan and Nair

25
(1989) reported that plant height, internodal distance, stem girth and leaf

length had high direct effects on yield.

Louis and Chopra (1993) studied the cause and effect relationship

of different characters on copra weight. They found that kernel weight,

length of petiole and thickness of shell had a positive direct effect on

copra weight, while thickness of shell had a positive direct effect on copra

weight, while thickness of kernel and pre-flowering period showed a

negative direct effect on copra weight.

Luke Rathna Kumar et al (1993) recorded high direct effects on

weight of copra percentage through the characters kernel weight, thickness

of kernel and number of nuts per palm.

Pam in and Asmono (1993) reported positive direct effects of nuts

per palm and nut weight on yield. They indicated that oil yield per palm

could be increased by direct selection for these traits or by simultaneous

progeny selection for fruit number per tree or oil content and number of

nuts per tree.

Ataga (1995) concluded that the number of bunches per palm and

bunch weight had high direct effect on yield.

Renuga (1999) reported that the characters viz., length of leaf,

thickness of husk, Weight of kernel, setting per cent, length of petiole and

plant height had maximum positive direct effects on yield. Weight of

dehusked nut, number of leaflets (left), number of female flowers, whole

26
nut weight and number of internodes for one metre had high and negative

direct effects on nut yield per palm.

2.5. Stability analysis

2.5.1. Genotypic x Environmental interactions

Stability in performance is an important property of a genotype,

which is desired for wide adaptation for cultivation. Johanson (1909)

explained the meaning of phenotype, which is related to the appearance or

form arising as a result of interaction of genotype with the environment in

which it is grown. He was the pioneer in considering the importance of

environment in the expression of a phenotype. Fisher and Mackenie

(1923) first reported the existence of genotype x environment interaction

from the results of varietal trials in potatoes.

The absolute phenotypic stability would be expressed by b = 0 and

the ideal variety in respect of adoption would the one having maximum

yield potential on the most favourable environment and maximum

phenotypic stability. According to Eberhart and Russel (1966), mean,

regression and deviation from regression provides an account of G x E

interactions.

Louis and Chandrasekaran (1976) reported that the growth and

yield attributing characters of coconut were highly influenced by

environment. Balakrishnan et al. (1998) analysed stability for 32 coconut

cultivars on yield. The results showed that WCT, Laccadive Ordinary,

27
Philippines Ordinary and Andaman Ordinary were stable for annual

production of nuts at Pilicode, Kerala.

Patil et al. (1991) evaluated nine coconut genotypes for nature and

degree of genotype x environmental interaction. They observed a

significant linear genotype x environmental interaction for all the traits.

Among the genotypes studied, Pratap was considered the best for

cultivation while, T x D and Banawali Yellow Round were found to adopt

better under fluctuating environmental condition.

Vijayaragahvan et al. (1993) showed the occurrence of seasonal

variation in the production of spadix in coconut hybrids and parents. The

results revealed that the hybrid combination MYD x ECT and its

reciprocal was found to be the best, producing highest number of spathes.

They also reported that Tall types produced maximum number of spathes

during March and August, while Dwarf types had single peak during

August.

Vijayalakshimi et al. (1962) observed the development of coconut

during different months of the year and expressed that the effect of season

was not operating the same extent to the different characters.

In Sri Lanka, Abeywardena and Fernando (1963) found that the

coconut filed consistently showed a symmetrical curve with a peak in

March - April. In Philippines (Menon 1964), peak spadix production was

from June - August.

28
 In Zanzibar, Tremlett (1964) reported that the number of

inflorescences and female flowers expressed higher number in August to

October and low in April.

Fremond et al. (1968) observed that in Ivory Coast, the percentage

of nuts within the annual total yield was more during May to October than

during November to April, while in Dahomey higher yield was estimated

during February to July.

Wickramasurya (1968) observed that the coconut nuts harvested

from November to February were lower in number and that the available

water, temperature and sunshine had influence on seasonal fluctuations in

coconut yield. Their action also influences the development of

inflorescence and fruit.

In Tamil Nadu, Louis and Annappan (1980) observed significant

variation in yield during the four different season’s of the year with higher

yield during the summer months (April- May) and lower yield during

South West monsoon period (May -June).

Jacob Mathew (1982) found seasonal variation in coconut

production in Kerala with about 62 per cent of the nuts harvested during

first six months of the year. Mandal and Mehta (1988) found that summer

rains have positive correlation with the yield in succeeding year and about

60 per cent of the variations in annual yield were due to changes in the

duration of dry spell in the proceeding two years.

29
 Renuga (1999) used the trait nut yield per palm over season for

classification of stable genotypes. The genotypes with high mean values

were resolved into four groups based on the stability parameters like

regression and deviation from regression. Those genotypes falling under

group I namely Laccadive Ordinary, East Coast Tall and West Coast Tall

were found to be stable for nut per yield palm.

2.6. Molecular marker studies

Genetic diversity study of 17 distinct South Pacific populations

was carried out by means of RAPD technique using 14 primers by

Ashburner et al. (1997b), The study revealed that over 60 per cent of the

observed variability occurred within populations. Genetic drift and a

possible bottleneck in the past of the species were suggested as reasons for

the low intra-population diversity. The low genetic diversity detected

indicates that coconut behaves as a self-fertilized species in the South

Pacific, despite having a mixed system of cross-pollination.

Rodriguez et al. (1997) analyzed 10 coconut populations (five tails

and five dwarfs) from the Philippines using RAPD technique using eight

polymorphic primers. Wadt et al. (1999) used RAPD technique to

determine genetic relationships between Tall coconut populations from a

germplasm bank in Brazil.

30
 A preliminary study was carried out by Cardena et al. (2003) to

identify RAPDs associated with resistance to lethal yellowing disease of

the coconut palm.

Upadhyay et al. (2004) used RAPD technique to analyze the

genetic diversity among 15 Indian and five exotic accessions with eight

polymorphic primers. In general, tall accessions were more heterozygous

as they had higher proportions of polymorphic bands and genetic

diversity. Similarly exotic accessions exhibited more variation. Dwarfs

from geographically distant regions did not cluster separately.

Perera et al. (1998) conducted AFLP analysis of 42 genotypes

indigenous to Sri Lanka using eight primer pairs. Overall, more variation

was detected in tall forms (typica), rather than intermediate (aurantica)

and dwarf (nana) forms. Teulat et al. (2000) used AFLP markers in

combination with SSR markers to analyze genetic diversity of 14

populations.

Molecular markers are among the promising and reliable tools to

describe genetic diversity since they are free from environmental effects

and the consequent interaction with the genotype, resulting to a large

amount of morphological variation polymorphism (Ford-Lloyd and

Painting 1996). One of these techniques is the simple sequence repeats

(SSRs) otherwise called microsatellites (Cregan et al. 1994). Devos et al.

1995; Groppe et al. 1995; Xiao et al. 1996)

31
 Rivera et al. (1999) characterized 40 coconut samples from the

Philippines using eight SSR primer pairs. Dwarfs grouped separately from

tails and showed less genetic diversity.

Using a pre-cloning enrichment procedure, Perera et al. (1999)

isolated eight coconut microsatellites. These eight microsatellites were

used to study the levels and patterns of genetic diversity of Sri Lankan

coconut populations. The results showed that the Sri Lankan tall coconuts

exhibit higher levels of diversity than the dwarfs and intermediates, and

the intermediate coconuts are more similar to dwarfs than tails. This was

in agreement with the results obtained using AFLPs in the same set of

genotypes in an earlier study (Perera et al., 1998).

Perera et al. (2000) used eight pairs of SSR primers to analyze the

genetic diversity in 130 individuals of coconut comprising 75 tall and 55

dwarf individuals representing 94 ecotypes from different geographical

regions. A phenetic tree based on genetic distance clustered individuals

into five groups, each mainly composed of either tails or dwarfs. Thirty-

three tall coconut populations from Sri Lanka were subjected to

microsatellite assay with eight SSR primer pairs in order to study the

levels and distribution of genetic variation for formulating future

collection strategies and selecting parents for breeding programmes

(Perera et al, 2001). A high level of genetic diversity was detected in all

the populations.

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 Merrow et al. (2003) used 15 simple sequence repeat (SSR)

microsatellite DNA loci to analyze genetic variation within Cocos

nucifera germplasm collections at two locations in South Florida,

representing eight cultivars.

Thirty-three coconut accessions from a world-wide coconut

collection at the International Gene Bank in India were analyzed using

ISSR markers with 19 primers (Manimekalai and Nagarajan, 2006).

Coconut accessions from Southeast Asia, South Asia and South Pacific

formed separate groups and this grouping was generally in accordance

with their origin and pattern of dispersal of coconuts from the centre of

origin.

2.7. Mitochondrial studies

Mitochondrial activity and its superiority with hybrid vigour was

studied by Me Daniel in 1973. Oxidizing and phosphorylating activity of

mitochondria was related with hybrid vigour in maize and soybean by

Kridel (3972) and Hanson 1975).

The oxidation of various substrates like succinate, malate and

glycine was studied in isolated mitochondria from spinach leaves (Douce,

1977).

Purification of mitochondria was done through discontinuous

percoll density gradient and the oxidative properties of washed and

gradient mitochondria were compared (Christopher Jackson, 1978).

33
 Joseph (1982) studied malate oxidation through Malate

Dehydrogenase and NAD linked malic enzyme.

Malate oxidation by plant mitochondria occurs via action of two

enzymes, MDH and NAD located in the matrix. The presence and absence

of one or the other was depends on the condition imposed based on co­

factors, pH etc. (Wiskich, 1980).

Malatedehydrogenase and the electron transport mitochondria for

the addition of Malate and NADH, different pools of ubiquinon act as

branch point between various dehydrogenases via mitochondrial oxidation

in cauliflower (Pierre, 1980).

The interaction of TCA cycle and respiratory chain with various

substrates with pea leaf and potato mitochondria was studied ((Day et

al., 1985).

Mitochondrial heterosis was worked out and it was related with the

bunch production and other morphological characters in oil palm by

Koume et al., (1978).

Based on the above information’s available on hybrids of coconut,

an attempt has been made presently to study the performance of some

coconut hybrids and also to develop certain techniques to identify hybrids

and confirm their hybrid status.

34