Observations on Sex Differentiation in the Oil Palm,

Elaeis guineensis L.
BY

C. N. WILLIAMS AND R. L. THOMAS'

University of Malaya, Kuala Lumpur, Malaysia

ABSTRACT
The morphology and frequency of occurrence of abnormal hermaphrodite in-
florescences in the oil palm is described for palms of varying age. Hermaphrodite
inflorescences are manifest during changes of the sex cycle in the normally monoe-
cious species, and gross morphological differences in the type of hermaphrodite
inflorescences occurred depending on whether the change of cycle was from male
to female or vice versa. The significance of the occurrence and morphology of the
abnormal inflorescences is discussed in relation to the accumulation of photo-
synthetic reserves and to the maturation process.

INTRODUCTION

T HE oil palm (Elaeisguineensis L.) is a predominately monoecious species,

producing generally either male or female inflorescences in alternate
cycles each of several months' duration. The frequency and duration of par-
ticular sex cycles appears to be strongly associated with environmental factors;
good growing conditions (adequate moisture and high sunshine, good soil,
etc.) tending to promote and increase the duration of female inflorescence
cycles. Production of male inflorescence cycles is associated with unfavour-
able seasonal periods such as cloudy weather or moisture stress (e.g. Broek-
mans, 1957; Hemptinne and Ferwerda, 1961; Sparnaaij, Rees, and Chappas,
1964). The sex of inflorescences appears to be determined in the primordial
stages, some 18 months to 2 years before emergence of the inflorescence from
the axil of its subtending frond (Broekmans, 1957). Thus it is the environ-
mental conditions some considerable time before the inflorescence emerges,
that presumably affects sex differentiation. Work with applied auxins, how-
ever, which may promote the development of abnormal inflorescence types,
as discussed later, indicates that secondary changes in sex can occur at a much
later stage.
At the time of change from cycles of one sex to another, inflorescences of
mixed sex may be produced for a varying period of time, particularly when
the palms are young. The present paper gives a description of the type and
frequency of occurrence of such hermaphrodite inflorescences, and discusses
the possible role of photosynthetic reserves and auxins in sex differentiation.
1
Permanent address: Michigan State University, Michigan.
Ann. Bot. 34, 937-63, 1970.]

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 958 Williams and Thomas—Observations on
OBSERVATIONS
Altogether the changes of sex cycle from male to female and vice versa, were
observed in seven plantings of oil palm in Western Malaya, ranging from 1 to
7 years in age (calculated from the time of field planting of approximately
i-year-old seedlings). In each age-group, 25 palms were observed which were
undergoing a change from male to female and 25 undergoing a change from
female to male. In each case the frequency of hermaphrodite inflorescences
TABLE I
Scores of inflorescence types from oil palm of different ages during
changes of sex cycle from male to female and vice versa
Twenty-five palms were observed in each of the 14 sub-categories
Mean
No. of
Mixed inflores- Mixed inflores- herma-
cence with cence with Male spike- Male spike- Total phrodite
Age of predominantly predominantly lets with lets with No. of inflores-
palms Cycle male spike- female spike- few female many female hermaphrodite cences
(years) change lets lets elements elements inflorescences per palm
i a. 6-9 6 27 o o 46 O.02
b. 9-6 o o 13 o
2 a. 6-9 o 33 o o 52 1*04'
b. 9-6 o 0 19 0
3 a. 6-9 o 43 o o 86 1.72
b. 9-<J o o 43 o
4 a. 6-9 2 34 o o 5o 1.00
b. 2-6 o 0 14 o
5 a. 6-9 2 24 o o 1.02
b. 9-6 O 24 z 5-
o
6 a. 6-9 o II o o 0.46
b. 9-6 o o 12 o
7 a. 6-9 o I O o
s O.IO
b. 9-6 o o 4 o

was recorded and the inflorescences were classified on the basis of certain
gross morphological differences (individual florets were not generally ob-
served). Hermaphrodite inflorescences showed differences in morphology,
depending on whether the change of cycle was from male to female or vice
versa. During a change from a male to a female cycle, inflorescences of a type
which bore both male and female spikelets were produced, generally with
more female spikelets than male, but sometimes with a higher proportion
of male spikelets. Examples of these types are shown in Figs. 1 and 2. (All
inflorescence types illustrated in the figures are from the 1-year-old group of
palms which were smaller and therefore easier to represent.) The inflorescence
of Fig. 1 is predominantly male while that of Fig. 2 possesses a larger number
of female spikelets surmounted by two male spikelets. Occasionally, in-
florescences which bore spikelets with both male and female elements were
produced during a change from male to female, as illustrated in Fig. 3. The
two predominantly male spikelets here have developed abortive female flowers
towards the base.
When the change of sex was from female to male, a different type of in-
florescence was formed in which predominantly male spikelets were produced,

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 Sex Differentiation in the Oil Palm 959

FIG. I FIG. 2
FIGS, I and 2. Inflorescences of a type produced during a change from a male to a female cycle.
More often a greater number of female spikelets are produced as shown in Fig. 2.

but with single female flowers or small truncated female inflorescences deve-
loped on them. Examples are shown in Figs. 4 and 5. The existence of this
type (predominantly male fingers) indicates that the male spikelet is not the
ultimate branch of the inflorescence since further flowers and even inflores-
cences can develop from it. The morphology of the inflorescence suggests
that the whole may be considered as a determinate system of branches, gener-
ally bearing sub-branches with maleflowers,but which can proliferate further.
Generally, there were a fewer number of female spikelets or female flowers than

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 960 Williams and Thomas—Observations on

FIG. 3. An inflorescence produced during a change from a male to a female

cycle. The male spikelets have abortive female flowers towards the base. This
type of inflorescence is somewhat unusual.

male, during a change from female to male. The results of all the observations
are shown in Table 1 where the presence or absence of four distinct groups of
hermaphrodite inflorescences is recorded.
DISCUSSION
One obvious feature of these results is that the frequency of occurrence of
all categories of abnormal inflorescences tends to decrease progressively with
age, as is seen from the total column of the table. It is interesting to speculate
on the morphological differences between the types formed during a change
from male to female and vice versa, and on the general reduction of frequency
of all abnormal types, in the light of the possibility that a change from male to
female in the oil palm is associated with a depletion of reserves and that
a change in the opposite direction with increasing reserves. Generally this

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 Sex Differentiation in the Oil Palm 961

Stigma
Br.act

Flower
FIG. 4 FIG. 5
FIGS. 4 and 5. Inflorescences of a type produced during a change from a female to a male cycle.
More often only a few female elements are developed on the male fingers as shown in Fig. 4.

assumption is made on the basis of various types of indirect evidence, and

reserves here are thought of as predominantly carbohydrates (although other
substances may also be involved). The evidence for the 'reserve' theory is as
follows:
1. Growing sites which are cloudy produce a higher number of male
flowers. For example, Hemptinne and Ferwerda (1961) compared yields of
palms (which is directly related to the sex of the inflorescences) from three
climatic regions of the Congo, and found a deleterious influence of excess
rain which they attributed to the accompanying low sunlight values.
2. Regions with a dry season produce variations in sex of inflorescences
which appear to be correlated with rainfall variations affecting photosynthesis
(Sparnaaij et ah, 1964). Rees (1958) has shown that such periods of mois-
ture stress affect stomatal functioning and therefore would lead to a lowering
of photosynthetic activity and a lower rate of accumulation of reserves.

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 962 Williams and Thomas—Observations on
3. Severe pruning of palm fronds (which reduces photosynthesis of the
palm) often results in a subsequent decrease in sex ratio (Sparnaaij et al.,
1964).
The occurrence of the two predominate types of hermaphrodite inflores-
cence ; namely that with numerous male fingers with a few female elements
developed on them (Fig. 4), and that in which a large number of separate
female spikelets is surmounted by a few male fingers (Fig. 2), also suggest that
availability of photosynthetic reserves may be involved in sex differentiation;
particularly when considered in conjunction with the general decline in fre-
quency of all abnormal types with increasing maturity. One of the most signi-
ficant changes in this regard is the massive increase in the storage regions of the
palm which takes place with trunk growth. (The trunk is the main storage
area of the palm.) Consequently, older palms process a larger absolute reserve
strength and a larger buffer capacity for reserves. Small palms, on the other
hand, could be quickly depleted of reserves and depend on day-to-day fluctua-
tions in photosynthetic level for growth and development of the inflorescences.
Predominantly male inflorescences occur when a change from female to
male is taking place (Fig. 4), and if it is assumed that reserves are low and
tending to decrease, it is significant that only small proliferations of female
elements are usually formed at this time (Fig. 4). In the young palm, which
is more dependent on the rate of assimilation throughout the period of develop-
ment of the inflorescence, minor fluctuations or increases in assimilate level
could give rise to these secondary proliferations. In the mature palm, however,
with significant reserve centres, these could themselves compete with the
inflorescence for assimilates during the commencement male cycle, thus
reducing the tendency for reversion to the female type and proliferation of
female elements. Furthermore, it is likely that these proliferations of female
elements can occur fairly late in development of the male inflorescence since
the application of auxins (see below) can produce inflorescences of this type
(Fig. 4) as close as a few months before anthesis or inflorescence maturity.
When a change from male to female is taking place, inflorescences with
predominantly female spikelets are formed (Fig. 2). These do not occur as
a result of auxin application at a late stage and it is likely that they are formed
some considerable time before emergence, as indicated by Broekmans's studies
referred to earlier on. In young palms, however, hermaphrodite inflorescences
are more frequent and here again the availability of photosynthates could affect
differentiation if it is assumed that female inflorescence development occurs as
a result of the accumulation of high levels of photosynthate, or a prolonged
favourable period for growth. The relative contributions of reserve photo-
synthate, which should supply a steady outflow, and daily assimilation, which
would be more variable, particularly at the beginning of a favourable climatic
period, would probably not be the same in young and mature palms, the latter
being more evident in the young palms with a lower reserve capacity. The
fact that male elements do not proliferate on the female spikelets (as in the
reverse situation) is probably related to the determinate nature of the female

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 Sex Differentiation in the Oil Palm 963
spikelet and to the general small size of the inflorescence at the primordial
stage.
The role of auxins in the creation of sinks in growing organs has been dis-
cussed by many workers and it is generally assumed that these are active in
sinks and may even be of central importance in affecting the flow of assimi-
lates to growing organs. If auxins cause expansion of tissue, for example, and
a dilution of cell contents, then the movement of assimilates from regions of
high concentration (the photosynthetic apparatus) to the growing organs, can
take place in accordance with the principle of Molish. In experiments with
auxins (2,4,D and 2,4,5—TP) applied at concentrations of 500 ppm, mg/
palm at 10-day intervals, it was found that these tended to increase the fre-
quency of abnormal inflorescences, particularly of the types illustrated in Figs.
4 and 5. These appear 2-3 months after application. Consequently, variations
in natural auxin level could also account for the occurrence of abnormal types,
and in this regard certain abnormal palm genotypes consistently produce
abnormal inflorescences, suggesting some hormonal imbalance. It may very
well be that the occurrence of female elements is associated with higher auxin
activity, and that fluctuations in auxin level or activity occur more frequently
in young palms; however, the question of which comes first, the sink or
the substrate, is one which has yet to be resolved in plants in general. The
present observations tend to suggest that the storage buffer capacity of the
palm (which changes with age), may be more central in the determination
of sex in the inflorescence under natural conditions, and that application of
auxins causes an abnormally high sink activity which may result in the pro-
duction of female elements in palms otherwise disposed to male inflorescence
production, at the expense of other growth processes.

ACKNOWLEDGEMENTS
We are indebted to Mr. Khew Yui Fooi for making the drawings and to
Mr. K. W. Chan for help in the collection of data.

LITERATURE CITED
BROEKMANS, A. M. F., 1957. Growth, flowering and yield of the oil palm in Nigeria. J. W. Afr.
Inst.for Oil Palm Res. 2, 187-220.
HEMPTINNE, J., and FERVVERDA, J. D., 1961. Influence des precipitations sur les production du
palmier a huile. Oleagineux, Lond. 16, 431.
REES, A. R., J958. Field observations on midday closure of stomata in oil palm. Nature, Lond.
182, 735.
SPARNAAIJ, L. D., REES, A. R , and CHAPPAS, L. C , 1964. Annual yield variation in the oil
palm. J. Nigerian Inst.for Oil Palm Res. 4, 111-25.

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