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Architecture and development of the oil-palm (Elaeis guineensis Jacq.) root

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Plant and Soil 189: 33–48, 1997. 33

c 1997 Kluwer Academic Publishers. Printed in the Netherlands.

Architecture and development of the oil-palm (Elaeis guineensis Jacq.) root

system

Christophe Jourdan and Hervé Rey1

Département des Cultures Pérennes et Unité de Modélisation des Plantes, Centre de Coopération Internationale
en Recherche Agronomique pour le Développement (CIRAD), B.P. 5035, F-34032 Montpellier Cedex 1, France.
1
Corresponding author

Key words: development, Elaeis guineensis, growth, oil palm, root architecture, root system

Abstract

The growth dynamics and architecture of the oil-palm root system are described. Following a transitional juvenile
phase, eight different morphological types of roots have been distinguished according to their development pattern
and state of differentiation: primary vertical and horizontal roots, secondary horizontal roots, upward growing
secondary vertical roots and downward growing secondary vertical roots, superficial and deep tertiary roots and
quaternary roots. The relative position of these types of roots determines a morphological and functional unit of
the root system called "root architectural unit" of the oil palm. This root polymorphism enabled us to define a
morphogenetic gradient, which reflected the oil-palm root-system ontogenesis.

Introduction to the concept of the root architectural unit (Atger,

Root architecture is a fundamental aspect of plant pro-
ductivity (Lynch, 1995) through its functional impor-
tance in the efficient acquisition of soil resources. It is
therefore a subject of considerable interest in agricul-
ture and ecology. Root system architecture is directly
linked to the various functions ensured by the roots.
Many authors have demonstrated links between archi-
tecture and plant anchoring (Coutts, 1983; Ennos et al.,
1993; Fitter, 1986), or with water and nutrient uptake
(Barley, 1970; Bosc and Maertens, 1981; Habib et al.,
1991; Hamblin and Tennant, 1987; Sattelmacher et al.,
1993).
The architecture of a branched system can be per-
ceived through architectural analysis. This analysis,
begun by Hallé and Oldeman (1970) on the aerial sys-
tems of tropical trees, is based on studying how the
meristems of each axis function and how hierarchical
relations are established between these axes. It also
makes it possible to carry out a complete diagnosis of
the different root types making up the branched sys-
tem, and show the growth, branching, differentiation
and mortality processes, which are characterized for
each category of axes identified. This analysis led on
 tural practices than to increase overall knowledge of
FAX No: +33467593858. E-mail: rey@cirad.fr
1992). Architectural analysis is therefore an appropri-
ate tool for measuring the root architecture of plants.
With practical applications in mind, it can also serve
as a back-up for quantification, modelling, then simu-
lation of the various processes identified.
The oil palm, Elaeis guineensis Jacq., has been
cultivated on a large scale since the turn of the cen-
tury for its high oil yield (triglycerides). According to
Oil World Annual (1995), palm oil production world-
wide in 1995 amounted to 17.2 Mt, placing it second
in the world rankings with 23.7% of edible vegetable
oil production, just behind soybean (27.3%). Howev-
er, knowledge of its root system is still only sketchy.
Paradoxically, the surroundings of the oil-palm root
system have been studied more than the roots them-
selves. In fact, earlier studies have often concentrated
on characterizing soil evolution under oil palm (Cal-
iman, 1990; Hartmann, 1991; Tinker, 1976) and the
hydric properties of these soils (Dufrêne, 1989; Olivin
and Ochs, 1978). Only a few studies have been made
of environmental effects on oil palm root system devel-
opment (Agamuthu and Broughton, 1986; Ruer, 1968;
Tailliez, 1971). Moreover, the isolated observations
carried out were more to solve problems linked to cul-

ICPC: PIPS No.: 127936 BIO2KAP

plso6193.tex; 2/07/1997; 16:33; v.7; p.1
34
root system development. For example, oil-palm root
system distribution and the position of the absorbing
parts were studied specifically to determine optimum
fertiliser application (Frémond and Orgias, 1952; Ruer,
1967a, 1968; Tailliez, 1971) or solve water supply
problems (Dufrêne, 1989; Ruer, 1969).
Although some authors (Frémond and Orgias,
1952; Purvis, 1956; Ruer, 1967b, 1968; Wright, 1951;
Yampolsky, 1922) have endeavoured to describe in
varying degrees of detail the different axes making up
the oil-palm root system, only a few (Purvis, 1956;
Ruer, 1968) have described its overall structure or
architecture and nobody has touched upon growth and
development dynamics.
Our initial aim was therefore to carry out a precise
analysis of oil-palm root system architecture. We shall
then show how architectural analysis enabled us to
reveal root polymorphism in the oil-palm root system,
along with its rules of formation and functioning during
ontogenesis. Lastly, knowledge acquired recently in
these fields (Atger, 1992; Hallé et al., 1978; Kahn,
1983; Kubikova, 1967) will enable us to reposition the
specificities of the oil-palm root system in relation to
those of other plants.
Material and methods
Planting material, study site and climate
The oil palm, Elaeis guineensis Jacq. (Arecaceae) is a
perennial monoecious monocotyledon that originated
in the Gulf of Guinea in West Africa. The variety cho-
sen comes from the family usually known as C1001F,
commonly used in commercial plantations.
The optimum conditions for oil palm cultivation
(IRHO, 1978) are (i) at least 2,000 mm of rain per
year, with a minimum of 150 mm per month and there-
fore no water deficit, (ii) temperatures of between 20
and 34  C and, (iii) 1,800 hours of sunshine per year.
Our study was conducted at the La Mé experimen-
tal station in south-eastern Côte d’Ivoire, where the
humid, subtropical climate with marked seasons is
characterized by (i) average annual rainfall of 1,400
mm with a 380 mm year 1 water deficit, (ii) average
temperatures of between 24 and 28  C and, (iii) around
1,800 hours of sunshine per year. The study site is
located on a low altitude plateau resulting from fluvio-
lagoon sedimentation in the tertiary era, at the origin
of loamy-sand detrital formations dominated by coarse
sand (Hartmann, 1991). These formations, commonly
known as "tertiary sands" (Olivin and Ochs, 1978), are
derived from ferrallitization processes (Perraud, 1971)
and give rise to deep, uniform soils with neither appar-
ent interruptions nor constraints to a depth of at least 6
metres.
Architecture and development analysis
Our study was based on concepts developed in plant
above-ground architecture studies (Barthélémy et al.,
1989; Edelin, 1977, 1984; Hallé and Oldeman, 1970;
Hallé et al., 1978). The architecture of a plant or root
system can be defined by the type (or category) and
relative position of the different axes making it up. It
results from the functioning of the above-ground and
below-ground apical meristems (Hallé and Oldeman,
1970).
The architectural analysis of an above-ground
branched system (Hallé and Oldeman, 1970) or of
a root system (Atger, 1992) is based on observing
and characterizing the way in which the different axes
grow, branch and morphologically differentiate at vari-
ous stages of development (typology). An architectural
analysis involves three main stages, described by Atger
(1992): (i) identification and characterization of the
various elements making up the system, (ii) character-
ization of the relative layout of the different axes, along
with their hierarchical relationships and (iii) character-
ization of the sequence in which the different compo-
nents of the system occur, and how they develop. The
architectural analysis that led on to the concept of the
architectural model (Hallé and Oldeman, 1970) does
not take plant dimensions or their spatial occupation
into account, whereas these criteria are crucial if plant
growth dynamics are to be accurately characterized
and especially if any agricultural applications are to
be found. In our approach, the notions of plant size
and spatial occupation are included in the architectural
analysis.
The results are summarized in a table and dia-
grams, which together constitute an architectural dia-
gram (Edelin, 1977), now called an architectural unit
(Barthélémy et al., 1989). An architectural unit is based
on a complete diagnosis of all vegetative axis cate-
gories in the plant. The axes are grouped into types,
associated or not with a topological order. For each cat-
egory of root axes, the characters used to identify them
are noted: growth and branching processes, direction of
growth, nature, topological order, presence or absence
of lignification, deciduousness, diameter, growth rate,
etc.
plso6193.tex; 2/07/1997; 16:33; v.7; p.2
 35

Figure 1. Photos of the oil-palm root system. a. Seedling 36 days after germination; b. 3-month-old root system; c. 1-year-old plant; d. Close-up
of 20-year-old palm root system in situ.

Given the absence of morphological traits as dis- ter, inter-lateral lengths, insertion angle, direction of
tinct as those for the above-ground parts (nodes, leaves, growth and growth rates.
buds, scars, etc.), we had to characterize the root axis We studied 4 growth phases: juvenile phase (0-1
using simple biometrical parameters: length, diame- year); field establishment phase (3 years), adult phase

plso6193.tex; 2/07/1997; 16:33; v.7; p.3

36
(11 years) and the limit of economic viability phase
(20 years). Two methodological approaches were taken
during these different phases: a static approach based
on a morphological and geometrical study of the root
system, and a dynamic approach based on a study
of development kinetics for the different axes making
up the system, along with their growth and branching
processes (Table 1).
Static approach
The study techniques were based on partial or total
excavation (Table 1). Static analysis was only car-
ried out for a given stage of development, since it
destroys the roots observed, but a reconstruction of
growth dynamics was possible by studying the differ-
ent development stages on several individuals from a
homogeneous population.
The investigation was carried out in relatively
homogeneous soil for the juvenile stages and the crop
management sequence was in accordance with IRHO
(1981) recommendations. Ten plants were observed
every fortnight for this juvenile phase. For the adult
stages, under plantation conditions and on homoge-
neous soils, a representative sample (30 to 40 roots) of
each root order was taken and analysed. The direction
of growth, angle of emission on the palm and reori-
entation (final direction of growth) were measured.
These observations were made on lateral profiles, i.e.
by digging a trench perpendicular to the planting row
and carefully removing the soil from around the roots
found. Root samples, were taken from the palm to a
distance of 2 m, down to a depth of 1.5 m. These mea-
surements were completed by frontal profiles (Böhm,
1979) carried out in the planting row from 0 to 2 m
down, in order to obtain vertical root density maps.
Spatial distribution of the roots was observed, along
with their number, type (young, growing, dead, etc.)
and their diameter. A few surface roots were also fol-
lowed from the stem to their apical tip, primarily to
characterize the life span of their apical meristem. Last-
ly, total root-soil plate excavations were carried out to
make an exhaustive count of the roots emitted, charac-
terize the mortality rate and locate new emissions.
Dynamic approach
For juvenile root systems aged 0 to 3 months, the
dynamic analysis (Table 1) mostly involved the use
of miniature root-growth chambers fitted with Nylon
mesh (Le Roux and Pagès, 1994) and measuring 100
cm high by 60 cm wide by 1.6 cm thick. Seven minia-
ture root-growth chambers, each containing 2 oil palms
grown from germinated seeds, were inclined at an
angle of 30 from the vertical. They were placed out-
side under slight shading, the substrate used was poor
forest topsoil (Jourdan et al., 1995), and the soil tem-
perature varied between 24  C at night and 32  C
during the day.
For juvenile root systems aged 3 to 12 months, two
wooden miniature root-growth chambers were built:
(i) one was 47 cm high, 59 cm wide and 8 cm thick,
containing three 3-month old plants and (ii) another
74 cm high, 54 cm wide and 26 cm thick, contain-
ing one 7-month old plant. The two miniature root-
growth chambers had 2 glass sides and were placed in
direct sunlight (growing conditions for nursery plants).
The Nylon mesh was not used in this case, due to the
volume occupied by the roots. The substrate and soil
temperatures were the same as before.
For root systems in the field, 20 "field rhizotrons"
were set up in two positions, near vertical and near
horizontal, and at different distances from the palms.
The vertical "field rhizotrons" were square, with 1 m
sides, and sloped 20 from the vertical. The Plexiglas
window was held against the soil by a metal frame
propped up by bamboo poles. The horizontal "field
rhizotrons" were rectangular (0.5 m1 m) and were
inclined 20 to 30 from the horizontal.
The progression of the roots was recorded using
different coloured indelible felt pens at 3-day intervals
for the minirhizotrons and every 7 days for the "field
rhizotrons". Once these multicoloured records were
processed, after a stage of computer coding (LADORA
software developed and described by Colin-Belgrand
et al. (1989)), various types of processes can be carried
out (growth kinetics, rhythm ratios, dates on which the
different roots appear, etc.).
Histology
Numerous root samples were taken, at all stages of
oil palm development, in order to characterize cell
differentiation in the various tissues, and validate the
root typology already discovered. The samples were
fixed in a solution comprising a phosphate buffer, 25%
glutaraldehyde, 10% paraformaldehyde and caffeine.
After embedding in resin, they were sliced with a 3
m thick microtome and stained with Schiff’s periodic
acid and aniline blue-black (Fisher et al., 1968).
plso6193.tex; 2/07/1997; 16:33; v.7; p.4
 37
Table 1. Overview of the various methods of investigation of the root system for the four growth phases studied in this work

Static approach Dynamic approach

Root system Lateral and frontal Root-soil plate Miniature root- Vertical and horizontal
Growth phases studied excavations profiles excavations growth chambers <<field rhizotrons>>

Juvenile (0-1 year) total no no yes no

Field establishment (3 years) partial yes yes no yes
Adult (11 years) partial yes yes no yes
Limit of economic viability (20 years) partial yes yes no yes

Table 2. Quantitative parameters of different root types of oil palms in the juvenile phase. RI1 5 -primary root emitted for a 1 to
5-month-old oil palm, RI5 12 -primary root emitted for a 5 to 12-month-old oil palm

Juvenile Primary roots Lateral roots

phase Radicle RI1 5 RI5 12 Long roots Medium roots Short roots

Average
growth rates 0.44 0.88 1.02 0.39 0.30 0.27
(cm day 1 ) 0.11  0.33  0.15  0.02 0.02 0.07
Maximum length
(cm) 50 100 180 50 10 1.5

?
Diameter ( )
(cm) ?
0.10< <0.20 ?
0.20< <0.30 ?
0.30< <0.40 ?
0.05< <0.10 ?
0.03< <0.06 ?
0.02< <0.05

Number of
vascular bundles 10 to 15 15 to 25 25 to 40 8 to 10 5 to 8 3 to 5

Average distance
between 0.14 0.24 0.84 0.11 0.11 Non-branched
lateral roots (cm) 0.015 0.02 0.20  0.01  0.01

Results constant length and remains unbranched. Several types

Juvenile phase from 0 to 1 year in the nursery
During the juvenile phase, from 0 to 1 year, we differ-
entiated the radicle (RAD) from the set of adventitious
roots emitted (RI).
Radicle (RAD)
The radicle is downwardly oriented, slightly undulat-
ing and ends in a pointed, white apex bearing a root cap
(Figure 1a). Its growth is continuous, with no apparent
rhythm, limited in time and the average growth rate
is around 0.44 cm day 1 (Table 2). It is rarely more
than 50 cm long with an average diameter of 1.5 mm.
Its ramification is diffuse, dense and covers the entire
axis, apart from the apical zone, which is of relatively
of lateral formations can be identified (Jourdan et al.,
1995); (i) "long", (ii) "medium" and (iii) "short", all
emitted at an angle of around 90 . The so-called "long"
roots are mostly located at the base of the radicle and
are emitted very soon after germination. They vary in
length and are usually over 10 cm long, growing at an
average rate of 0.39 cm day 1 (Table 2). These "long"
roots bear two types of roots: (i) "medium" and (ii)
"short". The so-called "medium" roots do not exceed
10 cm in length, they grow at an average rate of 0.3
cm day 1 (Table 2), and are distributed over the entire
branched zone of the radicle and "long" roots described
above. Irrespective of their topological order within
the branched system, these "medium" roots are mor-
phologically similar to each other. They bear "short"
lateral roots with the same density as above. The so-
called "short" roots do not exceed 1.5 cm in length,

plso6193.tex; 2/07/1997; 16:33; v.7; p.5

38
#
Figure 2. Variation in the cumulated average number of roots ( )
and leaves ( ) emitted per palm during the 0-1 year period. The
emission rate values for roots (Rr ) and leaves (Rl ), during each
period are expressed as roots or leaves emitted per year.
grow at an average rate of 0.27 cm day 1 (Table 2)
and are non-branching. They differ from one another
by their origin, i.e. the type of the axis by which they
are borne (Figure 1a).
Adventitious primary roots (RI)
On average, the first adventitious root is emitted a
month after germination (Figure 2). During the first
year, the subsequent RIs are emitted at an accelerating
rhythm, whereas leaf emission remains constant (21
leaves per year). During this period, the RIs take on
a branched structure similar to that developed by the
radicle, insofar as three branching orders are observed
(Figure 1b). Their direction of growth is vertical and
downwards. The average growth rate evolves from
0.88 cm day 1 (Table 2) for 1- to 5-month old palms
(RI1 5 ), to 1.02 cm day 1 for 5- to 12-month old palms
(RI5 12 ). The final length varies from 1 m at 5 months
to 1.8 m at 1 year. They are white when young, but
rapidly turn light to dark brown (Figure 1c). The diam-
eter remains constant for each root emitted but varies
from 0.2 to 0.4 cm depending on the date of emis-
sion between 1 and 12 months, the roots emitted at
12 months having a larger diameter. The number of
vascular bundles depends on root diameter and ranges
from 15 to 25 at 1 month and from 25 to 40 at 1 year.
Branching, which is comparable to that of the radi-
cle, is diffuse and dense. The lateral roots produced,
which are also of several types, are distributed along
the whole length and around the entire circumference
of the RIs, inserted at an angle of 90 . The average
distance between laterals increases from 0.24 cm for
RI1 5 to 0.84 cm for RI5 12 along with an increase in
the RI diameter (Table 2). The so-called "long", "medi-
um" and "short" roots described above are also found
here, but in different proportions. The proportion of
"long" roots increases, with a gradual disappearance
of "medium" and especially "short" roots. The "long"
roots are mostly distributed in the proximal quarter of
the RIs. The characteristics of these lateral formations
are similar to those borne by the radicle, but with a
slight increase in diameter.
Field growth phases from 1 to 20 years
Primary roots (RI) and secondary roots (RII)
New RIs are mostly emitted on the periphery of the
root-soil plate, often through the petiole bases of leaves
inserted at soil level. Two distinct types of RIs have
been described in the field: vertical, downward grow-
ing RIs, RI VDs, and horizontal RIIs, RII Hs (Figures
1D and 3). Their number increases steadily up to 11
years, then the number of RI VDs stabilizes where-
as that of the RI Hs continues to increase (Figure 4).
These two types of RI therefore differ in number and
direction of growth, but also in diameter and the type of
roots they bear. The RI VDs, with an average diameter
of 0.45 cm, mostly bear horizontal RIs (RII H) and are
only slightly ramified (Table 3). These RIIs are quite
short (less than 20 cm), highly branched and distribut-
ed around the entire circumference of the RIs. The RI
Hs, with an average diameter of 0.68 cm, bear both
upwardly growing vertical RIIs (RII VU), of medium
length (0.5 to 2 m) and highly branched, and down-
wardly growing vertical RIIs (RII VD), which are very
long (more than 3 m) and only half as branched as
the RII VUs (Table 3). The very precise direction of
these RIIs, either upward or downward, gives the RI
Hs a bilateral symmetry which marks a clear difference
from the radial symmetry of the RI VDs (Figures 1D
and 3). The average diameter of the RII Hs, RII VUs
and RII VDs is more or less the same, approx. 0.2 cm
(Table 3).
The RI VDs and RII VDs have the characteristics
of orthotropic axes (vertical growth, radial symme-
try, lateral formations emitted radially at an angle of
90 ), with positive geotropism; they are therefore said
to be positive orthogeotropic axes. Likewise, the RII
VUs, which are also orthotropic axes but with nega-
plso6193.tex; 2/07/1997; 16:33; v.7; p.6
 39

Figure 3. Oil-palm root architectural unit. Partial drawing of the root system on a 10-year-old oil palm with the 8 root types described. Apex

death indicated by . The harvesting pole measures 3.5 metres.

#
Figure 4. Variation in the average number of RI H ( ) and RI VD ( ) emitted per tree over time.

tive geotropism, are said to be negative orthogeotropic
axes. On the other hand, the RI Hs are plagiotrop-
ic axes (horizontal growth, bilateral symmetry, lateral
formations emitted at 90 in the same plane), without
reactions to gravity, and are said to be plagiogeotropic
axes. RII Hs retain generally horizontal growth, they
are ageotropic. These roots have radial symmetry and
the lateral axes (RIII) are emitted radially in all direc-
tions.

plso6193.tex; 2/07/1997; 16:33; v.7; p.7

40

Figure 5. Anatomy and morphology of oil palm roots. A. Histological cross-section of an RI; B. Histological cross-section of an RIV; C.
RIV seen under a scanning electron microscope; D. General view of a pneumathode under a scanning electron microscope. Abbreviations:
a–aerenchyma, c–cortex, e–endodermis, h–hypodermis, p–pith, rh–rhizodermis, s–sclerenchyma, v–medullary vessel.

plso6193.tex; 2/07/1997; 16:33; v.7; p.8

 41
Table 3. Quantitative parameters of different root types of oil palms in the << adult >> phase of field cultivation

<< Adult >> phase

of field RI VD RI H RII VU RII VD RII H sRIII dRIII RIV
cultivation

Average
growth rates 0.30 0.30 0.20 0.20 0.20 0.08 0.08 -
(cm day 1 ) 0.11  0.11  0.04  0.04  0.04  0.04 0.04
Maximum length
(cm) 600 2500 200 600 20 20 10 1.5

?
Diameter ( )
(cm) ? ? ? ? ? ?
0.41< <0.50 0.50< <0.80 0.15< <0.25 0.20< <0.25 0.13< <0.22 0.05< <0.14 0.07< <0.15 0.03< <0.07 ? ?
Number of
vascular bundles 34 to 45 34 to 45 15 to 25 15 to 25 15 to 25 8 to 12 8 to 12 3 to 5

Average distance
between lateral 2.20 2.90 0.23 0.54 0.23 0.11 0.15 Non-branched
roots (cm)  1.50  1.70  0.09  0.48  0.15  0.07 0.11

RI H, RI VD and RII VD growth is of the indefinite
type, their final length ranges from twenty or so metres
for the RI Hs to over 6 m (limit of observations) for
the RI VDs and RII VDs. RII VU growth is limited by
the soil surface, which they reach quite quickly (Figure
3). The average growth rate of RIs in the field evolves
from 0.63 cm day 1 ( 0.2) at 3 years to 0.3 cm day 1
( 0.11) at 11 years, remaining stable thereafter. For
RIIs, the rates decrease from 0.27 cm day 1 ( 0.1) at
3 years to 0.2 cm day 1 ( 0.04) at 11 years, stabilizing
thereafter.
RIs and RIIs in the field have a characteristic colour.
A few centimetres from the apical region they are usu-
ally white, becoming brown then purple and final-
ly black a metre from the apex. The aerenchyma of
RIs and RIIs is highly developed and may take up
almost 70% of root volume (Figure 5a). The differ-
ence between RIs and RIIs lies mainly in the size of
the stele, which is larger in RIs. An average of 20
vascular bundles were observed in each RII and 40 in
each RI. These roots can become considerably lignified
a few dozen centimetres from the apex. Such lignifi-
cation first affects the xylem vessels, then the sub-
hypodermic region and the pith, then the endodermis
and neighbouring regions. Eventually, only the phloem
poles and the aerenchyma remain non-lignified.
Under oil palms, at less than 50 cm from the root-
soil plate, the RIs (especially RI VD) bear numerous
relay axes emitted after the death and self pruning of
successive bearing axes (Figure 3). This is not a rare
occurrence, indeed it affects 12.7%, 41.3% and 58.8%
of primary roots emitted by palms aged 3, 11 and 20
years respectively.
Tertiary roots (RIII) and quaternary roots (RIV)
All RIIs in the field emit RIIIs at an average angle of
90 , which could only be distinguished according to
size and branching criteria. The RIIIs borne on RII Hs,
and the RIIIs borne on RII VDs are generally shorter
and much less branched than those borne by RII VUs.
The least branched RIIIs mostly found deep down,
are designated dRIII; the highly ramified RIIIs, found
mostly near the surface, are designated sRIII (Table 3
and Figure 3). These two types of RIII show definite
growth, with dRIII rarely exceeding 10 cm in length
and sRIII 20 cm. They are beige to brown in colour
and never turn very dark. They are around 0.1 cm in
average diameter and have ten or so vascular bundles.
There is no preferential growth direction; these roots
are ageotropic.
The branched system formed by RII VUs and sRI-
IIs sometimes forms a veritable "root mat" more than
20 cm thick. Numerous traumatic incidents can trigger
the emission of relay axes, which increase the branch-
ing potential of these roots (Figure 6). RII VUs and
sRIIIs even come out of the ground when conditions

plso6193.tex; 2/07/1997; 16:33; v.7; p.9

42
Figure 6. Drawing of an RII VU taken from the 0-10 cm horizon. Abbreviations: pn–pneumathode, r–reiterated RII VU.
are favourable (saturating humidity in the rainy season)
and may penetrate the remnants of decomposing peti-
oles and leaves left in the windrows. The existence of
numerous pneumathodes (Yampolsky, 1924) on these
roots (Figure 6) is indicative of saturating humidity.
When conditions become unfavourable, such as in
the dry season, the roots located on the surface die
and decompose. Thereafter, when abundant rainfall
returns, the surviving underground roots can start emit-
ting numerous relay axes again, which reach the sur-
face horizons that are rich in organic matter. All RIIIs
bear identical RIVs (Figure 3): ageotropic, white, less
than 1.5 cm long with an average diameter of 0.05 cm,
and very few vascular bundles (Table 3 and Figure 5b).
RIIIs and RIVs do not reveal any tissue lignification
similar to that occurring in RIs and RIIs, and they are
therefore classed as non-woody.
RIII growth is definite, continuous and regular, with
an average increase ranging from 0.14 cm day 1 (
0.07) for 3-year old palms to 0.08 cm day 1 ( 0.04)
for 11-year olds, stabilizing thereafter. RIV growth in
the field was not measured because their small size
prevented reliable values from being obtained with a
weekly observation rate.
Some characteristics are common to all types of oil
palm roots, such as the existence of calcium oxalate
precipitates in bundles of raphides in cells on the
periphery of the cortex, and endomycorrhizae fila-
ments. The epidermal cells (rhizodermis) located near
the root apex have a specific shape, similar to that of
a grain of maize and are regularly distributed on the
surface of each root (Figure 5c). Many pneumathodes
have also been observed, especially on surface roots
in the field (RIII and RIV), but also on the radicle, RIs
and RIIs in the juvenile phase. They are characterized
by a split of the rhizodermis and hypodermis, exposing
the internal cells of the cortex. On lateral roots, such
splitting only occurs in the few millimetres near the
zone of insertion, where a sleeve of intact cells persists
(Figure 5d).
To conclude, 6 types of roots were found in the
juvenile phase and 8 types in the adult phase using
simple morphological and geometrical criteria. These
results are shown in Table 4 which summarizes the
morphological characters and types of growth, branch-
ing and differentiation, but also the layout of the dif-
ferent axes making up the adult oil-palm root system.
This summary table characterizes the root architectural
unit of the oil palm.
Discussion and conclusion
Root polymorphism
Diversity of differentiation states
The oil-palm root system revealed the existence of
diversity in root diameter, growth rate and direction,
plso6193.tex; 2/07/1997; 16:33; v.7; p.10

type of branching processes and life span, among other
things. Despite this apparent diversity, our architec-
tural analysis tools enabled us to establish a typology
of different root axes. We detected 8 stable root types
which characterize the root architecture of the oil palm.
Each root type has morphological, morphogenetic and
functional properties which differentiate it from the
other root types (Atger, 1992). It therefore represents a
population of homogeneous roots with a set of similar
characters (structure, way of functioning, etc.). Then
it characterizes the specific state of differentiation of
the set of meristems of the plant (Hallé and Oldeman,
1970), which helps to maintain the polymorphism of
root systems (Le Roux and Pagès, 1994).
Relations between differentiation criteria
Close relations exist between the various differentia-
tion criteria of root system axes. In the literature, there
are many cases of positive correlations between the api-
cal diameter of roots and their growth rate, such as in
the rubber tree (Le Roux, 1994), the maple tree (Lyford
and Wilson, 1964), the pine tree (Wilcox, 1968) or
some cereals (Hackett, 1969a, 1969b), and with their
internal anatomical structure, particularly the number
of xylem poles (Charlton, 1967) or the diameter of
the primary xylem (Horsley and Wilson, 1971). Other
relations bringing into play root length and diameter,
branching density and the action of external factors on
root development are summarized by Coutts (1987).
As far as the oil-palm root system is concerned, roots
that have high growth rates, long life spans and low
density branching (such as primary roots), have both
a large diameter and a large number of woody poles
(Tables 2 and 3). On the other hand, unbranched roots,
with a low growth rate that is limited in time (such
as quaternary roots), have a very small diameter and
number of woody poles. Likewise, we showed (Jour-
dan, 1995) that there is a close relation between root
diameter and branching parameters (case of RIIs in
the field). Other correlations have also been found,
notably between the diameter of juvenile RIs and the
length of the unbranched apical zone, but also between
the growth rate and the length of the unbranched api-
cal zone of radicles. The diameter is therefore one of
the main criteria for differentiation of root types and
should be taken into account in architectural analysis
of the oil-palm root system.
43
Root system hierarchy
The existence of several levels of differentiation in
a root system was described at turn of the century in
terms of "heterorhizia" (Tschirch, 1905; Noelle, 1910).
Since then, several attempts at root axis classifica-
tion have been published and were summarized by
Kubikova (1967): anchorage roots and feeding roots,
long roots and short roots (Wilcox, 1964), pioneer
roots and feeding roots, or even woody roots and
non-woody roots (Lyford and Wilson, 1964). This
terminology has been clearly defined by Sutton and
Tinus (1983). Root systems were thus made up of 2
types of qualitatively different axes: macrorhizae and
brachyrhizae (Jenik and Sen, 1964). Macrorhizae are
the large roots that explore the soil and extend the rhi-
zosphere (Kahn, 1977); they are vertical with marked
positive geotropism in which case they represent the
"orthotropic state", or horizontal in which case they
represent the "plagiotropic state". These axes persist
within the system, their secondary growth can be con-
siderable, they have a large number of woody poles,
a highly developed pith, a long and pointed apex of
large diameter and, lastly they show strong reiteration
ability (Oldeman, 1974). According to many authors,
these roots also ensure the anchorage and the stabil-
ity of trees in the soil (Coutts, 1986; Quine et al.,
1991). Brachyrhizae are fine roots that are not part of
the root main structure but form the ephemeral roots
at the summit of the plagiotropic hierarchy, like leaves
in the above-ground system (Kahn, 1983). They are
characterized by a small number of woody poles, the
absence of pith, a substantial cortex, a short apex with
a small diameter, a limited life span and no reitera-
tion ability. Although their respective roles are clearly
established, conduction in macrorhizae and absorption
in brachyrhizae (Kubikova, 1967; Lyford and Wilson,
1964), the distinction between these two root cate-
gories is not always so clear. In fact, all the interme-
diate axes between macrorhizae and brachyrhizae can
be found within the same root system and a given axis
may switch from one morphological and functional
structure to another (Wilcox, 1964).
These more or less restrictive classifications, which
differentiate between large woody roots and short non-
woody roots, have mostly been described for woody
Dicotyledons and Gymnosperms. In the oil palm, the
macrorhizae would be the RIs and RIIs observed in
the field, and the brachyrhizae would be the RIIIs and
RIVs. Indeed, the RI VDs and RI Hs along with the
RII VDs, play a full role in the anchorage of the tree
plso6193.tex; 2/07/1997; 16:33; v.7; p.11
44
and in exploring the volume of available soil, whereas
the RIIIs and RIVs mostly exploit it. Nevertheless, oil
palm roots do not exactly satisfy all the macrorhiza and
brachyrhiza criteria, since, most importantly, they are
characterized by an absence of cambium, the existence
of pith in the fine roots and by reiteration ability in the
tertiary roots.
The oil-palm root system, and more generally that
of the Monocotyledons, reveals other analogies with
the root system of Dicotyledons or Gymnosperms. For
example, on germination the oil palm has an orthotrop-
ic tap root, or radicle, from which horizontal lateral
roots are emitted. The main difference is the absence
of radicle perenniality: it self-prunes 3 to 4 months
after germination. Contrary to the so-called primary
root system (Kahn, 1983), in which the single primary
root is orthotropic and governs the plagiotropic dif-
ferentiation of its lateral formations, the "orthotropic
state" in the case of oil palm is thus represented by a set
of vertical primary roots (juvenile RIs, then RI VDs),
which succeed each other whilst maintaining the same
behaviour. The “plagiotropic state” in the oil palm is
characterized by a set of horizontal primary roots (RI
H), emitted more than a year after germination, which
succeed each other in time and mostly bear vertical
lateral roots. This "plagiotropic state" was observed by
Granville (1974) in another palm, Mauritia flexuosa,
which has 40 m long horizontal roots that are flattened
at 30 m and beyond! The "plagiotropic" roots of the
oil palm do not seem to be directly under the influence
of the apical meristems of the orthotropic roots since
they do not grow from them, whereas that is the case
in most woody plants, such as the cocoa tree (Dyanat-
Nejad and Neville, 1972a, 1972b) or several species of
oak (Beissalah et al., 1988; Champagnat et al., 1974;
Riedacker et al., 1982). The neutral state of free growth
in a root axis is defined according to Kahn (1977) by
the "orthotropic state"; RI Hs do not therefore seem to
correspond to this definition. Lastly, the "plagiotropic
state" of the oil palm is complex insofar as the RI Hs
give rise to two clearly distinct types of lateral roots,
one growing upwards (RII VUs) and bearing the dense-
ly branched "sRIIIs + RIVs" combination making up
the root mat, and the other growing downwards (RII
VDs) and bearing much shorter and less branched roots
(dRIII). In addition, this "plagiotropic state" occurs late
in the root system, after a transitional juvenile phase
(Granville, 1974) lasting roughly a year.
On the whole, the oil-palm root system is thus char-
acterized by the succession in time of three phases: (i)
a very brief phase where the radicle reveals a marked
"orthotropic state", similar to the tap root of woody
plants, (ii) a transitional phase where juvenile RIs char-
acterize (still by analogy with the tap root system) an
"orthotropic state" and lastly, (iii) a much longer phase
where the RI VDs represent an "orthotropic state" and
the RI Hs a "plagiotropic state". The oil-palm root
system thus appears as a structured set of differen-
tiated and hierarchized roots, with clearly identified
corresponding functions: exploration and exploitation
of the soil. In fact, it constitutes a veritable morpho-
logical and functional unit that we have called "root
architectural unit" (Table 4) in reference to the work
by Edelin (1984) on the above-ground system, then by
Atger (1992) on the root system of trees.
Root growth and development
Observations of the oil-palm rooting process, from ger-
mination to the adult phase, show that the formation of
the different types of roots over time is not random and
disordered, but in accordance with a well determined
programme. Completion of this programme, which in
fact reflects development of the different meristems
according to their specific differentiation sequence,
governs root architecture development.
Establishment growth
The radicle of the oil palm, and of Monocotyledons in
general, has a limited life span, since it is a low diam-
eter axis, with exclusively primary growth (Hallé et
al., 1978). According to these authors, as the absorb-
ing zones are extended through branching, the point
of attachment of the radicle to the above-ground part
becomes a veritable "bottleneck" and the radicle alone
is unable to meet increasing demand from the above-
ground organs. The plant then emits numerous adven-
titious roots, which thus by-pass the "bottleneck".
Throughout the life of an oil palm, each root emit-
ted keeps the same morphological characteristics, par-
ticularly the same diameter, up to its death. More-
over, during the juvenile phase these characteristics
change from one primary root to another depending on
the emission date (Jourdan, 1995). The roots emitted
on a certain date have, among other things, a larger
diameter, faster growth, a lower branching density and
greater life span than those emitted before this date.
It is only when all the different root types inventoried
have been expressed that the characteristics of each
new primary root emitted remain identical to those
of the previous primary root. This growth process,
plso6193.tex; 2/07/1997; 16:33; v.7; p.12
 45
Table 4. Oil-palm root architecture unit. W–woody, NW–non woody, I–indefinite growth, D–definite growth,  –maximum observed value that can
be exceeded, DS–diffuse and sylleptic branching, NB–non-branched, R–radial symmetry, B–bilateral symmetry, L–long-term self-pruning, more
than 6 months, M–medium-term self-pruning, from 1 to 6 months, S–short-term self-pruning, less than or equal to 1 month

Morphological
parameter RI VD RI H RII VU RII VD RII H sRIII dRIII RIV

Woody axis W W W W W NW NW NW

Geotropism Orthogeotropic Plagiogeotropic Orthogeotropic Orthogeotropic Ageotropic Ageotropic Ageotropic Ageotropic

(positive) (negative) (positive)

Average 0.53 0.63 0.18 0.23 0.15 0.10 0.10 0.05

diameter (cm)  0.07  0.10  0.05  0.08 0.04 0.03 0.03 0.01
Growth I I I I D D D D

Maximum length 600 2500 200 600 50 20 10 1.5

(cm)

Branching DS DS DS DS DS DS DS NB

Symmetry R B R R R R R R

Self-pruning L L M M M M M S

where the organs develop more and more up to the
perduring ultimate stage, is similar to the "establish-
ment growth" defined by Tomlinson and Zimmerman
(1967) for the above-ground part of a palm, Rhapis
excelsa, and described since in other Monocotyledons
(Tomlinson and Esler, 1973) and certain Dicotyledons
considered to be "primitives" (Blanc, 1986).
The dynamics of space occupation
Kahn (1980, 1983) defined 4 phases, which cover all
the rooting processes of forest plants: (i) establish-
ment of the orthotropic axis which produces a few
root mats, (ii) production of plagiotropic macrorhizae
from the initial axis, (iii) considerable development of
these axes which extend the exploitation zone from
the trunk and thereby delimit a distal exploitation ring
and (iv) production of new plagiotropic macrochizae
from the base of the tap root or trunk, which exploit
previously abandoned proximal space. However, cer-
tain plants do not complete all 4 phases, e.g. single-
stem plants in the understorey, most shrubs and a
few trees in which adaptive reiteration is non-existent
(Kahn, 1983). When observing the different develop-
ment phases of the oil-palm root system, the different
phases described by Kahn (1983) are generally found.
Indeed, we have shown that the juvenile root system
(from 0 to 1 year) was characterized by an "orthotrop-
ic state" in the radicle and juvenile RIs. The following
phase is characterized by the appearance of RI Hs more
than a year after germination, corresponding to pla-
giotropic macrorhizae. These horizontal roots explore
and exploit superficial regions distant from the palms,
through RII VUs - sRIIIs - RIVs, but also deep regions,
through RII VDs - dRIIIs - RIVs. The final phase is that
of new exploitation of the proximal space by continual
emission of oil palm roots and total RI reiteration under
the palm (Jourdan, 1995). Consequently, the proximal
space is permanently exploited by the oil palm, unlike
the case of forest trees described by Kahn (1980) and
Atger (1992). The dynamics of space occupation by
the oil-palm root system can thus be expressed in dia-
grammatic form as in Figure 7.
Synthesis of root development
During its ontogenesis, the oil palm forms different
types of roots in an ordered sequence. The "transient"
radicle is rapidly replaced by numerous adventitious
roots that gradually produce increasingly developed
lateral formations towards the periphery of the system,
along the lines of growth by intercalation (Atger and
Edelin, 1994; Edelin, 1984). Eventually, this system

plso6193.tex; 2/07/1997; 16:33; v.7; p.13

46
Figure 7. Various stages in oil palm rooting. The radicle stage (A) is rapidly replaced by the juvenile RI stage (B), then by the adult stage
(side view: C and view from above: D). The RI VDs and RII VDs, along with the RI Hs extend the oil palm’s exploitation zone at depth and
superficially respectively. Continuous root emission and the RI total reiteration process result in continuous exploitation of the proximal space
(D).
comprises a finite number of root types that charac-
terize the root architectural unit of the oil palm. The
oil-palm root system, along with those of herbaceous
and other woody monocots presents a relative rigidi-
ty in its developmental patterns. The continuity in the
development characteristics of the different root types
inventoried means that a morphogenetic gradient can
be defined between these different types, independent-
ly of their branching order. They are mainly character-
ized, through the architectural analysis, by a whole set
of various morphological and morphogenetic parame-
ters (Table 4), essentially the root diameter.
With knowledge of this gradient, mathematical for-
malization is possible and quantitative architectural
models can be developed. These models can be used to
describe various architecture building processes, such
as branching (Jourdan et al., 1995) and to summarize
all the knowledge acquired through more complex for-
malization (Reffye et al., 1991). An appropriate digital
processing sequence (modelling, simulation, display,
postprocessing on digital 3-D mock-ups) can then be
envisaged to achieve practical applications of agro-
nomic interest. This will be made possible under the
condition of both acquiring supplementary knowledge
in the environmental effects on the root system devel-
opment and in the complex phenomena of intra- or
interspecific root competition.
Acknowledgements
We would like to thank Kouamé Brou, Director of the
IDEFOR-DPO station at La Mé in the Côte d’Ivoire,
for his hospitality and for the facilities made available
locally for carrying out the observations.
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Section editor: H Lambers
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