The Spread of Ganoderma From Infective Sources in The Field and Its Implications For Management of The Disease in Oil Palm
The Spread of Ganoderma From Infective Sources in The Field and Its Implications For Management of The Disease in Oil Palm
The Spread of Ganoderma From Infective Sources in The Field and Its Implications For Management of The Disease in Oil Palm
8
J.
Spread
FloodofetGanoderma
al. from Infective Sources in the Field
Introduction
Basal stem rot (BSR), caused by species of Ganoderma, has been recognized as a
serious disease of oil palm (Elaeis guineensis) for over 80 years and has caused
severe economic loss in Malaysia (Turner, 1981; Singh, 1991; Ariffin et al.,
1996) and North Sumatra, especially during the past 30 years (Hasan and
Turner, 1998). Initially, the disease was considered to affect only old palms (at
least 25 years old) but, with successive palm generations on the same land, a
higher disease incidence has been observed and the symptoms occur earlier
with each replanting. For example, in Sumatra, where replanting was con-
ducted by pushing over the old stand, with no attempt to remove BSR-infected
tissues, young replanted palms have died from the second year onwards after
planting. Economic loss may begin to occur within 10 years and severe loss
after 15 years; the normal life span of a planting of oil palm would be 25–30
years. Where this phenomenon has occurred, it has generally been accepted
that there has been an increase in available inoculum from the previous palm
planting (Turner, 1981). Thus, efforts to manage the disease have been
directed largely towards disease avoidance through reducing the amounts of
potential infection sources for the replanting at the time of clearing the old
stand (Singh, 1991). In Malaysia, the benefit of this clean-clearing approach
over no disease avoidance measures has been demonstrated (Hashim, 1991).
However, the total removal of all infective tissues from an old stand with a high
©CAB International 2000. Ganoderma Diseases of Perennial Crops
(eds J. Flood, P.D. Bridge and M. Holderness) 101
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102 J. Flood et al.
disease incidence is a practical impossibility, and so the aim has been to con-
centrate on removal of as many of the larger tissue sections as economically
feasible. To investigate the efficacy of sanitation in BSR management, a series
of trials was undertaken at Bah Lias Research Station (BLRS) of P.T.P.P.
London in North Sumatra, Indonesia over a period of several years. The trials
were designed to assess the relative importance of various tissue remnants
from the old palm stand as potential sources of inoculum at replanting (Hasan
and Turner, 1998) so as to make practical recommendations for management
of the disease in Sumatra.
The trials were set up as to be sufficiently large to overcome any variations
in BSR inoculum, with each treatment being replicated at different sites in the
plantation. Six-month-old seedlings were used to bait the Ganoderma-infected
material. External leaf symptoms developing on these bait seedlings were
recorded for the duration of each trial, while at the end of each trial, all
seedlings were examined internally for Ganoderma infection by destructive
sampling and plating to Ganoderma-selective medium (GSM) (Ariffin and Idris,
1991). Each experimental plot was isolated by a deep trench to increase the
likelihood that any infection recorded was derived from the tissue being tested
and not from an outside source, but more recently, molecular fingerprinting
techniques (Miller et al., this volume; Bridge et al., this volume; Rolph et al., this
volume) became available which allowed confirmation of the origin of the
pathogen in infected seedlings.
Stump Tissues
A stump comprises the base of the palm, or bole, and the thick crust of roots
immediately surrounding it. Stumps are usually recognized as major sources of
BSR. The first trial compared BSR stumps, prepared by felling diseased palms
about 20 cm above the ground (standard practice), as an inoculum source
with stumps derived from healthy palms. Around each stump eight bait seed-
lings were planted. An additional treatment, planting additional seedlings
immediately outside the plot isolation trench and isolating these by a further
trench 1 m from the inner trench, in order to emphasize disease origin, was
added. Each treatment was replicated eight times at different sites. Six months
after planting, a small number of seedlings began to exhibit disease symptoms
and by the end of the 28-month trial period, 76% of all bait seedlings showed
symptoms; Ganoderma was isolated from these seedlings. In comparison, seed-
lings planted outside the first trench and within the second isolation trench
perimeter showed very little infection – only 1.6% of these seedlings were
diseased and, at 80% of the replicate sites, these seedlings exhibited no
symptoms at all. No disease was recorded in bait seedlings planted around
healthy palm stumps within the period.
Another trial aimed to assess the effect of stump size on disease incidence.
Additional treatments in this trial were comparisons with stumps derived from
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Spread of Ganoderma from Infective Sources in the Field 103
Table 8.1. Effects of the removal of soil and palm tissues from around healthy
basal stem rot (BSR)-infected stumps on disease incidence after 24 months.
BSR 0 97 75
BSR 20 85 58
BSR 40 70 28
BSR 60 55 21
BSR 80 0 0
BSR 100 0 0
Healthy 60 0 0
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to be expected since the amount of infective tissue within stumps and its
location in relation to seedling root contact will differ considerably, as will the
rates of subsequent decay.
During the course of these trials, molecular fingerprinting techniques
became available for Ganoderma and were used to confirm the origin of the
Ganoderma from infected bait seedlings. Material was collected from diseased
seedlings, stump tissues and sporophores growing on the stumps and
isolations made on GSM. DNA was extracted from pure cultures (Miller et al.,
1999) and purified DNA samples tested with the ITS3/GanET primer (Bridge
et al., this volume) to check their identity. All isolates were positive with the
ITS3/GanET primer, confirming that the pathogen had been isolated from
the various tissues (Fig. 8.1). Mitochondrial profiles were generated using the
enzyme HaeIII as the restriction enzyme (Miller et al., 1999; Rolph et al., this
volume) and revealed that identical profiles were present in the BSR stumps
and the infected bait seedling material (Figure 8.2).
% Seedling infection
Months after
planting Site A Site B
6–8 8 11
9–12 45 32
13–18 20 35
19–24 20 14
25–28 7 8
Fig. 8.1. Confirmation of the presence of the pathogen from stump tissues and
infected seedlings.
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Spread of Ganoderma from Infective Sources in the Field 105
Trunk Tissues
Unless trunks of the old palm stand are destroyed at the time of replanting, they
are usually windrowed, i.e. placed in rows. Such trunks are colonized by many
species of fungi, including Ganoderma. Trunks will also remain following a
number of estate practices, e.g. following underplanting, those excavated as
low-yielding, palms removed for thinning or road construction and excavated
diseased palms, and palms affected by upper stem rot (USR) often remain
standing for long periods, as do palms killed by lightning. The trials summa-
rized below assessed the significance of trunk sections as sources of BSR
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Spread of Ganoderma from Infective Sources in the Field 107
Table 8.3. Basal stem rot incidence in bait seedlings around oil-palm residues.
Diseased Nil 9 38 2 3 2 2
Diseased P 13 34 3 5 2 3
Diseased C 19 34 4 6 1 2
Diseased PC 17 27 4 5 5 6
Healthy Nil 0 6 1 1 1 1
Healthy P 1 9 2 3 1 1
Healthy C 0 7 5 6 4 4
Healthy PC 4 12 1 1 1 3
become sources of disease at the end of 2 years (Table 8.4). However, from
the 2-year data alone, it is clear that under field conditions they will certainly
present a significant disease risk.
In another trunk treatment, pieces were cut to simulate shredding as a
clearing method, with and without poisoning prior to preparation. These were
either placed on the soil surface or buried at 20 cm deep. Both infected and
healthy trunk tissues were examined, with seedling baits used to detect BSR
in plots isolated by trenches. Both diseased and healthy shredded tissues can
give rise to disease after burial. Except in a single instance, superficially placed
tissues were not a disease hazard. In plots with buried tissues where disease
was recorded, sporophores of Ganoderma were produced on the soil surface.
Roots
The current recommendation for BSR sanitation procedure concentrates on a
1.5 m square centred on the point where the palm is planted. The assumption
has been that the remaining inter-space presents no serious disease hazard. In
a trial to examine this, areas between neighbouring diseased palms were each
divided into three equal parts and isolated by deep trenches. Bait seedlings
were then planted in each sector, as well as around the bases of the BSR-
affected palms. Similar sectors between apparently healthy palms were also
baited. In the BSR plots, Ganoderma fructifications developed on cut root
ends, signifying the presence of infected roots. The overall incidence of seedling
infection was low (4%) and was confined to the sectors closest to the diseased
palms, whereas 69% of bait seedlings planted around the main disease sources
became infected. No disease was recorded between healthy palms.
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Table 8.4. Comparison of basal stem rot (BSR) in bait seedlings around standing
BSR and healthy palms compared with stumps, 2 years after treatment.
BSR Nil 30 6 90 38
BSR P 40 10 95 34
Healthy Nil 20 3 20 6
Healthy P 30 7 40 9
P, Poisoned.
Discussion
It is apparent from these results that, when suitable disease sources are
present, oil-palm seedlings can be attacked by Ganoderma soon after planting.
Disease development and overt symptom appearance will depend on the size of
the palm when it becomes diseased, its continued growth vigour and the size
of the inoculum. Small seedlings close to large disease sources are killed
rapidly. Larger, rapidly growing plants are also affected, but frequently do
not die quickly. Numerous investigations have reported that many infected
palms continue to grow well, often for very long periods, before the internal
BSR lesion becomes so extensive that visible external symptoms develop. This
explains why so many cases of BSR occur long after planting and also after
obvious sources of primary infection have disappeared.
Once a few palms in a field are infected it has been considered that further
colonization of palms in the field is due to root-to-root contact by the palms or
mycelial spread. Both Singh (1991) and Hashim (1994) reported the disease as
occurring in patches or groups, which would support palm-to-palm infection,
but this view has been challenged recently by Miller et al. (1999). Studies of
somatic incompatibility and mtDNA profiling of isolates taken from many
adult palms within two oil-palm blocks (Miller et al., this volume) revealed con-
siderable variation between isolates, and led to the conclusion that isolates
occurred as numerous distinct genotypes, even within the same palm. Thus,
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110 J. Flood et al.
root masses can become significant sources of infection. Even where a field that
has been carefully cleaned of debris at replanting, as the new seedlings grow,
more and more root debris is produced. This will include a large amount of root
material from self-pruning (Hartley, 1988; Jourdan and Rey, 1996) and large
numbers of fine quaternary roots are present in the upper layers of the soil. The
hypothesis that this material could become the substrate for basidiospore
colonization requires further study.
The depth factor poses considerable problems from the practical viewpoint
of sanitation at the time of clearing for replanting. Breaking up deeply located
root masses requires deep tining, for which equipment is not always available.
If seedlings are planted at the same points of former BSR palms, there is a
distinct possibility that their roots will soon encounter infective sources of
Ganoderma, and thus as much of the diseased stump tissue as possible should be
removed. However, further baiting using seedlings showed that these potential
BSR sources were less of a disease hazard after 2 years. This means that their
importance could be expected to be very much reduced, or even negligible, if
new palms are planted as far as possible from the old planting points. Their
disease potential would have greatly diminished by the time the roots of the
new planting reach the hazard sources, provided the old stand had been
poisoned before felling. Alternatively, delayed planting could be a useful
method of disease avoidance.
Windrowed palm trunks represent another significant problem, and the
same considerations apply to the necessity for planting as far away as possible
from windrows. The lateral extent of root development during immaturity
reaches roughly the edge of the canopy, meaning that it should take 2–3 years
before reaching this particular disease source if planted at the furthest possible
distance. An important observation is that the period over which windrows
remain a disease hazard is greatly reduced when palms of the old stand are
poisoned by paraquat prior to felling, and this effect is further enhanced
when they are cut into sections and with a thick overgrowth of legume cover.
Where there has been no poisoning, the tissues remain a disease hazard for
years. In such situations older palms of the replant become infected, with overt
disease symptoms only appearing long after the original infection sources have
disappeared.
One solution is to shred palm tissues so that they do not become BSR
sources over long periods, which is already a common practice in Malaysia but
not in Sumatra. However, even this does not provide a total answer to the
problem. Occurrence of BSR in bait seedlings, arising from buried, shredded
diseased and healthy trunk segments, was limited, but illustrated that the
technique still contains a degree of disease risk. Disease arising from
superficially placed segments was very slight and unexpected. It was in some
ways remarkable that in such segments, buried or superficially placed, disease
occurred at all, since many attempts at artificial inoculation of seedlings in
polybags using such tissues have failed. The appearance of Ganoderma sporo-
phores on the soil surface above buried BSR sections indicated that a sufficient
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Spread of Ganoderma from Infective Sources in the Field 111
Acknowledgements
This chapter is published with the permission of P.T.P.P. London, Sumatra,
Indonesia. The considerable assistance of field staff in the execution of trials is
gratefully acknowledged. The authors would like to thank the Crop Protection
Programme (CPP) of the Department for International Development (DFID) for
funding some of the research reported here, which was administered through
NRI (RNRRS Project 6628).
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