Australasian Plant Pathol.
(2012) 41:59–68
DOI 10.1007/s13313-011-0090-6
Phosphonate applied as a pre-plant dip controls
Phytophthora cinnamomi root and heart rot in susceptible
pineapple hybrids
J. M. Anderson & K. G. Pegg & C. Scott & A. Drenth
Received: 4 May 2011 / Accepted: 5 September 2011 / Published online: 6 October 2011
# Australasian Plant Pathology Society Inc. 2011
Abstract The effectiveness of pre-plant dips of crowns in
potassium phosphonate and phosphorous acid was investi-
gated in a systematic manner to develop an effective
strategy for the control of root and heart rot diseases
caused by Phytophthora cinnamomi in the pineapple
hybrids ‘MD2’ and ‘73-50’ and cultivar Smooth Cayenne.
Our results clearly indicate that a high volume spray at
planting was much less effective when compared to a pre-
plant dip. ‘Smooth Cayenne’ was found to be more resistant
to heart rot than ‘MD2’ and ‘73-50’, and ‘Smooth Cayenne’
to be more responsive to treatment with potassium
phosphonate. Based on cumulative heart rot incidence over
time ‘MD2’ was more susceptible to heart rot than ‘73-50’
and was more responsive to an application of phosphorous
acid. The highest levels of phosphonate in roots were
reached one month after planting and levels declined during
the next two months. Pre-plant dipping of crowns prior to
planting is highly effective to control root and heart rot in
the first few months but is not sufficient to maintain health
of the mother plant root system up until plant crop harvest
when weather conditions continue to favour infection.
J. M. Anderson (*) : K. G. Pegg
Agri-Science Queensland, Department of Employment,
Economic Development and Innovation,
Ecosciences Precinct, 41 Boggo Rd,
Dutton Park, QLD 4102, Australia
e-mail: Jay.Anderson@deedi.qld.gov.au
C. Scott
11 Galleon Court,
Taroomball 4703, Australia
A. Drenth
Centre for Plant Science, University of Queensland,
Brisbane, QLD, Australia
Keywords Ananas comosus . Phytophthora cinnamomi .
Root and heart rot . Phosphonate . Metalaxyl
Introduction
Pineapple (Ananas comosus), an edible member of the
Bromeliaceae which originated in South America, has been
grown in Australia since 1838 when German missionaries
in Brisbane introduced plants of the fresh fruit cultivar
Queen from India. In 1858 plants of the canning cultivar
Smooth Cayenne were imported from Kew gardens, the
original stock having come from French Guiana (Collins
1960). Commercial processing of ‘Smooth Cayenne’ started
in Queensland in the 1920s. Many growers are now
changing from ‘Smooth Cayenne’ to grow new pineapple
hybrids for the fresh fruit market and it is expected that in a
few years the volume grown for fresh fruit will exceed that
for processing. Hybrids currently grown include ‘MD-2’
(developed by Del Monte Fresh Produce Hawaii Inc.) and
‘73-50’ (Pineapple Research Institute, Hawaii).
The damaging effect of root and heart rot disease of
pineapple in Queensland, found to be caused by Phytoph-
thora cinnamomi (Simmonds 1929), was first observed at
Nundah (Brisbane) in 1887 (Tryon 1905) as an undeter-
mined fungal disease dependent on wet soil conditions.
Tryon commented that the disease had probably occurred in
southeast Queensland some fifteen years earlier in 1872. To
minimize losses by improving soil drainage he astutely
suggested planting “along the summit of an elevation
caused by throwing the furrows in ploughing.” Despite
many years of investigation there is still no completely
satisfactory method for controlling the disease. ‘MD-2’ has
been reported as more sensitive to Phytophthora than
‘Smooth Cayenne’ (Chan et al. 2003). The root rot phase
60
caused by Phytophthora cinnamomi reduces plant growth
and yield and in subtropical Queensland may result in the
complete loss of the ratoon crop. It is an established
principle that ratoon crop yield in pineapple is determined
by the health of the mother plant root system at plant crop
harvest (Malézieux et al. 2003). Heart rot results in reduced
plant densities due to plant mortality. As well as P.
cinnamomi, heart rot in the more tropical regions of
Queensland can also be caused by P. nicotianae (Pegg
and Anderson 2009).
Serious losses caused by P. cinnamomi have occurred
recently in plantings of the new hybrids, particularly in
‘MD-2’, due to high rainfall, greater host susceptibility,
reduction in bed height and a move towards using increased
plant densities. Leaf and canopy transpiration rates in
pineapple are significantly lower than in other crops and
in high density plantings this gives rise to a build up of
water in the topsoil, even in well drained soils, which
accentuates the Phytophthora problem. The low transpira-
tion rate is due to the thick cuticle, the water storage
capacity of the leaf tissue and the presence of few stomata
which are sunken and in addition are located below
multicellular trichomes (Malézieux et al. 2003).
Pineapple has many other residual epiphytic features
which include root initials in lower leaf bases which
produce the axillary root system which winds around the
lower stem and does not reach the soil. These axillary roots
absorb water and nutrients that run down the leaves. They
can become infected by P. cinnamomi if infested soil lodges
in the leaf bases. The adventitious soil root system, which
originates from the boundary between the cortex and stele
in the lower stem, is extremely susceptible to root
pathogens such as P. cinnamomi, as well as pests and
anoxia when it first emerges (Rohrbach and Schmitt 2003).
If these young vigorously growing roots are killed up to the
stem they do not regenerate. If older major roots are
damaged new lateral roots may form behind the damaged
main root tips. Root growth commences soon after planting
and there is steady continuous growth until flowering when
root growth ceases until after plant crop harvest (Black
1962).
Measures to reduce the impact of P. cinnamomi in
pineapple include planting in double rows on uncompacted,
well drained and aerated raised beds. Soil applications of
elemental sulphur also reduce disease if the pH is depressed
to or below 3.8 (Pegg 1977). At this pH germination of
chlamydospores, the primary inoculum source, is inhibited.
In field experiments it was occasionally observed that
sulphur failed to completely control heart rot even though
root rot control was excellent. Most heart rot caused by P.
cinnamomi is an extension of root rot where infection
progresses from the root to the stem apex, but in these field
experiments infection occurred through the leaf bases. Even
J.M. Anderson et al.
though the soil pH remained below 4.0, the soil which had
accumulated in leaf bases always showed a pH greater than
5.5. This rise in pH was hypothesised to be due to the
exudation of plant nutrients (R.N. Allen and K.G. Pegg
unpublished data). Chlamydospores had apparently sur-
vived in the soil in the leaf bases, then germinated and
infected the axillary roots when the pH was more
favourable. Sulphur treatment of soil does have limitations
in that it can have an adverse effect on plant nutrition. Low
pH limits availability of important nutrient elements such as
calcium, magnesium and potassium (Pegg 1977).
These limitations with sulphur led to the evaluation of
new fungicides for root and heart rot management. Allen et
al. (1980) found that high volume sprays with metalaxyl or
fosetyl-Al were effective for disease control in ‘Smooth
Cayenne’ and the utilisation of these chemicals has become
an established industry practice in Queensland. Soil
populations of P. cinnamomi in Queensland pineapple
fields have not developed resistance to metalaxyl (Soo et
al. 1998) and populations of microorganisms which rapidly
degrade metalaxyl and reduce its persistence and effective-
ness apparently do not occur in high enough population
numbers in low pH pineapple soils. There has been a
decline in phosphonate sensitivity in the P. cinnamomi
population in pineapple soils due to prolonged use. A study
by Soo et al. (1998) found that the range of effective
concentrations to limit the growth of the pathogen by half
(EC50) for P. cinnamomi isolates obtained from pineapple
fields was 7 ppm to 358 ppm with a mean value of 99 ppm.
Base line sensitivity data for isolates not exposed to
phosphonate gave a mean value of 22 ppm. The loss of
sensitivity after 18 years did not differ from the reduction
after two years of potassium phosphonate applications (Soo
et al. 1998).
Rohrbach and Schenck (1985) demonstrated that pre-
plant dips containing metalaxyl, fosetyl-Al or phosphorous
acid controlled heart rot caused by P. nicotianae in ‘Smooth
Cayenne’. They found that phosphorous acid gave compa-
rable control to fosetyl-Al. Phosphorous acid also con-
trolled root rot caused by P. cinnamomi in their glasshouse
trials. They clearly showed that high volume sprays of
fosetyl-Al (4.48 kg in 2805 L/ha) to newly planted material
not yet actively growing were less effective than pre-
plant dip applications. As Queensland growers did not
have dipping facilities and potassium phosphonate was
compatible with the fruit ripening agent ethaphon a
single preharvest spray to fruit crowns of ‘Smooth
Cayenne’ was evaluated for root and heart rot control
(de Boer and Pegg 1990). Sprays of potassium phospho-
nate (0.1 to 0.6% w/v) provided two years protection,
increased plant weights by 40% and increased plant crop
fruit weight (mean 1.5 kg) compared with untreated
controls (mean 0.65 kg).
Control of root and heart rot in pineapple using phosphonate
In a glasshouse experiment where potassium phosphonate
sprays were applied to one month old ‘Smooth Cayenne’
plants, Drenth et al. (2001) showed that a low phosphonate
level (3 to 8 μg/g) in root tips resulted in a lower root rot
rating following inoculation with low inoculum levels of P.
cinnamomi isolates with a range of EC50 values (13 to
302 ppm). At high inoculum levels the root rot rating was
reduced where the isolate had an EC50 value of 9.9 ppm but
not when the isolates had values of 298 and 302 ppm. Field
trials to corroborate these results were not conducted.
With increased susceptibility in the currently grown
hybrids and concern about fruit residues from preharvest
sprays, pre-plant dips in potassium phosphonate were again
considered. The distribution of phosphonate in plants is
source/sink related and it accumulates in the organ with the
greatest sink strength at the time of application (Whiley 1994).
The stem of the pineapple crown has a starch reserve of
some 20% dry matter and after planting the dominant
carbohydrate sinks are the young roots and leaves. In
pineapple there is a period of 4 to 8 weeks when roots will
have resource priority over leaves (Black 1962). As phospho-
nate will be translocated with the carbohydrates in the stem,
pre-plant dips should give rise to phosphonate levels effective
for disease control in young vigorously growing roots.
Because of the serious impact P. cinnamomi has on the
production of highly susceptible pineapple hybrids our
overall objective was to develop an effective and sustain-
able way to control root rot and heart rot in pineapples.
Specifically we sought to address in a temporal order the
following questions; (1) determine the most effective rate
and method for applying potassium phosphonate or
phosphorous acid to protect pineapple plants, (2) study
the accumulation and persistence of phosphonate in
pineapple roots, (3) assess the relative susceptibility of the
hybrids MD2 and 73-50 and cultivar Smooth Cayenne
based on the cumulative incidence of heart rot, (4)
determine if there has been any further loss of phosphonate
sensitivity in P. cinnamomi populations from extensive and
continuous use of the chemical and (5) demonstrate to
growers the most effective and environmentally innocuous
method of applying phosphonate to pineapples. The
outcome of this research will enable the development and
implementation of effective and environmentally sustain-
able disease control methods for the pineapple industry.
Materials and methods
Phosphonate uptake in ‘Smooth Cayenne’ and ‘MD2’
in the glasshouse
To determine the most effective rate and method of
application for the uptake of phosphonate applied as a
61
pre-plant dip in pineapple a concentration profile over time
was conducted using fresh crowns (tops) from fruit of
‘Smooth Cayenne’ which were dipped for two minutes in
potassium phosphonate solutions (0.1% and 0.2% w/v Fos-ject
200®, U.I.M. Agrochemicals (Aust.) Pty.Ltd.) and allowed to
dry. After two days half the crowns from each treatment were
rinsed for two minutes under running water to remove surface
deposits of potassium phosphonate to simulate heavy rain after
planting. They were then planted in 1 L pots in a pasteurised
peat-sand mix and plants were harvested one and three months
later and the white root tips and tops (leaves plus stem) were
collected to determine the level of phosphonate through
chemical analysis.
Phosphonate concentrations in the plant material were
determined using the method of Hargreaves and Ruddle
(1990). Root samples were washed to remove and soil and
then roots, leaves and stem were dried and ground to a fine
powder. Phosphonate was extracted from samples using
dilute sulphuric acid. Phosphonate was reacted with diazo-
methane and the dimethylphosphite formed was injected
into a gas chromatograph equipped with a glass chromato-
graphic column (Carbowax 20 M, Agilent Technologies)
and a flame photometric detector. Dimethylphosphite was
detected as a peak on the chromatogram and the concen-
tration determined by comparison with a known standard.
This method was fast, specific, sensitive (<0.1 μg/g fresh
weight (f.w.)) and reproducible (Hargreaves and Ruddle
1990).
Fresh crowns from fruit of the hybrid ‘MD2’ were dipped
for two minutes in potassium phosphonate (0.5% w/v–pH 6.5,
Agri-Fos 600®, Agrichem, Australia ) or technical grade
phosphorous acid (0.5% v/v–pH 1.4) and allowed to
dry. After two days half the crowns from each treatment
were rinsed under running water as above. Crowns were
then planted in pasteurised potting mix in 1 L pots and grown
at 29°C day/20°C night temperatures for six weeks. Plants were
then harvested and roots were analysed for phosphonate by
SGS Agritech, Toowoomba, Australia (sensitivity level of the
commercial analytical method ≤5.0 μg/g). This method was
used as the more sensitive method was not commercially
available at the time this experiment was conducted.
Effective rate of phosphonate for a pre-plant dip
To determine the rate of phosphonate which effectively
controls Phytophthora root rot fresh crowns from fruit of
‘MD2’ were dipped for two minutes in a range of
potassium phosphonate concentrations (0.1%, 0.25%,
0.5%, and 1.0% w/v, Agri-Fos 600®, Agrichem, Australia).
Crowns were then planted into 1 L pots containing a sandy
loam collected from beneath root rot affected pineapple
plants. One month after planting they were subjected to
cyclic water logging for two weeks (three days wet
62
followed by four days dry by placing pots in saucers for the
wet cycle and inverting the saucers during the dry period).
Plants were then harvested and roots visually assessed for the
percentage of healthy root tips in major and lateral roots.
‘MD2’ crowns dipped in water were used as a control.
Application method for phosphonate under field conditions
To determine the most effective way of applying potassium
phosphonate to pineapples, crowns of ‘MD2’ were planted in
September 2009 into a poorly drained site on a pineapple farm
at Glasshouse Mountain (26°53′ S 152°57′ E) where there had
recently been high plant mortality due to P. cinnamomi heart
rot. The soil was a red earth developed over sandstone and
had a chlamydospore population density of 1–2 g−1 of soil.
Crowns were planted in a randomised block design and
received one of three treatments viz. a pre-plant dip in 0.5%
(w/v) potassium phosphonate (Agri-Fos 600®, Agrichem,
Australia) for two minutes, an immediate post-plant drench
(100 mL of 0.5% potassium phosphonate poured into the
heart of each crown) and untreated control. There were 10
crowns per treatment. After 11 weeks plants were assessed
for the incidence of heart rot and five plants of each
treatment were harvested and roots removed for analysis of
level of phosphonate as described for the previous experi-
ment. The remaining plants were again assessed for heart rot
nine weeks later and were then harvested for analysis of their
phosphonate levels in the roots and leaves.
Rates of potassium phosphonate required to control heart rot
in ‘73-50’, ‘MD2’ and ‘Smooth Cayenne’ under field
conditions
To determine the most effective concentration of potassium
phosphonate required in a preharvest dip to control heart rot
in ‘MD2’, ‘73-50’ and ‘Smooth Cayenne’, a field trial was
planted in the same location as the field trial described
above in March 2010.
Crowns of ‘Smooth Cayenne’, ‘MD2’ and ‘73-50’, were
dipped in various concentrations of potassium phosphonate (0,
0.1, 0.25, 0.5 and 1.0% w/v, Agri-Fos 600®, Agrichem,
Australia) and planted in a field site heavily infested with P.
cinnamomi four weeks later. The layout was a 3×5 randomised
block design with five replications per treatment. Each trial plot
consisted of four data plants of each cultivar. Counts of plants
which had developed heart rot were made at regular intervals
with a final assessment conducted 14 weeks after planting.
Validation of pre-plant phosphorous acid dip to control
Phytophthora under field conditions
To examine the effect of phosphorous acid and metalaxyl
on the development of heart root in ‘MD2’ and ‘73-5’ a
J.M. Anderson et al.
field trial was planted in February 2009 at the same location
as the other field trials. As the availability of planting
material was limited, different planting material was used
for each variety; crowns were used for ‘MD2’ and slips for
‘73-50’. Each treatment was replicated twice in a rando-
mised block design with 30 plants of each hybrid per plot.
There were three treatments; untreated control, pre-plant
dip in phosphorous acid (0.5% v/v Agri-Fos 600®,
Agrichem, Australia) and the commonly used industry
practice of metalaxyl incorporated into the soil (500 g a.i. /ha).
The incidence of heart rot was recorded as percentage of dead
plants out of 30 plants per plot seven times from 10th June
2009 to 13th January 2010.
Percentage heart rot, both untransformed and with an
angular transformation applied, was analysed using a
repeated measures analysis of variance. The analysis
assessed the effects of hybrid, treatment and time and all
their interactions. The repeated measures analysis adjusts
for the correlation between times caused by the repeated
measures using the method of Greenhouse and Geisser
(1959).
Sensitivity to phosphonate of P. cinnamomi isolates
from the field trial location
Since the field trials were conducted over a number of years
and to be able to better compare the data with previous
research findings we sought to determine the sensitivity of
P. cinnamomi isolates from the field trial location to
phosphonate. A composite root and soil sample was
collected from beneath treated plants in the first field trial
and was baited using three day old New Zealand Blue
Lupin radicles (Chee and Newhook 1965). Infected lupin
roots were surface sterilised in 70% ethanol and cut
sections blotted dry and placed onto P10VP media (Tsao
and Ocana 1969) amended with 50 ppm hymexazol
(Tachigaren). Four P. cinnamomi isolates were subcultured
onto water agar (15 g agar/L Difco Agar Technical) and a
hyphal tip from each isolate was subcultured onto corn
meal agar (CMA, BBL Corn Meal Agar) (17 g CMA/L).
Phosphonate sensitivity tests were conducted on CMA
amended with the following concentrations of potassium
phosphonate (pH 6.2): 0 ppm, 1 ppm, 5 ppm, 10 ppm,
50 ppm, 100 ppm, 500 ppm and 1000 ppm. Plugs of four
day old cultures on CMA were used to inoculate each test
plate. There were three replications for each isolate.
Colony diameter measurements (subtracting 0.5 cm for
the plug) were used as an indication of growth at each
concentration. The growth at the various concentrations
was used to generate the EC50 for each isolate. EC50
values were calculated with the Half-Maximal Response
curve for estimating drug responsiveness in STATISTICA
(Version 4.5).
Control of root and heart rot in pineapple using phosphonate 63

Results Application method for phosphonate under field conditions

Phosphonate uptake in ‘Smooth Cayenne’ and ‘MD2’
in the glasshouse
For ‘Smooth Cayenne’ the 0.2% concentration of potassium
phosphonate provided the highest root levels after one and
three months but they were not significantly different to
those produced by the 0.1% concentration (Table 1). The
stem and leaf levels were significantly different between the
0.1% potassium phosphonate (unwashed) and the 0.2%
potassium phosphonate (unwashed) three months after
treatment. With both rates there was a considerable decline
in the phosphonate concentration level in roots after three
months growth in the glasshouse which coincided with the
appearance of new leaves. Rinsing crowns under running
water two days after treatment to simulate heavy rain after
planting, removed about 40% of the phosphonate from the
crowns treated with the 0.2% level and significantly
reduced root concentrations at one and three months.
For the ‘MD2’ crowns potassium phosphonate gave a
significantly higher level of phosphonate in the roots
compared to phosphorous acid and this level was reduced by
>50% if treated crowns were washed under running water to
remove surface deposits (Table 2). Neither treatment was
phytotoxic, as evidenced by the absence of leaf tip burn and
necrosis of white basal leaf tissue (data not shown).
Effective rate of phosphonate for a pre-plant dip
The 0.5% treatment of dipping crowns for two minutes in
potassium phosphonate was very effective in producing
healthy root tips and preventing heart rot and significantly
better than all other treatments (Table 3). The 1.0% and
0.25% dip treatments were as effective for heart rot control
but plants had less healthy white root tips.
The pre-plant dip treatment (Table 4, Fig. 1) resulted in no
losses due to heart rot and gave a concentration of
14.8 ppm in the root tips after 11 weeks. The high volume
post-plant treatment was less effective; growth was reduced
(Fig. 1) and two plants developed heart rot. Phosphonate
levels in the roots were 5 ppm or less after 20 weeks. Nine
of the 10 untreated control plants died.
Rates of potassium phosphonate required to control heart
rot in ‘73-50’, ‘MD2’ and ‘Smooth Cayenne’ under field
conditions
Significant cultivar effects were apparent in this study (P<
0.001) (Table 5). ‘Smooth Cayenne’ was more resistant to
heart rot than hybrids ‘MD2’ and ‘73-50’. Also ‘Smooth
Cayenne’ responded significantly (P=0.03) to a lower level
of potassium phosphonate (0.1%) than the hybrids (0.25%
for ‘73-50’; 0.5% for ‘MD2’). The 0.5% and 1.0% pre-
plant dip rates were the most effective treatments when all
cultivars are considered.
Validation of pre-plant phosphorous acid dip to control
Phytophthora under field conditions
The pre-plant dip in phosphorous acid resulted in signifi-
cantly less heart rot in both hybrids compared to the
untreated control, however ‘MD2’ had significantly less
heart rot than ‘73-50’ by the end of the recording period
(Fig. 2).
The pre-plant soil incorporation of metalaxyl was
ineffective for both hybrids and was no better than the
control. ‘MD2’ was significantly more susceptible than ‘73-
50’ in both the control and the metalaxyl incorporation
treatment from the third recording time.

Table 1 Mean phosphonate concentrations (ppm dry weight) in crowns (stem+leaves) and roots of ‘Smooth Cayenne’ one and three months after
a pre-plant dip of potassium phosphonate. Data were analysed using ANOVA (n=3)

Treatment Crown pre-plant Mean phosphonate concentration of tissue (ppm)

Post-plant (1 month) Post-plant (3 months)

Stem and leaves Roots Stem and leaves Roots

Control (unwashed) 7.9 da 8.6 d 4.1 c 8.2 c 1.8 d

Control (washed) 10.9 d 7.9 d 9.8 bc 7.6 c 2.6 cd
Potassium phosphonate 0.1% (unwashed) 77.2 bc 28.0 ab 22.35 ab 17.6 bc 5.75 ab
Potassium phosphonate 0.1% (washed) 58.6 c 14.4 cd 9.2 bc 23.3 b 6.1 ab
Potassium phosphonate 0.2% (unwashed) 137.1 a 33.6 a 33.6 a 36.9 a 8.0 a
Potassium phosphonate 0.2% (washed) 82.4 b 22.8 bc 16.4 bc 28.5 ab 4.4 bc
a
means within a column followed by the same letter are not significantly different at P=0.05
64 J.M. Anderson et al.

Table 2 Mean phosphonate

concentration in roots of ‘MD2’ Treatment Mean phosphonate concentration in roots
six weeks after crowns were (μg/g f.w.) six weeks after planting
dipped in potassium phosphonate
or phosphorous acid. Data was Control—no treatment applied 6.5 da
analysed using ANOVA (n=5) 0.5% phosphorous acid 59.8 b
0.5% phosphorous acid+rinse after 2 days 41.0 c
a
means followed by the same 0.5% potassium phosphonate 97.5 a
letter are not significantly 0.5% potassium phosphonate+rinse after 2 days 43.6 bc
different at P=0.05

Sensitivity to phosphonate of P. cinnamomi isolates the crop cycle. A high volume post-plant foliar spray was
from the field trial location shown to be much less effective in controlling root and
heart rot in pineapple when compared to a single pre-plant
The EC50 values for four P. cinnamomi isolates recovered dip. A major advantage of the pre-plant dip treatment is the
from this site were 93.3, 64.1, 35.5 and 56.4 ppm. protection afforded to emerging roots when crowns or slips
are planted into P. cinnamomi infested soil. Potassium
phosphonate applied prior to root emergence makes a
Discussion significant difference as this is a critical period for the
establishment of a good root system as there is no
Our experiments clearly demonstrate that dipping pineapple regeneration of these young new roots if they are infected
planting material in 0.5% potassium phosphonate is highly and colonised to the stem.
effective for preventing Phytophthora heart rot and delaying Due to the low transpiration rate of pineapples root rot
root rot for an extended period in fresh fruit pineapple symptoms are slow to express visible above ground
hybrids. Our data also demonstrates that the pineapple symptoms in contrast to heart rot which occurs mainly in
hybrids ‘MD2’ and ‘73-50’ are more susceptible to heart rot the first few months after planting. Most heart rot is an
than ‘Smooth Cayenne’ and do not respond to a pre-plant extension of root rot with P. cinnamomi progressing up the
dip in 0.1% potassium phosphonate which gives effective stem to its apex. This is in contrast to heart rot caused by
control in ‘Smooth Cayenne’. A 0.25% pre-plant dip was Phytophthora nicotianae where infection mainly occurs
required to give effective control in ‘73-50’ and 0.5% was through the leaf bases (Rohrbach and Schmitt 1994). P.
required for ‘MD2’. It is well documented that the nicotianae only causes limited root disease in the pineapple
effectiveness of phosphonate to control Phytophthora cultivar ‘Smooth Cayenne’ and where heart rot occurs plant
depends on the sensitivity of the pathogen to phosphonate mortality is compensated for by increased exposure to
and the capacity of the defence responses in the host (Guest sunlight in adjacent plants and a subsequent increase in fruit
et al. 1995). size (Rohrbach and Johnson 2003). This does not occur to
Cumulative heart rot data in the pre-plant dip validation the same extent in fields with heart rot caused by P.
field trial shows that ‘MD2’ was more responsive to a cinnamomi because root rot in surviving plants will still
phosphorous acid pre-plant dip than ‘73-50’, a hybrid with result in reduced growth and fruit size.
a much less developed root system over the total length of When the roots of treated and control plants of both
hybrids were visually assessed for root health in the
Table 3 Effect of pre-plant dips in potassium phosphonate on validation of a pre-plant phosphorous acid dip trial after
percentage healthy root tips and heart rot incidence of ‘MD2’ planted 13 months growth (data not presented), it was found that all
in soil infested with P. cinnamomi. Data were analysed using ANOVA root systems were unhealthy. Soils had become quite
(n=7)
anaerobic and it is likely that root death was due to a
Treatment Percentage Percentage combination of anoxia and P. cinnamomi. Pineapple roots
healthy root tips heart rot affected by waterlogging may be more susceptible to
infection by P. cinnamomi as zoospores are attracted by
Control—no treatment applied 17.1 da 42.9 a
ethanol, a product of anaerobically respiring roots (Allen
0.1% potassium phosphonate 39.3 c 14.3 ab
and Newhook 1973). Despite this, anchorage was adequate
0.25% potassium phosphonate 60.0 b 0.0 b
in the treated plants undoubtedly due to the establishment
0.5% potassium phosphonate 80.0 a 0.0 b
of a healthy root system in the months following planting.
1.0% potassium phosphonate 59.4 b 0.0 b
The consequence of an unhealthy root system at 13 months
a
means within a column followed by the same letter are not significantly would be the production of an acceptable plant crop but a
different at P=0.05 failed ratoon crop as the success of a ratoon crop depends
Control of root and heart rot in pineapple using phosphonate
Table 4 Heart rot incidence and phosphonate levels in ‘MD2’ 11 and 20 weeks post planting after a pre-plant dip or high volume drench with
potassium phosphonate
Treatment Heart rot incidence
11 weeks
Pre-plant dip 0.5% Potassium phosphonate 0/10
Post-plant drench 100 mL 0.5% Potassium phosphonate 1/10
Untreated control 5/10
a
n—number of samples used to determine average
b
n.d.—not determined
on how well root rot is controlled in the plant crop
(Malézieux et al. 2003). It appears that once roots lose
resource priority and export phosphonate to the leaves root
rot control using potassium phosphonate or phosphorous
acid is diminished and the root system becomes susceptible
to attack by P. cinnamomi. Thus, it is clearly evident that
dipping crowns at planting time is not sufficient to protect
roots up until plant crop harvest and drench applications of
chemicals may be needed once leaves have resource
priority.
The incorporation of metalaxyl into the soil before
planting is a common industry practice. In this study it
failed to control heart rot. Heavy rain fell immediately after
planting and due to its high water solubility metalaxyl may
have leached down the soil profile away from the emerging
roots. Sharom and Edgington (1982) showed that 87% of
applied metalaxyl was leached from a 20 cm column of
sandy soil in the presence of 30 cm of simulated rainfall.
Metalaxyl has been used as a high volume spray continu-
ously in many pineapple fields for some 30 years. It is still
effective and there is no evidence to suggest that P.
cinnamomi populations are less sensitive to the chemical
(Soo et al. 1998). The continuing efficacy of multiple
applications of metalaxyl to control Phytophthora root rot
in pineapple may be because populations of microorgan-
isms which degrade metalaxyl (Al-Sa’di et al. 2008) do not
build up to high population levels in low pH (pH 4.5–5.6)
pineapple soils. Another possible explanation is that
Fig. 1 Twenty-week-old ‘MD2’
plants growing in P. cinnamomi
infested soil. Untreated control
plants have been killed. The
plants subjected to the pre-plant
dip do not show any visible
signs of disease while the
growth of the plants treated with
a post-plant drench show re-
tarded growth compared to the
dip treated plants
65
Mean phosphonate concentration (ppm f.w.) (na)
20 weeks Roots 11 weeks Roots 20 weeks Leaves 20 weeks
0/5 14.8 (5) 5.6 (5) 76.2 (5)
1/5 <5 (4) <5 (4) 52.0 (5)
4/5 5.7 (3) n.d.b 30.2 (1)
enhanced biodegradation of metalaxyl does occur but with
the method of application used by growers the pineapple
roots take up metalaxyl before it has time to degrade which
may in some soils be as short as one to two weeks (Davison
and McKay 1999). For pineapple metalaxyl is applied in
high volume sprays (2000–5000 L/ha containing 500 g a.i./ha)
to ensure that the chemical reaches the soil. The long tapering
leaves of the pineapple plant form a funnel which channels the
spray directly to the base of the plant and down into the root
zone.
Our results indicate that there was a strong accumulation
of phosphonate in roots four weeks after planting treated
crowns, followed by a sharp decline in concentration after
three months, which coincided with the appearance of new
leaves. In the field phosphonate could no longer be detected
in the roots of five month old plants. This suggests that
phosphonate in the roots is remobilized and translocated to
the leaves. Thus the weak sink strength of the roots at that
time provides only limited opportunities to increase their
phosphonate levels. This is in marked contrast to avocado
where root flushes alternate with shoot flushes and
following injections phosphonate concentrations in the
feeder roots peak after two months then decline slightly
but remain relatively constant for at least one year (Whiley
1994). It is surprising that in this study we were unable to
detect phosphonate in older pineapple roots. Roots of
pineapple, in comparison to avocado feeder roots, are very
close to the phosphonate reservoir in the leaves and stem.
66 J.M. Anderson et al.

Table 5 Effect of different rates

of potassium phosphonate on Treatment Percentage mortality due to heart rot
the percentage mortality of
‘73-50’, ‘MD2’ and ‘Smooth Cultivar
Cayenne’ due to heart rot 73-50 32 aa
MD2 45 a
Smooth Cayenne 17 b
Potassium phosphonate concentration (%)
0 68.33 a
0.1 45 b
0.25 26.67 c
0.5 6.67 d
1 10 cd
Cultivar x Potassium phosphonate concentration (%)
73-50 0 80 ab
73-50 0.1 40 cde
73-50 0.25 20 def
73-50 0.5 0f
73-50 1 20 def
MD2 0 70 abc
MD2 0.1 85 a
MD2 0.25 50 bcd
MD2 0.5 10 ef
MD2 1 10 ef
Smooth Cayenne 0 55 abc
Smooth Cayenne 0.1 10 ef
a
Means followed by the same Smooth Cayenne 0.25 10 ef
letter in each group of treat- Smooth Cayenne 0.5 10 ef
ments are not significantly Smooth Cayenne 1 0f
different at P=0.05

Although older roots do not have resource priority they of phosphonate to the white root tips would be expected.
grow continuously and steadily up until flowering (Black This may be important as only very low levels of
1962) and some photosynthate partitioning and movement phosphonate are reported to induce phosphate deficiency
in Phytophthora and reduce its pathogenicity and inhibit
70
sporangial production (Daniel and Guest 2006). The
inability to detect phosphonate in older roots may reflect
60
the limitations of the commercial analytical method used in
m ality
y

50 these studies (detection limit 5 ppm).

e morta

40 High volume foliar applications when applied prior to

P centtage

30
root emergence were limited in their effectiveness and did
not give measureable increase in phosphonate levels in the
Perc

20
10
0 axillary root system which does not grow into the soil, and
100 120 140 160 180 200 220 240 260 280 300 320 340 360
Days after planting
Fig. 2 Percentage mortality (angular transformed) of ‘MD2’ and ‘73-
50’ field trial. Treatments were: ‘MD2’ untreated control ( ),
‘MD2’ pre-plant dip in phosphorous acid ( ), ‘MD2’ metalaxyl
incorporated into the soil pre-plant ( ), ‘73-50’ untreated control
( ), ‘73-50’ pre-plant dip in phosphorous acid ( ) and ‘73-50’
metalaxyl incorporated into the soil pre-plant ( ). The six means
within each date have been compared (LSD (P=0.05)=12.5)
roots. We postulate that this may be because some of the
phosphonate applied by this method is taken up by the
the remainder leaches down the soil profile where it is not
available to emerging adventitious roots. It is thus an
ineffective method of application and also environmentally
undesirable as the chemical is leached into soil and water
systems. Leaching of phosphonate through the soil profile
has occurred following soil drenching of avocado trees (A.W.
Whiley unpublished data). Soil applications of potassium
phosphonate for Phytophthora root rot control have proved
Control of root and heart rot in pineapple using phosphonate
ineffective in established avocado trees (C.Kaiser and A.W.
Whiley unpublished data 1997). The lack of phosphonate
persistence in the roots and the consequent necessity for
frequently reapplying the chemical at high rates made soil
application uneconomical. This may also be the situation in
pineapple.
Our phosphonate sensitivity study shows that the P.
cinnamomi isolates from our field trial do not differ in
sensitivity levels from those reported by Soo et al. (1998).
Phosphonate chemicals have been used on the farm where
the experiments were conducted for thirty years and
despite this phosphonate is still highly effective in the
current field trials depending on application method.
Phosphonate has been found to select for the least
sensitive isolates from the naturally occurring diversity
within P. cinnamomi populations in Australian avocado
orchards following years of application by trunk injections
(M. Weinert, pers. comm., Brisbane, 1997). As in
pineapple the sensitivity shift was in the order of ten fold
and not one thousand fold which occurs with the
emergence of resistance as typically is the case for single
mode of action fungicides (Hewitt 1998). It appears that as
long as an effective concentration can be maintained in the
roots to prevent rapid colonisation of the root and to
induce defence, disease control will be achieved despite
the sensitivity shift in P. cinnamomi populations.
This study has demonstrated that a pre-plant dip
treatment of crowns in potassium phosphonate is extremely
effective for controlling Phytophthora root and heart rot in
susceptible pineapple hybrids. We assume that a defence
response plays a role in the practical efficacy of the dip
treatment but do not know what level of phosphonate is
required in the roots to induce resistance to P. cinnamomi.
Whether there is any downward systemic movement of
phosphonate from leaves and stem to roots when roots are
no longer the priority sink is unknown. This will require
studies using an analytical method with a low detection
limit. We need to determine precisely when roots loose
resource priority and retranslocate phosphonate to leaves
and stem and find if the roots then become susceptible to
infection due to depletion of available phosphonate. In the
current study the site was poorly drained and plants were
frequently subjected to anoxia and besides P. cinnamomi
low oxygen levels may also have killed or damaged roots.
On free-draining sites the pre-plant treatment may be
sufficient to produce a healthy root system up until
flowering and consequently an acceptable ratoon crop.
When conditions are more conducive to disease develop-
ment chemicals such as metalaxyl may need to be applied
as foliar drenches. Studies are in progress to evaluate
combining other defence activators such as acibenzolar-S-
methyl or potassium silicate with phosphonate to prolong
resistance in pineapple roots.
67
Acknowledgements We thank Murray Pike, Glasshouse Mountain,
for making land available for field trials and Dr Vivienne Doogan for
the statistical analysis of data from the field experiments. This
research was originally funded by Rural Industries Research and
Development Corporation, Golden Circle Pty Ltd and more recently
by Horticulture Australia Ltd. The fresh fruit marketing companies
Piñata and Tropical Pines funded the phosphonate analyses.
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