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South African Avocado Growers’ Association Yearbook 1987. 10:106-108
Proceedings of the First World Avocado Congress
JJ BEZUIDENHOUT, JM
DARVAS and JC TOERIEN
Westfalia Estate, PO Box
14, Duivelskloof 0835, RSA
Avocado root rot caused by Phytophthora cinnamomi Rands
was controlled by phosetyl-Al and phosphorous acid. Boric acid and potassium
aluminium sulphate injections improved tree condition in a five-year
trial, although no significant improvement could be detected in an 18-month trial. Trees
injected with potassium hydroxide responded positively and less roots were infected with P. cinnamomi. Trees
injected with hydrogen peroxide, hypophosphorous or dibasic-ammonium
phosphate did not improve significantly.
Root rot of
avocado caused by Phytophthora
cinnamomi Rands is the most damaging disease in South African avocado
orchards. The use of metalaxyl against this pathogen was the first significant step in the control of this disease and the next major breakthrough
was the application of phosetyl-Al
through stem injection (Darvas, Toerien & Milne, 1984) which
resulted in a commercially
acceptable control programme.
Although differences of opinion exist in
explaining the mode of action of phosetyl-Al,
it is generally accepted that
phosphorous acid is the active compound
(Fenn & Coffey, 1984). Besides phosphorous acid, a few inorganic
compounds, for example copper and sulphur, are applied in the
control of plant diseases.
In the present study, some of the results obtained with
phosetyl-Al and phosphorous acid are reviewed and the tests with different inorganic compounds
(potassium hydroxide, hydrogen
peroxide, boric acid, the potash of alum, dibasic-ammonium phosphate and hypophosphorous acid) are presented in the control of root rot.
MATERIALS AND METHODS
Chemicals
- Water solutions of 10 per cent ai phosetyl-Al, 3 per cent
phosphorous
acid, 3 per cent potassium hydroxide, 3 per cent boric acid, 5 per cent hypophosphorous
acid, 10
per cent
dibasic-ammonium phosphate, 10 per cent aluminium potassium sulphate (alum), or 5 per cent
hydrogen peroxide.
Zinc was evaluated in 10 per cent chelate form (Farmchem), or as 5 per cent zinc sulphate.
Tree
injection - Trees were injected as
described by Darvas et al, 1984. With potassium aluminium sulphate, double the normal number of
syringes was used per tree.
Assay for
P.
cinnamomi - Feeder root tips were collected after surface
sterilisation with 0,1 per cent mercuric chloride and a triple wash in sterilised water, plated out on PARP-medium (Kannwishcher & Mitchell, 1978). Percentage infection was taken as the number of P. cinnamomi colonies
grown out from the roots per total number of root tips plated out.
Evaluation of tree condition - Trees were rated on a scale
from 0 (healthy) to 10 (dead), as described by Darvas et al, 1984.
Statistical analysis - One-way analysis of variance with Duncan's multiple range test (Steel
& Torrie, 1960) was used to compare the treatments according to the number of
feeder roots infected with P. cinnamomi.
Wilcoxon's signed rank test (Steel
& Torrie, 1960) was applied in
analysing changes in tree condition.
An improvement in tree condition could be noted within one
year after three injections with either phosetyl-Al or phosphorous acid (Figure 1). The tree usually responded faster with phosphorous acid than with phosetyl-Al.
No loss in the efficacy of these compounds was. detected where trees were treated by injection for seven years, as illustrated in Figure 2.
Changes in tree condition after six injections during an
18-month trial with different
inorganic compounds, are compared in Figure
3. Tree condition improved significantly after injection of potassium
hydroxide, phosetyl-Al or phosphorous acid.
The condition of the control trees and trees injected with alum or hydrogen peroxide deteriorated. No
significant improvement was noted with the
application of boric acid, dibasic-ammonium
phosphate or hypophosphorous acid.
In a five-year trial, the tree condition improved slightly after
injecting either potassium hydroxide or hypophosphorous acid (Figure 4). The condition of trees treated with alum, boric acid or phosphorous acid improved significantly.
In trees treated with phosetyl-Al, potassium hydroxide, alum, hypophos-phorous acid or phosphorous acid, the percentage of feeder roots infected with P. cinnamomi
was significantly lower than in the
control trees (Figure 5).
Trees injected with phosphorous acid
supplemented with boric acid and zinc chelate, responded significantly
better than trees injected with phosphorous
acid only (Figure 6). A similar effect was observed when phosetyl-Al was injected with boric acid and zinc sulphate.
Indications are that a synergistic effect exists
between phosphorous acid and zinc chelate in both the tree condition
and the incidence of P, cinnamomi isolated
from the roots. Zinc sulphate reacts with phosphorous acid to form insoluble
zinc phosphite, which is not suitable for
injections. Aqueous solutions of phosetyl-AI can be mixed with zinc sulphate without any detrimental
effect (Darvas, 1984).
Although the application of phosetyl-Al lead to a new dimension in orchard management in which
Phytophthora root rot is no longer a
limiting factor in production, the danger
that resistance could develop cannot be
ignored. Phosetyl-Al has been injected
at Westfalia Estate since 1980 and no
relapse in root rot control has been
detected. Indications are that resistance can develop with regular application (Vegh, Leroux, Le Berre & Lanen, 1985), making the need
for an alternative compound for root
rot control more urgent.
These results indicate that the injection of some inorganic
compounds
(potassium
hydroxide, alum and boric acid)
can control P. cinnamomi. This control
is not as drastic as that obtained with
phosetyl-Al or phosphorous acid. An
improvement of one unit in tree rating
per year can be observed with phosetyl-Al and even more so with phosphorous acid. With potassium hydroxide
it is 0,7 units per year with very diseased
trees (rating 7,5) and 0,1 units per year for trees with an initial rating of two. In the long term (five years) trees responded very well to the injection of boric acid or alum, while during the 18-month period, the change detected
was not significant.
Hydrogen peroxide was included in the experiments, due to
its reducing capacity analogous to the phosphite ion. In contrast with potassium hydroxide, no
significant improvement in tree condition was observed with the injection of
hydrogen peroxide. Therefore no simple relation between the chemical reducing
capacity of a compound and its potential in controlling root rot exists.
Hypophosphorous acid would theoretically be oxidised
to phosphite after injection. In these experiments, hypophosphorous acid was not able to improve tree condition.
No phosphite could be detected in the leaves or roots of trees injected with hypophosphorous acid (Bezuidenhout,
Darvas & Kotze, unpublished).
Injections with dibasic-ammonium phosphate did not improve tree condition and there was even a
tendency of increased root rot, According
to Darvas (personal communication), phosphoric acid was also not able to control this disease.
Hypothetically potassium hydroxide, aluma and boric
acid could act in the tree as follows:
Essential
nutrients - Avocado soils in Tzaneen are low in available zinc
and boron.
In experiments where phosphorous acid was injected with boric acid and zinc chelate, the authors observed a
much better response in comparison to phosphorous acid only.
Boron has been found to confer resistance to powdery
mildew in wheat (Schutte, 1967) and Fusarium oxysporum was more severe on
boron deficient
flax than on boron adequate plants (Keane & Sackston, 1970)
Our results suggest the existence of a
synergistic effect between zinc and phosphorous
acid on the disease. This agrees with
the observation made by Darvas (1984) on
the more rapid improvement in tree
condition when phosetyl-Al is
supplemented with zinc sulphate.
There are some reports of beneficial
effects of added zinc increasing host
resistance (Singh &
Aggarwal, 1979), but there are also publications showing the opposite effect (Graham, 1983). Welsh, Webb & Loneragan (1982) found that zinc
deficient roots lost membrane
integrity with a corresponding leakage
of soluble organic substates into
the environment, which may attract
motile spores.
In the experiments the two potassium compounds, potassium
hydroxide and potash alum, improved tree condition. Effects of potassium on diseases
are often
reported in relation to fungal root rots such as Phytophthora
and Pythium. In these cases, yield responses to potassium appear to be
due to compounding
effects of concurrent increases in growth of
the host
and decreases
in the levels of infection (Graham, 1983). In this context the adjustment of
phosphorous acid by potassium hydroxide in some of the root rot formulations available in Australia, is
significant.
Plant
defence - A number of inorganic compounds stimulate phyto-alexin production
(Bailey & Mansfield, 1982) and it can be expected that injection of potentially
phytotoxic compounds, such as boric
acid, may induce phyto-alexin synthesis. Preliminary observations by Labuschagne (personal communication)
indicate that the production of water
soluble antifungal compounds are
stimulated by the injection of phosphorous acid.
Direct
effect - Boric acid is a classical fungistatic compound with a wide spectrum of activity against microorganisms. Whether the concentration of
boric acid is of such magnitude at the infection
site that it will inhibit P. cinnamomi, remains
to be investigated.
Should the mode of action of potassium hydroxide, alum
and boric acid resemble that of phosphorous acid, there is a high probability that cross-resistance could occur
against these compounds. This possibility should be investigated further.
REFERENCES
1
Bailey. JA & Mansfield, JW, 1982. Phytoalexins. Blackie: London.
2
Darvas, JM, 1984. Zinc supplemented to avocado trees in conjunction with root rot control injections, S Afr Avocado Growers Assoc Yrb, 7, 79
3
Darvas, JM, Toerien, JC & Milne, DL, 1984 Control of avocado root
rot by trunk injection with fosetyl-Al. Plant disease, 68, 691-693.
4
Fenn, ME & Coffey. MD, 1984. Studies on the in vitro and in vivo antifungal activity of fosetyl-Al and phosphorous
acid. Phytopathology, 74, 606-611.
5
Graham, RD, 1983 Effect of nutrient stress on susceptibility of
plants to disease with particular reference to the trace elements Advances Botanical Research,
10, 222-276
6
Kannwischer, ME & Mitchell, DJ, 1978. The influence of a
fungicide on the epidemiology of black shank of tobacco. Phylopathology,
68,
1760-1765.
7
Keane, EM, & Sackston, WE, 1970. Canadian
Journal of Plant Science, 50, 415-422.
8
Schutte, KH, 1967. Plant soil, 27, 450-452.
9
Singh, P & Aggarwal, RK, 1979. Indian
Journal of Agricultural Science, 49, 459-462.
10 Steel, RGD &
Torrie, JH, 1960. Principles and procedures of statistics. McGraw-Hill Book Company, Inc, New York.
11
Vegh, I, Leroux, P, Le Berre, A
& Lanen, C, 1985. Detection sur Chamaecyparis lawsoniana
'Ellwoodii' d'une souche de Phytophthora cinnamomi Rands resistante au phosethyl-Al. PHM - Revue Horticole, 262, 19-21.
12 Welch, RM, Webb, MJ & Loneragan, JF, 1982. Proceedings 9th International Plant Nutrition Conference, 2, 710-715.




