Ann. appl. Biol.

(2002), 141:225-236
Printed in Great Britain 225

Integrating cultural control methods for tomato late blight (Phytophthora

infestans) in Uganda
By J TUMWINE 1, H D FRINKING 2 and M J JEGER3*
1
Kawanda Agricultural Research Institute, PO Box 7065, Kampala, Uganda
2
Laboratory of Phytopathology, Wageningen Agricultural University, PO Box 8025, 6700 EE Wageningen,
The Netherlands
3
Department of Agricultural Sciences, Imperial College at Wye, Wye, Ashford, Kent,
TN25 5AH, UK

(Accepted 5 September; Received 2 April 2002)

Summary

Cultural control measures against tomato late blight (Phytophthora infestans) were evaluated in six
field experiments over 3 years in Uganda. Each experiment included sanitation (removal of diseased
plant tissues), fungicide (mancozeb) application, and an untreated control, as standard treatments.
Late blight incidence and severity were greatly reduced by sanitation, without reducing the number of
healthy leaves; however, tomato growth and production were adversely affected. Fungicide treated
plants retained the highest numbers of flowers and attached fruits and gave the highest yields. Three
cultural practices were evaluated in repeated experiments for their effectiveness in alleviating the
adverse effects of sanitation. Tomatoes grown within plastic shelters early in the production cycle
were taller, and had more healthy leaves than those grown late. The numbers of diseased leaves and
disease severity were equally low in sanitation alone and plastic shelter/with sanitation treatments.
Flower and fruit production were significantly higher when tomatoes were grown under early shelters
with sanitation than with sanitation alone. Planting density was increased without significant effects
on late blight and tomato growth and production. Intercropping tomato with soybean (Glycine max) or
sesame (Sesamum indicum), with sanitation, limited late blight development, but taller intercrops
suppressed tomato growth and production. Integrated treatments (combining plastic shelters, a sesame
intercrop and high tomato planting density) were evaluated, with and without sanitation, against the
fungicide mancozeb. The mean numbers of healthy leaves in the integrated treatments were not
significantly less than with fungicide treatment. Late blight incidence and severity were higher in the
integrated plots without than with sanitation. The numbers of flowers and attached fruits were not
significantly less in integrated treatments than in fungicide treated plots, but tomato yield was highest
with fungicide treatment.

Key Words: Phytophthora infestans, tomato late blight, phytosanitation, cultural control, Uganda, plastic
shelters, planting density, integrated control

Introduction susceptible to late blight (Tumwine et al., 2002). Late

blight can be controlled by chemical means, although
Late blight (Phytophthora infestans (Mont.) de a less than optimal number of sprays is applied on
Bary), bacterial wilt (Pseudomonas solanacearum potato in Uganda, according to an analysis of GIS-
(Smith) Smith), early blight (Alternaria solani linked disease forecast models (Hijmans et al., 2000).
Sorauer), root knot nematodes (Meloidogyne spp.) and On tomato, fungicides, predominantly mancozeb
some virus diseases are major constraints to tomato (Dithane M-45), are used but are expensive, often
cultivation in Uganda, with late blight the most adulterated, have expired label dates or have lost
important disease (Tumwine et al., 2002) as elsewhere efficacy, and alternatives are urgently required. In the
in Africa (Fontem, 1995). In Uganda, host adaptation literature there has been little evaluation of non-
of P. infestans on potato and tomato is not determined chemical control of tomato late blight.
by the ability to cause disease, but by quantitative
differences in pathogenicity (Vega-Sánchez et al., Sanitation
2000). Although there has been some testing of local Sanitation, in the broad sense, refers to the exclusion
and exotic tomato germplasm in E. Africa (Mlungu of inoculum sources both within and outwith a crop.
et al., 1996) all cultivars grown in Uganda are This may be achieved by a range of different practices

*Corresponding Author E-mail: [email protected]

© 2002 Association of Applied Biologists

226 J TUMWINE ET AL.

ranging from basic crop hygiene to the separation from were improved (Hanada, 1988). Tomatoes grown in
alternate hosts. Field sanitation has been low plastic tunnels were less affected by late blight
recommended for the control of fungal diseases of than uncovered plants (Kumar & Srivastava, 1998).
perennial crops (Emebiri & Obiefuna, 1992), Optimal planting density is determined by a number
damping-off diseases (Gutierrez et al., 1997), and in of factors, including cultivar, soil fertility and moisture
combination with fungicides for fruit rots in and crop purpose. Host density affects disease
strawberry (Legard et al., 1997). Sanitation forms an incidence (Burdon & Chilvers, 1982). Depending on
integral part of the management of Botrytis in the disease, high crop density can reduce (Jakhar et
greenhouses (Hausbeck & Moorman, 1996). al., 1994), increase (Kostandi, 1992) or have no effect
Sanitation is recommended for late blight control in on disease levels (McEwen & Yeoman, 1990). High
tomato and potato in home gardens (Inglis et al., tomato plant density increased incidence of P.
1996). Excluding early P. infestans inoculum by field infestans, S. lycopersici and A. solani (Silva-Junior
sanitation and growing tomatoes far from potatoes et al., 1992). High tomato density resulted in increased
were recommended for late blight management in yields (Al-Maslamani & Suwwan, 1987) and less
commercial production (Sherf & Macnab, 1986; unmarketable yield. Tomatoes cultivated at high
Cohen, 1987). Destroying potato foliage 2 wks prior densities were protected effectively provided standard
to harvesting prevents tuber infection by P. infestans fungicide treatments were made (Jovancev, 1998).
(Cohen, 1987). Weekly removal of diseased leaves Intercropping of crop plants is common in the
controlled late blight in potatoes (Fontem, 1995). tropics with effects on pest population dynamics that
Sanitation delayed late blight development in potato minimise crop damage (Perrin & Phillips, 1978).
by about 8 days (Van der Plank, 1963). The removal Intercropping also ensures food security and
of diseased plants or tissues can in some cases be additional income to farmers. Intercropping tomatoes
compensated for by enhanced growth in tomato with onion or garlic reduced the levels of Bemisia
(Ricker & Riedel, 1993), although some deleterious tabaci, Myzus persicae and Phthorimaea operculella
effects on growth and productivity may also be (Afifi et al., 1990). Intercropping tomato with beans
expected. Sanitation has been practised for control of reduced the damage of tomatoes by Heliothis spp.,
bacterial disease of tomato (Cazelles, 1992). Spodoptera spp. and Liriomyza satiae (Rosset et al.,
In this paper a more limited use of sanitation is 1987) and pest incidence was low in tomato
adopted, referring only to the removal of diseased intercropped with maize (Picanco et al., 1996).
plant parts from within a crop. It is hypothesised Intercropping tomatoes with sorghum controlled
that sanitation during plant growth will reduce the whiteflies through effects on arthropod predators
incidence and severity of late blight, but will also (Gravena et al., 1984) and with cucumber or pepper
have adverse effects on tomato growth and delayed the development of Tomato yellow leaf curl
production. It is possible that by supplementing virus (Al-Musa, 1986). Intercropping can also give a
sanitation with other cultural practices, the need for reduction in fungal disease incidence and severity
sanitation will be reduced and the adverse effects (Boudreau & Mundt, 1992; Boudreau, 1993),
avoided. Cultural practices previously proposed although there are few reports on intercropping with
include the use of plastic shelters, high density tomatoes (Zavaleta-Mejia & Gomez, 1995), or with
planting, and intercropping. late blight (Sharaiha et al., 1989). Intercropping
tomato with cowpea reduced bacterial wilt
Cultural practices (Pseudomonas solanacearum) on tomato (Michel et
Plastic shelters have been used for various purposes al., 1997).
in tomato cultivation (Al-Maslamani & Suwwan,
1987; Randhawa, 1990; May, 1991; Porporato et al., Integrated cultural control
1993). Shelters enable more efficient use of fertiliser, Integrated control of late blight has been practised
increase yield and protect fruits from direct rain in potato production (Perley, 1994; Duvauchelle &
damage. Seedlings of tomato emerged and developed Dubois, 1998) with a range of promising late blight
faster in plastic tunnels than in the open (Mittra et al., control methods identified and utilised. Development
1990). Tomato seedlings raised under cover had better of durable resistant cultivars has been promoted as a
yields than those raised without protection major element in the control of late blight (Leach et
(Randhawa, 1990). The use of unheated plastic tunnels al., 1986; Mackay, 1996). Late blight resistant potato
proved economical in tomato and pepper production cultivars can be grown with fewer fungicide
(Gent, 1991). Tomatoes grown in plastic greenhouses applications (Fry, 1978; Mackay, 1996; Kankwatsa
were protected against frost, and had low disease et al., 2002). The integration of fungicides, resistant
incidences caused by A. solani and Septoria cultivars and the use of forecasting systems has been
lycopersici Speg. (Bhatnagar et al., 1990). Plastic claimed as the best strategy for potato late blight
shelters reduced tomato late blight infection, fungicide control (Stevenson, 1983; Sherf & Macnab, 1986;
sprays were minimal, and tomato yields and fruit size Gujer, 1991; Perley, 1994; Kirk, 1996). Integrated
 Cultural control of tomato late blight 227

methods for controlling late blight on tomato have promising cultural practices were integrated and
been evaluated in the USA (Bolkan, 1997; Bauske et evaluated for the control of tomato late blight,
al., 1998) but are less commonly practised. Although compared with the standard fungicide mancozeb.
there are difficulties in developing resistance and the
frequency of resistant lines remains low, disease General Procedures
control based on resistant tomato cultivars and Experimental designs
fungicides may counteract the threats posed by the The seven experiments were conducted sequentially
presence of both mating types A1 and A2 of P. over the years 1996-98 at Kawanda Agricultural
infestans. At present only A1 has been found in Research Institute, Uganda. In Expts 1-6 there were
Uganda (Vega-Sánchez et al., 2000; Tumwine et al., three standard treatments: sanitation, fungicide
2002). The introduction and promotion of resistant application with mancozeb (as Dithane M-45 80%
and acceptable tomato cultivars is not an immediate w/w wp Rohm & Haas) and untreated control, and
prospect for Ugandan farmers (Tumwine et al., 2002). four further treatments depending on the experiment
It is hypothesised that cultural practices can be (see Table 2). The treatments for Expt 7 (Table 3)
integrated, together with sanitation, to give control were determined after the completion of Expts 1-6.
of late blight equivalent to that provided by mancozeb, Experimental units were plots of 20 m2 (5 m ´ 4 m)
the most commonly used fungicide in Uganda. arranged in a randomised block design with five
blocks. Each plot was surrounded by a 3 m space.
Materials and Methods Procedures common to all experiments are first

Experimental programme
The effectiveness of sanitation in combination with Table 1. Dates of planting and harvesting periods in
other cultural practices for the control of tomato late sanitation experiments 1996-98. Expts 1-6 were pre-
designed with three standard treatments (sanitation,
blight was studied in Uganda during 1996-1998. In
fungicide and control) and other treatments based
six pre-designed field experiments the effectiveness on one of the three cultural practices (Table 2). For
of sanitation as a sole means of control for late blight each cultural practice there were two treatments I
was compared with routine application of mancozeb and II. The design of Expt 7 was designed based on
and an untreated control. Additionally, in each of the results from Expts 1-6
these experiments there were treatments based on
Expt Planting date Harvesting time Other treatments
the use of cultural practices in combination with
sanitation. For each of three cultural practices the 1 24 March 1996 May 1996 High density I
effe cts on late blight and tomato growth and 2 14 April 1996 May 1996 Intercropping I
production, compared with sanitation alone, were 3 28 October 1996 January 1997 Intercropping II
evaluated in two experiments (denoted I & II), giving 4 4 November 1996 February 1997 Plastic shelters I
six experiments in total (Table 1). The intention was 5 4 April 1997 July 1997 Plastic shelters II
to establish whether cultural practices could alleviate
6 10 October 1997 January 1998 High density II
the hypothesised adverse effects of stringently applied
sanitation. In a final experiment (Expt 7), the most 7 27 March 1998 June 1998 Integrated

Table 2. Treatments and linear contrasts used in experiments comparing the use of cultural practices with
sanitation with sanitation alone. A standard fungicide treatment and an untreated control at ‘normal’ planting
densities and without shelters/intercrops were included in all experiments
Cultural practice
Treatment Plastic shelter Planting density (per plot) / spacing (m) Intercrop
number* (Expts 4 & 5) (Expts 1 & 6) (Expts 2 & 3)
1 0-2 months 77 / 0.35 ´ 0.70 sorghum
2 1-3 months 110 / 0.35 ´ 0.50 sunflower
3 2-4 months 154 / 0.35 ´ 0.35 soybean
4 0-4 months 80 / 0.25 ´ 1.0 tomato + sesame
5 Sanitation alone 55 / 0.35 ´ 1.0 sanitation alone
* Linear 1 - 4 vs 5, All shelters vs sanitation 1 - 4 vs 5 All high density plantings vs 1 - 4 vs 5 All intercrops vs sanitation
contrasts alone; ssanitation alone (normal planting alone;
1 & 2 vs 5, Early shelter vs sanitation density). 1 & 2 vs 3 & 4 Tall vs short intercrops.
alone;
1 vs 3, Early shelter vs late shelter;
shelter vs no early shelter.
228 J TUMWINE ET AL.

Table 3. Treatments and linear contrasts used in the spreader plants were transplanted within the interstices
integrated control of tomato late blight experiment of the experimental plots and the edges (a total of 48
(Expt 7). All plots were planted at high density (96 plants) about 2 m distant from the corners of each
plants per plot) plot and inoculated in the evening 2 wk after
Treatment Description transplanting using a 1.5-litre hand sprayer.
1 Plastic shelter, sesame intercrop, sanitation
Field management
2 Plastic shelter, sesame intercrop, no sanitation
In sanitation treatments, plants were inspected every
3 Sanitation alone 2 days and diseased leaves, flowers, fruits, branches
4 Fungicide (mancozeb) and entire plants (though exceptional) were carefully
Linear contrasts used to compare treatment effects were: removed with scissors, placed in containers and
1 vs 2, effect of sanitation;1 vs 3, effect of integration; 1 vs 4
and 2 vs 4, effects relative to standard fungicide treatment disposed of away from the experimental field. In the
fungicide treatment, mancozeb was sprayed to run-
off at a dosage of 2.5 g litre-1 water at intervals of 7
described and then procedures specific to experiments days, using a 15-litre capacity knapsack sprayer. In
concerned with each of the cultural practices. case of insect pest attack, chlorpyrifos (as Dursban
400 g litre-1 ec Dow) at a dosage of 18.5 ml / 15 litres
Soil characteristics of water was applied to run-off. Weeding was done
Prior to establishment of the experimental plots, regularly by hand.
soil samples were analysed at the Faculty of
Agriculture, Makerere University. Based on the Assessments
results, the experimental field was partitioned into two Two weeks after transplanting, six plants (main
areas with 80, 200 and 60 kg ha-1 of nitrogen (N), stems) per plot were tagged for field assessments made
phosphorus (P2O5) and potassium (K 2O) respectively at 14-day intervals over periods ranging from 8-10
applied to one area; and 80, 100 and 40 kg ha-1 applied wk from first observation of disease. The variates
to the other area. All P2O5 was applied at planting recorded were plant height, numbers of healthy (in
with split application of N and K2O before and 7 wk Expts 3-7 only) and diseased leaves, disease severity
after transplanting tomatoes into experimental plots. (proportion of leaf area diseased) on the 5th and 9th
leaves, and numbers of flowers and attached fruits.
Field establishment Assessments of diseased fruits were not made. All
Tomato line MT55 (susceptible to late blight, variates were expressed as mean values for the
resistant to bacterial wilt) seedlings were raised in assessment period. Disease incidence (proportion of
nurseries for c. 4 wk before each experiment. plants diseased) was observed on 30 randomly chosen
Nurseries were protected from late blight infection plants per plot. At the end of the experiment, healthy
by spraying to run-off with mancozeb at the fruits from the plots were harvested and weighed.
recommended dosage of 2.5 g litre-l at intervals of 7 Disease incidence and severity were transformed to
days. Four-week-old tomato seedlings of about 15 angles to improve normality.
cm average height were transplanted into the plots at
a normal density range of 44-66 plants per plot, except Cultural practices
in Expts 1 and 6 where high planting density Six experiments (two experiments for each of the
treatments were included. The experimental fields three cultural practices) were carried out with the order
were enclosed within guard rows of maize planted 4 of experiments determined randomly (Table 1).
m from the outside plots. All experimental plots were Procedures for field establishment, inoculations,
mulched with locally available spear grass, Imperata management and disease assessments were standard
cylindrica. for all experiments. In each of the experiments the
effects of the cultural practice with sanitation were
Field inoculation compared with the sanitation alone using analysis of
Inoculum was prepared by incubating late blight variance and pre-determined linear contrasts defined
infected tomato leaves of cv. Moneymaker for 48 h according to the experiment.
at room temperature in 94 mm diameter Petri dishes
containing moist cotton wool. Original isolates were Plastic shelters
obtained from tomato in Uganda as described by Tomatoes were grown for different periods under
Tumwine et al. (2000), and characterised as A1 in plastic shelters (Table 2). Transparent plastic shelters
compatibility tests with known A1 and A2 isolates at (4 m long, 1.2 m wide and 1.5 m high) constructed
Wageningen University (Tumwine et al., 2002). from eucalyptus poles and transparent plastic
Sporangia were retrieved by washing with 500 ml of sheeting were positioned above the transplanted
water and adjusted to a concentration of 2.5 ´ 105 seedlings. Three such units were necessary to cover
sporangia ml -1 using a haemocytometer. Tomato each plot in the shelter treatments. At the top of the
 Cultural control of tomato late blight 229

tunnels there were five 3-cm diameter holes to allow Integrated cultural practices
ventilation and reduce resistance to wind. A total of Four treatments were included (Table 3) in a
66 tomato seedlings per plot, spaced at 0.9 m inter- randomised complete block design with five
row and 0.35 m intra-row distances, and 48 spreader replications. Experimental plots, their configuration,
plants, were transplanted into the experimental plots. and other procedures were as described in Expts 1-6.
Expt 1 was planted on 4 November 1996 and Expt 5 Plastic shelters constructed as previously were placed
on 4 April 1997. A digital thermohygrometer was in position above the transplanted tomatoes
used to measure morning, afternoon and evening throughout the experiment. An intercrop of sesame
shelter temperatures and relative humidity. Linear was planted in March 1998 at an intra-row spacing of
contrasts were designed to test whether the effects 10 cm. Tomato seedlings, including spreader plants,
of specified sheltering treatments with sanitation were transplanted between the sesame rows in two
differed significantly from sanitation alone (Table rows at spacings of 1.2 m inter-row and 0.25 m intra-
2). row, giving an intermediate high planting density of
96 plants per plot. Expt 7 was planted on 27 March
High planting density 1998 and harvested in June, 1998.
The spacings used and the corresponding densities
of tomato plants per m2 are shown in Table 2. The Statistical analysis
‘normal’ mid-range density of 55 seedlings per plot In all experiments treatments were compared by
was obtained by 1 m between and 0.35 within-row analysis of variance using the statistical package SAS
spacings. Expt 1 was planted on 24 March 1996 and version 6.
Expt 6 on 10 October 1997. One linear contrast was
made to test whether the effect of high planting density Results
combined with sanitation differed significantly from
sanitation at normal planting density (Table 2). Sanitation
In Expts 1-6 there were three standard treatments
Intercropping of sanitation, fungicide treatment and an untreated
The intercrop ping treatments (Table 2) were control. The effect of sanitation compared with
separated into tall (sorghum and sunflower) and short fungicide treatment and the untreated control across
(sesame and soybean) classes. Intercrops were experiments is first described, with the treatment
planted 3 wk before transplanting tomatoes at 120 means for each experiment regarded as replicates in
cm between rows and within-row spacings of 10 cm, the overall analysis (Table 4). Comparisons of
25 cm, 5 cm and 10 cm for sorghum, sunflower, sanitation with the fungicide treatment and the
soybean and sesame respectively. Four-week tomato control within experiments can be made from the
seedlings of about 15 cm average height were results presented for each cultural practice, but are
transplanted between the intercrop rows giving a plot not commented on further. The comparisons of
density of 0.44 seedlings/plot. Spacings for tomatoes interest within experiments are those between the
were 0.35 m intra-row and 1.2 m inter-row. Expt 2 cultural practice with sanitation and sanitation alone.
was planted on 14 April 1996, and Expt 3 on 28
October 1996. Linear contrasts were made to test the Tomato growth
effects of intercropping with sanitation against Across the six experiments fungicide treatment
sanitation alone, and whether the effects of tall consistently gave the tallest plants and sanitation the
intercrops were significantly different from short shortest (Table 4). Overall, plants in sanitation plots
intercrops (Table 2). were shorter than with the fungicide treatment (c. 8
cm, P < 0.01) and control (c. 5 cm, P < 0.05).

Table 4. Effect of sanitation on tomato growth, late blight incidence and severity, and production. Values are mean values for
Expts 1-6 in Table 1. In each experiment there were standard sanitation, fungicide and control treatments at normal planting
density and without shelters or intercrops
Disease Severity
Height Healthy Diseased incidence leaf 5 Flowers Fruits Yield
Treatment (cm) leaves leaves (angles) (angles) /main stem /main stem (kg plant-1)
Sanitation 33.7 4.33 0.57 28.2 4.8 3.70 1.38 0.21
Fungicide 42.2 6.68 2.76 38.9 15.8 6.92 2.83 0.89
Control 39.3 4.25 4.11 56.1 29.4 4.00 2.02 0.40
SED (df 10)a 2.06 0.480b 0.430 6.11 2.12 1.284 0.316 0.230
a
Equivalent to the standard error for the linear contrasts: sanitation vs fungicide; sanitation vs control
b
Based on 6 df as this variate was recorded in Expts 3-6 only
230 J TUMWINE ET AL.

Disease incidence and severity Table 5 shelter treatment is used as shorthand for
Fungicide treatment gave the greatest mean number shelter treatment with sanitation.
of healthy leaves on the main stem in the four
experiments in which this variate was recorded. Tomato growth
Overall, there were fewer healthy leaves (c. 1.2 per In Expt 4 with shelter treatment I, plants were
main stem) in the sanitation compared with the shorter in sanitation alone plots (P < 0.001) than in
fungicide treatment (P < 0.01) but there was no all shelters combined or in early shelters, a difference
difference from the control. The stringently applied of about 20 cm. Early sheltering (0-2 months) gave
sanitation resulted in fewer (P < 0.001) diseased leaves taller tomato plants (P < 0.001) than late sheltering
per main stem than the fungicide treatment (c. 2) and (2-4 months) or where there was no early shelter (P <
the control (c. 3). Overall, mean disease incidence 0.001). In Expt 5 with shelter treatment II, plants in
was lowest in the sanitation treatment, about half of sanitation alone plots were again shorter (P < 0.001)
that in the control, but was not significantly less than than in all shelters combined and in early shelters, a
in the fungicide treated plots. Mean severities on the difference of about 12 cm. Early shelters gave tomato
5th leaf (as on the 9th) were lower with sanitation plants taller (P < 0.001) than those in late shelters or
than with fungicide treatment or the control (P < where no early sheltering was applied (P < 0.001).
0.001).
Disease incidence and severity
Tomato production In Expt 4 with shelter treatment I, plants in sanitation
Fungicide treatment gave the highest numbers of alone plots had about two fewer healthy leaves per
flowers and attached fruits in all experiments. In Expts main stem (P < 0.001) over the period of assessment
1 and 2 fruit set and consequently mean numbers than plants in all shelters combined or in early shelters.
fruits/main stem and yield were very low. The mean Tomato plants in early shelters had more healthy
numbers of flowers (P < 0.05), attached fruits (P < leaves (P < 0.001) than where no early shelters were
0.001) and yield (P < 0.001) in the sanitation used. Numbers of diseased leaves remained low (<
treatments were lower than in the fungicide treated 0.5) and there were no significant differences between
plots but did not differ significantly from the control. shelter treatment and sanitation alone. In Expt 5 with
shelter treatment II, overall there were again about
Plastic shelters two more healthy leaves per main stem for the shelter
Expts 4 & 5 included shelter treatments I & II treatments than sanitation alone (P < 0.01), with about
(Table 1). The effects of the shelter combined with the same difference between early sheltered and late
sanitation compared with sanitation alone are shown sheltered plants (P < 0.01). Again numbers of diseased
in Table 5, in cases where significant differences leaves were low (< 0.3) and there were no significant
were found. For completeness mean values for the differences among the shelter treatments. Sheltering
fungicide treatment and control and SEDs are shown, either overall or early (P < 0.05) combined with
but the comparisons of interest are those given in sanitation reduced mean disease incidence compared
Table 2 with standard errors calculated according to with sanitation alone. In Expt 5 with shelter treatment
the specific contrast made. For brevity, in presenting II weather conditions were only conducive to late

Table 5. Effect of plastic shelters combined with sanitation on tomato growth, late blight incidence and severity,
and production. Values are means of five replicates in each of two shelter treatments (Expts 4 and 5)
Height Healthy Disease incidence Flowers/main Fruits/main Yield
Treatment (cm) leaves (angles) stem stem (kg plant-1)
I II I II I I II I II I II
Plastic
(0-2 months) 55.1 56.9 4.74 10.9 19.0 4.16 4.18 2.97 5.41 0.74 1.16
(1-3 months) 48.0 55.3 5.69 9.38 16.5 2.83 3.83 2.89 6.64 0.68 1.46
(2-4 months) 37.5 43.6 3.61 8.69 23.8 2.35 3.00 1.45 4.88 0.30 1.13
(0-4 months) 53.7 56.8 5.63 10.8 15.1 4.18 4.25 3.76 5.03 0.85 1.18
Sanitation only 35.5 43.2 3.42 8.37 23.6 2.11 3.18 1.14 4.32 0.23 0.78
Fungicide 50.4 45.8 7.14 11.3 40.9 3.72 3.18 3.78 5.52 1.98 1.56
Control 46.0 47.4 2.53 8.28 58.7 2.79 3.04 2.32 5.03 0.46 1.00
a
SED (df = 24) 3.35 3.39 0.550 0.640 2.65 0.502 0.432 0.532 0.856 0.212 0.234
a
Equivalent to the standard error for the linear contrasts: sanitation only vs fungicide; sanitation only vs control. Plastic shelter with
sanitation treatments were compared with the sanitation only treatment using linear contrasts (Table 2) and their corresponding
standard errors
 Cultural control of tomato late blight 231

blight late in the season and there were no significant main stem with sanitation, irrespective of the planting
differences in disease incidence between shelter density. In Expt 6 sanitation ensured few diseased
treatments and sanitation alone or among the shelter leaves and c. 3.5 healthy leaves on average per main
treatments. Disease severity, where sanitation was stem. Plot yields were c. 0.1 kg plant-1 where sanitation
practised, remained low in both experiments and there was practised, less than the mean for sanitation
were no significant differences between the shelter treatments over all experiments (Table 4).
treatments and sanitation alone, or between shelter
treatments. Intercropping
Expts 2 & 3 included intercropping treatments I
Tomato production & II (Table 1). The effects of intercropping combined
Flower (P < 0.01) and fruit (P < 0.001) production with sanitation compared with sanitation alone are
were greater in the shelter treatments in Expt 4 with shown in Table 6, in cases where significant
shelter treatment I, whatever the period compared, differences were found. For completeness, mean
than with sanitation practised alone. Early sheltering values for the fungicide treatment and control and
gave more flowers and fruits (P < 0.001) compared SEDs are shown, but the comparisons of interest are
with no early sheltering. The same trend was observed those given in Table 2 with standard errors calculated
for the yield of harvested tomatoes with increases in according to the specific contrasts made. For brevity,
yield (more than doubled) in shelter plots compared in presenting Table 6 intercrop treatment is used as
with sanitation alone (P < 0.05) and with early shorthand for intercrop treatment with sanitation.
sheltering compared with no early sheltering (P <
0.01). Flower production was greater in Expt 5 in Tomato and intercrop growth
treatments with early shelters compared with At all stages of crop development, and in both
sanitation alone (P < 0.05), late shelters (P < 0.05) experiments, the sunflower and sorghum intercrops
and no early shelters (P < 0.01). The only significant were taller than tomato plants with sesame and
difference in mean numbers of attached fruits (P < soybean intercrops of about the same height. Tomato
0.05) was between early shelters and sanitation alone. plant height was not significantly affected by any of
Harvested yield was greater (P < 0.05), about 50%, the intercrops in either year compared with sanitation
for the mean of all sheltered treatments compared with alone.
sanitation alone. Plot yields in the sanitation only
treatment were much higher in Expt 5. Disease incidence and severity
There were no significant differences in the mean
High planting density numbers of diseased leaves between the intercrop
Expts 1 & 6 included high density treatments I & II treatments and sanitation alone in Expt 2; on average
(Table 1). There were no significant differences in between 1 – 1.5 diseased leaves per main stem where
either year for any of the growth, disease or production sanitation was practised. In Expt 3, tomatoes
variates assessed when comparing the mean of high intercropped with sunflower and sorghum had fewer
planting densities with sanitation and sanitation alone healthy leaves (P < 0.05) than those intercropped with
using the specified linear contrast (Table 2). In Expt sesame and soybean. Tomatoes in sanitation alone
1, there were c. 1.5 diseased leaves on average per plots had more healthy leaves (P < 0.05) than the mean

Table 6. Effect of intercropping combined with sanitation on tomato growth, late blight incidence and severity,
and production. Values are means of five replicates in each of two intercropping treatments (Expts 2 and 3)
Intercrop Healthy leaves Flowers/main stem Fruits/main stem
II I II II
Intercrop
Sorghum 1.10 2.51 1.89 0.84
Sunflower 0.87 1.97 1.68 0.77
Soybean 1.20 4.42 2.87 1.28
Sesame 1.80 4.67 3.18 1.77
Sanitation only 1.82 3.97 2.80 1.31
Fungicide 3.27 13.4 5.47 3.44
Control 2.18 2.67 4.03 2.64
SED (df 24)a 0.268 1.320 0.218 0.278
a
Equivalent to the standard error for the linear contrasts: sanitation only vs fungicide; sanitation only vs control. Intercropping with
sanitation treatments were compared with the sanitation only treatment using linear contrasts (Table 2) and their corresponding
standard errors
232 J TUMWINE ET AL.

of the intercrop plots. There were no significant Tomato growth

differences in number of diseased leaves, disease Plants in integrated plots with sanitation were not
incidence or severity between the intercrop treatments significantly shorter than in integrated plots alone,
and sanitation alone; or between the tall and short nor different in height from fungicide treated plots.
intercrop treatments in either experiment. Disease They were, however, taller than plants in sanitation
incidence reached 100% at the end of Expt 2 for all alone plots (P < 0.001). Sanitation alone reduced
treatments. tomato growth but this effect was ameliorated by the
integrated treatment. Plants in integrated plots without
Tomato and intercrop production sanitation were taller than those in fungicide treated
Tomatoes intercropped with short plants in Expt 2 plots (P < 0.01).
had more flowers (P < 0.05) than those with tall
intercrops, but fruit set was sparse in all treatments Late blight incidence and severity
including fungicide treatment and yield data were There were about the same number of healthy
unreliable. Tomatoes intercropped with tall plants had leaves, seven to eight per main stem, on plants in
fewer flowers (P < 0.001) and fruits (P < 0.01) than integrated plots irrespective of whether sanitation was
those intercropped with short plants in Expt 3. There practised. There were no significant differences in
were no significant differences between the sanitation numbers of healthy leaves between integrated and
alone and intercrop treatments in flower and fruit fungicide treated plots. The integrated treatment with
production, or in crop yield. Intercrops provide sanitation gave more healthy leaves than sanitation
additional yields to the farmer, and reduce the risk of alone (P < 0.05); again the adverse effect of sanitation
crop failure by providing an alternative crop harvest was ameliorated by the integrated treatment. There
and additional income (Table 7). The average market were very few diseased leaves per main stem with
price of tomato in Uganda was 1.25 US$ per kg in sanitation. Plants in integrated plots without sanitation
1998, although this fluctuates with production levels. had considerably more diseased leaves than with
sanitation (P < 0.001). Plants in integrated plots with
Integrated cultural control sanitation had fewer diseased leaves than in the
One experiment (Expt 7) was designed following fungicide treated plots (P < 0.001), with no difference
the results of Expts 1-6 (Table 1). The use of plastic in the number of healthy leaves. There were no
shelters, planting at high density and intercropping significant differences in diseased leaves per main
with sesam e, were combined in an integrated stem between integrated plots without sanitation and
treatment and compared, with and without sanitation, fungicide treated plots. Disease incidence was always
with the standard fungicide treatment. The effects highest in the integrated treatment without sanitation,
of the integrated treatments are shown in Table 8, in and in the fungicide treatment plots and by the end of
cases where significant differences were found. the experiment approached 100%, whereas in plots

Table 7. Intercrop yield (kg ha-1) and their crop values (1998) from Expts 2 and 3
Intercrop Expt 2 Expt 3 Value (US$ kg-1) Remarks
Sorghum 449 1230 0.3 Drought tolerant, severely attacked by birds at maturity.
Sunflower 515 2304 0.6 Severely attacked by birds at germination, emergence and maturity.
Soybean 595 290 0.7 Severely attacked by birds at germination and emergence.
Sesame 210 192 0.7 Small seeded, requires extra care at planting by covering lightly.

Table 8. Effect of integrated cultural control with and without sanitation (Expt 7) on tomato growth, late blight
incidence and severity, and production. Linear contrasts were made as listed in Table 3
Height Healthy Diseased Severity leaf 5 Flowers Fruits Yield
Treatment (cm) leaves leaves (angles) /main stem /main stem (kg plant-1)
Integrated
+ sanitation 67.9 7.81 0.34 2.8 4.15 5.77 0.88
Integrated
Î sanitation 74.7 7.63 5.05 15.6 4.44 5.85 1.35
Sanitation alone 50.5 6.38 0.06 4.0 2.89 4.35 0.69
Fungicide 62.8 8.55 3.63 9.1 3.74 7.77 2.14
SED (df = 16) 3.42 0.630 0.810 3.09 0.404 0.656 0.344
 Cultural control of tomato late blight 233

where sanitation was practised (integrated or alone) physical act of removing leaves and also to the
incidence never exceeded 50%. Where sanitation was removal of healthy photosynthetic areas on diseased
practised, late blight severity was very low: less in leaves, i.e. diseased leaves may still be contributing
the integrated treatment with sanitation (P < 0.001) to yield assimilation. The adverse effect of sanitation
than with fungicide treatment; and higher in the was reflected in fewer flowers and attached fruits and
integrated treatment alone than with sanitation (P < harvested yield compared with fungicide treatment.
0.001) or with fungicide treatment (P < 0.05). Integrating sanitation with other control measures
such as fungicides (Legard et al., 1997) could possibly
Tomato production reduce the intensity and frequency of sanitation and
Both integrated treatments gave as many flowers thus any adverse effects. Fungicide treatment with
as the fungicide treatment (Table 8) and more than mancozeb, however, did not completely contain late
sanitation alone (P < 0.01). There were more attached blight epidemics in these studies.
fruits with fungicide treatment than either of the Although sanitation had an adverse effect on tomato
integrated treatments (P < 0.01). The integrated growth, development and production, it reduced the
treatment with sanitation treatment had more attached level of late blight disease. The intensity and frequency
fruits than sanitation alone (P < 0.05). There were no of sanitation required as a sole control means that the
differences between the integrated treatments in practice must be combined with other control practices
numbers of flowers, attached fruits or yield, although to avoid some of the adverse effects. Although
there was a consistent trend of reduction when combining sanitation with other cultural practices
sanitation was combined with integration. Plants in proved effective for stem rot in tobacco (Gutierrez et
both integrated treatments gave yields less (P < 0.01 al., 1997) such integration has not been tested
with sanitation; P < 0.05 without sanitation) than those adequately for foliar diseases. The use of resistant
from fungicide treated plots. Intercrop yields in the varieties could play an important role in integrated
integrated treatments were 140 kg ha-1 of sesame. control of tomato late blight by again reducing the
need for sanitation, but these are not currently
Discussion available in Uganda (Tumwine et al., 2002).

The aims of the experimental programme were Combining cultural practices with sanitation.
firstly to determine the effectiveness of sanitation The cultural practices used to supplement sanitation
as a control measure against late tomato blight as an improved late blight control and reduced to some
alternative to use of the commonly used fungicide extent adverse effects of sanitation practised alone
mancozeb; secondly to determine whether cultural under conditions of high disease pressure. In the two
practices could alleviate the need for sanitation and plastic shelter experiments, late sheltering combined
avoid adverse effects on the crop. It should be noted with sanitation maintained low late blight incidence
that in all experiments rainfall was sufficient and and severity, but resulted in reduced plant growth,
ambient temperatures within a suitable range for late and low numbers of flowers, attached fruits and yield.
blight development for at least part of the growing By contrast early sheltering combined with sanitation
period. had a positive effect on plant growth and maintained
relatively high numbers of healthy leaves on plants.
Sanitation as a sole control measure The use of plastic shelters early in the growth period
Sanitation significantly reduced late blight would exclude P. infestans sporangia (Hanada, 1988;
compared with fungicide treatment. The high numbers Bhatnagar et al., 1990) and effectively reduce late
of diseased leaves in fungicide treated (and untreated) blight development on tomatoes. The most
plots demonstrated the high potential for sporulation, encouraging aspect of the use of early shelters is the
dispersal and infection under favourable conditions. significantly improved tomato growth and production
However, fungicide treatment maintained the highest compared with sanitation alone. Although not
number of healthy leaves in each of the four formally assessed, there was also a general impression
experiments where this was recorded, which in terms of less insect damage in sheltered plots than those in
of the available photosynthetic area might have had the open. Fruits from sheltered plots were generally
significant consequences for yield. On average, smaller than those grown in open plots, suggesting
sanitation gave the same number of healthy leaves that possibly the sheltered plants received less water.
per plant as the control. Removal and destruction of The maximum temperatures recorded inside the
diseased tissues five times per week also reduced late shelters (often 10°C higher than those outside) were
blight in potato (Fontem, 1995), due presumably to well above the optimum range for P. infestans and
the reduction of inoculum and prevention of focus there was no evidence that the shelters created
establishment (Sherf & Macnab, 1986; Cohen, 1987). sufficiently humid conditions to enhance late blight
However sanitation used as a sole means of control development (Jebari, 1989).
retarded growth of tomato plants, possibly due to the Increased tomato densities, combined with
234 J TUMWINE ET AL.

sanitation, did not aggravate late blight as previously treatment. Sanitation alone, without the benefits of
reported (Silva-Junior et al., 1992). Flower and fruit the integrated practices, gave the shortest plants,
formation were about the same at the various tomato lowest numbers of healthy leaves, and reduced
crop densities when combined with sanitation, numbers of flowers and fruits, confirming the adverse
indicating that increased tomato density would be effects on plant growth and production in the first
compatible with other cultural controls for late blight. series of six experiments. The integrated practices
Late blight was reduced with intercropping alleviated the adverse effects of sanitation by reducing
combined with sanitation as reported previously for the intensity of leaf removal and gave reasonable
other diseases (Sharaiha et al., 1989; Michel et al., yields. The integrated treatment without sanitation
1997), although this was mostly due to the sanitation gave the most diseased leaves and also the tallest
practised. Tomatoes intercropped with sesame and plants possibly due to excessive etiolation caused by
soybean had more healthy leaves on the main stem the growth environment. Fruit yield was reasonably
than those in tall intercrops. The intercrop ped high in the integrated treatment without sanitation
tomatoes could have gained from the enhanced although fungicide treatment gave the highest yield
growing conditions noted for other polyculture of harvested fruits per plot. The tomatoes harvested
systems (Boudreau & Mundt, 1992; Boudreau, 1993). from the integrated plots appeared clean with few
Intercropped tomatoes were slightly shorter than blemishes, but were of smaller size, possibly due to
sesam e and soybean, and these may still have plants inside the shelters receiving insufficient water
provided a barrier to spore dispersal (Gretna, 1983). for yield assimilation. This problem could be
Intercropping also increases the distance between overcome by irrigation where necessary. The yield
susceptible tissues and decreases wind velocity data may also be misleading if some of the fruits were
responsible for spore release and dispersal (Boudreau, latently infected, but assessment of this at the time of
1993). Tomatoes with short intercrops had harvesting was not possible in this study.
significantly more flowers than with the tall intercrops
presumably because of shading effects at the intercrop Conclusions
spacing used. Row spacing between intercrops Integrated control of late blight in tomatoes is
influences plant competition for light and nutrients possible, as has been the case for potato (Gujer, 1991;
and inappropriate intercrops or spacings can suppress Perley, 1994; Mackay, 1996; Duvauchelle & Dubois,
growth and yield (Afifi et al., 1990). Soybean and 1998), and should be developed further, recognising
sesame, and possibly other crops of similar stature the constraints present for smallholder production in
and growth habit, are compatible for intercropping Uganda (Tumwine et al., 2002). The key conclusions
with tomatoes, help to reduce late blight levels and from this study are:
provide additional yield which could give additional 1) Sanitation alone, or combined with sheltering and
income to growers. Significantly better tomato yields intercropping, largely prevented late blight
were obtained with the compatible intercrops but epidemics, but sanitation alone adversely affected
further studies in Uganda are necessary to optimise tomato growth and production. Sanitation can be
yields and financial returns through a proper economic a component of integrated cultural control, but
analysis. sanitation thresholds to avoid adverse effects need
to be developed.
Integrated cultural control 2) Integrated methods of control combining plastic
The use of plastic shelters, intercropping with shelters and intercropping alleviated the adverse
sesame and planting at high density were combined effect of sanitation and promoted good tomato
as an integrated control treatment and compared, with growth and reasonable yields at high population
and without sanitation, with a standard fungicide density. Further work on optimising the
treatment of mancozeb. Late blight was reduced most compatibility of shelters and intercropping, and
where sanitation was practised alone or when economic analysis of the different production
combined with the integrated treatment and the systems, will be required to achieve integrated
disease was highest where there was no sanitation. control.
The fungicide treatment and the integrated treatment 3) Treatment with the fungicide mancozeb did not
without sanitation led to high disease levels prevent late blight epidemics but still achieved
throughout the experimental period. The integrated higher yields than controls based on sanitation
treatment with sanitation gave greater numbers of and cultural practices. Cultural control without
flowers and fruits than sanitation alone, which we sanitation is unlikely to achieve good disease
attribute to the beneficial effects of sheltering and control compared with fungicide treatment.
intercropping reducing the requirement for sanitation. Further research on the use of reduced fungicide
The integrated treatment, with or without sanitation, application and/or partially resistant cultivars in
gave tomato plants of about the same height and with combination with cultural practices merits further
the same number of healthy leaves as did fungicide consideration. Problems arise however in the
 Cultural control of tomato late blight 235

limited access to such cultivars and availability late blight. Phytopathology 68:1650-1655.
of appropriate fungicides. Gent M P N. 1991. High tunnels extend tomato and pepper
production. Bulletin of the Connecticut Agricultural
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