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Stella Melugin Coakley
Department of Botany and Plant Pathology, Oregon State University, Corvallis,
Oregon 97331; e-mail: coakleys@bcc.orst.edu
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Harald Scherm
Department of Plant Pathology, University of Georgia, Athens, Georgia 30602;
e-mail: scherm@uga.edu
Sukumar Chakraborty
CSIRO Tropical Agriculture, CRC for Tropical Plant Pathology, University of
Queensland, Queensland 4072 Australia; e-mail: sukumar.chakraborty@tag.csiro.au
INTRODUCTION
Global climate change is a major topic of discussion within both scientific and po-
litical forums. By mid-1998, evidence was mounting that 1998 would set a record
high for global mean temperature, with 1997 holding the current record. Combined
0066-4286/99/0901-0399$08.00 399
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ocean and land temperatures were 0.25 C warmer from January to May 1998 than
previously recorded (154). September 1998 was the hottest on record (104 years)
in the United States, with a mean of 20.6 C; the earlier record of 20.2 C was set in
1931 (11). Increasingly, scientists are convinced that human activity, primarily
in the form of increased emissions of carbon dioxide (CO2) and other greenhouse
gases (predominantly methane and nitrous oxide), is a major contributor to the
warming trend. Indeed, the Second Assessment Report of the Intergovernmental
Panel on Climate Change (IPCC), which was edited in 1995 (59), concluded that
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(i) greenhouse gas concentrations have increased and will continue to do so given
their long life in the atmosphere; (ii) climate has changed over the past century in
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response to increasing atmospheric CO2, with the global mean surface air temper-
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ature increasing between 0.3 and 0.6 C; (iii) anthropogenic aerosols such as sulfur
dioxide tend to produce negative radiative forcing (i.e. a cooling trend) but are
relatively short-lived; and (iv) climate model simulations using future emission
scenarios of greenhouse gases and aerosols suggest an increase in global mean
temperature between 1 and 3.5 C by the year 2100 (67). Although the IPCC re-
port represents the broad consensus among many scientists from a wide range of
disciplines, alternative interpretations of the evidence presented in the report do
exist (61, 138).
The IPCC report was the basis for discussion at the Kyoto Conference in
December 1997 that led to a climate treaty ratified by more than 150 countries.
The United States signed the treaty in November 1998 but ratification by Congress
is not expected within the next two to three years. The Kyoto Protocol could result
in reductions in six greenhouse gases including CO2, methane, and nitrous oxide.
It calls for an average 5.2% reduction (from 1990 levels) of CO2 emissions by
38 industrialized nations by 2012, although increasing emissions from developing
countries could offset these reductions (85). Curbing of CO2 emissions is a very
charged political issue in the United States (73), in part because of the position
that any reduction in fossil fuel consumption would be damaging to the economy.
Nevertheless, scientists optimistically provide a road map and recommendations
of the technology necessary to achieve a reduction to below 1990 CO2 levels
(102, 115).
Is global warming an absolute? As recently as 1994, Time magazine carried
a feature article The Ice Age Cometh? (74), describing the January blizzard that
resulted in record low temperatures in dozens of cities in the United States. This
extreme expression of the weather is not inconsistent with the notion that cli-
mate change would lead to increased climate variability and more frequent or
more severe climatic extremes (39, 55, 92, 113, 162). Underlying all discussions
is the expectation that as climate changes, so does the human response to it. With
this in mind, there have been numerous attempts to predict the potential impacts of
climate change on human enterprises, including agriculture (1, 38, 101, 117, 118).
Despite the important role of plant pathogens in limiting agricultural production
and food supply (100), there has been limited research to assess the potential im-
pacts of climate change on plant diseases (22, 24, 26, 28). For example, a recent
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monograph entitled Climate Change and the Global Harvest: Potential Impacts
of the Greenhouse Effect on Agriculture (117) devoted only two pages (out of 324
total) to plant diseases. In contrast, potential impacts on infectious diseases of
humans have received considerably more attention, including a monograph pub-
lished by the World Health Organization (91) and a stand-alone chapter in the
1995 IPCC report (90). Suggestions in these publications that a warmer climate
will bring increases of human disease have been challenged as premature and
with little basis in fact, based on examination of historical patterns and the general
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absence of research in this area (147). It is with this example that we enter into
a review of what appears probable, what may be possible, and what is unlikely in
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Mahlman (84) commented on and updated the summary of IPCCs 1995 report.
He classified the relative certainty of change into several categories. Virtually
certain facts include that atmospheric concentrations of greenhouse gases would
continue to rise, largely caused by human activities such as burning of fossil fuel
and changes in land use. These gases, which may remain in the atmosphere from
a decade to centuries, act to heat the planet because of absorption and re-radiation
of infrared radiation. Changes in other radiatively active substances (e.g. sulfur
aerosols) and increased cloudiness caused by greater evaporation in a warmer
climate may offset some of the greenhouse effect. Another virtually certain fact
is that earths surface has warmed about 0.5 0.2 C during the past century. The
expected rate of increase is now at 0.1 C per decade (69).
Under virtually certain projections (99% likely to happen), Mahlman (84)
included the forecast that the stratosphere would continue to cool as CO2 levels
rise and that global mean concentrations of water vapor in the lower troposphere
would increase (approximately 6% per 1 C warming).
A very probable projection would have a greater than 90% chance of being
correct; under this category, Mahlman (84) predicted that a doubling of atmo-
spheric CO2 (from a current concentration of 360 ppm) would result in a warming
between 1.5 and 4.5 C. The rate of evaporation would increase in a warmer cli-
mate, which would lead to an increase in global precipitation of 2 0.5% per 1 C
warming. Higher latitudes in the Northern Hemisphere are expected to experience
above-average increases in both temperature and precipitation.
Mahlmans final category (probable projections, which have a greater than
two thirds chance of occurring) included the forecast that there would be decreases
in soil moisture because of increased temperatures, although this could be offset by
simultaneously increased precipitation. Further, changes in mean climate would
probably be accompanied by changes in the frequency and magnitude of climate
extremes; globally, this would include an increased probability of warm events
and decreased probability of cold events.
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What is much less certain is the magnitude of climate change and its impacts on
biological and ecological processes and on human enterprises.
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IMPACTS ON CROPS
Climate change will influence the geographical distribution and growth of plant
species around the world. The magnitude of these impacts would vary depending
on the species involved and their growth patterns (e.g. annuals vs perennials;
agricultural crops vs natural vegetation). Natural vegetation would be affected
by factors such as competition, migration, and recovery from disturbances, and
therefore new combinations of species are likely to occur (41). In managed systems
(agriculture and forestry), any effects of climate change have important socio-
economic implications for the countries involved, hence considerable efforts have
been directed toward agricultural impact assessment (1, 38, 101, 117, 118). How
a particular agroecological region will fare under climate change will depend
largely on societys ability to use available knowledge, technology, and financial
resources.
Nicholls (98) analyzed historical trends in Australian wheat yields and estimated
that recent climate changes were responsible for as much as 30 to 50% of the
observed yield increase, with most of the variation explained by an increase in
minimum temperature. Goudriaan & Zadoks (44) summarized a large body of
evidence suggesting that elevated CO2 would result in increased photosynthesis
and water use efficiency. This would lead to yield increases in most crops in most
production conditions. Idso & Idso (60) reviewed hundreds of CO2 enrichment
studies done over ten years and reported that in most cases, there was evidence
of growth-enhancing effects. IPCCs 1995 report (112) predicted that a doubling
in CO2 could increase yields of several major crops by an average of 30%. In
contrast, recent data taken in field-scale experiments with elevated CO2 suggested
only an 8 to 12% wheat yield increase under optimal growing conditions (45).
Further, evidence is now accumulating that crop production would be affected
differentially depending on latitude. Although increases in yields are expected at
mid and high latitudes, there may be decreases at lower latitudes where food needs
are greatest (112, 117).
Any direct yield gains caused by increased CO2 or climate change could be off-
set partly or entirely by losses caused by phytophagous insects, plant pathogens,
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IMPACTS ON PLANT PATHOSYSTEMS
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The focus of this review is on how climate change manifested by elevated CO2,
increased temperature, and changes in precipitation may affect plant diseases and
their management. Considerable literature is available on the impacts of other as-
pects of global change, particularly air pollutants (26, 31, 82). For example, the
effects of ozone on plant metabolism, crop yield and productivity (54, 72), plant
health (87, 124, 156), host-pathogen interactions (53, 125), host defense mech-
anisms, and interactions with pathogenic and saprophytic organisms have been
extensively studied. Similarly, studies on the effects of increased UV-B on plants
(119, 153), pathogens (4, 71, 111), phyllosphere yeasts (46), and disease develop-
ment (2, 37, 47, 87, 104) are well represented in the literature.
Most research on how climate change influences plant diseases has concentrated
on the effects of a single atmospheric constituent or meteorological variable on the
host, pathogen, or the interaction of the two under controlled conditions (21, 2426,
28, 83, 87). Interactions are clearly more complex in the real world, however, where
multiple climatological and biological factors vary simultaneously in a dynamic
environment. Climate change has the potential to modify host physiology and
resistance and to alter stages and rates of development of the pathogen. The most
likely impacts would be shifts in the geographical distribution of host and pathogen,
changes in the physiology of host-pathogen interactions, and changes in crop
loss. Another important impact may be through changes in the efficacy of control
strategies.
ecology in the new environment will largely determine how quickly pathogens
become established in a new region.
Changes may occur in the type, amount, and relative importance of pathogens
and affect the spectrum of diseases affecting a particular crop. This would be more
pronounced for pathogens with alternate hosts. For perennial species in plantation
and natural forests and orchards, plants may continue to be grown in existing
regions under marginal climatic conditions. The need for large tracts of suitable
land, infrastructure such as fruit- or timber-processing facilities, and the high
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cost of establishing a new plantation could limit the mobility of perennial crops.
Plants growing in marginal climate could experience chronic stress that would
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predispose them to insect and disease outbreaks. Warming and other changes could
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also make plants more vulnerable to damage from pathogens that are currently not
important because of unfavorable climate. Infection of eucalypts by Phytophthora
cinnamomi is favored in wet soils at temperatures of 12 to 30 C (107), hence the
pathogen does not pose a serious threat to the susceptible Eucalyptus spp. grown in
southeastern Australia. This situation may change with an increase in temperature.
Similarly, poplar clones in northern Europe may experience increased damage from
the thermophilic rust, Melampsora alli-populina (76).
If the frost-line moves north in the Northern Hemisphere, higher winter tem-
peratures could be accompanied by increased survival of insects (109, 145). For
virus-vector aphids, this could lead to higher incidence of virus diseases, espe-
cially in those regions where the timing of virus arrival is linked to winter survival
and spring flight of aphids (51). Barley yellow dwarf potyvirus (BYDV) is an
example of a virus that causes more severe disease following mild winters. Since
BYDV is exclusively vectored by aphids, increased survival of pathogen reservoirs
could greatly increase the economic losses caused by infection. Similar increases
in viruses of potato and sugar beet have also been observed following warmer
winters (19, 51, 81, 150).
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plants under increased CO2 (23). Similar effects in other pathosystems include a
reduction in the rate of primary penetration in Erysiphe graminis (barley powdery
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Elevated Temperature
Increases in temperature can modify host physiology and resistance. Considerable
information is available on heat-induced susceptibility and temperature-sensitive
genes (33, 42, 123). For example, a rise in temperature above 20 C can inacti-
vate temperature-sensitive resistance to stem rust in oat cultivars with Pg3 and
Pg4 genes (89). In contrast, lignification of cell walls increased in forage species
at higher temperatures (165) to enhance resistance to fungal pathogens (140).
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lodgepole pine (65, 76). Such projections, however, do not consider other factors
that can enhance the resilience of forest ecosystems to climate change, which led
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Loehle (75) to conclude that there is a systematic bias toward alarmist predictions
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Crop Loss
At elevated CO2, increased partitioning of assimilates to roots occurs consistently
in crops such as carrot, sugar beet, and radish. If more carbon is stored in roots,
losses from soilborne diseases of root crops may be reduced under climate change.
In contrast, for foliar diseases favored by high temperature and humidity, increases
in temperature and precipitation under climate change may result in increased
crop loss. The effects of enlarged plant canopies from elevated CO2 could further
increase crop losses from foliar pathogens (23, 87). Unfortunately, canopy char-
acteristics have not featured prominently in plant pathology research, despite their
influence on microclimate and pathogen dispersal (3). Recent developments in
three-dimensional modeling of plant architecture (virtual plants) offer new op-
portunities to integrate canopy architecture with microclimate effects and pathogen
dispersal (166).
Chakraborty et al (23) studied dispersal of and infection by C. gloeosporioides
under ambient weather conditions in the field on S. scabra plants that had been
raised under 1 or 2 CO2 in controlled environment chambers. Plants from
the two CO2 environments were exposed to naturally occurring inoculum in the
field on different dates, and conidial dispersal and infection were monitored. The
enlarged canopy of plants grown under elevated CO2 trapped more conidia that,
together with increased humidity in the denser canopy, led to more severe anthrac-
nose than on plants grown under 1 CO2 (Figure 1). In a separate experiment,
disease was reduced when plants were grown under elevated CO2 and inoculated
with C. gloeosporioides inside the controlled environment (23). The contrasting
results help to highlight dangers in extrapolating effects observed under controlled
environments to field situations.
It is difficult to arrive at realistic predictions of yield losses that may result
from a slow and gradual climate change by extrapolating findings from controlled
environment studies which generally considered only two contrasting CO2 levels.
Indeed, host and pathogen populations are apt to adapt to gradual changes in CO2
concentration. Long-term field studies utilizing free-air CO2 enrichment (FACE
technology) (136) or similar open-air facilities (99) offer the best approach to
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Figure 1 Cumulative number of lesions (a) and disease severity expressed as per-
cent leaf area affected (b) caused by Colletotrichum gloeosporioides on susceptible
Stylosanthes scabra plants that had been raised under 1 or 2 CO2 in controlled
environment chambers before being exposed to naturally occurring inoculum in three
field plots for 48 h on five separate occasions. Data are means and standard errors
from five exposures with three plants from each field plot. Source: S Chakraborty, IB
Pangga, PM Room & D Yates (unpublished data).
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and tomato but increase sugarcane root rot caused by Pachymetra chaunorhiza.
Increases in summer temperature and the frequency of precipitation would favor
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Host Resistance
Cultivar resistance to pathogens may become more effective because of increased
static and dynamic defenses from changes in physiology, nutritional status, and
water availability (see Physiology of Host-Pathogen Interactions). Durability of
resistance may be threatened, however, if the number of infection cycles within
a growing season increases because of one or more of the following factors: in-
creased fecundity, more pathogen generations per season, or a more suitable mi-
croclimate for disease development. This may lead to more rapid evolution of
aggressive pathogen races. In a pilot study, Chakraborty et al (unpublished data)
monitored evolution of C. gloeosporioides on S. scabra under elevated CO2. A
susceptible cultivar was grown in a controlled environment under 1 or 2 CO2
and inoculated with three isolates of the pathogen. For each isolate, conidia col-
lected from infected host tissue were used to inoculate a second group of plants of
the same cultivar. Successive groups of plants were inoculated with conidia arising
from the previous infection cycle to simulate polycyclic disease development and
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pathogen evolution over time. After each cycle, measurements were made on com-
ponents of pathogen aggressiveness, such as fecundity, lesion size, lesion number,
and disease severity. Preliminary results suggested a significant trend toward in-
creased disease severity (Figure 2); further, two of the three isolates showed a
gradual increase in fecundity under elevated CO2 after eight infection cycles.
Chemical Control
Climate change could affect the efficacy of crop protection chemicals in one of
two ways. First, changes in temperature and precipitation may alter the dynam-
ics of fungicide residues on the crop foliage. Globally, climate change models
project an increase in the frequency of intense rainfall events (39), which could
result in increased fungicide wash-off and reduced control. The interactions of
precipitation frequency, intensity, and fungicide dynamics are complex, and for
certain fungicides precipitation following application may result in enhanced dis-
ease control because of a redistribution of the active ingredient on the foliage (128).
Neuhaus et al (96) applied simulated rain to potato foliage at two intensities (6 and
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30 mm h1) and found that the higher rate significantly reduced the fungicide
residue that could be measured with a chemical assay, but that there was no dif-
ference in disease between the two treatments when the leaves were challenged in
a bioassay with Phytophthora infestans.
Second, morphological or physiological changes in crop plants resulting from
growth under elevated CO2 could affect uptake, translocation, and metabolism
of systemic fungicides. For example, increased thickness of the epicuticular wax
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layer on leaves (13, 167) could result in slower and/or reduced uptake by the host,
whereas increased canopy size could negatively affect spray coverage and lead to
a dilution of the active ingredient in the host tissue. Both factors would suggest
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Microbial Interactions
Climate change may alter the composition and dynamics of microbial communities
in aerial and soil environments sufficiently to influence the health of plant organs
(4, 46, 122). Changed microbial population in the phyllosphere and rhizosphere
may influence plant disease through natural and augmented biological control
agents. A direct effect of elevated CO2 is unlikely in the soil environment as
the microflora there is regularly exposed to levels 10 to 15 times higher than
atmospheric CO2.
Trees grown in soils of poor nutrient status, especially nitrogen, favor colo-
nization of roots by arbuscular mycorrhizal fungi (70). The relationship between
elevated CO2 and mycorrhizae is not well understood (139), and there are con-
flicting reports on how it may be influenced by the nutrient status of the plant and
soil. If a lower nitrogen status of plant tissue under increased CO2 results in more
mycorrhizal colonization, this could improve plant health through improved nutri-
ent uptake. Similar confusion exists on the potential role of vesicular-arbuscular
mycorrhizae and ectomycorrhizae in the suppression and biological control of
plant pathogens. Mycorrhizae can have positive, negative, or neutral effects on
plant disease, and their role is not well understood despite numerous studies on the
subject (105). Clearly, the influence of mycorrhizae on plant health under climate
change requires further research.
Changes in temperature may have highly nonlinear effects on tri-trophic interac-
tions of host, pathogen, and biocontrol agent. In wheat (152), a rise in temperature
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Management of climate change will put additional pressure on agencies responsi-
ble for exclusion as a plant disease control strategy (64). In some regions, certain
diseases of economic concern do not currently occur because the climate has
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precluded the causal agents from becoming established. Use of Geographical In-
formation Systems and climate matching tools may assist quarantine agencies in
determining the threat posed by a given pathogen under current and future cli-
mates. This approach was used by Sansford & Baker (126) to assess the risk of
establishment of Karnal bunt in the cereal-growing regions of the European Union.
IMPACT MODELS
Most of what has been said about plant disease in relation to climate change is
based on qualitative, rule-based reasoning. This approach seems attractive because
of the substantial body of knowledge already available regarding the environmen-
tal requirements of plants and their pathogens (25, 28, 36). For example, it seems
plausible that increased air temperature would result in a poleward expansion of
the geographical range of pathogens and in more generations per year (109); that
elevated winter temperatures would increase survival and hence the amount of
initial inoculum in many pathosystems (28); and that greater continental dryness
during summer (6) would reduce risk of infection by pathogens that require leaf
wetness or saturated soils for infection. But will this really be what happens? The
answer will depend on complex interactions of atmospheric, climatic, and biolog-
ical factors with technological and socioeconomic changes that are exceedingly
difficult to predict (24). It would appear, therefore, that in all but the simplest
cases these interactions are not amenable to qualitative analyses. Hence, quanti-
tative (modeling) approaches, which allow one to investigate multiple scenarios
and interactions simultaneously, will become more important for impact assess-
ment (28). Guidelines for such model-based assessments are needed, and Sutherst
et al (146) and Teng & Yang (149) have given a framework. With few exceptions
(28, 146, 148), not enough attention has been given to modeling approaches and
the analytical tools needed for quantitative impact assessment in plant pathology.
Climate Matching
Climate matching involves the calculation of a match index to quantify the
similarity in climate between two or more locations. The match index is based on
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locations, for example based on disease distribution maps (161), would allow
predictions to be made about future disease risk at the location of interest.
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Booth et al (10) used climate matching to identify regions suitable for Cylin-
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drocladium leaf blight on Eucalyptus spp. in Southeast Asia and around the world.
They first established a simple rule for presence or absence of the disease based
on long-term means of temperature and precipitation. This rule was then imple-
mented in a climate matching program to identify high-risk regions in Africa,
Australia, Latin America, and Southeast Asia under current climate. Further, two
climate change scenarios were run for locations in Southeast Asia. The results sug-
gested an increase in disease risk in northern Vietnam, southern Laos, and eastern
Thailand. These predictions are consistent with limited field observations indi-
cating that severe disease can occur in these regions during years with extreme
weather.
Possible effects of climate change on Phytophthora cinnamomi, a soilborne
oomycete with an extremely wide host range, were considered by Brasier (15)
and Brasier & Scott (16). This pathogen requires warm, wet soils and is hence
limited primarily to tropical and subtropical regions (76, 88, 107). More recently,
P. cinnamomi has been associated with oak declines in southern and Mediterranean
Europe. It was hypothesized (14) that this may be an early indication of climate
warming as the pathogen may have become more active because of higher soil tem-
peratures and/or increased host susceptibility caused by stress (e.g. more frequent
winter droughts in the region). For a more formal impact assessment, Brasier &
Scott (16) used the CLIMEX climate matching program (143) to map regions in
Europe favorable or unfavorable for this pathogen under present and future cli-
mate scenarios. The climate change simulations suggested that the pathogen could
extend its range further north, although it appeared unlikely that it could become
established in those regions where winter temperatures are low such as central and
eastern Europe (15). It was further hypothesized that the pathogens host range
could increase if spread occurs into regions where it is currently absent.
Empirical Models
Four diseases of two major crops in China, wheat and rice, were examined by
regression analysis to determine how they have varied through time and whether
this may relate to recent increases in mean and minimum temperatures (169). Rice
blast and wheat scab have increased sharply since the 1970s. The wheat acreage
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infected with powdery mildew has become more extensive, whereas stripe rust
has decreased steadily (169). This may be related to the increased spring and
early summer temperatures and would be consistent with the changes in stripe rust
observed in the Pacific Northwest associated with climate variability (27).
Boag et al (8) used data from soil samples collected during the European Plant-
Parasitic Nematode Survey to assess the possible impacts of climate warming on
the geographical range of virus-vector nematodes. Initial analyses of nematode
presence-absence data suggested a close association between mean July soil tem-
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perature and nematode distribution. Based on this result, the authors predicted that
climate change could result in increased nematode and virus problems in northern
Europe; they estimated that a 1 C warming would allow the species in study to mi-
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grate northward by 160 to 200 km (95). Although nematodes migrate very slowly,
humans are credited with efficiently disseminating them. Hence, nematode spread
into new regions could put a wide range of crops at risk; additionally, introduction
of new crops into a region could also expose them to infestation by nematode
species already present. Changes in precipitation, which were not considered in
these analyses, could influence nematode distribution on a large scale, although
previous findings had suggested that soil moisture would not affect nematode
distribution in most agricultural soils in northern Europe (8, 95).
In a similar study, Jahn et al (62) utilized long-term plant disease monitor-
ing records collected by the State Plant Protection Service in the former German
Democratic Republic (GDR) to develop empirical climate-disease models for 15
individual host-pathogen combinations. These models were then used with vari-
ous climate change scenarios to predict possible changes in infestation levels in
a future climate. Calculations with the most realistic scenario (a temperature in-
crease of 1 C combined with a decrease in precipitation of 30%) indicated that
leaf rusts of wheat and barley and powdery mildew of sugar beet could increase
substantially, reaching levels between two and five times as high as under the cur-
rent climate. Infestation levels on small grains by powdery mildews would remain
virtually unchanged, whereas those caused by foot rots and leaf blotch diseases
would decrease. Most notable was a decrease in potato late blight to a mere 16%
of its current level. The authors cautioned against over-interpreting their results,
which were based on calculations with data from only 1 of 14 regions in the former
GDR.
Population Models
A very different conclusion regarding the importance of potato late blight under
climate change was reached by Kaukoranta (68). This author developed degree-
day models for the emergence of potatoes and the date of late blight outbreaks
in Finland. The two models were coupled and extended by including leaf area
expansion of the crop as a function of thermal time; calculating radiation inter-
ception as a function of leaf area; transforming the intercepted radiation to tuber
dry matter; and simulating the effects of late blight on tuber dry matter through a
P1: FHP/FGO P2: FHP/FGP/FGM QC: FHP/arun T1: FDX
June 29, 1999 14:33 Annual Reviews AR087-16
reduction in green leaf area, assuming that disease reduced leaf area to zero within
14 days after the predicted outbreak. Model parameters were obtained and model
validation was done using data from a three-year field and greenhouse study. The
combined model was then used with various temperature change scenarios to pre-
dict possible changes in potato yield and yield losses caused by late blight in a
warmer climate. The results suggested that tuber yield could increase by 2 t ha1
per 1 C warming in the absence of late blight. This potential yield gain was al-
most completely offset when late blight was considered, chiefly because late blight
?
outbreaks occurred 4 to 7 days earlier and the period during which the crop was
susceptible was lengthened by 10 to 20 days per 1 C warming. This study did not
Annu. Rev. Phytopathol. 1999.37:399-426. Downloaded from www.annualreviews.org
Simulation Models
Simulation models have been used extensively to predict yields of various crops
in different agroecological zones under climate change (43, 113). Biotic yield-
reducing factors such as insects, pathogens, and weeds have, however, been largely
ignored in these simulations (148). Because of this shortcoming, the development
of linked disease-crop models is an important objective within the overall goal
of developing a predictive capability for agricultural impact assessment and mit-
igation (130, 146, 148). For at least one key crop, rice, preliminary analyses con-
sidering the combined effects on yield of increased temperature, elevated UV-B
radiation, and rice blast disease (Pyricularia grisea) have been done using a cou-
pled simulation model (7780). The model consisted of a physiological rice growth
model and a leaf blast epidemic simulator, linked via the quantitative effects of
leaf blast on photosynthesis and biomass production (78). Climate change was im-
posed by increasing mean temperature in fixed increments and by either including
or omitting effects of UV-B on the host and pathogen (77). The results suggested
that elevated UV-B could result in direct yield losses of 10%. Impacts of increased
temperature varied by agroecological zone, with an increase in blast and associ-
ated yield losses in cool, subtropical rice production regions (e.g. Japan) and a
decrease in humid tropics and subtropical regions (e.g. the Philippines). The au-
thors cautioned that the results must be considered preliminary as the simulations
did not include neck and panicle blast, two other important symptom types caused
by P. grisea. Further, increased CO2 was not considered, nor were changes in
precipitation as preliminary analyses had indicated that the combined model was
insensitive to changes in rainfall (77).
?
A proposed integrated case study of impacts of global change on vector-borne
diseases of crops, livestock, and humans was outlined by Sutherst et al (142).
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On the other hand, numerous challenges remain regarding the use of models for
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?
1995 IPCC report (67). Under the fuzzy scenario, climate changes were expressed
as triangular fuzzy numbers, utilizing the lowest and highest predictions in addition
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to the best estimates. Simulations with the crisp climate change scenario suggested
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only minor changes in environmental favorability for the two species compared
with the current climate. When simulations were conducted with the fuzzy climate
change scenario, important changes in environmental favorability emerged. In
some regions, the possibility of considerably increased winter precipitation led to
greater environmental favorability. The simulations further showed that this result
harbored considerable uncertainty, with a very broad range of possible outcomes
(Figure 3). Fuzzy logic could provide a simple but formal means of expressing
uncertainty that could be used across different levels of model complexity.
?
Climate cycles such as the El NinoSouthern Oscillation (ENSO) are major mech-
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anisms of interannual atmospheric variability (17, 106). Indeed, the ENSO is con-
sidered second only to the seasonal cycle in its impacts on climate variability
(103). It has been suggested, therefore, that ENSO-related climate extremes could
serve as an analogue for assessing possible impacts of long-term climate change
(103, 168). This approach seems particularly appropriate as ENSO events may
become more frequent in a high-CO2 climate (134).
Numerous studies in veterinary and medical epidemiology have established re-
lationships between ENSO and outbreaks of infectious disease (12, 29, 48, 97, 103,
159). There is a conspicuous lack of published work on plant disease in relation
to ENSO. In an early paper, Zhao & Yao (170) reported that the occurrence of
El Nino coincided with increased wheat scab and that observation of sea surface
temperature (a proxy for the strength of El Nino) could be used to predict the disease
along the median and lower reaches of the Yangtze River in China. More recently,
Scherm & Yang (132, 133) established similar associations between ENSO events
and other climate cycles (teleconnection patterns) with rust diseases of wheat in
various production regions.
Mitigation measures developed in response to natural climate extremes such
as ENSO may be useful in adapting to long-term climate change (93, 94, 116).
Hence, promotion of research on climate analogues in relation to plant disease
constitutes a true no-regrets approach; it would result in concrete benefits to
society regardless of the magnitude of climate change.
CONCLUSIONS
Climate change can have positive, negative, or neutral impact on individual patho-
systems because of the specific nature of the interactions of host and pathogen.
As a result, it has been difficult to decipher rules of thumb that may be used for
specific impact assessment. Three factors are largely responsible for this apparent
lack of general principles. First is a serious lack of knowledge of the effects of
some important factors such as CO2. The role of pathogens in the response of
plants to increased CO2 has not been well studied, hence its effect on disease is not
currently considered in crop simulation models. Second, there is only rudimen-
tary information on the interactions of individual factors that collectively influence
P1: FHP/FGO P2: FHP/FGP/FGM QC: FHP/arun T1: FDX
June 29, 1999 14:33 Annual Reviews AR087-16
plant disease in a changing climate. For example, recent studies showed that the
impacts of ozone in the field cannot be estimated without considering the pre-
disposing effects of fungal infections and the compensating effects derived from
elevated CO2 (156). Third, impacts on plant disease have largely been consid-
ered in small-scale experiments. Given that climate change operates at a global
scale, a lack of understanding of epidemic processes at relevant environmental and
spatial scales has hampered progress. The uncertainties associated with climate
change projections and the difficulty in extracting epidemiologically meaningful
?
environmental variables such as surface wetness from GCMs have contributed
to this.
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?
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