Managing Crop Diseases Under Water Scarcity: Annual Review of Phytopathology

PY58CH16_Swett ARjats.
cls June 15, 2020 19:1
Annual Review of Phytopathology

Managing Crop Diseases
Under Water Scarcity
Cassandra L. Swett
Department of Plant Pathology, University of California, Davis, California 95616, USA;
Access provided by University of Uppsala on 07/01/20. For personal use only.
Annu. Rev. Phytopathol. 2020.58. Downloaded from www.annualreviews.org
email: clswett@ucdavis.edu
Annu. Rev. Phytopathol. 2020. 58:16.1–16.20 Keywords

The Annual Review of Phytopathology is online at
irrigation management, water use efficiency, water sustainability, drought,
phyto.annualreviews.org
soil moisture, plant disease, soilborne disease, plant pathology, fungi,
https://doi.org/10.1146/annurev-phyto-030320-
oomycete, irrigation–pathogen interaction
041421
Copyright © 2020 by Annual Reviews. Abstract

All rights reserved
The significance of water scarcity to crop production and food security has
been globally recognized as a pivotal sustainability challenge in the UN Sus-
tainable Development Goals (86). The critical link between water scarcity
and sustainability is adaptation. Various changes in water use practices have
been employed to alleviate production constraints. However, the potential
for these changes to influence crop diseases has received relatively little at-
tention, despite the circumglobal importance of diseases on agricultural sus-
tainability. This article reviews what is known about the realized effects of
scarcity-driven alterations in water use practices on diseases in the field in or-
der to raise awareness of the potential for both increased disease risk and pos-
sible beneficial effects on crop disease management. This is followed by con-
sideration of the primary mechanistic drivers underlying disease outcomes
under various water use adaptation scenarios, concluding with a vision for
disease–water co-management options and future research needs.
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16.1
, Review in Advance first posted on

June 24, 2020. (Changes may still
occur before final publication.)
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INTRODUCTION
The direct effects of water scarcity and associated changes in water use on crop production have
been well reviewed, but little attention has been paid to interactions with crop diseases despite
the fact that diseases are among the most significant drivers of crop loss worldwide. Accordingly,
the purpose of this review is to synthesize the multifaceted ways in which water scarcity is im-
pacting plant diseases in managed crop systems circumglobally. Although this review considers
water insecurity drivers such as climate change, increased demand fueled by population growth,
periodic drought, and cumulative water-source depletion, the thematic driver of the narrative is to
examine how on-farm changes in water use in response to water scarcity influence disease dynam-
ics. Specifically, we examine how disease dynamics are changing (a) in response to the adoption
of irrigation in historically rainfed systems that no longer have sufficient rainfall and (b) follow-
ing adjustments in irrigation methods and management strategies aimed at reducing irrigation
inputs in response to surface and groundwater scarcity. A subsequent goal of this review is to ar-
ticulate needs in responding to the negative outcomes of these changing water use practices on
plant pathosystems. This includes developing water–disease co-management strategies as well as

mechanism-based platforms that can be used to form predictive hypotheses and interpret impacts
of water scarcity adaptation on crop diseases. The review concludes by highlighting strategies for
conducting rigorous transdisciplinary crop–disease–water research aimed at achieving sustainable
cropping systems under increasing water insecurity. Direct effects of climate change and periodic
droughts on plant-pathogen dynamics are not covered here, as they have been extensively dis-
cussed in more than 100 reviews; the reader can refer to the Juroszek et al. (40) compilation of
reviews for further reading.
WATER SCARCITY CHALLENGES IN AGRICULTURE: AN OVERVIEW

To consider how to manage diseases under water shortage, it is necessary to understand the sources
of water used in crop production and how they are applied as well as the vulnerabilities of each
water source that result in water insecurity. This is not intended to be a comprehensive review
of agricultural water use, as many excellent reviews exist (15, 27, 63, 67, 71, 77, 79), but instead
functions as a primer for non-specialists.
Globally, an estimated 84% of all agriculture relies primarily or solely on rainwater that falls
directly on crops during the growing season (27, 77, 79); this is referred to as dryland agriculture
in areas where rainfall amounts are low and/or occur erratically (77). Advantages to solely rainfed
production include minimal cost, lack of water resource competition, and lack of restrictive regu-
lations on use. However, because of cyclic droughts coupled with climate changes, rainfall events
are becoming increasingly unpredictable and many regions that have historically relied solely on
rainfall for crop production can no longer meet crop water demands with rainfall alone (15, 27,
29, 71, 77); in response, many growers are adopting supplemental irrigation practices.
Globally, approximately 16% of agricultural land is managed with irrigation water, but this
land produces approximately 40% of the world’s food supply (15, 27). Irrigation water comes from
surface water, groundwater, and recycled water (79). Surface water is generally a more reliable and
high-quality source of water but is becoming increasingly scarce because it is highly vulnerable
to changes in precipitation patterns, such as reductions in annual rainfall and snowpack; many
of these changes relate to normal cyclic droughts and/or climate changes (71, 78). Surface-water
allocations are typically also the most tightly regulated because of competition with municipal
and industrial groups, and allocations to agriculture face routine cuts (27, 29, 71, 78). In many
regions, surface water is being overused, resulting in the closure of river basins, which significantly
·.•�-
reduces water available for agriculture (27). Many growers use groundwater (underground aquifer
16.2 Swett

water) to adapt to surface-water scarcity. In countries such as the United States, groundwater
is controlled directly by the grower if they own the farmland and water rights and is therefore
not under the same restrictions as surface water. However, overuse of groundwater is leading
Evapotranspiration
to aquifer depletion, and without active efforts to balance use with recharge, depletion allows (ET): water losses due
saltwater intrusion, increasing soil salinity when applied to the crop (29, 71, 77). Grey, or recycled, to the combined
water can help alleviate surface and groundwater scarcity but problematically allows recirculation processes of water
and potentially biomagnification of plant pathogens and phytotoxic pesticides, causing damage to evaporation from soil
and canopy surfaces
crop plants (37, 65, 79).
and transpiration of
To respond to increasing scarcity of surface water and high-quality groundwater, irrigation water through the
methods (delivery systems) can be altered to improve water use efficiency. Irrigation delivery sys- crop canopy
tems can be either gravitational (e.g., furrow, flood, border, or basin irrigation) or pressurized
Irrigation method:
(sprinkler and micro-irrigation) (15, 77). Irrigation water inputs can be conserved by converting any water application
from non-site-specific application methods, such as furrow and sprinkler irrigation, to site-specific method to the soil
irrigation, such as surface or subsurface drip, which more precisely deliver water to the rootzone and plants (e.g.,
(77). These systems can reduce applied water volumes without reducing yields (60, 77, 78) and are gravitational methods
such as flood, basin,
often adopted to improve water use efficiency as well as yield and quality traits (2, 51, 55, 61, 82).
and furrow irrigation,
In addition to changes in irrigation method, changes in irrigation management strategies (e.g., or pressurized
alterations in irrigation volume and timing) can also be used to conserve water by more precisely methods such as
meeting plant water needs (1, 30, 50, 73). This is a very active area of research and technology sprinkler and
development, full coverage of which is beyond the scope of this review; for recent comprehensive micro-irrigation)
reviews, see GAO and FAO reports (28, 29, 31). The primary irrigation management strategies Irrigation
which have been examined for impacts on plant diseases include deficit irrigation and irrigation management
management using soil moisture–based scheduling. Deficit irrigation entails the application of strategies: strategies
that aim to apply water
irrigation water in amounts that are lower than the full satisfaction of maximum crop water re-
to cropped fields with
quirements [evapotranspiration (ET)] (10 13, 26, 30). Using soil moisture–based scheduling, water specific amounts,
inputs are reduced in reference to soil moisture (volumetric soil water content) or water availabil- timing, and frequency
ity in the soil (matric potential) (1, 44, 50, 53). Of note, irrigation management strategies are not
Deficit irrigation: the
solely aimed at reducing water usage and can also be used to improve crop quality traits (e.g., application of water
increased sugar content) (6, 83), manage diseases (35, 62), or achieve cultural management targets below full crop-water
(70). requirements
There are many additional strategies that growers use to increase on-farm water use efficiency (evapotranspiration)
in response to increasing scarcity of rainwater, surface water, and high-quality groundwater. These
include (a) improvements to soil characteristics using methods such as tillage, cover crops, and
soil amendments, which increase root system access to water and improve water-holding capacity;
(b) site alterations such as land leveling to improve water use efficiency; (c) use of crops that are
more drought and/or salt tolerant (crop rationalization); (d) changes in planting practices, such as
avoiding times of year with greater rainfall uncertainty and altered plant spacing, to enable crop
productivity when water is limited; and (e) improvements to the water delivery system infrastruc-
ture to reduce losses during transport (15, 27, 63, 67, 71, 77, 79). Although important, these adap-
tations are not emphasized here because of limited available information on disease interactions.
REALIZED EFFECTS OF ADAPTIVE WATER USE PRACTICES

ON DISEASES IN THE FIELD
To review the realized effects of water use adaptations on disease, we focus here on field studies,
starting with the simplest adaptation, conversion from rainfed to irrigated production, moving to
alterations in irrigation method (specifically conversion to drip), and, lastly, reviewing the effects
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of changes in irrigation management strategies (specifically use of deficit irrigation and soil
www.annualreviews.org • Plant Disease and Water Scarcity 16.3

Disease Disease No
enhancement suppression effect
Aboveground pathogens Belowground pathogens
Rainfed irrigation
Gravitational/sprinkler drip irrigation (Water- (Dry-

loving) loving)
Full reduced irrigation

(including deficit irrigation)
Figure 1
Summarizing trends in disease outcomes associated with common water use adaptations, indicating disease enhancement (up arrow),
disease suppression (down arrow), or no effect (bar); pathogens were separated into water loving and dry loving for drip irrigation
conversion because of disparate effects. Note that exceptions can be found to every described trend.
moisture-based precision irrigation). Any bias in the literature toward certain kinds of crops
(e.g., perennial, agronomic) or certain regions is highlighted. Each section is separated into
aboveground and belowground diseases because of the consistent divergence in the responses of
these different types of pathosystems (Figure 1). Although effort was made to include all relevant
bacterial and viral disease studies, this review is focused primarily on fungal and oomycete
pathogens, reflecting their dominance in the literature. If no taxonomic designation is given, the
reader can assume that the pathogen is in the kingdom Fungi. The Latin binomials (or virus
name) for pathogens are those given by the authors of each cited study; it should be noted that
some names have changed.
This review does not address the effects of soil salinity accumulation resulting from the use
of groundwater or drip irrigation. Despite clear potential to influence diseases, there is a paucity
of true field studies to specifically evaluate salinity effects on plant diseases. Recycled water also
has significant effects on disease, but these interactions have been well reviewed elsewhere (37,
65). Effects of adaptive water use practices on diseases in controlled laboratory and containerized
(in-pot) greenhouse settings were excluded in this section because results cannot be confidently
extrapolated to crop performance in the field. However, controlled studies can provide valuable in-
sight into the underlying mechanisms that account for disease outcomes and are emphasized in the
section titled Mechanisms Underlying Interactions Between Water Use Adaptation and Disease.
How Conversion from Rainfall-Only to Supplemental Irrigation Affects

Disease Development
Rainfed production is largely used for agronomic (field) crops and thus the majority of disease
studies are on these crops (e.g., wheat, barley, sunflower, soybean); additional studies are primarily
in orchard crops. There are studies examining effects on both aboveground and belowground
pathosystems, with better coverage of aboveground diseases; studies have been conducted across
a wide range of geographic regions worldwide.
Aboveground. Diseases affecting aboveground plant parts tend to increase in severity following
·.•�-
irrigation adoption, although in some cases there are suppressive or neutral effects (Figure 1).
16.4 Swett

The following are examples of increased disease risk in agronomic crops: In tobacco fields in
Zimbabwe, powdery mildew (Erysiphe cichoracearum) severity was greater under irrigation com-
pared to rainfall only (19), and in sunflower production in Brazil, supplemental use of sprinkler
irrigation increased the severity of Alternaria and Septoria leaf blights (caused by Alternaria he-
lianthi and Septoria helianthi, respectively) compared to solely rainfed production (47). Beyond
agronomic crops, in traditionally rainfed cherry orchards in Spain, irrigation increased posthar-
vest fruit rot (etiology not included) (88). Disease-enhancing effects have been attributed to in-
creased canopy moisture, changes in stomatal conductance, and/or development of microcracking
or wounds under irrigation, all of which are suggested to facilitate pathogen infection.
In some cases, irrigation adoption did not increase the risk of aboveground diseases (Figure 1),
such as in Kansas (USA) cornfields, where irrigation adoption increased yield and reduced corn
stalk rot caused by Fusarium moniliforme (25), and in Argentina dryland wheat production, where
adoption of sprinkler irrigation reduced tan spot caused by Drechslera tritici-repentis (48). As an
orchard crop example, fungal leaf scorch (caused by Glomerella cingulata and a Phomopsis species)
in pecan orchards in Georgia (USA) was less severe under irrigation compared to solely rain-
fed production (93). As an example of a neutral effect, in Brazil coffee plantations, the shift from
rainfall-only to supplemental irrigation did not increase the progression of brown eye spot (caused
by Cercospora coffeicola) (8). Researchers commonly postulated that disease-suppressing or neu-
tral effects were due to reduced plant stress and thus enhanced disease resistance under irrigated
conditions.
Belowground. Diseases affecting belowground plant parts can be less severe, unaffected, or en-
hanced following a shift from rainfall-only to irrigated production (Figure 1). As an example of
reduced disease severity, in barley production in the Northern Great Plains (USA), Grey et al. (34)
found that converting from dryland to irrigated production (200 versus 400 mm cumulative mois-
ture) reduced yield losses from common root rot in barley (caused by Fusarium culmorum and Bipo-
laris sorokiniana). As an example of a neutral effect, in spring wheat production in Nepal, Burlakoti
et al. (12) did not see any significant effects on the development of root rot and spot blotch (B.
sorokiniana) when adopting flood irrigation compared to solely rainfed production. The authors of
both studies attributed these neutral-to-suppressive effects to improved disease resistance under
reduced water stress (12, 34). In contrast, as an example of soilborne disease enhancement follow-
ing irrigation adoption, Piccinni & Rush (61) found that in conversion from dryland to irrigated
sugar beet in Texas (USA), more frequent irrigation increased impacts of Beet necrotic yellow vein
virus (BNYVV), transmitted within spores of the soilborne plasmodiophoroid Polymyxa betae; the
authors speculated that this was due to enhanced dispersal of the plasmodiophoroid vector under
increased soil moisture.
How Changes in the Water Delivery System Can Affect Disease Development:
Conversion to Drip
The most common irrigation system adaptation to save water is to shift from flood, furrow, basin,
or sprinkler irrigation to drip irrigation (either surface or buried) (77). Most field studies looking
at drip irrigation–pathogen interactions are focused on annual vegetable crops with fairly even
coverage of aboveground and belowground diseases and with a notable bias toward Phytophthora
diseases in the latter. Although this body of literature represents production systems in diverse
regions worldwide, a predominance of studies are in the United States.
Aboveground. Generally, drip irrigation poses little risk and provides possible management
·.•�-
advantages for water-loving pathogens but can increase risk of diseases caused by dry-loving

pathogens (e.g., powdery mildew) (Figure 1). As water-loving pathogen examples, in strawberry
fruiting fields in Brazil, conversion from sprinkler to drip irrigation typically had a neutral (non-
effect) or a suppressive effect on strawberry anthracnose (caused by Colletotrichum acutatum) or
reduced both flower blight and fruit rot (18). Similarly, in commercial greenhouse tomato pro-
duction in Algeria, conversion from flood to drip irrigation suppressed Botrytis fruit and stem rot
caused by Botrytis cinerea (2). Similarly, in California (USA), downy mildew of lettuce (Bremia lac-
tucae) developed at a slower rate under drip versus furrow irrigation (82), and incidence of downy
mildew of spinach (Peronospora effusa) was lower under drip versus sprinkler irrigation (51). In all
the above, reductions in plant surface moisture under drip irrigation (documented in most studies)
were postulated to be suppressive to pathogen sporulation, dispersal, and infection (2, 18, 51, 82).
In contrast, the dry-loving tomato powdery mildew pathogen (Leveillula taurica) was more severe
under drip compared to overhead sprinkler irrigation in organic fresh market tomatoes in Brazil;
this was attributed to lower leaf wetness under drip, which enhanced pathogen infection (43).
Belowground. There is a common theme for drip irrigation to have either no effect or a sup-
pressive effect on belowground diseases, with some exceptions (Figure 1). Compared to sprin-
klers, drip irrigation of apples in British Columbia (Canada) over eight years reduced incidence
of tree mortality caused by Phytophthora root rot and reduced the percent of trees sustaining
root infections by the pathogen Phytophthora cactorum (87). Similarly, drip irrigation reduced the
incidence of Phytophthora root rot (Phytophthora capsici) in chili peppers compared to furrow ir-
rigation in New Mexico (USA) (95) and basin irrigation in Turkey (32), and in commercial New
Mexico fields, disease incidence was 40% lower on drip- versus furrow-irrigated farms (75, 76).
Beyond Phytophthora, Subbarao et al. (82) found that converting from furrow- to drip-irrigated
lettuce could suppress lettuce drop (caused by Sclerotinia minor) and corky root (caused by the
bacterium Rhizomonas suberifaciens) in California (USA) lettuce fields, increasing yields (82). In
onion production in Argentina, drip irrigation either reduced bacterial bulb rot caused by Pec-
tobacterium carotovorum or had no effect when compared to sprinkler irrigation (17). Similarly, in
Nairobi, losses from Erwinia soft rot (caused by the bacterium Erwinia carotovora subsp. carotovora)
in greenhouse-grown Zantedeschia (calla lily) tubers were reduced under drip compared to over-
head irrigation (54). These neutral to beneficial effects were commonly hypothesized to be due to
reduced plant stress (not demonstrated in most studies) and thus greater disease resistance under
higher soil moisture (demonstrated in most studies) as well as changes in root system architecture,
elimination of excessive moisture periods, and/or changes in soil moisture distribution, which po-
tentially limited pathogen reproduction, dispersal, and/or infection opportunities (17, 32, 54, 75,
76, 82, 87, 95). As an exception to the trend above, Bryla & Linderman (11) found that blueber-
ries in Oregon (USA) suffered more severe Phytophthora cinnamomi root infections and root rot
under drip versus sprinkler irrigation, reducing shoot vigor; this effect was attributed to increased
localized soil moisture within the rootzone under drip, which favored disease development.
How Changes in Irrigation Management Strategies Can Affect Disease

Development: Reductions in Applied Irrigation Volume
Changes in irrigation management strategies can be used to achieve water use reductions by more
precisely meeting plant water needs (1, 26, 30, 50, 73). The primary irrigation management strate-
gies examined for impacts on plant diseases are deficit irrigation and soil moisture–based irrigation
management. These studies span a wide breadth of crops, with the greatest emphasis on hor-
ticultural crops. There is even representation of aboveground and belowground diseases in the
·.•�-
literature, with studies primarily focused on crop production in the United States.
16.6 Swett

Aboveground. Overall, irrigation management strategies aimed at reducing water inputs tend
to reduce severity or have no effect on aboveground diseases, with some exceptions (Figure 1).
For example, in grapevines in Australia, Austin & Wilcox (6) found that reducing water inputs
from 8 L/h (two drip lines) to 4 L/h (single drip line) resulted in less severe powdery mildew
(Erisphye/Unicula necator). In California almond orchards, Teviotdale et al. (84) found that reduc-
ing water inputs to near-stress levels using deficit irrigation (in reference to base ET) reduced the
incidence of dead leaf clusters, amount of dead wood, and incidence of hulls infected by Rhizopus
stolonifer or Monilinia fructicola. In nectarine orchards in Turkey, Atay et al. (5) found that deficit
irrigation up to 75% of full ET (drought stress-inducing) either suppressed or did not influence
fruit rot development associated with species of Monilinia, Rhizopus, and Botrytis (species names
not provided). Beyond woody crops, in Florida (USA) fresh market tomato production, severity
of bacterial spot (Xanthomonas campestris) was not affected by deficit irrigation (70% and 85% ET
versus 100% ET) (56); however, in this same study, early blight (Alternaria solani) was more severe
under deficit irrigation (56). Generally, suppressive to neutral effects were hypothesized to be re-
lated to reduction in infectable plant tissue and suppression (or lack of an effect) on sporulation
and infection under lower canopy humidity (5, 6, 56, 84). In the unusual instance of increased dis-
ease severity under deficit irrigation, drought stress was postulated to compromise host resistance
to early blight but not bacterial spot (56).
Belowground. Belowground disease risk is generally increased or unaffected when using wa-
ter use–reducing irrigation management strategies, with some cases of disease-suppressive effects
(Figure 1). As an example of disease enhancement, deficit irrigation–based reductions from 100%
ET to 60% ET and 40% ET both reduced soil water content and increased the severity of an-
thracnose foliar blight and basal rot (Colletotrichum cereale) in annual bluegrass turf in New Jersey
(USA); of note, moderate reductions to 80% ET had no soil moisture effect and did not affect
disease (70). In commercial poinsettia greenhouses in Maryland (USA), soil moisture–based irri-
gation scheduling reduced water inputs up to 29% without affecting Pythium root rot (Pythium
aphanidermatum), although further soil moisture reductions in commercial plants under controlled
conditions increased root rot severity (22). In both of the above studies, the authors attributed in-
creased disease risk to reduced soil moisture (demonstrated), which compromised plant resistance
under water stress (not demonstrated) (22, 70).
Illustrating neutral to suppressive effects, in California cauliflower fields reductions in soil
moisture reduced incidence of Verticillium wilt and root system colonization by the pathogen
Verticillium dahliae (94). Similarly, a 50% reduction from standard (80%) ET did not alter Verti-
cillium wilt (V. dahliae) in Texas cotton fields (92). In melons in Israel, Pivonia et al. (62) found
that reducing irrigation inputs reduced incidence of decline caused by Monosporascus cannonballus.
In these studies, neutral-to-suppressive effects were connected with reduced pathogen movement,
growth, and infection in drier soils as well as reduced root system growth and thus reduced infec-
tion opportunities (62, 92, 94).
MECHANISMS UNDERLYING INTERACTIONS BETWEEN WATER

USE ADAPTATION AND DISEASE
As illustrated above, there is potential for adaptive irrigation practices to enhance disease im-
pacts, have no effect, or suppress disease (Figure 1). Every adaptive water use strategy reviewed
above has been shown to enhance certain diseases, although there is no one irrigation method
that consistently poses a risk to all diseases (Figure 1). Furthermore, every adaptive practice has
·.•�-
been shown to either suppress or have no effect on diseases in certain cases; the conversion to

drip irrigation is the most consistent in suppressing diseases (Figure 1). The disease outcomes
are dictated by abiotic changes resulting from altered water use practices and the biology of the
pathogen(s) involved and are a result of both direct effects on the pathogen (survival, reproduc-
tion, and dispersal) and indirect effects on plant resistance to the pathogen. It is important to note
that the effects of a given irrigation practice on pathogen population size and infection success
can be independent from disease and yield outcomes. For instance, Del Castillo et al. (21) found
that tomato root colonization by P. capsici was greater under higher soil moisture, but root rot
severity was greater under lower soil moisture. In addition, increased disease risk does not always
translate into reduced yields. Changing practices may have direct positive effects on plant growth
and yield, which can override the lesser impact of disease enhancement. For example, Maldaner
et al. (47) found that supplemental irrigation of sunflower both increased Alternaria and Septoria
leaf blight and significantly increased yields. Conversely, just because a practice does not enhance a
given disease does not mean that yields are unaffected, as is shown in numerous reduced irrigation
studies such as those by Obreza et al. (56), Atay et al. (5), and Wheeler et al. (92).
Understanding the production risks associated with a given change in water use methodology
thus requires a holistic assessment of effects on pathogen populations, infection, disease develop-
ment, and yield outcomes. The above section provides hypothesized mechanisms proposed by the
various authors to account for treatment effects. In this section, we review the studies that test
these hypotheses to address the possible mechanistic bases for enhanced, suppressive, and neutral
effects on diseases associated with the different water adaptation scenarios reviewed above. Most
of these studies were conducted under controlled conditions, with field studies included where
possible.
Factors That May Favor Disease

Adaptive practices that increase plant stress, compromising disease resistance. Changes in
irrigation management strategies that aim to optimize water use, such as deficit irrigation, com-
monly enhance soilborne diseases and, less frequently, aboveground diseases (Figure 1). In some
of the case studies noted above, applying water deficit incurred a drought-stress penalty. It is well
known that drought stress can enhance diseases of a wide array of crops, including Fusarium wilt
of cotton (64), Fusarium ear rot of corn (58), Botryosphaeria blight of pistachio (46), and Sphaerop-
sis sapinea canker in Pinus halepensis (57), among others. It is likely that in some cases, water use
reduction methods enhance diseases because of drought stress–mediated suppression of plant dis-
ease resistance, which enhances colonization, disease incidence, and symptom severity. Studies
of response mechanisms have shown that when plants upregulate abscisic acid drought-stress re-
sponse pathways, disease defense–related hormones such as salicylic acid and pathogenesis-related
proteins are suppressed (9). This effect, referred to as predisposition, can increase disease suscepti-
bility even when plants are returned to well-watered conditions (9). Although there is a paucity of
evidence for predisposition under reduced irrigation methods, as an indication of predisposition,
Del Castillo et al. (21) found that when tomatoes were first exposed to a reduced soil moisture
treatment for different intervals and then returned to well-watered conditions and inoculated,
Phytophthora root rot (P. capsici) was more severe when exposed to longer periods of reduced soil
moisture. Because the plants were re-watered prior to inoculation, the authors speculated that
changes in disease severity were due to stress-induced alterations in host resistance.
When adaptive water use increases soil or canopy moisture, favoring pathogen survival,
population growth, dispersal, and/or infection. Changes in water use practices that increase
·.•�-
canopy or soil moisture commonly enhance aboveground or soilborne diseases, respectively. This
16.8 Swett

includes adoption of supplemental irrigation in rainfed systems and conversion to drip irrigation.
These changes can enhance pathogen reproduction, dispersal, infection, and survival. As an ex-
ample of canopy-related changes, Cole (19) found that shifting from rainfed to irrigated tobacco
increased infection severity and associated spore load of powdery mildew (Erysiphe cichoracearum).
Increased soil moisture under adaptive irrigation tends to have the greatest effect on waterborne
pathogens that produce motile spores, including Phytophthora and Pythium species, as well as
on pathogens that are vectored by motile spore-producing organisms such as oomycetes, plas-
modiophoroids, and chitrids. As an example of soil moisture facilitation of oomycete pathogens,
Phytophthora root rot of highbush blueberry was greater under surface drip irrigation, which
increased soil moisture around the roots compared to sprinklers and microsprinklers (11); cor-
roborating this, Ristaino et al. (68) found that both soil moisture and P. capsici inoculum density
were highest around drip lines. As an example of both pathogen and soilborne vector-enhancing
scenarios, Domfeh & Gudmestag (24) found that conversion from rainfed (lower soil moisture) to
supplemental irrigation (higher soil moisture) increased incidence and severity of powdery scab
on potato tubers, which is caused by the zoospore-producing plasmodiophoroid Spongospora sub-

terranea f. sp. subterranea, and increased incidence of tuber necrosis, caused by the Potato-mop top
virus, which is vectored in the former’s motile spores.
Factors That May Suppress or Have No Effect on Disease

If the change reduces water needed for pathogen sporulation, dispersal, or survival. Meth-
ods that reduce soil water content can suppress the dispersal of soilborne pathogens. As long
as plant-available water is not significantly reduced, the overall outcome can be suppressive (or
neutral) to disease development. Such soil moisture reductions occur following the shift from
furrow/flood irrigation to drip/microsprinkler irrigation and when using irrigation management
strategies that reduce water inputs (e.g., deficit irrigation). These practices can suppress disease
development of waterborne pathogens by suppressing zoospore production and dispersal, as has
been well documented for P. capsici soil populations (32, 68, 69, 95). In addition, pathogen coloniza-
tion and subsequent survival within dead host tissue in the soil can be suppressed. For example, the
shift from rainfed to irrigated soybeans increased soil matric potential (less negative matric poten-
tial, higher soil moisture) and reduced the density of Macrophomina phaseolina microscelerotia in
root tissue compared to nonirrigated (rainfed) plants (42). Likewise, deficit irrigation treatments
reduced the formation of microsclerotia by V. dahliae in cabbage tissue (94).
Many aboveground pathogens require humid conditions for sporulation and infection and are
suppressed under low canopy humidity. Canopy humidity can be decreased following conversion
to drip from furrow, flood, or sprinkler irrigation, conversion from surface to buried drip, or re-
ductions in irrigation amounts, timing, or frequency. For example, shifting from flood to drip
irrigation reduced release of Mycosphaerella nawae ascospores from fallen leaves in a persimmon
orchard (89).
If the change in irrigation practice does not affect plant stress and thus disease resistance.
Reducing the amount of water applied to the crop does not mean that drought stress will develop.
Many methods to reduce water inputs do not significantly reduce plant-available water (soil matric
potential) and thus do not impose stress on the plant. For example, Teviotdale et al. (84) found that
certain deficit irrigation methods did not significantly influence stem water potential in almond
and did not enhance M. fructicola and R. stolonifer branch infections (84). Similarly, Del Castillo
et al. (22) demonstrated that use of a soil moisture sensor network irrigation system in a greenhouse
·.•�-
floriculture crop could reduce water inputs without influencing plant-available water or Pythium

root rot. In the latter study, the authors suggested that reductions to 50% VWC (volumetric
water content) did not incur a negative effect on the crop because under the reference grower
standard (60% VWC), water was being applied in excess of crop consumption abilities; reductions
to 50% merely reduced the application of excess water (22). This is not an uncommon outcome
of adopting precision irrigation techniques and highlights the opportunities to save water without
negatively impacting crop production (1, 30, 50, 73).
If the change in irrigation practice reduces the total volume of infectable plant tissue. Some
studies indicate that reduced irrigation management strategies, such as deficit irrigation, may be
beneficial to disease management by reducing the infectable aboveground or belowground plant
tissue, thus reducing the chances for infection. For example, reduced irrigation inputs decreased
both cauliflower root system area and the severity of Verticillium wilt (94). Similarly, application of
irrigation deficit in almond could reduce branch growth, and this corresponded with a reduction
in branch infections by M. fructicola and R. stolonifer (84). Both of these analyses were correlative,
not causal, and lacked proper controls to account for other drivers of treatment effects.
TOWARD A DISEASE–WATER CO-MANAGEMENT FRAMEWORK

A disease–water use co-management framework is required to enable sustainable water use under
pathogen pressure. Although in some cases water-saving irrigation practices can be exploited to
help manage diseases, every adaptation scenario comes with certain disease risks (Figure 1). Below,
we review common, available mitigation strategies and some potential management opportunities
that merit further study.
Incorporating Parameters of Irrigation Management into Plant Disease

Recommendations to Reduce Disease Risks or Exploit Adaptations
that Promote Disease Management
As discussed above, there are potential disease risks associated with adapting water use practices to
respond to water insecurity. One common theme in irrigation adaptation is the increased above-
ground and belowground disease risk associated with the shift from dryland to irrigated produc-
tion (Figure 1). To address this risk, several studies indicate that altering irrigation methods can
significantly reduce disease risk. For example, Domfeh & Gudmestag (24) found that shifting
from full-season irrigation to irrigation for only part of the season (either for the first 50 days
after planting or starting 50 days after planting) mitigated disease-enhancing effects of irrigation
adoption on powdery scab (caused by Spongospora subterranea f. sp. subterranea) and tuber necrosis
(caused by the Potato mop-top virus) in potato. Similarly, Piccinni & Rush (61) found that irriga-
tion every four weeks, compared to every two or three weeks, offered optimal suppression of the
fungal-transmitted Beet necrotic yellow vein virus, with better yields than plants irrigated every five
weeks.
Studies of irrigation management strategies suggest that the impacts of some belowground dis-
eases can be enhanced when irrigation inputs are reduced, but this effect can be avoided by curbing
the severity of irrigation reductions. For example, Del Castillo et al. (22) found that although re-
ductions to 25% VWC increased Pythium root rot incidence in potted poinsettias, reductions to
35% VWC had no effect on disease and resulted in ∼25% water savings. Likewise, whereas sever-
ity of anthracnose in annual bluegrass increased under both 40% ET and 60% ET, a reduction to
80% ET had no effect (versus 100% ET) (70). In addition, alterations in irrigation timing can mit-
·.•�-
igate disease-enhancing effects. For instance, Teviotdale et al. (84) reported a significant reduction
16.10 Swett

in hull rot of almonds when deficit irrigation was applied for only a portion of the growing season,
compared to application of deficit irrigation across the whole season. Similarly, Del Castillo et al.
(23) found that incidence of Phytophthora root rot in tomato was significantly lower under reduced
soil moisture irrigation for two versus six weeks.
Adoption of drip irrigation can also increase disease risk in rare cases. To manage this risk in
blueberries, Bryla & Linderman (11) reduced drip irrigation inputs from 100% ET to 50% ET,
thereby reducing colonization of oomycete root rot pathogens. However, most studies suggest
that conversion to drip irrigation (from furrow, basin, or sprinkler) is itself a disease management
tool, suppressing diseases favored by high canopy moisture, such as anthracnose flower blight
and fruit rot in strawberry (18), Botrytis fruit and stem rot of greenhouse tomatoes (2), and downy
mildew of spinach (51), and also suppressing diseases favored by high belowground moisture, such
as Phytophthora root rot of pepper (32, 95).
Pesticide and Biological Product Use to Manage Effects of Water

Use Adaptation on Disease

Pesticides (fungicides, bactericides, oomycides, fumigants) can be an effective way to manage dis-
eases affecting higher-value crops (e.g., vegetable, fruit, nut, floricultural, and landscape crops) and
can be used to mitigate disease risks associated with water use adaptation. For example, Maldaner
et al. (47) found that when converting from dryland to irrigated sunflower, fungicide applications
could overcome the disease-enhancing effects of irrigation adoption on Alternaria leaf spot and
Septoria leaf spot.
In many regions, there is a concerted effort to reduce synthetic pesticide inputs through aug-
mentation or replacement with biopesticides or other biological-based products. Furthermore,
biostimulants are gaining increased attention for application in plant stress mitigation (33, 90),
and some studies suggest that microbes can be used to mitigate stresses associated with water
use adaptation. For example, Kavroulakis et al. (41) reported that an endophytic Fusarium solani
strain increased the net photosynthetic rate, yield, plant water content, and stomatal conductiv-
ity of tomatoes grown under reduced water inputs (41). Biological compounds may thus have
utility in suppressing negative effects of adaptive water use practices on plant diseases by reduc-
ing plant stress and/or through direct pathogen suppression. This could be achieved by either
applying treatments to favor activity of endogenous beneficials or adding beneficial microbes to
the crop environment. In support of water–disease co-management potential, several bacterial
strains were shown to enhance tolerance of corn to a high salt environment and a subset of strains
also inhibited the soilborne pathogens Sclerotium rolfsii and Fusarium oxysporum, showing poten-
tial for co-management of salt stress and disease (66). Similarly, Vyas & Kaur (91) characterized
several bacterial endophytes that conferred salt-stress tolerance to pea and maize and many of
these endophytes also suppressed growth of the pathogens Fusarium verticillioides, Curvularia lu-
nata, and Alternaria alternata in culture (91). As a non-salt example, Del Castillo et al. (22) found
that Pythium root rot enhancement under reduced soil moisture could be mitigated in poinsettia
through the application of Bacillus subtilis. Of note, for many crops, including most agronomic
crops, and production regions, pesticides and/or biological products are not typically economical
because of small profit margins.
Shifting Crop and Cultivar use to Manage Effects of Water Use

Adaptation on Disease
When adaptive strategies increase disease risk, a simple risk-reduction practice is to shift from
·.•�-
a highly susceptible cultivar to a more resistant cultivar. For example, Sanogo (74) found that

Phytophthora root rot development under high salt conditions could be managed by using a resis-
tant rather than susceptible pepper cultivar. Of course, resistant cultivars are not always available,
and, when available, they can come with other production challenges, such as reduced yield and
quality and sometimes increased susceptibility to other diseases. Of concern, some of the changes
associated with water use adaptation (e.g., increased soil salinity) also appear to compromise single-
gene resistance to some pathogens, such as Fusarium wilt race 3 (I3 gene) resistance in processing
tomato; this may be similar to the well documented effects of high temperature on resistance gene
expression (e.g., the N gene conveying Tomato mosaic virus resistance and the Cf-4 and Cf-9 genes
conveying Cladosporium fulvum resistance in tomato) (36, 39). To address this concern, breeding
programs are encouraged to consider screening candidate resistant lines under a range of irri-
gation conditions to establish field stability of resistance traits. These efforts could be aided by
mechanistic studies to understand the basis for compromised resistance gene expression, which
may be similar to high-temperature suppression mechanisms, including temperature-dependent
changes in transcription factor RNA structure or activation of calcium channels in the plant (39).
There is a large body of literature describing the need and current progress in breeding for
crops with improved resistance to drought and salt tolerance (4, 59, 71). However, these efforts
do not typically include evaluations of disease resistance or selection of cultivars with pathogen
tolerance/resistance traits (7). Where single-gene resistance exists, breeders are encouraged to
include the resistance gene as part of the cultivar development process. Where there is not a well-
characterized resistance gene, breeders may consider adding pathogen screenings as part of their
cultivar development process to avoid releasing drought-/salt-tolerant materials with high disease
susceptibility.
ADVANCING THE FIELD OF PLANT DISEASE–IRRIGATION

INTERACTION SCIENCE: CONSIDERATIONS AND NEEDS
Extending Management Studies to Understand Interactions Between
Water Use Adaptations and Latent/Endophytic Infection
Latent pathogen infections represent a situation in which the pathogen has colonized the host
but has not yet developed symptoms. Latent infections of nursery stock are one of the primary
means by which pathogens are introduced to new regions and countries and can be drivers of
significant losses when the plants are later planted in fields or the landscape. Very few studies
have investigated how water use management influences latent infection development. An excel-
lent study conducted by Daugovish et al. (20) revealed that drip (versus sprinkler) irrigation in
nursery production of strawberry plants reduced the percentage of nursery stock plants asymp-
tomatically infected with Colletotrichum acutatum, the cause of anthracnose fruit and crown rot
in fruiting fields. They also found less C. acutatum DNA in nursery stock plants produced un-
der drip-irrigated conditions. This in turn reduced anthracnose impacts when they followed the
transplants to fruiting fields, increasing canopy development and reducing the frequency of plant
mortality and stunting. Similarly, Del Castillo et al. (21) found that reduced soil moisture regimes
reduced asymptomatic infection of tomato roots by P. capsici. Although these studies suggest that
the adaptive water use practices under study (drip and reduced soil moisture) reduced risk of la-
tent infection (20, 21), such trends need to be evaluated across a wider range of pathosystems,
irrigation methods, and nursery and field crops. Further studies of interactions between water
use adaptation and latent pathogen infection can help inform strategies to minimize regional and
global pathogen movement and resulting crop loss.
The vast majority of studies consider how drought-stress responses in the host influence micro-
·.•�-
bial infection processes (and resulting disease); it may also be important to consider how microbial
16.12 Swett

infection could influence drought-stress responses in the host. There is some limited evidence
that asymptomatic plant-associated microbes, including non-pathogenic endophytes and latent
pathogens, may alter the ability for plants to tolerate drought stress. For instance, symptomless
infection of Pinus radiata roots by the pitch canker pathogen Fusarium circinatum significantly re-
duced stem water potential in plants following two weeks without water compared to plants that
were not infected (C.L. Swett & T.R. Gordon, unpublished data). Similarly, in some perennial
ryegrass (Lolium perenne) genotypes, the endophyte Neotyphodium lolii has a detrimental effect on
drought-stress recovery (16). There may be diverse, heretofore unlooked for, microbes that are
significantly modulating the ability of crops to tolerate stresses associated with adaptive water
use practices; identifying these interactions has the potential to further advance effective water
resource management.
Understanding How Changes in Water Use May Drive Emergence

of New Pathogens and Influence Food Safety
As discussed above, many adaptations pose increased risk to diseases through alterations in the
canopy or soil environment and by suppressing host resistance. These same changes have the
potential to both enable previously non-pathogenic organisms to exploit crops as pathogens and
enhance the activity of opportunistic pathogens that did not previously cause economically signif-
icant yield loss. Community-scale studies offer opportunities to simultaneously evaluate effects on
multiple pathogen groups and enable researchers to forecast how certain adaptive water use prac-
tices might drive pathogen emergence. For example, a community-scale study by Liu et al. (45) re-
vealed that although conversion from rainfed to irrigated wheat reduced Fusarium propagule loads
in soils and Fusarium haplotype diversity, irrigation use also increased Pythium propagule loads and
recovery of Pythium species from roots, suggesting that this adaptation may pose a greater risk of
losses from Pythium than Fusarium diseases. Recent advances in affordable metagenomics-based
community analyses offer new opportunities to predict how water use practices can change com-
munity profiles and pathogen dynamics, favoring emergence of new or opportunistic pathogens,
as illustrated by Del Castillo et al. (23). In addition, water scarcity adaptations can have signifi-
cant effects on connected food safety issues, including mycotoxin levels in livestock feed, pesticide
residues on plant products (associated with changes in pest pressure), and presence of human
pathogens on food (49, 58). Although critical to food security and human health, these aspects of
water use adaptation have been largely neglected, and impacts are unknown.
Need for Field-Based Studies Looking at Effects of Increased Soil Salts

Associated with Groundwater and Drip Irrigation Use
Irrigation practices, such as drip irrigation, deficit irrigation, and heavy reliance on saline ground-
water come with a risk of accumulating salts in the rootzone. Soil salinization can have direct
negative effects on plant growth and can also influence disease. Many controlled laboratory and
greenhouse studies have demonstrated that salt levels can enhance disease as well as pathogen
sporulation and survival (3, 14, 38, 72, 80). There are a few field studies to examine how diseases
are affected by increased soil salinity resulting from altered irrigation practices; most focus on
saline groundwater use and consistently indicate that saline groundwater increases impacts of var-
ious diseases, including Fusarium crown and rot of tomato (85), Fusarium crown rot of wheat (81),
and early blight of potato (A. solani) (52). Furthermore, as noted above, there is some indication
that high soil salt may compromise single-gene resistance to diseases such as Fusarium wilt of
·.•�-
tomato (36). Given the importance of soil salinity in the context of water scarcity and adaptation,

there is a major need to expand salinity–pathogen studies to assess realized effects of salt accu-
mulation associated with water use practices on a wider array of diseases and crops under field
conditions.
Improving the Rigor in Irrigation–Pathogen Interaction Science

Meaningful analysis of the impact of water use adaptation on diseases in the field requires an
accurate diagnosis of the disease or diseases affecting the crop under study and quantification of
incidence and/or severity of each disease separately. It is also important to report the currently used
name of each pathogen and disease. Disease and pathogen identity should never be inferred based
on symptoms alone. These efforts will likely require assistance from a diagnostician or extension
pathologist. Sequence-based analysis should be used to identify the pathogen, as morphology-
based identification is no longer sufficient to identify most fungi and sequence-based analysis has
been long required for bacteria and viruses. It can often be advantageous to employ controlled
studies to explicitly examine the effects of water use practices on a given disease. In this case, it
is important that crops be inoculated with the pathogen using an appropriate method and that
proper controls are included (e.g., mock-inoculated plants).
Many of the studies evaluating disease risks of deficit irrigation do not determine the extent to
which plants are subjected to water stress. A reduction in irrigation input does not necessarily mean
an induction of plant stress, as discussed above. It is critical to measure water stress in irrigation
studies because it provides valuable insight into treatment effects on host infectability and disease
resistance. For instance, Austin & Wilcox (6) found that, based on their findings that there was
no effect of reduced irrigation on canopy air temperature, grapevine leaf water potential or leaf
temperature, reductions in powdery mildew under deficit irrigation could not be explained by the
proposed mechanism of reduced stomatal conductance. Water stress can be measured at the plant
level (e.g., stem water potential or stomatal conductance) or the field level (e.g., Crop Stress Water
Index, Normalized Difference Vegetation Index). To understand water stress outcomes in turn
requires assessing the effect of the treatment on available soil water, because the effect of a given
reduction in irrigation inputs on plant stress is related directly to the soil water that is available
to supplement crop ET needs (30). Soil moisture can be measured gravimetrically, volumetrically,
or based on soil matric potential, with soil matric potential being the most informative indicator
of plant water availability. If the objective is to reduce water inputs, each treatment should also be
assessed for changes in applied irrigation volume (typically using a flow meter).
Although physiological plant stress metrics provide inferential support for effects on plant dis-
ease resistance, it cannot be assumed that reductions in water inputs translate to changes in plant
defense responses (e.g., changes in plant biochemistry, protein production, or expression of genes
which regulate host defenses). To understand the mechanistic basis for disease development out-
comes, future research efforts would benefit from more in-depth studies of plant defense regula-
tion under relevant water use adaptation scenarios. It is often necessary to conduct such studies un-
der controlled conditions to ask more finessed questions and partition the effects of pathogens and
water stress more readily. However, it is always desirable to validate results in field studies to under-
stand how pathogen–water use interactions play out in their effects on plant health and crop yields.
Lastly, there is a need for more studies assessing the long-term, cumulative effects of adap-
tive water use practices on diseases. The vast majority of studies have been conducted over sin-
gle growing seasons and thus cannot account for the cumulative effects of changes in water use
practices over time. For example, cumulative effects of conversation from flood to buried drip
irrigation of California vegetables include soil salt accumulation and repeat planting of the same
·.•�-
crops in multiple years to avoid the high costs of more frequent installation associated with annual
16.14 Swett

crop rotation; these effects likely influence disease and cannot be captured in single-year studies.
Thus, multi- and single-year analyses likely result in highly disparate outcomes. As one example
illustrating this, in an eight-year study of drip, microjet, and sprinkler irrigation of apples, there
were no significant effects of irrigation treatment on Phytophthora root rot severity in years one
and two, but in years three to eight, disease severity was significantly lower under drip irrigation
(87). Such long-term studies will be critical to accurately identifying disease risks as well as dis-
ease management opportunities associated with changes in water resource management, thereby
strengthening the sustainability of regional and global food systems.
SUMMARY POINTS
1. Complexity underlies interactions between adaptive water use and plant diseases, but, in
many cases, disease outcomes can be explained once physiological effects of the water
use practice and biology of the pathosystem are understood.
2. To assess and interpret disease–water use interaction outcomes requires assessing the
effects on both plant and soil water status as well as the effects on both pathogen biology
(dispersal, survival, infection) and plant health (disease incidence and severity, yield). To
properly translate to water use adaptation, changes in applied irrigation volumes should
also be measured. To capture realized disease impacts, long-term study of cumulative
water-use adaptation effects are critical.
3. Some adaptive water use practices can increase disease risk; these effects are currently
not considered in the vast majority of efforts to transition to water-efficient practices,
posing significant production risk.
4. Methods are needed to mitigate disease risks while enabling water resource management;
this may involve alterations in irrigation practice, cultivar use, and use of pesticides/
biopesticides, biostimulants, or other biological-based products. In certain scenarios,
there is potential for adaptive water use practices to also suppress disease, providing an
exciting water–disease co-management framework.
5. There remain critical research gaps in understanding interactions between adaptive
practices and the more cryptic components of the phytobiome, including latent pathogen
dynamics, effects in driving emergence of new pathogens, effects of non-pathogenic
members of the phytobiome on plant stress tolerance, and possible effects on food safety
(toxin production, human pathogens on food).
6. Research and extension efforts to serve stakeholders can be optimized through collabo-
rations between growers and diverse specialists, including plant pathologists, entomolo-
gists, irrigation scientists, soil scientists, and plant physiologists.
DISCLOSURE STATEMENT
The author is not aware of any affiliations, memberships, funding, or financial holdings that might
be perceived as affecting the objectivity of this review.
ACKNOWLEDGMENTS
I would like to thank Tom Gordon for his content and editorial assistance, Rick Bostock for pre-
·.•�-
disposition edits, Daniele Zaccaria for his input on water use concepts and terminology, Mallika

Nocco for input on irrigation concepts, and Thor Harris for editorial support. I would also like to
thank my past collaborators within the SCRI Clean Water group for initiating and greatly con-
tributing to my irrigation science education. I am grateful to the Annual Review of Phytopathology
editorial board, and notably Karen Garrett, for inviting me to contribute this work.
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cls June 15, 2020 19:1

Annual Review of Phytopathology

Annu. Rev. Phytopathol. 2020. 58:16.1–16.20 Keywords

Copyright © 2020 by Annual Reviews. Abstract

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plant pathosystems. This includes developing water–disease co-management strategies as well as

WATER SCARCITY CHALLENGES IN AGRICULTURE: AN OVERVIEW

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REALIZED EFFECTS OF ADAPTIVE WATER USE PRACTICES

www.annualreviews.org • Plant Disease and Water Scarcity 16.3

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Aboveground pathogens Belowground pathogens

Gravitational/sprinkler drip irrigation (Water- (Dry-

Full reduced irrigation

How Conversion from Rainfall-Only to Supplemental Irrigation Affects

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www.annualreviews.org • Plant Disease and Water Scarcity 16.5

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How Changes in Irrigation Management Strategies Can Affect Disease

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MECHANISMS UNDERLYING INTERACTIONS BETWEEN WATER

www.annualreviews.org • Plant Disease and Water Scarcity 16.7

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Factors That May Favor Disease

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on potato tubers, which is caused by the zoospore-producing plasmodiophoroid Spongospora sub-

Factors That May Suppress or Have No Effect on Disease

www.annualreviews.org • Plant Disease and Water Scarcity 16.9

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TOWARD A DISEASE–WATER CO-MANAGEMENT FRAMEWORK

Incorporating Parameters of Irrigation Management into Plant Disease

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Pesticide and Biological Product Use to Manage Effects of Water

Use Adaptation on Disease

Shifting Crop and Cultivar use to Manage Effects of Water Use

www.annualreviews.org • Plant Disease and Water Scarcity 16.11

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ADVANCING THE FIELD OF PLANT DISEASE–IRRIGATION

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Understanding How Changes in Water Use May Drive Emergence

Need for Field-Based Studies Looking at Effects of Increased Soil Salts

www.annualreviews.org • Plant Disease and Water Scarcity 16.13

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Improving the Rigor in Irrigation–Pathogen Interaction Science

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www.annualreviews.org • Plant Disease and Water Scarcity 16.15

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Work. Pap. 12–02

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www.annualreviews.org • Plant Disease and Water Scarcity 16.17

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www.annualreviews.org • Plant Disease and Water Scarcity 16.19

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