D ROU G HT

REVIEW (8). In most instances, crops experience mod-

erate droughts caused by prolonged precipi-
The physiology of plant responses to drought tation deficits, reduced groundwater levels,
and/or limited access to water supplies, lead-
Aditi Gupta, Andrés Rico-Medina, Ana I. Caño-Delgado* ing to substantial losses in overall yield. There-
fore, investigating the mechanisms of how a
Drought alone causes more annual loss in crop yield than all pathogens combined. To adapt to moisture plant sustains its growth during moderate
gradients in soil, plants alter their physiology, modify root growth and architecture, and close stomata drought and devising strategies to improve
on their aboveground segments. These tissue-specific responses modify the flux of cellular signals, resulting plant health during such periods can provide
in early flowering or stunted growth and, often, reduced yield. Physiological and molecular analyses of the solutions for future food security. Understand-
model plant Arabidopsis thaliana have identified phytohormone signaling as key for regulating the response ing the response of cellular signaling to water
to drought or water insufficiency. Here we discuss how engineering hormone signaling in specific cells and shortage is key for shedding light on these
cellular domains can facilitate improved plant responses to drought. We explore current knowledge and modern agricultural problems (12). Here we
future questions central to the quest to produce high-yield, drought-resistant crops. explore how water availability cues cell and
tissue growth patterns and how these patterns

D
are coordinated in the whole plant to improve
rought is a misfortune for agriculture, referred to as the drought survival rate. From drought resistance without loss of yield. Over-
humanity, and livestock alike (1). Climate the perspective of molecular biology, cellular expression of drought-responsive genes often
change is leading us toward a hotter, water loss marks the first event of drought results in growth deficits and yield loss. Tissue-
more parched world (2). There is an stress. At the cellular level, drought signals or time-specific expression of drought-response
urgent need to produce high-yielding promote production of stress-protectant metab- traits may improve drought response without

Downloaded from http://science.sciencemag.org/ on April 16, 2020

plants that use water more efficiently than olites such as proline and trehalose, trigger the depressing yield. A combination of strategies
their present-day counterparts (Fig. 1A). In the antioxidant system to maintain redox homeosta- may boost agricultural yields despite increased
past decade, global losses in crop production sis, and deploy peroxidase enzymes to prevent water insecurity.
due to drought totaled ~$30 billion. Global pop- acute cellular damage and membrane integrity.
ulation rose from 5 billion inhabitants in 1990 Factors such as the extent of water stress and Traits for improving drought resistance
to more than 7.5 billion presently and is pre- the plant organ in which the stress is sensed also During drought spells, plant systems actively
dicted to rise to 9.7 billion to 10 billion by 2050 trigger specific signaling responses, including but maintain physiological water balance by (i) in-
(3), at which time 5 billion people are projected not limited to abscisic acid, brassinosteroids, and creasing root water uptake from the soil, (ii)
to be living in water-scarce regions (Fig. 1B) (4). ethylene phytohormone pathways (8–11). reducing water loss by closing stomata, and
Despite the moderate increase in global arable Drought’s impact on agriculture depends (iii) adjusting osmotic processes within tissues
land, an additional 1 million ha will be needed on the degree and duration of the reduced (13). Activated stress response pathways include
to ensure food security (Fig. 1C) (5). In addition, precipitation and soil water gradients, as well phytohormone signaling as well as antioxidant
water demand for agriculture could double as on plant species and developmental stages and metabolite production and mobilization (11).
by 2050, whereas the availability of fresh water
is predicted to drop by 50%, owing to climate
change (Fig. 1D) (6). Certainly, plant biotech-
nology holds one of the promises to meet A
the societal demand for increased global crop Duplicate
production. crop production
Water is crucial for plant survival, and water Year 2020 Year 2050
deficits limit plant growth. However, plants
have strategies to prevent water loss, balance
optimal water supply to vital organs, maintain
cellular water content, and persevere through B C D
periods of drought. The ability of a plant to 10 2.5 5
Billions of cubic meters

sense the water-deficiency signal and initi-

Billions of hectares
Billions of people

8 2 4
ate coping strategies in response is defined as
drought resistance. Drought resistance is a 6 1.5 3
complex trait that proceeds through several
mechanisms: (i) escape (acceleration of plant 4 1 2
reproductive phase before stress that could 2 0.5 1
hinder its survival), (ii) avoidance (endurance
with increased internal water content and 0 0 0
1990 2020 2050 1990 2020 2050 1990 2020 2050
prevention of tissue damage), and (iii) tolerance
(endurance with low internal water content Fig. 1. Past, present, and future of global climate, agriculture, and food security. (A) Most scenarios
while sustaining growth over the drought predict that water scarcity will increase in the coming years. With the world’s population continuously
period) (7). After a period of drought, the per- growing, crop production must also increase to fulfill civilization’s basic needs. For this purpose, plants must
centage of viable plants upon rewatering is become more water efficient. (B) Estimated world population for the 1990–2050 time period. The arrow
indicates the estimated number of people living in water-scarce areas. (C) Global arable land for agriculture
for the 1990–2050 time period. The arrow indicates the predicted demand for arable land to ensure food
Department of Molecular Genetics, Centre for Research in security, given the current rates of crop production per hectare. (D) Global freshwater demand for agriculture
Agricultural Genomics (CRAG), CSIC-IRTA-UAB-UB, Campus
UAB (Cerdanyola del Vallès), 08193 Barcelona, Spain. for the 1990–2050 time period. The arrow indicates the predicted decline in freshwater availability for
*Corresponding author. Email: ana.cano@cragenomica.es agriculture, given the current trends for climate change and precipitation.

Gupta et al., Science 368, 266–269 (2020) 17 April 2020 1 of 4

 Roots respond to changes in soil moisture ways (Fig. 3) (21). Many existing schemes to ciency and drought resistance in A. thaliana,
at the cellular scale and with the entire root improve water use efficiency and drought re- tomato, and wheat (27). Thus, computational
system architecture. The root stem cell niche, sistance engage the ABA pathway. design combined with experimental biology
meristem, and vasculature each coordinate re- Genetic engineering to improve the function led to identification of a small molecule that
sponses to drought (Fig. 2, A and B). During of PYR/PYL/RCAR (Pyrabactin Resistance 1/ can mitigate the effects of drought on crop
periods of water scarcity, the root system ar- PYR1-Like/Regulatory Component of ABA Re- yields.
chitecture undergoes morphological changes ceptors) and SnRK2 (SNF1-related protein Brassinosteroid hormones also regulate
to enhance its ability to absorb water and nu- kinase 2) and repress the negative regulator drought response through signaling com-
trients (9, 10). These modifications can be PP2C (clade A type 2C protein phosphatase) ponents linked to the ABA response pathway
traced to coordinated cell division, elongation, has resulted in improved water use efficiency in (Fig. 3) (28, 29). Brassinosteroid signaling
and differentiation events in the root apex. In plants such as A. thaliana and wheat under negative regulator BRASSINOSTEROID-
the pursuit of moisture, root systems grow controlled laboratory growth conditions and INSENSITIVE 2 (BIN2) is dephosphorylated
differentially and adapt their architecture to greenhouses (22–25). A regulatory network by ABA INSENSITIVE1 (ABI1) and ABI2. ABA
be either deep or shallow (Fig. 2C). Longer and of ABA pathway genes, a hierarchy of ABA- activates BIN2 by inhibiting the activity of
deeper roots with reduced branching angles related transcription factors, and signaling ABI1 and ABI2 (30). BIN2 phosphorylates
can efficiently capture water from soil that is
dry at the surface but retains moisture in deep
layers. By contrast, shallower root architec- A C E
tures are more beneficial for maximizing water
capture from the soil surface in regions of low
precipitation (9). Roots that encounter a soil

Downloaded from http://science.sciencemag.org/ on April 16, 2020

environment with nonhomogeneous water dis-
tribution display hydropatterning by favoring
lateral root emergence toward soil patches
with higher water content, a process that is also
mediated by auxin signaling (9, 14). Another
adaptive response to nonhomogeneous distribu-
tion of moisture through soil is hydrotropism
(Fig. 2D), in which root tips grow toward zones 50 µm 100 µm
with higher water content to optimize the root
system architecture for water acquisition (15). B D F
Stomatal closure is a more rapid defense
against dehydration (Fig. 2, D and E). Stomatal
pores on leaf surfaces open or close accord-
ing to the turgidity of the surrounding guard
cells. The turgor-driven shape changes of guard
cells are affected by the cell wall structure,
plasma membrane, tonoplast properties, and
ia
ed
m

cytoskeletal dynamics (16). Plant vascular tis-

ic
ot

sues, the xylem and phloem, transmit water

m
os

availability signals from roots to shoots and

gh
Hi

transmit photoassimilates from shoots to roots, 50 µm 1 cm 20 µm

respectively (17). Development of these inner
vasculature tissues also affects drought re- Fig. 2. Root and shoot traits that account for drought resistance. (A and B) Plants initially sense
sistance. Crop yield becomes most vulnerable drought through their roots, where particular cell types (shown in blue)—such as stem cells, cortical cells,
if the drought occurs during a plant’s repro- and vascular cells—mediate adaptive responses toward water limitations. Roots can modulate their
ductive phase. In Arabidopsis thaliana, early system architecture to (C) maximize access to superficial humidity or delve into deep humid soil layers,
flowering associated with drought escape is as well as to (D) bend toward more humid soil zones (hydrotropism). (E and F) In aboveground plant organs
linked to phloem loading and transport of the such as leaves and stems, stomata work actively against dehydration. In water-limiting conditions,
photoperiod-dependent protein FLOWERING stomata remain closed to reduce water loss.
LOCUS T (FT) from leaves to the shoot apical
meristem (18).

Phytohormones to combat drought feedback were identified among ABA-mediated SnRK2s and activates the downstream path-
The hormone abscisic acid (ABA) regulates stress responses to drought (26). Engineering way (31). ABA signals can also converge with
plant responses to dehydration and optimizes the ABA receptor PYR1 for heightened sen- the brassinosteroid pathway at the level of
water use. Dehydration signals stimulate local sitivity toward the preexisting agrochemical downstream transcription factors (Fig. 3).
production of ABA in different plant organs. mandipropamid resulted in improved drought BRI1-EMS-SUPPRESSOR 1 (BES1) inhibited
However, ABA production is more efficient resistance in A. thaliana and tomato (22). Vir- ABA induction of a drought-related transcrip-
in the leaf mesophyll cells than in the root tual screening for ABA receptor agonists led to tion factor RESPONSIVE TO DESICCATION 26
tissues (19). The accumulated ABA then ac- the identification of a bioactive ABA mimic (RD26) (32). RD26 shows reciprocal antag-
tivates downstream signaling components called opabactin. This small molecule can onism with brassinosteroid by modulating
(20). ABA executes its function during stress enhance ABA receptor activation and down- BES1-regulated transcription and inhibiting
by mediating signal cross-talk with other path- stream signaling to improve water use effi- brassinosteroid-regulated growth (33). WRKY46,

Gupta et al., Science 368, 266–269 (2020) 17 April 2020 2 of 4

 D ROU G HT

-54, and -70 belong to another class of tran- ulates local auxin transport by affecting the for drought adaptation. In rice, the auxin-
scription factors that interact with BES1 to pro- homeostasis of the auxin efflux carrier PIN- inducible gene DEEPER ROOTING1 provides
mote plant growth while repressing drought FORMED 4 in root columella cells (Fig. 3). drought resistance by promoting a more ver-
responses (34). BIN2 can phosphorylate and Natural variation in EXO70A3 was correlated tical and deeper root system architecture (41).
destabilize WRKY54 to negatively regulate its with seasonal precipitation and conferred dif- Although auxin modulates root architecture
effect on the BES1-mediated brassinosteroid ferent adaptive root system architecture con- under stress (40, 41), hydrotropic root re-
response (35). BIN2 phosphorylates and acti- figurations under different rainfall patterns. sponses function relatively independent of
vates the ubiquitin receptor protein DSK2, In areas with high temperatures and irrigated auxin and involve ABA signaling in root elon-
which leads to BES1 degradation via autoph- soils, deeper root architectures proved better gation zones. Coordinated activity of ABA-
agy and coordinates plant growth and inducible MIZU-KUSSEI1 (MIZ1) and
survival under drought conditions. (36). SNF1-RELATED KINASE 2 (SnRK2.2)
An AP2/ERF transcription factor called ABA in root elongation zone cortical cells
TINY is another candidate that balances interprets water potential gradients in
brassinosteroid-mediated stress adap- soil environments (15, 42).
tation with growth. TINY interacts with PYR/PYL Brassinosteroid receptors regulate
RCAR Stomata
BES1 and antagonizes brassinosteroid- root hydrotropic responses (Fig. 3).
closure
regulated growth. BIN2, on the other Overexpression of the vascular-enriched
hand, phosphorylates and stabilizes brassinosteroid receptor BRI1-Like3
SnRK2 ABA-responding
TINY to promote ABA-induced stomatal genes (BRL3) promotes root hydrotropic bend-
closure and drought resistance (37). ing. The brl1brl3bak1 triple mutant of
Thus, brassinosteroids as well as ABA the BRL3 signalosome shows a reduced

CL

Downloaded from http://science.sciencemag.org/ on April 16, 2020

E2
aid drought resistance. hydrotropic response, suggesting a role
BIN

5
2
for the vascular BRL3 receptor complex
Tissue-specific responses BRASSINOSTEROIDS
in regulating hydrotropic responses (43)
for drought resistance (Fig. 3). Activation of the BRL3 pathway
BIN2 BRL3
Stomatal closure preserves water in the Hydrotropism, in vasculature triggers accumulation of
plant. ABA content in leaves regulates osmoprotectant osmoprotectant metabolites such as
stomatal movement in response to metabolites proline, trehalose, and raffinose family
accumulation
water availability (25) (Fig. 3). Because oligosaccharides in plant roots in re-
BES1 WRKYs/ RD26
stomatal movements control CO2 in- TINY
sponse to water withdrawal, which
flux and transpiration, efforts to reduce improves drought resistance without
water loss via stomatal closure occur at penalizing growth (43) (Fig. 3). Phloem-
the cost of photosynthesis, growth, specific localization of BRL3 is likely to
5

AUXIN
E2

and yield (13). Therefore, most strat- be the determining factor for promot-
CL

egies to improve water efficiency and ing drought resistance without impair-
EXO70A3
drought resistance in plants focus on IAA ing yield (29, 43).
fine-tuning stomatal conductance In drought conditions, roots sense
and manipulating ABA signaling via Root system water scarcity from soil. The above-
stomata-specific promoters (38). With architecture ground segments of plants respond by
optogenetics, scientists have improved closing stomata in leaves, implicating
the responsiveness of the stomata and PIN4
CLE25 a systemic communication system. In
overcome the coupling of CO2 uptake times of drought, the CLE25 peptide is
with water vapor loss. Upon introduc- Drought produced in the roots and moves through
ing BLINK1 (a light-activated synthetic sensing the vasculature to plant leaves to drive
K+ ion channel) into guard cells, sto- ABA production by activating the bio-
mata became more synchronized with Fig. 3. Hormone signaling events underpinning drought. Sche- synthetic enzyme NCED3. This burst of
fluctuating light conditions (39). This matic representation of hormone signaling modules that control ABA synthesis leads to stomatal closure
manipulation improved the performance drought adaptation. Plants work against dehydration in organs and improved water balance, thereby
of the stomata and, consequently, growth such as leaves, vasculature, and roots. ABA, through SnRK2, promoting drought survival (44) (Fig. 3).
and productivity of the plant. Thus, activates a variety of genes that trigger stomata closure and This insight into small-peptide signaling
water use efficiency was improved by improve water balance. When roots sense drought, the CLE25 in A. thaliana may help with identifica-
engineering the stomata to maximize peptide moves through the vasculature to the leaves, where it tion of similar mechanisms in crop plants
the amount of carbon fixed per unit of locally controls ABA biosynthesis and stomata closure. Brassinosteroids for root-to-shoot mobilization of stress
water lost. also play roles in regulating plant drought response. Brassinosteroid signals.
Improving water acquisition by roots pathways converge with ABA by activating SnRK2 through
can also improve plant performance downstream pathway component BIN2 and vice versa. Indepen- A view to the future
upon drought. In A. thaliana, the auxin dently of ABA, brassinosteroid receptors (BRI1, BRL1, and BRL3) Genetic traits that sustain crop plant
pathway modulator EXOCYST SUB- modulate hydrotropic responses in the roots. The vascular BRL3 growth under moderate drought may
UNIT EXO70 FAMILY PROTEIN A3 receptor coordinates plant growth and survival under drought come from multiple sources, including
(EXO70A3), which regulates root system stress by promoting the accumulation of osmoprotectant metabo- natural genetic variation in wild relatives
depth, was identified through genome- lites in the root tissues. Furthermore, noncanonical auxin responses or bioengineering. Traditional breeding
wide association mapping (40). EXO70A3, a via EXO70A3 and PIN4 can modulate root architecture patterning has been the main strategy for exploiting
component of the exocytosis system, is and depth to boost water absorption from the soil, thereby the genetic diversity of adaptive traits
expressed in root tips. EXO70A3 reg- improving drought tolerance. in natural alleles. The advent of genomic

Gupta et al., Science 368, 266–269 (2020) 17 April 2020 3 of 4

technologies and gene mapping tools such as egies to combat water scarcity. Discovering 7. S. Basu, V. Ramegowda, A. Kumar, A. Pereira, F1000Research
genome-wide association study (GWAS) and ways to ameliorate agriculture’s “thirst” will 5, 1554 (2016).
8. F. Tardieu, T. Simonneau, B. Muller, Annu. Rev. Plant Biol. 69,
precision genome editing with the CRISPR- ease competition for freshwater resources, 733–759 (2018).
Cas9 system has been instrumental for the even as the world’s population grows. 9. J. R. Dinneny, Annu. Rev. Cell Dev. Biol. 35, 239–257 (2019).
10. R. Rellán-Álvarez, G. Lobet, J. R. Dinneny, Annu. Rev. Plant Biol.
generation of alleles that can improve plant
67, 619–642 (2016).
yield and performance under various stresses. RE FERENCES AND NOTES 11. J. Bailey-Serres, J. E. Parker, E. A. Ainsworth, G. E. D. Oldroyd,
Molecular studies that use tissue- or cell- 1. Food and Agriculture Organization of the United Nations (FAO), J. I. Schroeder, Nature 575, 109–118 (2019).
“The impact of disasters and crises on agriculture and food 12. Y. Eshed, Z. B. Lippman, Science 366, eaax0025 (2019).
specific promoters coupled with live micros- 13. J. Rodrigues, D. Inzé, H. Nelissen, N. J. M. Saibo, Trends Plant
security” (FAO, 2018).
copy techniques for real-time visualization of 2. FAO, “Proactive approaches to drought preparedness – Where Sci. 24, 652–663 (2019).
cellular processes are paving the way for anal- are we now and where do we go from here?” (FAO, 2019). 14. N. E. Robbins 2nd, J. R. Dinneny, Proc. Natl. Acad. Sci. U.S.A.
3. United Nations, Department of Economic and Social Affairs, 115, E822–E831 (2018).
ysis of drought response networks that can be 15. D. Dietrich et al., Nat. Plants 3, 17057 (2017).
Population Division, “World population prospects: the 2010 revision,
targeted by various approaches (Fig. 4). Small volume I: comprehensive tables” (ST/ESA/SER.A/313, United Nations, 16. T. N. Buckley, New Phytol. 224, 21–36 (2019).
molecules such as peptides or hormone ago- 2011); www.un.org/en/development/desa/population/publications/ 17. J. D. Scharwies, J. R. Dinneny, J. Plant Res. 132, 311–324 (2019).
18. F. Andrés, G. Coupland, Nat. Rev. Genet. 13, 627–639 (2012).
nists may be useful for fine-tuning drought pdf/trends/WPP2010/WPP2010_Volume-I_Comprehensive-Tables.pdf.
19. S. A. M. McAdam, T. J. Brodribb, Plant Physiol. 177, 911–917 (2018).
4. E. Koncagül, M. Tran, R. Connor, S. Uhlenbrook, “World water
response pathways while preserving yield in 20. R. Kalladan et al., Proc. Natl. Acad. Sci. U.S.A. 114, 11536–11541 (2017).
development report 2020 – Water and climate change” 21. T. Kuromori, M. Seo, K. Shinozaki, Trends Plant Sci. 23, 513–522 (2018).
agriculture. Together, research efforts aimed (SC-2018/WS/5, UNESCO WWAP, 2018). 22. S. Y. Park et al., Nature 520, 545–548 (2015).
at uncovering the physiology of plant responses 5. FAO, FAOSTAT Land Use Module, Arable land data, updated 23. P. Wang et al., Mol. Cell 69, 100–112.e6 (2018).
4 December 2019; www.fao.org/faostat/en/#data/EL. 24. R. Mega et al., Nat. Plants 5, 153–159 (2019).
to drought in model systems and translating 6. P. H. Gleick, Ed., in The Worlds Water 2000-2001: The Biennial 25. M. Okamoto et al., Proc. Natl. Acad. Sci. U.S.A. 110, 12132–12137
these findings to crops will deliver new strat- Report on Freshwater Resources (Island Press, 2000), p. 53. (2013).
26. L. Song et al., Science 354, aag1550 (2016).
27. A. S. Vaidya et al., Science 366, eaaw8848 (2019).

Downloaded from http://science.sciencemag.org/ on April 16, 2020

28. T. M. Nolan, N. Vukašinović, D. Liu, E. Russinova, Y. Yin,
Plant Cell 32, 295–318 (2020).
Natural Genetic 29. A. Planas-Riverola et al., Development 146, dev151894 (2019).
variation/ editing 30. H. Wang et al., Mol. Plant 11, 315–325 (2018).
GWAS 31. Z. Cai et al., Proc. Natl. Acad. Sci. U.S.A. 111, 9651–9656 (2014).
Traditional 32. Y. Chung, S. I. Kwon, S. Choe, Mol. Cells 37, 795–803 (2014).
breeding Bioengineering 33. H. Ye et al., Nat. Commun. 8, 14573 (2017).
34. J. Chen, Y. Yin, Plant Signal. Behav. 12, e1365212 (2017).
35. J. Chen et al., Plant Cell 29, 1425–1439 (2017).
36. T. M. Nolan et al., Dev. Cell 41, 33–46.e7 (2017).
37. Z. Xie et al., Plant Cell 31, 1788–1806 (2019).
38. F. Rusconi et al., J. Exp. Bot. 64, 3361–3371 (2013).
Optogenetics 39. M. Papanatsiou et al., Science 363, 1456–1459 (2019).
40. T. Ogura et al., Cell 178, 400–412.e16 (2019).
Chemicals/ 41. Y. Uga et al., Nat. Genet. 45, 1097–1102 (2013).
hormone 42. D. Shkolnik, R. Nuriel, M. C. Bonza, A. Costa, H. Fromm,
mimetics Proc. Natl. Acad. Sci. U.S.A. 115, 8031–8036 (2018).
43. N. Fàbregas et al., Nat. Commun. 9, 4680 (2018).
44. F. Takahashi et al., Nature 556, 235–238 (2018).
Biostimulants AC KNOWLED GME NTS
Small
peptides Funding: A.I.C.-D. has received funding from the European Research
Council (ERC) under the European Union’s Horizon 2020 research
and innovation programme (grant agreement 683163). A.I.C.-D. is a
Fig. 4. The promise of overcoming drought in agriculture. Genetic strategies provide solutions to counteract recipient of a BIO2016-78150-P grant funded by the Spanish Ministry
of Economy and Competitiveness and Agencia Estatal de Investigación
drought and can be used to develop drought-smart crops. Natural allelic variations in plants can be selected to (MINECO/AEI) and Fondo Europeo de Desarrollo Regional (FEDER). A.G.
improve drought resistance and yield. Traditional breeding approaches have selected drought characteristics to has received funding by a postdoctoral fellowship from the “Severo
obtain more-resistant crops. Advancements in gene mapping tools such as GWAS can explore the genetic diversity Ochoa Programme for Centers of Excellence in R&D” 2016–2019 from
the Ministerio de Ciencia e Innovación (SEV-2015-0533). A.G. and A.R.-M.
of drought resistance traits in natural alleles with nucleotide-level precision. Genetic engineering of drought
have received funding from ERC-2015-CoG–683163 granted to the
response markers at the spatiotemporal scale and precise genome editing with tools such as CRISPR-Cas9 have A.I.C.-D. laboratory. A.R.-M. is a predoctoral fellow from Fundación
opened new horizons for developing crops with improved drought resistance, without sacrificing yield. Emerging Tatiana Pérez de Guzmán el Bueno. This work was supported by the
CERCA Programme/Generalitat de Catalunya. Competing interests: The
techniques such as optogenetics allow fine manipulation of cell- and tissue-specific responses to signaling and
authors declare no competing or financial interests.
therefore increase growth and plant resistance to drought. Small peptides, hormone mimics, and receptor agonists
can be used to design better agrochemicals and fine-tune drought resistance while preserving yield. 10.1126/science.aaz7614

Gupta et al., Science 368, 266–269 (2020) 17 April 2020 4 of 4

The physiology of plant responses to drought
Aditi Gupta, Andrés Rico-Medina and Ana I. Caño-Delgado

Science 368 (6488), 266-269.

DOI: 10.1126/science.aaz7614

Downloaded from http://science.sciencemag.org/ on April 16, 2020

ARTICLE TOOLS http://science.sciencemag.org/content/368/6488/266

RELATED http://science.sciencemag.org/content/sci/368/6488/254.full
CONTENT
http://science.sciencemag.org/content/sci/368/6488/226.full
http://science.sciencemag.org/content/sci/368/6488/256.full
http://science.sciencemag.org/content/sci/368/6488/261.full
http://science.sciencemag.org/content/sci/368/6488/270.full
http://science.sciencemag.org/content/sci/368/6488/274.full
http://science.sciencemag.org/content/sci/368/6488/238.full
http://science.sciencemag.org/content/sci/368/6488/250.full
http://science.sciencemag.org/content/sci/368/6488/251.1.full
http://science.sciencemag.org/content/sci/368/6488/251.2.full
http://science.sciencemag.org/content/sci/368/6488/252.full
http://science.sciencemag.org/content/sci/368/6488/314.full
REFERENCES This article cites 38 articles, 14 of which you can access for free
http://science.sciencemag.org/content/368/6488/266#BIBL

PERMISSIONS http://www.sciencemag.org/help/reprints-and-permissions

Use of this article is subject to the Terms of Service

Science (print ISSN 0036-8075; online ISSN 1095-9203) is published by the American Association for the Advancement of
Science, 1200 New York Avenue NW, Washington, DC 20005. The title Science is a registered trademark of AAAS.
Copyright © 2020 The Authors, some rights reserved; exclusive licensee American Association for the Advancement of
Science. No claim to original U.S. Government Works