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Chapter 5 Impact of drought stress on the medicinal and aromatic plants’ biochemistry 213
ter, longest lateral shoot length, root length, shoot yield, and dry leaf yield were observed under T1 (unstressed conditions). In contrast, the highest levels of proline, soluble carbohydrates, and root development were recorded under T4 (40% field capacity).
While drought stress negatively affected most morphological traits and flowering shoot
yield, it led to an increase in root length, proline accumulation, and soluble carbohydrates in flowering shoots.
The effect of drought stress on the quality and quantity yield of Thymus vulgaris
was evaluated under field and laboratory conditions [88]. The study that included five
different stress treatments showed that drought stress significantly affected plant
height, flowering shoot yield, oil percentage, oil yield, thymol percentage, carvacrol
percentage, chlorophyll a amount, chlorophyll b, proline, soluble sugars, sodium,
magnesium, iron, and relative water content. The highest (2.22%) and lowest (0.74%)
essential oil percentages were observed at 20% and 100% of the field capacity, respectively. The maximum thymol percentage was at 80% (42.37%), 60% (42.52%), and 40%
(41.4%) of the field capacity.
Irrigation causes significant changes in the morphological and biochemical properties of Ocimum species [89]. The same researchers reported that while fresh and dry
biomass yields increased in irrigated plants, there was no significant change in essential oil yield and composition of main compounds among the treatments. However, a
slight increase was observed in camphor, nerol, and trans-β-caryophyllene ratios. On
the other hand, drought stress increased EO content, polyphenol content, and antioxidant capacity. Moreover, drought stress had a positive effect on 1,8-cineole and eugenol ratios. Morphological and biochemical variations were also detected among basil
species. Accordingly, higher biomass and essential oil yield among the species were
obtained from O. basilicum and O. × africanum, respectively.
Plants respond to various abiotic and biotic signals that affect their growth and
development. Although the responses vary from plant to plant, the growth and development of medicinal plants, in short, their responses to environmental stresses depend on the genotype. In their study with 10 different fennel genotypes, Poudineh
et al. [90] stated that water stress has different effects on different varieties and
causes various physiological and biological changes in fennel plants, one of which is
the accumulation of reactive oxygen species (ROS) in the cell.
Torun et al. [91] investigated the physiological and biochemical responses of
ninety-day-old Hypericum perforatum seedlings by exposing them to three weeks of
drought. The results revealed that it decreased leaf length, relative water content, osmotic potential, chlorophyll fluorescence, increased lipid peroxidation, hydrogen peroxide, proline content, superoxide dismutase, catalase and glutathione reductase, and
decreased peroxidase and ascorbate peroxidase activities.
Basil plants changed the number and size of stomata, depending on the severity
of drought they were exposed to for 3 months, and accordingly, partial changes occurred in their phytochemical contents [62]. When drought levels were compared
with the control subject, obvious phytochemical changes were observed.
214 Gülen Özyazıcı and Negar Valizadeh
Baudoin et al. [92] applied five different irrigation regimes (severe over-irrigation,
moderate over-irrigation, standard irrigation, moderate under-irrigation, and severe
under-irrigation) to Oregano (thyme) and rosemary plants, and while Oregano phytochemical ratios increased significantly under moderate under-irrigation, there was no
change in rosemary phytochemical ratios. Researchers reported that the fact that the
phytochemical ratios of rosemary did not change under drought stress may be due to
some morphological characteristics of rosemary, and the plant’s ability to keep stomatal
opening under tight control throughout the day and its ability to develop tolerance to
stress by activating some mechanisms in its leaves.
In a study evaluating three irrigation regimes in ten black cumin (Nigella sativa L.)
genotypes, water stress increased the activities of carotenoids, proline, total soluble carbohydrates, malondialdehyde, hydrogen peroxide, and catalase and ascorbate peroxidase, but decreased the relative water content and chlorophyll content. These physiological changes varied according to the genotypes [93].
In a separate study, under lysimeter conditions at Shahid Sadoughi’s combating
desertification research station, Rad et al. [94] investigated the effect of three different
water constraints (100%, 70%, and 40% of field capacity) on Eucalyptus camaldulensis
Dehnh. Results revealed that mild drought stress resulted in increased essential oil
yield, water use efficiency, and 1,8-cineole production, but reduced or stopped the
production of many other compounds.
In a water stress study of Mexican marigold (Tagetes minuta L.) at 100%, 75%, 50%,
and 25% of field capacity, growth responses, oxidative stress indicators, and phytochemical variations were recorded in stressed and unstressed plants. Photosynthetic pigments and relative water content decreased in stressed plants, but malondialdehyde,
osmolyte compounds, and total phenol contents increased with increasing water limitation. Catalase, guaiacol peroxidase, ascorbate peroxidase, and polyphenol oxidase activities were also increased in stressed T. minuta plants, in response to drought stress.
Drought stress did not have a significant effect on the essential oil content of T. minuta,
but the essential oil composition was significantly affected. Drought stress changed the
proportions of essential oil components and induced the synthesis of new components,
including 1,8-cineole and germacrene D. T. minuta can resist water stress up to 75% of
its field capacity [26]. Thakur and Thakur [95] tested Chlorophytum borivilianum, Stevia
rebaudiana, Withania somnifera, and Andrographis paniculata plants under 50% water
deficit and different stress periods and showed different potentials in terms of growth,
yield, and physiological characteristics. The negative effect of stress on growth, yield,
photosynthetic rate, canopy temperature decrease, and chlorophyll fluorescence (Fv/
Fm) ratio was higher in
Stevia rebaudiana and Andrographis paniculata compared to
Chlorophytum borivilianum and Withania somnifera.
Zhang et al. [96] investigated the effects of different water stress levels on root
biomass, secondary metabolites and endogenous hormones in roots, relative water
content, and tissue density in leaves of Stellaria dichotoma L. var. lanceolata Bge. The
findings showed that in root biomass, total saponin content first increased and then
Chapter 5 Impact of drought stress on the medicinal and aromatic plants’ biochemistry 215
decreased with increasing drought intensity. The researchers reported that moderate
water stress (60–70% or 80–90% field capacity) was suitable for root biomass formation and secondary metabolite accumulation, which were influenced by endogenous
hormones and water status.
In anise plant from the Umbelliferae family, drought stress decreased yield and
yield components (seed yield, number of branches per plant, number of seeds, number of umbels, and thousand seed weight) and physiological traits such as chlorophyll
content, relative water content, quantum efficiency of photosystem II, and cell membrane stability, while increasing leaf temperature. Otherwise, moderate drought severity increased anise essential oil content, while severe drought decreased it [97].
Drought stress is a major environmental constraint that severely limits crop productivity. Water scarcity caused significant decreases in umbels per plant, umbels per
umbel, fruits per umbel, 1,000 fruit weight, biological yield, and finally fruit yield, despite increases in fruit essential oil content under moderate drought stress conditions
[98]. In their greenhouse study, Soltanbeigi et al. [99] investigated the effects of different irrigation regimes and nutrient sources on the growth parameters and essential
oil components of Salvia officinalis. Yield decreased significantly as drought stress increased, and essential oil content was observed in moderate and then severe drought
stress.
In another study, Mirniyam et al. [100] investigated the seed yield, essential oil
constituents, polyphenolic composition, and antioxidant capacity of ajowan (Trachy-
spermum ammi L.) populations under three (normal, moderate, and heavy) irrigation
regimes. The results revealed that both essential oil and seed yield showed significant
decreases as a result of water stress, while total phenolic and flavonoid contents increased under drought stress treatment.
An effective method to solve the water deficit problems in arid regions, which have
increased in recent years as a result of global climate change, is the development of
drought-resistant species. Shams et al. [101] conducted a study to develop droughttolerant ecotypes in Lallemantia royleana (Benth.) plants collected from Kalat in Khorasan Razavi province, Zakheh in Kurdistan province, Kondor in Alborz province, and
Jupar in Kerman province. Their studies revealed that drought-tolerant ecotypes produced greater dry matter and seed yields under drought conditions. Relative water content, photosynthetic pigment content, seed yield, seed oil amount, and omega-6 fatty
acid contents decreased under drought conditions in all ecotypes, while ascorbate peroxidase, catalase, superoxide dismutase and peroxidase activities, and phenol and proline amounts increased. Tavosi et al. [102] they examined the effect of plant characteristics of coneflower (Echinaceae purpurea) in drought conditions. Their findings showed
that drought stress caused a significant decrease in the growth characteristics of different coneflower, chlorophyll a, carotenoid, and chlorophyll b content. In addition, severe
drought stress (40% field capacity) caused a significant decrease in the phytochemical
compounds of coneflower; secondary metabolites were affected not only by genetics
but also by changing environmental factors.
216 Gülen Özyazıcı and Negar Valizadeh
Antioxidant activity of Cuminum cyminum L. seeds, one of the most common aromatic plants of Mediterranean cuisine, increased under dry conditions. While the essential oil content increased at moderate drought severity, it decreased as the drought
severity increased, and total phenol content also increased under drought conditions.
Moderate drought improved the number of umbels per plant and the number of umbels per umbel and seed yield of cumin seeds compared to normal conditions, but severe drought reduced it. This showed that cumin plant is moderately resistant to
drought [103].
Leaf area, and dry and fresh leaf weight were significantly decreased in Hibiscus
esculentus L under different irrigation regimes. On the other hand, protein content
decreased as a result of drought-affecting protein biosynthesis and degradation. Application of salicylic acid and ascorbic acid to plants under drought stress alleviated
the effects of stress [104]. Protein and sugar content of Satureja hortensis grown
under three different irrigation regimes were negatively affected. Drought stress affected protein biosynthesis, decreasing the amount of protein and sugar content due
to the photosynthetic process [105]. Similar to Hibiscus esculentus, Satureja hortensis
also alleviated the negative effects of drought. Antioxidant enzyme activities, essential
oil yield, and abscisic acid content of hyssop (Agastache foeniculum [Pursh] Kuntze)
were found to be high under drought conditions [106].
In Aloe vera (L.) Burm.f., under severe drought stress, the impairment of the ability of leaves to synthesize assimilates caused growth suppression, while mild drought
stress increased total phenolic and flavonoid content. Increasing leaf thickness, leaf
biomass, and gel production of the plant associated with mild drought severity increased. It also increased the photochemical activity in the leaves and changed the
amount of all secondary metabolites of vanillic acid produced. Mild water restriction
can be applied for secondary metabolite productivity and for better growth of aloe
plant [107].
Increased water stress in Chrysanthemum morifolium caused an increase in phenolic compounds such as chlorogenic acid, rutin, ferulic acid, quercetin, apigenin, and
luteolin. Investigating the expression of genes that play a role in the formation of
these metabolites under drought conditions and understanding the accumulation
mechanism of polyphenols against water stress may create new perspectives [108].
On the other hand, Mustafavi et al. [109] reported that many of the biochemical
properties of the valerian plant were significantly affected by water stress. The potassium, zinc, and iron contents of the leaves increased as the amount of available water
decreased to 70%, and the amount of these elements decreased as the level of drought
increased further. Interestingly, while the aboveground biomass and root biomass of
the valerian plant decreased with drought, its essential oil content increased. The fact
that belowground organ development and essential oil production are affected differ
ently by drought levels requires caution in irrigation in production.
The less studied S. dolomitic species of sage plant has gained importance due to
its antiplasmodial and anti-inflammatory properties. Moderate and severe drought in-
-
Chapter 5 Impact of drought stress on the medicinal and aromatic plants’ biochemistry 217
creases the production of sesquiterpenes, an important class of terpenoids for their
intended use, which has led to the application of controlled drought in the production
of secondary metabolites of this plant [110].
Drought and/or heat stress induced the accumulation of proline, sugars, glycine
betaine, and sugar alcohols (osmolites), including inositol and mannitol, in M. piperita
and C. roseus plants, while total phenol, flavonoid, and saponin contents decreased in
response to drought and/or heat stress, but the levels of other secondary metabolites
(including tannins, terpenoids, and alkaloids) increased under stress in both plants.
Researchers have emphasized that the application of abiotic stress (drought and/or
heat stress) could be a strategy to increase the content of therapeutic secondary metabolites of these plants [111].
Nanoparticles (NP) and growth regulators are increasingly being used to reduce
the negative effects of drought stress. In coriander (Coriandrum sativum L.) plants exposed to drought stress, chlorophyll content decreases, whereas total soluble sugar,
superoxide dismutase, and peroxidase activities increase [112]. When coriander plants
were sprayed with salicylic acid and silicon-NPs to reduce the negative effects of
drought, increases in chlorophyll content, total soluble sugar, and activity of antioxidant enzymes occurred. Moderate drought significantly increased total phenolic content and total flavonoid content, essential oil content, and essential oil yield with SiNPs. Foliar application of silicon nanoparticles was determined to be more effective
than salicylic acid for improving the antioxidant potential and EO efficiency of coriander plant. Similarly, Mahmoud et al. [113] reported that the application of silicon, zinc,
and zeolite nanoparticles only positively affected the morphological, physiological,
and biochemical properties of coriander plant under drought stress. In a study investigating the combined effects of drought stress and nanosilicon application on the
morphological traits and essential oil content and composition of hemp (Cannabis sat-
iva L.), maximum plant height, number of nodes, and number of flowering branches
were recorded in 1.5 mM nanosilicon and 100% field capacity application, while the
lowest fresh and dry above-ground biomass was recorded in severe drought stress
(40% field capacity [114]. Mild water stress (80% field capacity) and foliar application
of 1.5 mM nanosilicon provided the maximum essential oil content, while the highest
cannabidiol content in essential oil was detected in severe water stress (40% field capacity) and 0.5 mM nanosilicon application. The findings showed that nanosilicon application improved the morphological characteristics of the cannabis plant and
changed its biochemical content and components under dry conditions.
In cichory (Cichorium intybus), root growth and cumulative inulin yield decreases
as drought duration increases [115]. The percentage of total inulin in roots increased
under mild drought stress and decreased under severe drought stress. Bat et al. [116]
stated that drought stress decreased the leaf area, relative water content in leaf tissues, and membrane durability index of echinacea (Echinacea purpurea L.) plant, and
increased malondialdehyde level and ion leakage in leaf tissues, while it did not affect
the leaf chlorophyll ratio.
218 Gülen Özyazıcı and Negar Valizadeh
It was determined that water stress and temperature increase negatively affected
seed production in Fagopyrum tataricum [117]. The use of mycorrhiza and vermicompost is recommended under stress conditions. Mycorrhiza, applied to buckwheat
under different stress conditions, was effective on phytochemicals, while worm compost increased aboveground biomass and seed yield.
In chamomile (Matricaria recutita L.), drought conditions caused a decrease in
plant height, flower yield, shoot weight, and apigenin content, but had no significant
effect on oil content or oil composition, maintaining the potential for biomass production. Despite the decrease in the agronomic properties of chamomile, the phytochemical properties of the plant did not change, indicating that chamomile is a moderately
drought-resistant medicinal plant [118]. Shoot fresh and dry weight, root fresh weight,
and shoot length of rosemary (Rosmarinus officinalis) plant did not show any difference under drought stress conditions (75% field capacity) compared to normal conditions (100% field capacity) [119]. On the other hand, under drought stress conditions
at 75% of field capacity, root length increased, and root dry weight, leaf area, and leaf
number decreased significantly under drought conditions. In contrast to these
changes in roots and leaves, quercetin, trans-ferulic acid, hesperidin, eugenol, hesperetin, and rosmarinic acid amounts increased under drought stress at 25% field capacity. Kharazi and Asgharzadeh [120] reported that in Nigella sativa L., plant growth
traits decreased with increasing drought severity, but foliar salicylic acid application
alleviated the adverse effects of drought stress.
In Rosa damascena Herrm., grown in drought conditions, flower yield decreased,
and irrigation regime significantly affected essential oil yield and some components
in essential oil. Drought stress increased the amount of citronellol and geraniol in essential oil, and decreased the amount of nonadecane, eicosane, and heneicosan [121].
This situation proves that the components in the essential oil of R. damascena can be
changed and managed by water stress. The response of Damascus rose to drought and
the mechanisms that mediate this response are unknown. In a study conducted by
water-restricted R. damascena, it was determined that water stress significantly reduced the fresh and dry weights of the plant and all photosynthetic parameters, except leaf temperature [122]. Apoplastic water fraction did not change significantly in
response to water stress. R. damascena underwent an osmotic adjustment in response
to water stress, resulting from active accumulation of soluble carbohydrates and, to a
lesser extent, proline under mild stress and tissue dehydration (passive osmotic adjustment) under severe stress. Farahani et al. [123] showed that the quality and quantity of Rosa damascena could be increased by foliar application of potassium silicate
under water deficit stress equal to 50% and 25% of plant water requirement. Plant
biomass of Stevia rebaudiana, an economically important medicinal plant, decreased
after drought treatments [124]. The photosynthetic properties decreased by drought
included intercellular CO
, net photosynthesis, chlorophylls, carotenoids, and water
2
use efficiency, followed by the decrease in carbohydrates. Under water stress, reactive
oxygen species accumulated and hydrogen peroxide production increased in plants.
Chapter 5 Impact of drought stress on the medicinal and aromatic plants’ biochemistry 219
Drought stress also caused the accumulation of proline and glycine betaine. The results showed that carbohydrates and plant growth were reduced. These results indicate that water deficiency is an important criterion for Stevia plants used in vegetative parts.
In Lycoris aurea, grown under drought conditions, increased plant growth is restricted, causing an increase in the fresh weight of the bulb, a decrease in the chlorophyll content, and a decrease in the maximum net photosynthesis rate of the leaves
[125]. In contrast, galanthamine and lycorine alkaloids in the bulb increased. Mild
water stress increased the galanthamine and lycorine contents to the maximum level.
These results indicate that L. aurea has a water requirement during vegetative growth
periods, and plants should be subjected to mild water restriction in order to increase
their alkaloid content in advanced growth stages.
Interestingly, in the saffron plant (Crocus sativus L.), one of the most expensive
spice plants in the world, the increase in the severity of drought stress caused an increase in secondary metabolites (crocin, picrocrocin, and safranal) [126–128]. In contrast, the dry weight of corm decreased due to drought stress. Methyl jasmonate and
auxin applications were reported to have the potential to reduce the negative effects
of drought stress. However, more research is needed on this subject to understand
these effects comprehensively.
5.7 Conclusions
Drought affects the morphological, physiological, and biochemical characteristics of
medicinal plants. Medicinal plants respond to drought not only by decreasing yields
but also by changing the amount and content of secondary metabolites. This situation
causes medicinal plants to lose their economic importance. In drought or water deficiency, signals are transmitted from roots to leaves via xylem vascular bundles and
stomata partially close. This causes gas exchange in the cell to slow down, free radicals such as hydrogen peroxide and superoxide, and increase reactive oxygen species.
As a result, the biosynthesis of secondary compounds (such as volatile oil content,
phenols, terpenoids, alkaloids, glycosides, and flavonoids) decreases, cells and tissues
are damaged and, depending on the severity of drought, the death of the plant occurs.
Therefore, the development of drought-resistant varieties should be the primary goal
in breeding programs for medicinal, aromatic, and spice plants, whose secondary metabolites are of economic importance.
220 Gülen Özyazıcı and Negar Valizadeh
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