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Chapter 4 Medicinal and aromatic plants that are toxic 193
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Part II: Effect of stress factors on medicinal
and aromatic plants
Gülen Özyazıcı✶ and Negar Valizadeh
Chapter 5 Impact of drought stress on the medicinal and aromatic plants’ biochemistry
Abstract: Today, medicinal plants have a wide range of uses not only in the field of
health but also in different sectors such as perfumery, cosmetics, food industry and phytotherapy. In addition to their potential to treat diseases, these plants are also valued for their contributions to different industries. Medicinal plants with various phytochemical compounds such as secondary metabolites, for example, al kaloids, terpenoids, phenols, steroids, flavonoids, tannins, glycosides, volatile oils and aromatic compounds are exposed to abiotic stress such as drought. Drought, one of the abiotic factors, causes a decrease in plant height, plant leaf area, number, and such other decreases, thus not only changing the plant structurally and anatomically, but also leading to fluctuations in its chemical components. The quality and quantity of the components of secondary metabolites synthesized by the plant help to cope with the harmful effects of stress for adaptation and defense. Numerous studies have shown that drought affects the accumulation of secondary metabolites in different organs of the plant and causes an increase or decrease in different plant species and even in different species of the same genus. Since the main aim in medicinal plants is not only to increase the yield of seeds, leaves, and flowers but also to increase the production of active ingredients such as essential oils, the cultivation and manage ment of medicinal plants under stress conditions is different from other crops. This study provides a summary of recent literature covering the studies on the morphol ogy, physiology, and biochemistry of medicinal and aromatic plants under drought stress.
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Keywords: water deficit, drought stress, plant secondary metabolites, essential oil, morphology, plant physiology
Corresponding author: Gülen Özyazıcı, Department of Field Crops, Faculty of Agriculture, Siirt
University, Siirt, Türkiye, e-mail Negar Valizadeh, Research Division of Natural Resources, East Azerbaijan Agricultural and Natural Resources Research and Education Center, Agricultural Research, Education and Extension Organization (AREEO), Tabriz, Iran
: gulenozyazici@siirt.edu.tr, https://orcid.org/0000-0003-2187-6733
200 Gülen Özyazıcı and Negar Valizadeh

5.1 Introduction

Drought, which occurs as a natural result of climate change and is one of the abiotic stress factors, is one of the most important stress factors affecting agricultural produc­tion all over the world. When usable areas on earth are classified, drought, which is a natural stress factor, comes first with a share of 26% [1], followed by mineral stress with 20%, cold and frost stress with 15%, while 10% of the area is not exposed to any stress factor [2].
Drought is an important factor affecting the growth and metabolic activities of plant species. Drought conditions inhibit plant growth, disrupt plant-water balance, reduce water use efficiency, and negatively affect the physiological processes in the plant. Drought stress in plants results from disruption of water flow in the xylem. Ob­struction of water flow leads to a decrease in cell turgor pressure and, therefore, af­fects cell elongation and expansion. Important physiological developments such as cell growth and division, enzyme synthesis, and protein synthesis slow down [3]. As a result, growth and development are also significantly affected, and yield is reduced. Drought stress causes a decrease in plant development parameters such as plant height, leaf number, leaf size, root length, leaf area, etc. and these are the first visible symptoms of drought [4–6]. Research has shown that the responses of medicinal and aromatic plants to salinity and drought stress have common points [1]. In order to ac­cess water resources in drought stress situations, plants accelerate root development in the early stages of stress and, conversely, slow down shoot development [7]. Under drought conditions, reduction of plant leaf surface and reduction of transpiration in­crease tolerance to drought stress. In many plants, dry conditions accelerate the aging process and abscission of old leaves, while roots adapt by developing the root system to reach water in the deep layers of the soil [8, 9]. Moreover, responses such as short­ening of height, less fruit/seed/biomass yield, and changes in the reproductive process occur [10]. Examples of water stress reducing plant growth in medicinal and aromatic plants include Hypericum brasiliense Choisy [11]. Catharanthus roseus [12] and Bupleu- rum chinense DC plant species [13].
While drought generally has a negative effect on plant growth and development, there are studies on the positive effect of water deficit when it comes to the biosynthe­sis of secondary metabolites, enzyme activities and solute accumulation [14]. Some of these responses may be related to the plant’s ability to survive under restrictive con­ditions. Although the metabolic responses of most cultivated plants to water deficit or drought have been studied, studies on this subject in medicinal and aromatic plants are few or relatively new. Water is an important factor in the yield and development of medicinal and aromatic plants, as well as other plants. Unlike traditionally culti­vated crop plants, water and plant relationships in medicinal and aromatic plants are not fully understood. This section focuses on the effects of drought stress or water deficiency on the biochemistry of plants.
Chapter 5 Impact of drought stress on the medicinal and aromatic plants’ biochemistry 201

5.2 Effect of drought or water deficiency on the morphology of medicinal plants

Drought stress is one of the most important abiotic stresses that show dramatic changes in plant growth and yield. Drought occurs when the available water in the soil decreases. It also occurs due to inadequate rainfall and continuous water loss through the process of transpiration and evaporation [15]. Water availability is re­garded as a critical determinant influencing plant growth. The commercial medicinal value of an aromatic plant depends on the presence of secondary metabolites, which are impacted by water scarcity [16, 17]. Drought first manifests itself with reduced cell development, especially in the stem and leaves. Stem growth stops and leaf growth is also restricted. One of the first signs of water deficiency is a decrease in turgor pres­sure, which leads to inhibition of cell growth and development, especially in stems and leaves. In addition, nutrient uptake decreases under drought stress conditions due to decreased soil moisture, which causes a decrease in the diffusion rate of nu­trients from the soil matrix to the absorbent root surface, limiting leaf growth and development. As a result, leaf area, light interception, and overall photosynthetic ca­pacity are reduced [18]. The degree of stress tolerance varies from one plant species to another. Water stress is an important factor that limits the growth of the plant in the initial stages. It negatively affects the plant morphology of medicinal and aromatic plants. Dry matter accumulation decreases in all plant organs under drought stress, but different organs are affected to varying degrees. Therefore, plants growing in drought conditions are morphologically smaller than plants growing under normal conditions.
Drought stress leads to plant dehydration, stomatal closure [19], and restricted gas exchange, subsequently resulting in metabolic inhibition, reduced photosynthetic rates, and ultimately plant death [12]. However, a plant’s ability to survive under such stressful conditions depends on its species, growth stage, and the duration and inten­sity of water deficit [12]. Moisture deficiency triggers various structural changes in plants that are crucial for responding to drought stress. These include morphological adaptations (such as reduced growth rates, development of deeper root systems, and alterations in the root-to-shoot ratio to avoid desiccation [20] as well as physiological and metabolic responses (such as stomatal closure, accumulation of antioxidants, and the expression of stress-specific genes).
Drought stress reduces plant height, leaf length, leaf weight, leaf area, and fresh and dry weight in lemongrass species [21]. In a study investigating the effect of water deficit on the morphology of Salvia sclarea populations, it was reported that yield and yield components decreased under stress conditions [22]. In their study on various levels of drought stress on Plantago psyllium L., Achillea millefolium L., Salvia officina- lis L., Calendula officinalis L., and Matricaria chamomilla L. plants, it was reported that increasing drought stress led to a decrease in shoot weight and plant height com-
202 Gülen Özyazıcı and Negar Valizadeh
pared to nonstress conditions [23]. They reported that growth and yield decreased in thyme (Thymus vulgaris L.), Japanese mint (Mentha cordifolia Opiz.), and Mexican marigold (Tagetes minuta L.) under drought stress [24–26]. When water stress was ap­plied to Ocimum basilicum L. (sweet basil) at 100%, 85%, 70%, and 55% of field capac­ity, shoot branching was limited under severe drought stress (55% of field capac­ity) [27].
Water limitation reduces the yield of medicinal and aromatic plants in three ways: First, drought-induced leaf area expansion can be limited by temporary leaf wilting or premature leaf senescence. Secondly, it can limit the grain yield of medici­nal and aromatic plants by reducing radiation utilization efficiency, and thirdly, by reducing the harvest index. This can occur even without a strong reduction in total medicinal and aromatic plant dry matter accumulation if a short stress period coin­cides with the critical developmental stage around flowering [28]. Numerous studies have shown that grain yield can be significantly reduced as a result of water deficit during the reproductive period in coriander [29] and Mexican marigold [30]. Simi­larly, Petropoulos et al. [31] noted that leaf and root weight, and leaf number (35% of field capacity) were significantly reduced due to water stress. In the studies of Miao et al. [32], irrigation treatments in Tulipa edulis L., an important medicinal plant with anticancer properties, were control (80% of field capacity), continuous drought (50% of field capacity) and alternating wetting and drying (50% and 80% of field capacity, AWD). They found that persistent drought significantly inhibited plant growth and yield compared to the control. Khorasaninejad et al. [33] reported that mint plant is moderately tolerant to water stress (85% field capacity) because it does not have ad­verse effects on some growth parameters of this plant.
5.3 Effect of drought or water deficiency on
the physiology of medicinal plants
The response of plants to drought depends significantly on the drought severity, plant developmental stage, genotype, and physiological status [34, 35]. Under water stress, plants attempt to maintain the water potential of their tissues by closing their stomata and reducing water losses through transpiration. Stress-induced growth decline may result from a decrease in meiosis and photosynthesis as a result of stomata closure, or from a decrease in cell development brought on by a drop in turgor pressure [ Furthermore, plant growth and photosynthesis are reduced due to decreased CO els and increased formation and accumulation of ABA, proline, mannitol and radical scavenging compounds (ascorbate, glutathione, α-tocopherol, etc.), stress proteins, and mRNAs [ naked eye. The changes caused by drought in plants above and below the ground are presented in Figure 5.1.
37]. However, these are at the cellular level and cannot be seen with the
2
36]. lev-