The genus Ocimum L. belongs to the family Lamiaceae. It exhibits large morphological groups, comprising of 30 to 160 species owing to the ease of cross-pollination which has led to a large number of subspecies and varieties. Ocimum species are annual and perennial herbs/shrubs that are indigenous to Africa, Asia, Central, and South America, but extensively disseminated worldwide.
1. Introduction
These highly aromatic plants of the genus have long been established as economically important medicinal plants due to their essential oils that have medicinal, culinary, and perfumery applications [4][1]. In medicinal and aromatic plants, growth and essential oil production are influenced by various environmental factors, drought stress being one [5,6,7][2][3][4]. Apart from environmental conditions other multiple factors such as; geographical area of cultivation [8][5], cultivation cycle [9[6][7],10], cultivation condition [11][8], harvest year [12[9][10],13], cultivars or varieties [14,15,16][11][12][13], age and part of the plant part [17[14][15][16],18,19], salinity [20][17], essential oil extraction methods [21[18][19],22], and storage conditions [23[20][21],24], can also influence essential oil production and its composition. Moreover, a significant reduction of plant growth under drought stress is a well-known and widely reported phenomenon, Ocimum species are not exceptions. Thus, the effect of drought stress on fresh and dry herb weights of basil has been reported by many scholars. Water shortage conditions provoked lower shoot fresh (−48.3%) and dry weight (−50.6%) compared with non-stressed sweet basil plants [25][22]. A 34% dry mass reduction due to drought stress was also reported [26][23]. Furthermore, a reduction of the total herb fresh and dry weights of American basil due to drought stress were observed [27][24]. Similarly, several authors have reported a significant reduction of yield in sweet basil due to drought stress [27,28,29,30,31][24][25][26][27][28]. In addition, water supply also modifies polyphenol as well as the antioxidant capacities of plants. In line with that, a moderate water deficit enhanced the antioxidant capacity and the total polyphenol content in medicinal and aromatic plants [5,32][2][29]. However, its effect on essential oil content, essential oil composition, antioxidant activities, and polyphenol content often remains controversial. On one hand, increased essential oil content and enhanced antioxidant activities under drought treatment was reported on O. basilicum, O. americanum, and O. x africanum [5,33,34][2][30][31]. On the other hand, a review on ‘whether drought stress increases the biosynthesis and accumulation of plant volatiles,’ concluded that the available scientific evidence was not adequate to generalize [35][32]. Thus, further holistic studies to determine the optimal water supply for different species are paramount. The effect of drought stress on the growth characteristics and essential oil content of sweet basil (O. basilicum) is relatively well studied, however, its effect on different species of basil is not well known.
2. Influence of Drought Stress on Growth and Essential Oil Yield of Ocimum Species
Drought stress is one of the most important abiotic stresses which can affect the growth and accumulation of bioactive compounds in medicinal and aromatic plants. These changes are mainly related to altered metabolic functions, physiological, and morphological characteristics. Thus,
tresearch
is study ers aimed to describe the drought-triggered physiological, morphological, and biochemical changes on selected
Ocimum species. The measured relative water content (RWC) to evaluate the water status of
Ocimum species showed a reduction under the drought-stressed conditions. Similar findings were reported on sweet basil
[25[22][23][25],
26,28], lemon thyme
[43][33], and summer savory
[44][34]. The reduction of RWC resulted in turgor loss which strongly influences the plant growth and biomass production through its effect on cell expansion
[45][35]. A higher SPAD value was observed in the severe drought intensity. Previous studies also showed a higher SPAD value under water-deficit conditions in two
Plantago species
[46][36], summer savory
[44][34], and Jerusalem artichoke
[47][37]. The SPAD value is based on the light absorbance of leaves and thus turgor, leaf thickness, or leaf hairiness might influence the results. A higher SPAD value does not necessarily mean an increase in the chlorophyll content
[16][13].
Drought stress leads to turgor loss, trim down in photoassimilation, and metabolites that are required for cell division. As a consequence, impaired mitosis, cell elongation, and expansion result in reduced growth
[48,49][38][39]. Likewise,
wresearche
rs observed a significant reduction in all the measured production parameters (plant height, canopy diameter, leaf area, fresh root weight, and biomass production) under drought stress uniformly across all the tested species. Numerous scholars also reported the negative impact of drought stress on the production of sweet basil
[16[13][22][25][26][40],
25,28,29,50], O. x
africanum [33][30], and
O. americanum [27][24].
The influence of drought stress on secondary compound accumulation is highly debatable and a lot of contradictory results are reported. In this experiment,
wresearche
rs observed that, regardless of the drought intensity, the total polyphenol content (
O. x
africanum and
O. americanum), essential oil content of
O. x
africanum, and the essential oil composition ratios of the major compounds remained unchanged, whereas drought stress led to lower EO yield and antioxidant capacity in all species. Drought stress slightly enhanced the glandular hair densities of the basil species and the TPC of
O. basilicum ‘Genovese’. Other scholars, however, have stated that trichome density is mainly determined by ontogenesis and species
[51,52][41][42].
WResearcge
rs observed that water supply can also significantly influence the glandular hair density. On one hand, no changes in the essential oil content on five Lamiaceae species (
Lavandula latifolia, Mentha × piperita, Salvia lavandulifolia, Thymus capitatus, and
Thymus mastichina) were demonstrated
[53][43]. On the other hand, a higher EO content in a drought condition was observed on sweet basil
[7,26,54,55][4][23][44][45]. Moreover, a water-deficit also increased the essential oil content of
O. x
africanum while the essential oil production tended to be lower in the drought treatments
[33][30]. Although a slight essential oil content enhancement was reported by many scholars, drought stress significantly influences the essential oil production as a result of lower biomass production in water-deficit treatments.
The genus
Ocimum is characterized by great variability in both morphology and chemical composition due to polyploidy, aneuploidy, and inter-and intraspecific hybridizations
[15,56,57][12][46][47]. The studied species also showed variation in morphology and chemical compositions under optimum water supply. The sweet basil cultivar Genovese had an upright growth, large leaf area, higher biomass yield, and more linalool.
O. x
africanum produced more essential oil, glandular hair density, TPC, and AOC. The African basil accumulated more 1,8-cineole and camphor. Whereas the third species,
O. 10mericanum, had a spreading growth habit, narrow leaves, and higher ratios of neral and geranial.