Effect of Different Abiotic Elicitors on the Production of Various Secondary Metabolites in Plants
Abstract
Plant secondary metabolites are having the great application in human health and nutritional aspect. Plant cell and organ culture systems are feasible option for the production of secondary metabolites that are of commercial importance in pharmaceuticals, food additives, flavors, and other industrial materials. The stress, including various elicitors or signal molecules, often induces the secondary metabolite production in the plant tissue culture system. The recent developments in elicitation of plant tissue culture have opened a new avenue for the production of secondary metabolite compounds. Secondary metabolite synthesis and accumulation in cell and organ cultures can be triggered by the application of elicitors to the culture medium. Elicitors are the chemical compounds from abiotic and biotic sources that can stimulate stress responses in plants, leading to the enhanced synthesis and accumulation of secondary metabolites or the induction of novel secondary metabolites. Elicitor type, dose, and treatment schedule are major factors determining the effects on the secondary metabolite production. The number of parameters, such as elicitor concentrations, duration of exposure, cell line, nutrient composition, and age or stage of the culture, is also important factors influencing the successful production of biomass and secondary metabolite accumulation. This chapter reviews the various abiotic and biotic elicitors applied to cultural system and their stimulating effects on the accumulation of secondary metabolites.
Keywords
- Cell culture
- elicitor
- organ culture
- secondary metabolites
- stress
1. Introduction
The total mankind is dependent on plants as a source of carbohydrates, proteins, vitamins, food, and shelter. Plants are studied for their important constituents and the nutritional factors for over decades. Along with the essential primary metabolites, higher plants are also capable to produce a number of low molecular weight compounds. A diverse group of organic compounds that are produced by plants to facilitate interaction with the biotic environment and the establishment of a defense mechanism are called as plant secondary metabolites [1–3]. The production of these metabolites is very low (less than 1% dry weight) and depends greatly on the physiological and developmental stage of the plant [4,5]. Plant natural products have been an important part of medicine throughout human history. In recent years, the use of herbal medicines has steadily increased worldwide [6]. With this increasing demand comes growing concerns about the safety and efficacy of herbal medicines. Although the potential for medicinal plants seems almost limitless, there are a few major obstacles that hinder large-scale utilization by the western medical system. Among them is the lack of reproducibility common in testing many plant extracts (up to 40%), which has limited the enthusiasm for developing plant-based pharmaceuticals [7]. Unlike standardized single-entity pharmaceutical drugs, herbal medicines consist of complex mixtures with multiple compounds responsible for therapeutic activity, making standardization difficult [8]. Further complicating the issue is the fact that plants, unlike synthetic medicines, are living organisms, with inherent biological variation [9]. Just because plant material originates from the same species, it does not necessarily mean that the chemical content will be identical. This lack of reproducibility may be due to two main factors, genetic variability and differences in growing conditions.
In addition, plants are a valuable source of a wide range of secondary metabolites, which are used as pharmaceuticals, agrochemicals, flavors, fragrances, colors, biopesticides, and food additives. Plants are producing new compounds and in future new chemical models are drawing for new drugs because the most of the plants chemistry is yet to be explored [10]. The characterization of molecular structures and chemical analysis helped us to pinpoint the activities of plants under controlled conditions. Although all these advancements, we still depend on the secondary metabolites of biological sources including pharmaceuticals [11].
Due to various agro alimentary, perfumes, flavors, colors, and pharmacological effects, the secondary metabolites are having extensive demand and various commercial preparations are available in the market. Besides, the appeal of using natural products for medicinal purposes is increasing, and metabolic engineering can alter the production of pharmaceuticals and help to design new therapies. The evolving commercial importance of secondary metabolites has in recent years resulted in a greater interest in secondary metabolism, particularly in the possibility of altering the production of bioactive plant metabolites [12]. Secondary metabolites are separated into nitrogen compounds (alkaloids, nonprotein amino acids, amines, alcamides, cyanogenic glycosides, and glucosinolates) and nonnitrogen compounds (monoterpenes, diterpenes, triterpenes, tetraterpenes, sesquiterpenes, saponins, flavonoids, steroids, and coumarins).
The plant tissue culture plays an important role in the rapid clonal propagation, regeneration of genetically manipulated superior clones, conservation of germplasm, production of secondary metabolites, and ex vitro conservation of valuable phytodiversity [13,14]. The plant, cell, tissue and organ culture techniques have come up with an escapable tool with the possibilities of acclaiming and supplementing the conventional method in plant breeding, plant improvement, and biosynthetic pathways. This technique has several potential applications in crop improvement, and efficient regeneration is a prerequisite in such improvement programs. The biotechnological production of secondary metabolites in plant cell and organ cultures is an attractive alternative to the extraction of the whole plant material [15]. In particular, plant–specific important compounds are obtained by using the plant cell and organ cultures [2]. The faster proliferation rates and shorter biosynthetic cycle of cell and organ cultures leads to have a higher rate of metabolism when compared to field grown plants [16]. Further, plant cell/organ cultures are under controlled conditions proliferates at their optimum growth rates when compared to the cultivated plants, which are facing environmental, ecological, and climatic variations. In recent years, various strategies have been developed for use in biomass accumulation and the synthesis of secondary compounds, such as strain improvement, optimization of medium, and culture environments, elicitation, precursor feeding, metabolic engineering, permeabilization, immobilization, and biotransformation methods, bioreactor cultures, and micropropagation [17]. The focus of the present chapter is the influence of abiotic and biotic elicitors on the secondary metabolite production in the in vitro cultured medicinal plants.
2. Classification of elicitors and secondary metabolite production via in vitro culture of medicinal plants
Stress is an important factor in determining the chemical composition and therapeutic activity of medicinal plants. Actively stimulating, or eliciting, the plant stress response to induce the desired chemical response is called elicitation, harnessing the connection between plant stress and phytochemistry. “Elicitor may be defined as a substance for stress factors which, when applied in small quantity to a living system, it induces or improves the biosynthesis of specific compound which do have an important role in the adaptations of plants to a stressful conditions” [18]. Elicitation is the induced or enhanced biosynthesis of metabolites due to addition of trace amounts of elicitors [18]. Several biotechnological strategies have been hypothesized and applied for the productivity enhancement, and elicitation is recognized as the most practically feasible strategy for increasing the production of desirable secondary compounds from cell, organ, and plant systems [19–21].
On the basis of nature, elicitors can be divided into two types abiotic and biotic (Figure 1). Abiotic elicitors comprise of substances that are of nonbiological origin and are grouped in physical, chemical, and hormonal factors. Biotic elicitors are the substances of biological origin that include polysaccharides originated from plant cell walls (e.g. chitin, pectin, and cellulose) and micro–organisms.
3. Abiotic elicitors
As mentioned above, the abiotic elicitors are categorized into physical, chemical, and hormonal elicitors. Abiotic elicitors have wide range of effects on the plants and in the production of secondary metabolites (Table 1).
|
|
|
|
|
Ozone (O3) |
|
Shoot | Rosmarinic acid | [163] |
|
Cell suspension | Hypericin | [164] | |
|
Cell suspension | Puerarin | [165] | |
pH |
|
Shoot | Bacoside A | [166] |
|
Hairy root | Withanolide A | [167] | |
|
Cell suspension | Withanolide A | [168] | |
Sucrose |
|
Seedling | Hypericin and pseudohypericin | [41] |
|
Cell suspension | Paclitaxel | [169] | |
|
Shoot | Bacoside A | [166] | |
|
Cell suspension | Withanolide A | [168] | |
Ultraviolet C |
|
Cell suspension | Stilbene | [34] |
Proline |
|
Callus and suspension | Steviol glycoside | [40] |
Polyethylene glycol |
|
Callus and suspension | Steviol glycoside | [40] |
|
Seedling | Hypericin and pseudohypericin | [41] | |
Jasmonic acid |
|
Shoot | Bacoside A | [170] |
|
Hairy root | Plumbagin | [88] | |
|
Cell suspension | Plumbagin | [86] | |
Methyl jasmonate |
|
Hairy root | Tanshinone | [94] |
|
Adventitious roots | Cryptotanshinone and tanshinone IIA | [78] | |
|
Cell suspension | Stilbene | [34] | |
|
Shoot | Bacoside | [96] | |
|
Shoot | Diterpenoid | [171] | |
|
Cell suspension | Silymarin | [172] | |
|
Hairy root | Tanshinone | [173] | |
|
Cell suspension | Gymnemic acid | [148] | |
|
Hairy root | Withanolide A, withanone, and withaferin A | [95] | |
|
Cell suspension | Andrographolide | [97] | |
|
Cell suspension | trans-Resveratrol | [91] | |
|
Root | Glycyrrhizic acid | [135] | |
Gibberelic acid |
|
Hairy root | Tanshinones | [174] |
|
Hairy root | Caffeic acid derivatives | [175] | |
Salicylic acid |
|
Hairy root | Tanshinone | [94] |
|
Cell suspension | Stilbene | [34] | |
|
Shoot | Digitoxin | [176] | |
|
Shoot | Hypericin and pseudohypericin | [177] | |
|
Cell suspension | Gymnemic acid | [148] | |
|
Hairy root | Withanolide A, withanone, and withaferin A | [95] | |
|
Root | Hyoscyamine and scopolamine | [178] | |
|
Adventitious root | Glycyrrhizic acid | [179] | |
Sodium salicylate |
|
Shoot | Carnosol | [180] |
Sodium chloride |
|
Embryogenic tissues | Vinblastine and vincristine | [55] |
Sorbitol |
|
Adventitious roots | Cryptotanshinone and tanshinone IIA | [78] |
Silver (Ag) |
|
Adventitious roots | Cryptotanshinone and tanshinone IIA | [78] |
|
Cell suspension | Resveratrol | [67] | |
|
Hairy root | Tanshinone | [173] | |
|
Hairy root | Atropine | [150] | |
Cadmium (Cd) |
|
Cell suspension | Resveratrol | [67] |
|
Root | Sesquiterpenoid | [79] | |
Cobalt (Co) |
|
Cell suspension | Resveratrol | [67] |
Copper (Cu) |
|
Shoot | Xanthotoxin | [181] |
|
Shoot | Bacoside | [170] | |
|
Root | Sesquiterpenoid | [79] |
3.1. Physical elicitors
Physical elicitors include light, osmotic stress, salinity, drought, and thermal stress.
3.1.1. Light
The light is a physical factor that can affect the metabolite production. Light can stimulate such secondary metabolites include gingerol and zingiberene production in
3.1.2. Osmotic stress
Osmotic stress (water stress) is an abiotic physical elicitor [35] and is one of the important environmental stresses that can alter the physiological and biochemical properties of plants and increase the concentration of secondary metabolites in plant tissues [36]. Proline acts as an osmolyte, as protective agent for cytoplasmic enzymes, as a reservoir of nitrogen and carbon sources for post–stress growth, or even as a stabilizer of the machinery for protein synthesis, regulation of cytosolic acidity and scavenging of free radicals [37]. However, the various roles of proline have been proposed, but the main role could be the osmotic adjustment in osmotically stressed plant tissues and the protection of plasma membrane integrity [38]. Polyethylene glycol (PEG) is an osmotic agent (nonpenetrating osmoticum) that has been used for induction of water stress in many plants [39]. The proline and PEG enhanced the production of steviol glycosides content in both callus as well as suspension culture of
3.1.3. Salinity
Salinity reduces plant growth and development and alters a wide array of physiological and metabolic processes [44,45]. Plants have developed complex mechanisms for adaptation to the osmotic, ionic, and oxidative stresses that are induced by the salt stress. Exposure to salinity is known to induce or stimulate the production of secondary plant products, such as phenols, terpenes, and alkaloids [46–48].
3.1.4. Drought stress
One of the most important abiotic stress is drought, which affect plant growth and their developmental process [56]. The available water in the soil is reduced to such critical levels, and atmospheric conditions add to the continuous loss of water; the situation is called drought stress. The high temperature in the environment and solar radiations add up the water deficit in the soil, which leads to drought stress. Drought stress tolerance is observed in all types of plants, but its extent varies from species to species [56]. Drought stress, which can also greatly reduce plant growth, can increase secondary metabolite content. Mild water stress significantly increased the content of the anti–inflammatory saikosaponins in
3.1.5. Thermal stress
Although thermal stress can greatly reduce plant growth and induce senescence, elevated temperatures (heat stress) or low temperatures (cold stress) have also been shown to increase secondary metabolite production. Temperature strongly influences metabolic activity and plant ontology, and high temperatures can induce premature leaf senescence [62]. Elevated temperatures increase leaf senescence and root secondary metabolite concentrations in the herb
4. Chemical elicitors
Heavy metals have become one of the main abiotic stress agents for living organisms because of their increasing use in the developing fields of industry and agrotechnics and high bioaccumulation and toxicity [67]. Although a lot of information is available concerning the effects of heavy metals on plant growth and physiology, much less is known regarding their effects on the production of secondary metabolites. Heavy metal–induced changes in metabolic activity of plants can affect the production of photosynthetic pigments, sugars, proteins, and nonprotein thiols. These effects can result from the inhibition of enzymes involved in the production of these natural products, likely through impaired substrate utilization [68]. Metals may alter the production of bioactive compounds by changing aspects of secondary metabolism [2]. Metals including Ni, Ag, Fe, and Co have been shown to elicit the production of secondary metabolites in a variety of plants [69].
An increased oil content up to 35% in
5. Hormonal elicitors
Various plant hormones have been extensively used in elicitation studies. The most studied, because of their key roles in the plant defense response, are jasmonic acid (JA) and SA and its derivatives.
5.1. Jasmonates
Jasmonates, including JA and MeJA, are a family of cyclopentanone compounds that modulate a wide range of plant responses [80,81] and act as effective elicitors to enhance secondary metabolites in in vitro cultures. They constitute an important class of elicitors for many plant secondary metabolic pathways, which are typically manifested by the elicitation of secondary metabolite biosynthesis when plants face particular environmental stresses [82]. JA is an important signal molecule of plant in response to wound and pathogen attack [83]. JA and its more active derivative MeJA can induce the production of a wide range of plant secondary metabolites such as rosmarinic acid, terpenoid indole alkaloid, and plumbagin in various cell cultures [84–86]. JA elicitation are reported to induce the production of rosmarinic acid in
5.2. Salicylic acid
Salicylic acid, well known for the systemic acquired resistance it induces in the plant response to many pathogens, can also elicit the production of secondary metabolites in plants [101,102]. SA with transgenic technology highly enhanced the production of tanshinones in
5.3. Gibberellic acid
Gibberellin (GA), a phytohormone, is also well known as an effective elicitor for the production of secondary metabolites [109].
6. Biotic elicitors
In the production of secondary metabolites from plants, the use of biotic elicitors had an important role (Table 2).
|
|
|
|
|
Chitin |
|
Shoot | Hypericin and pseudohypericin | [182] |
|
Cell suspension | Phenylpropanoid and naphtodianthrone | [183] | |
|
Cell suspension | trans-Resveratrol and viniferins | [91] | |
Pectin |
|
Shoot | Hypericin and pseudohypericin | [182] |
Dextran |
|
Shoot | Hypericin and pseudohypericin | [182] |
Yeast extract |
|
Adventitious roots | Cryptotanshinone and tanshinone IIA | [78] |
|
Cell suspension | Plumbagin | [86] | |
|
Cell suspension | Silymarin | [172] | |
|
|
Hairy root | Tanshinone | [184] |
|
|
Hairy root | Thiarubrine A | [134] |
|
|
Hairy root | Azadirachtin | [136] |
|
|
Root | Glycyrrhizic acid | [135] |
|
|
Cell suspension | Phenylpropanoid and naphtodianthrone | [183] |
|
|
Cell suspension | Phenylpropanoid and naphtodianthrone | [183] |
|
|
Cell suspension | Phenylpropanoid and naphtodianthrone | [183] |
|
|
Cell suspension | Gymnemic acid | [149] |
|
|
Cell suspension | Gymnemic acid | [149] |
|
|
Cell suspension | Gymnemic acid | [149] |
|
|
Cell suspension | Gymnemic acid | [149] |
|
|
Cell suspension | Gymnemic acid | [149] |
|
Hairy root | Atropine | [150] | |
|
|
Hairy root | Atropine | [150] |
|
|
Hairy root | Atropine | [150] |
|
|
Root | Glycyrrhizic acid | [135] |
6.1. Polysaccharide
The biotic elicitors have been utilized to increase secondary metabolite production in medicinal plants. In a
6.2. Yeast origin
For decades, scientists are using yeast extract as one of the biotic elicitors. Yeast extracts stimulated ethylene biosynthesis in tomato [115] and bacterial resistance in bean (
6.3. Fungal origin
Biotic elicitors produced by pathogens have mainly been used to induce the plant defense response. In the past, biological mixtures were prepared from pathogens without identification of the active compounds. The use of pathogenic and nonpathogenic fungal preparations as elicitors has become one of the most effective strategies to induce phenylpropanoid/flavonoid biosynthetic pathways in plant cells [117,118]. Necrotrophic pathogens such as
The content of thiarubrine A was enhanced 3–fold in
6.4. Bacterial origin
The bacterial elicitors stimulated the biosynthesis of scopolamine in adventitious hairy root cultures of
7. Parameters of elicitors
Elicitation has been widely used to increase the production or to induce de novo synthesis of secondary metabolites in in vitro plant cell cultures [141]. This opened up a new area of research that could have important economic benefits for pharmaceutical industry. Several parameters such as elicitor concentration and selectivity, duration of elicitor exposure, age of culture, cell line, growth regulation, nutrient composition, and quality of cell wall materials are also important factors influencing the successful production of secondary metabolite [142]. Some of these parameters were highlighted on elicitation of some medicinal plants for the production of secondary metabolites.
7.1. Elicitor concentration
Elicitor concentration plays a very important role in elicitation process. High dosage of elicitor has been reported to induce hypersensitive response leading to cell death, whereas an optimum level was required for induction [143–145]. At 0.1% (w/v) sodium chloride, ginseng saponin content and productivity were increased to approximately 1.15 and 1.13 times control values, respectively [146]. In the cell culture of
7.2. Duration of elicitor exposure
The cell suspension culture of
7.3. Age of culture
Age of culture plays is an important parameter in the production of bioactive compounds by elicitation. The treatment with MeJA and SA in the hairy root culture of
7.4. Nutrient composition
The composition of the medium or selection of medium also played a vital role in elicitation process. In the callus culture of
8. Conclusion
The development of plant tissue cultures for the production of secondary metabolites has been underway for more than three decades. Although there are well–established plant tissue culture techniques, their application to large scale production is still limited to a few processes. Various stimulation and process strategies have been exercised to improve secondary metabolite production in plant tissue cultures. Elicitation has been widely applied for enhancement of secondary metabolite production in plant cell and organ cultures. The effects of various abiotic and biotic elicitors on secondary metabolite production in plant tissue cultures are dependent on the specific secondary metabolites. The exploration of the production of useful secondary metabolites through regulation of biosynthetic pathway of the various plant cell and tissue cultures of medicinal plants has been carried out by a group of plant scientists in several countries during the last decade. Although, elicitation enhances secondary metabolism in plant cells in vitro, but the exact mechanism is not exactly understood. There is a tremendous scope for the large–scale production of secondary metabolites in the plant tissue culture system by using the elicitors as an agent.
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