Abstract
Abiotic stresses such as increase in daily mean temperature, changed patterns of precipitation, increase in episodes of drought and floods in future are faced by the plants and pose threats to crop production and food security. Induction of secondary metabolites by several abiotic stress conditions can be helpful in the crop protection against biotic stress and can be a major link between biotic and biotic stress. Plants also face threats by injury caused by herbivores and insects that chew on plants. Plant develops, coordinates and combines defence mechanism to cope with the challenges caused by the injuries. The plant family Brassicaceae (or Cruciferae) includes some of the world’s most economically important crops; especially members of the genera Brassica L. Brassicaceae vegetables are a good source of secondary metabolite that is Glucosinolates. Which are responsible for characteristic flavour and odour, when degraded. Glucosinolates and their degradation products play important roles in stress tolerance, plants respond to abiotic and abiotic stress by systematically accumulating higher levels primary and secondary metabolites for increasing their resistance. Glucosinolates play important role and have a relation with biotic and abiotic stress in Brassica plant family, as they can act as a signalling molecules and affect the physiology of plant.
Keywords
- abiotic stresses
- biotic stress
- Brassicaceae
- glucosinolates
- stress tolerance
- human health
1. Introduction
Plants are attached to the ground by the roots for water and nutrient supply and fixed at one place, so they have to face various abiotic stresses such as temperature, water availability, salt concentrations, etc. This is a common environmental stress which impacts productivity of the plants worldwide. Along with abiotic stresses, plants also face injuries by herbivory and insects that chew on plants. Once the plant faces mechanical damage by insects, a lot of changes occur in the plant [1]. In case of biotic stress, immediately after wounding by pest changes occur in the damaged area, which can be systematic (in the non-wounded areas) or local (in the tissues directly damaged). Once the injury occurs, metabolic changes, physiological changes and induction of gene expression started taking place [2]. The response to stress is not simple, rather involves different mechanisms and molecules. The strength and type of response depend upon plant species, developmental stage, nature of stress and environmental conditions. Plant develops, coordinates and combines all the defence mechanisms to cope with the challenges caused by the injuries [1, 3, 4].
The plant family
Various biotic and abiotic factors activate the defence system in plants, and these result in significant variation in the plant metabolome. These factors result in enhanced production of certain metabolites, e.g., amino acids, sugars, indoles, phenolics including glucosinolates. Glucosinolates production in stressed plants is seen to be enhanced, and they have a crucial role to play in mitigating the negative effect of stress. The concentration and type of glucosinolates and their hydrolysis in plants are affected by many factors, such as age, abiotic stress such as heavy metal, water availability, postharvest storage signalling molecules [10, 11]. These play an important role in abiotic stress. Their levels change in response to a variety of abiotic stress tolerances [12]. They also play an important role in plant-insect interactions. Their levels also change in response to herbivory in plants [13].
2. Glucosinolates and abiotic stress
Plants synthesise a variety of secondary metabolites that do not seem to have any essential role in plants, but presence of these compounds plays a very important role in plants and has many advantages, especially in defence. Glucosinolates, a class of secondary metabolites mainly found in Brassica family, are nitrogen and sulphur-containing compounds derived from amino acids such as methionine, alanine, valine or leucine (aliphatic); phenylalanine or tyrosine (aromatic) and tryptophan (indolic glucosinolates). These glucosinolates are usually increased in different biotic or abiotic stress conditions and play an important role in plant defence in these unfavourable conditions [14].
During growth, plants are exposed to various biotic (herbivory, fungal, bacterial and/or viral infection) and abiotic (metals, UV, temperature) stresses. It leads to upregulation of various gene and biochemical changes, which finally results in an enhancement of the synthesis of primary and secondary metabolites. Plants have acquired various stress tolerance mechanisms, involving physiological and biochemical changes that result in adaptive or morphological changes [15]. It was also found that plants respond to abiotic stress by accumulating higher or lower levels of Glucosinolates. The intensity and duration of the abiotic stress, as well as the developmental stage of the plant at the moment of the imposed stress, are important factors in the accumulation of each specific Glucosinolates [16, 17].
The mechanism by which the plant resists these stresses with help of metabolites is unclear as their regulation is a complex process because glucosinolates synthesis pathways are linked to various other pathways making it more complex to understand its direct role. There are studies in elucidating a relationship between the metabolites and stress, but there is a strong need to study underground mechanisms as well.
2.1 Salt stress
One of the major abiotic stresses affecting plant growth and development is salt stress [18]. There are various ways by which plants respond to this stress conditions either by osmotic adjustments or by accumulating secondary metabolites, such as Glucosinolates [19].
In one of the studies, Broccoli was grown in greenhouse condition and was treated with NaCl (0, 40, 80, 100 mM) to study the effect of Glucosinolates in salt-stressed Broccoli. Glucosinolates were determined at three different time points (1, 3, 6 days upon treatment), and it was found that the total aliphatic and indoles were lower at 40, 80 and 100 mM NaCl applications compared with control plants determined at three different time points [20]. A study was conducted with two Broccoli cultivars
Greater accumulation of proline and glucosinolates was found with increased salinity level in Canola or oilseed rape (
Six types of Brassica rocket genotypes used for salad belonging to the
2.2 Water stress
Drought is unavailability of adequate amount of water to a plant, required for its proper growth and development. Drought in agriculture field can occur due to a variety of reasons majorly including low precipitation, inadequate ground water. When
2.3 Heat stress
Abnormally high temperature during hot weather is again one kind of abiotic stress. Effect of high temperature depends on the duration and its intensity. Heat stress may cause various physical, physiological and molecular damage to the plants and in turn affects its productivity. In a study on
2.4 Heavy metal stress
Use and need of metals in plants are important as they act as cofactors in various enzymes such as Mg2+, Mn2+, Co2+, Zn2+, but with increase in industrialisation use of heavy metals such as Hg, Pb, Cd, Ni Ba Cr which are highly toxic even at a very low concentration. There are micronutrient essential metals such as Mn, Mo, Zn, Cu, Co which are required by plants in low amounts, but they also become toxic at high concentrations. Effect of cadmium toxicity was evaluated on cabbage and kale, and it was found that glucosinolates accumulate in stems as compared with other organs such as root or leaves, which indicate that glucosinolates can play role in heavy metal stress tolerance as plants have shown no metal toxicity physically [35]. In another study,
2.5 UV stress
Plant growth and development are mainly determined by sunlight. Sunlight is used in germination, development, photomorphogenesis and various other important functions in plants. But as sunlight is composed of UV-A, UV-B and UV-C, these radiations cause stress in plants. As a result of these radiation stresses, DNA damage, generation of reactive oxygen species (ROS) and many other cellular changes take place in plants. Glucosinolates also get affected by stress caused by UV light, as proven with the studies done on broccoli sprouts which were exposed to various low- and high-intensity UVA and UVB lamps for 120 min which resulted in high levels of glucosinolates in broccoli sprouts [39]. Even the combined effects of Methyl Jasmonate (MJ), UVA or UVB lights enhance the glucosinolates in broccoli sprouts [40]. As it is known from the studies that glucosinolates levels are upregulated during the UV stress [41], and it was also associated with increased biotic stress tolerance too. Low-to-medium UV-B doses were applied to sprouts of broccoli, and it was found that there was an increase in glucosinolate levels of 4-methylsulfinyl butyl GS and 4-methoxy-indol-3-ylmethyl GS. These glucosinolates in turn affect the genes related to jasmonic acid and salicylic acid signalling pathway, which ultimately leads to overexpression of bacterial and fungal pathogen-responsive genes [42].
3. Glucosinolates and biotic stress
Biotic stress in plants is caused by any living organisms, such as insects, pests, viruses, bacteria, fungi, nematodes, etc. Microorganisms damage plants by causing symptoms such as plant wilt, leaf spots, root rot, etc. Insects can cause physical damage to plants as they either chew or suck on the plants, including the leaves, stem, bark and flowers. They can also act as a vector of viruses and bacteria from infected plants to healthy plants. As plants do not have any immune systems to deal with this kind of stress, they respond to this stress by developing many changes at morphological, chemical and protein levels. The main aim of these changes is to protect the plants from foreign invaders. One such chemical is glucosinolates which get elevated in response to biotic stress for the purpose of defence in plants.
Chemically, glucosinolates are composed of thiohydroximate-O-sulfonate group linked to glucose and an alkyl, aralkyl or indolyl side chain (R). More than 200 glucosinolates have been identified in Brassica crops, and they are characterised mainly by the variable R group, which can be aromatic, indolic or aliphatic derivatives of the amino acid precursors methionine, tryptophan and phenylalanine, respectively [43, 44].
Glucosinolates play a variety of biological functions in plants [45] and get converted into various degradation products (isothiocyanates, thiocyanates, indoles, etc.), when vegetables containing them are cut or chewed because during this process, they come in contact with the enzyme myrosinase which hydrolyses them, as discussed earlier in this chapter. A sudden outbreak of these insecticidal compounds in field is called as
Studies have determined the effect of insect pest and glucosinolates levels in
Several reports exist on the toxicity of Glucosinolates hydrolysis products to bacteria and fungi. The activity of Glucosinolates and their degraded products against various strains of microorganisms has been documented by many investigators being present in the leaves of
3.1 Glucosinolates and pest
When generalist herbivores feed on Brassica, degradation product of Glucosinolates in them, the isothiocyanates have toxic effects and function as repellents. However, specialist herbivores use Glucosinolates as attractants, and they can survive on plants containing them. For example,
It was also found that plants respond to herbivore or insect damage by systematically accumulating higher levels of glucosinolates and thus presumably increasing their resistance. Usually it is the indole glucosinolates which become induced [58, 59]. Sometimes volatiles produced by Glucosinolates provide indirect protection to plants by attracting natural enemies of herbivores such as parasitoids [60].
Lepidopteran that feeds almost exclusively on
Genetic engineering of glucosinolates biosynthesis pathway for the production of glucosinolates in non-Glucosinolates producing plants was successfully employed by [62]. Genetic transformation of
It was found that
4. Conclusion
Abiotic stress such as heat, drought and high salt and biotic stress including various attacks on plant from insects, pest or other bacterial viral disease can affect the plant in terms of physical, physiological and many other biochemical and molecular aspects, which can retard its growth and productivity.
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