Open access peer-reviewed chapter

Agronomic Factors Influencing Brassica Productivity and Phytochemical Quality

Written By

Cristine Vanz Borges, Santino Seabra Junior, Franciely S. Ponce and Giuseppina Pace Pereira Lima

Submitted: 08 November 2017 Reviewed: 31 January 2018 Published: 24 October 2018

DOI: 10.5772/intechopen.74732

From the Edited Volume

Brassica Germplasm - Characterization, Breeding and Utilization

Edited by Mohamed Ahmed El-Esawi

Chapter metrics overview

1,402 Chapter Downloads

View Full Metrics


Agronomic practices and climatic factors affect the content and profile of phytochemicals. The effects of the environment, such as salinity, climate, and other abiotic factors, promote biochemical responses, inducing changes in the quantity and quality of polyphenol compounds, carotenoids, vitamins, glucosinolates, and polyamines, which are bioactive compounds. In plants, among the various functions, some phytochemicals can protect against biotic factors. Brassica vegetables are a source of several primary and secondary metabolism compounds, and they might be responsible for disease prevention. In addition, the increase of bioactive compounds in plant-based foods is important to the diet and consequently for the improvement of public health. In this chapter, we will point out the abiotic factors that affect the productive performance, quality, and chemical composition of different Brassica species and cultivars. We will also discuss its implications on plant protection and human health.


  • environmental factors
  • cultivation conditions
  • polyphenol
  • carotenoids
  • glucosinolates

1. Introduction

The Brassica ceae family, previously known as Cruciferae, is composed of 338 genus and around 3700 species. The family includes many plants of economic importance, for the production of edible oil, such as the canola, forage rape (Brassica napus), and seasoning plants, as the mustard and also various species for consumption, in which the leaf, stem, roots, and tubercles are edible parts. The main genus, Brassica, is formed by 37 species, which can be annual and biannual, including even weeds, wild plants, and domestic crops. Brassica vegetables originate from regions between the Mediterranean and the Sahara, where the climate consists of mild winters followed by hot and dry summers. Besides that, there are species inside the genus that are well adapted to colder regions, and many species are now considered naturalized in the entire world and are commonly observed in Western Europe, in the Mediterranean, and in temperate regions of Asia. In addition, many species also grow in as invasive weeds in the Americas (North and South) and Australasia [1].

The species of the Brassica genus were widely modified and domesticated by human beings and are vegetables cultivated worldwide [2], especially the varieties belonging to the species Brassica oleracea, which includes cabbage, tronchuda cabbage (Brassica oleracea L. var. costata DC), mustard, rocket, and Brussels sprout (Brassica oleracea L. var. gemmifera), among others (Figure 1). Among these species, we also found broccoli, where the most consumed part is the inflorescence, the cauliflower, from which the floral peduncle is consumed and the tubercles, as the radish and the turnip.

Figure 1.

Main brassicas cultivated worldwide.

Brassica vegetables have attracted great attention due to the presence of phytochemicals with recognized beneficial functions in the human organism, reducing the risk of diseases [3]. These vegetables are potential sources of anticarcinogenic and antioxidant compounds as the glucosinolates (GLS), vitamin C, phenolic acids, flavonols, anthocyanidins, carotenoids, and amino acids [4]. Most of the researches are focused on the content of secondary metabolites, mainly the glucosinolates. The benefits to human health from the ingestion of these vegetables, such as the reduced risk of degenerative diseases, are in great part attributed to the content of secondary metabolites substances of the plants [5]. Besides the human health aspects, these metabolites play a fundamental role in the plants’ defense against microorganisms, raising the interest in higher quantities of these secondary compounds as a strategy of increasing the protection of the cultures and reducing the use of agrochemicals.

Variations in the agronomic conditions (e.g., vegetal species, cultivars, development stage, plants organs, fertilization, and soil pH) and climatic factors (e.g., light intensity and water availability) are known for significantly affecting the phytochemical content and profile. The understanding of the effects of climatic and agronomic factors is necessary for increasing the predictability of the desired compounds, increasing the benefits related to the human health and to the plants’ protection (plague control) [6]. Although there is little information on the real influence of cultivation on the contents of glucosinolates and other important phytochemicals in Brassicas, it appears that the use of ecological practices can induce a rise in the content of these molecules. In this chapter, we provide a general view about the roles of the glucosinolates and other phytochemicals present in Brassicaceae and their implications in the plants’ protection, productivity, and human health, as well as emphasize the factors that affect the contents of these compounds in Brassica vegetables.


2. Agronomic factors and production

The photosynthetic activity is the base for the production of reserves in the plant, which will constitute the biomass, a factor that can determine the vegetal development limits. The production depends on the interaction between the productive potential and the environmental factors. The edaphoclimatic factors are directly related to productive responses that influence the flowering, hydric balance, respiration, and absorption of minerals. Latitude, altitude, rainfall, topography, and soil physics act indirectly on the production and other factors, such as solar radiation, temperature, water, and chemical elements of the soil act directly on the photosynthesis. The environmental factors, such as water, temperature, quality, and quantity of light hours, will determine the plants’ growth rate.

The temperature is a climatic factor that can limit the production of determined species in tropical and equatorial regions. Besides determining the growth and development, it establishes the end of the vegetative stage and the beginning of the productive stage in the biennial species, such as broccoli, cauliflower, Brussels sprout, among other species. The broccoli has a better productive development under average temperatures between 60 and 65°F, with a maximum of 75°F [7]. Prolonged periods of temperature above 77°C can retard the formation of inflorescence in plants that are in phase of vegetative growth, reducing the size and causing the development of leaves and bracts in the floral peduncles [8].

The temperature strongly influences the plants’ metabolic activity, and the stress caused by high and low temperatures can induce effects in the primary and secondary metabolism (Table 1). The heat or cold can affect the membrane fluidity, metabolism, and cytoskeleton rearrangement, consequently affecting the vegetative and reproductive tissues [9]. Abrupt increases of temperature can provoke excessively fast growth of the inflorescence and elongating the peduncle in certain cultivars [10]. Cultivations in conditions of high temperatures, where there are only few days with ideal temperatures for vernalization, the plants can continue to vegetate or to not produce commercial inflorescences, which means uneven bunches, the presence of bracts, and low compactness of the head and yellow coloration. Temperatures below the ideal level can prolong the cycle of provoke premature flowering in some species, as in the case of summer cauliflowers submitted to low temperatures [11]. Among the brassicas more tolerant to the cold, the minimum temperature for the germination and cultivation for species is 40°F and the maximum tolerated temperature can reach 105°F for turnip and kohlrabi, which are the most tolerant species to temperature in the germination phase. However, in the cultivation phase, the maximum temperature is situated around 75°F. In Brazil, some regions with mild temperatures and with mensal averages varying from 66 to 88°F, there are reports of commercial cultivation of broccoli, kale, rocket, and watercress [12]. However, cauliflower, cabbage, and radish are also cultivated in tropical regions. These cultivations are favored by the utilization of thermotolerant cultivars obtained by genetic improvement.

Stress factor Productive and/or biochemical response Citations
Temperature Heat Affects the glucosinolates content
Higher glucosinolates production
Cold Higher glucosinolates production
Higher glucosinolates production in broccoli under low temperatures
Thermal amplitude Higher glucosinolates production in broccoli plants under temperatures between 53.6 and 89.6°F, compared to plants under temperature 71.6F [61]
Luminosity Competition (population density and consortium) Reduced levels of trypsin inhibitors in Brassica napus seedlings
Reduced levels of glucosinolates in the leaves and roots in B. napus
Excess Photoinhibition, thermal stress, and stomatal closing, leading to a reduction of net photosynthesis in brassicas, included B. napus. [62]
Protected cultivation (light diffusion and shading) Mustard plants (B. juncea) cultivated under shading screens of 50% showed lower quantities of ascorbic acid, larger foliar area, chlorophyll, carotenoids, N, NO3, and a higher content of mineral nutrients in comparison to the complete solar luminosity [63]
Photoperiod Affects the glucosinolates content
Spring broccoli growth in intermediate temperatures, high luminous intensity, longer days, and dry conditions have the highest total GLS content
Glucosinolates levels in kale are not influenced by the photoperiod
Water Hydric restriction Growth reduction, lower yield of cabbage heads, and an increase of dry mass
Kale: growth reduction, biomass reduction, increase of sorbitol, sucrose, verbascose and kestose levels, and a decrease of manitol
An increase in the sugar content in the phloem sap of broccoli submitted at hydric stress
Lower glucosinolate content in broccoli in hydric restriction
Biomass reduction, an increase of nitrogen in the leaf, and a darker green leaf in Chinese cabbage
Salinity Growth reduction, Na or Cl accumulation, and lower productivity
A decrease of fresh matter in the aerial part of broccoli
An increase in GLS content and phenolic compounds
A drastic decrease in the vitamin C content in old broccoli leaves
A significant decrease in the vitamin C content in young broccoli leaves;
Loss of flavonoids in old broccoli leaves;
Loss of turgor
Accumulation of glucosinolates in B. napus L.;
Reduction in the nitrate content
Increase or no effect in the nitrate content
Fertilization Nitrogen (N)
Sulfur (S)
Nitrogen fertilization influences the GLS metabolism in broccoli
High level of sulfur provided an increase of polyphenol contents (flavonoids and phenolic acids) in B. rapa ssp. sylvestris
An increase of the total glucosinolates with the increase of fertilization with sulfur
Higher quantities of sulfur and nitrogen combined did not provide higher contents of glucosinolates

Table 1.

Effect of environmental factors in the production and biochemical compounds in brassicas.

The plants can be modified to some degree, tolerating light stresses from either low or high temperatures when slowly submitted to the stress, leading to acclimatization. By contrast, plants that survive the exposition to conditions above the ideal temperature can produce chaperones, molecules that are related to the antioxidant activity, and solutes accumulation [9]. Low temperatures cause reduction in the enzymatic activity, rigidity of membranes, destabilization of protein complexes, compromise of photosynthesis, and rupture of the membranes. Cellular alterations associated to the tolerance to cold and/or freezing include the accumulation of sugar or compatible solutes, changes in the membrane composition, and synthesis of dehydrin-like proteins [9].

The stress by temperature can cause changes in the plants’ chemical constitution. Broccoli sprouts present increased glucosinolate contents when cultivated under high (84.2 or 89.6°F) or low temperatures (51.8 or 60.8°F), comparing the cultivated sprouts under ideal temperature (70.7°F) [13]. Similar to the sprouts, the broccoli leaves showed the highest glucosinolate level when cultivated under 53.6 or 89.6°F. This effect was also observed in younger cabbage plants. Under low temperatures, there is an increase of the glucosinolate levels in broccoli and watercress (Table 1). The combination of temperature and precipitation influences the glucosinolate content in brassicas (white cabbage, red cabbage, savoy cabbage, Brussels sprouts, cauliflower, kale, kohlrabi, turnip, red radish, black radish, and white radish). High and low precipitations induce higher contents of glucosinolates, when compared to the same vegetables cultivated in a year with mild temperatures and higher precipitation [14].

The light is a factor that influences the brassicas’ performance. The increase in the luminous intensity corresponds to a rise in the photosynthetic activity (within certain limits), while the decrease promotes a higher cellular elongation, resulting in etiolated plants. However, this response depends on the species’ susceptibility and on the plants’ density, producing a competition for light or on excessive shading obtained by the use of screens. The amount of energy intercepted is dependent on the characteristics of the cultivation system, row spacing, consortium, and even the architecture characteristics of each genotype, such as the leaf inclination. In the same plant it is possible to occur leaves exposed or not to the sun and with different quantic necessities, with different photosynthetic performance. In this context, there are species that show lower or higher stress when cultivated in a lower spacing or cultivated under consortium. In Ethiopian kale (B. carinata) and African nightshade (Solanum scabrum), cultivated in consortium and in ideal condition of irrigation and under hydric stress, there was an increase in the glucosinolates content in kale and the maintenance of the biomass production and nutritional characteristics. In addition, low irrigation induced the carotene levels in African nightshade, both under hydric stress and stress provided by the consortium. In opposition, hydric stress did not affect the glucosinolate content in B. carinata and the indole glucosinolates in B. rapa ssp. Rapifera [15]. These results suggest that the responses to drought and glucosinolates concentration can vary depending on the genotype.


3. Brassicas’ phytochemical composition

Brassica vegetables are important sources of fibers, vitamins, and minerals. In addition, these vegetables are potential sources of anti-carcinogenic and antioxidant compounds, such as the glucosinolates, vitamin C, phenolic acids, flavonols, anthocyanidins, carotenoids, and amino acids [4]. Most of the current researches are focused on the content of secondary metabolites, mainly of glucosinolates. Many epidemiologic studies indicate that a high ingestion of brassicaceous vegetables is associated to a reduced risk of cancer [5, 16], cardiovascular diseases [17], gut diseases (e.g., colite) [18], [19], and diabetes [20].

Many epidemiologic studies do not differ among the types of cruciferous vegetables, but the most common studies in the entire world include the broccoli, cauliflower, cabbages, bok choy, kale, watercress, turnip, and rocket [21]. Besides human health aspects, these metabolites play a fundamental role in the plants’ defense against microorganisms; thus, there is an increasing interest in raising the content of these secondary compounds as a strategy of increasing the protection to cultures and reducing the use of agrochemicals.

Even though there is little information on the real influence of the cultivation in the levels of glucosinolates and other important phytochemicals in brassicas, it seems that the use of ecological practices can induce a raise of these molecules. In addition, in most cases, it is fundamental to also study the impact of storage and cooking in these compounds, since brassicas are not consumed immediately after the harvest, in order to know the real benefits of these vegetables to the human health.

3.1. Glucosinolates

Brassica vegetables are the major source of glucosinolates that has been associated to its bioactivity, and these compounds may be responsible for their observed protecting effects. The glucosinolates are found in 16 families of dicotyledonous plants and in at least 120 different chemical structures that have been identified until now [21]. Depending on the chemical structure of the precursor amino acid, they are now classified into three groups: aliphatic, indole, and aromatic glucosinolates.

The glucosinolates profile and its modifications, together with specific products of hydrolyzation, are being discussed as a plant defense mechanism to deal with various abiotic and biotic stresses. Recent studies showed that glucosinolates, for example, breakdown products of 1-methoxy-indol-3-ylmethyl glucosinolate and 5-phenylpentyl isothiocyanate, exert mutagenic or genotoxic effects in mammalian and bacterial cell studies [22]. Studies indicate that broccoli sprouts are sources of GLS (varying from 679.01 to 554.90 mg/100 g FW), and the predominant GLS is the glucosinolate glucoraphanin (GRA) (33% of the total GLS) [23].

Glucosinolates and isothiocyanates (products of glucosinolate hydrolysis) are produced by some plants in response to biotic stress. They are important as protective agents of the plants, due to their toxic or repelling effects against potential plagues (herbivores, bacteria, and fungi) [24]. Even though these compounds can be used as protection agents in plants, with a great importance in agriculture and horticulture, they are significantly important to human nutrition, due to the preventive effects on human health [24, 25]. These compounds are known for protecting against cancer in humans [26] and, in plants, these secondary metabolites and/or their breakdown products have different biological functions, like fungicidal, bactericidal, nematocidal, and allelopathic properties [27].

Thus, factors influencing phytochemical content and profile in the production of brassicaceous plants are worth considering for both plant and human health. There are studies showing that the consumption of brassica vegetables has a direct relationship with cancer incidence reduction [28]. Besides GLS, these vegetables contain myrosinase, a thioglucoside glycohydrolase (EC which is released from intracellular compartments when the vegetable tissue is damaged by cutting or chewing and induces GLS hydrolysis into isothiocyanates and nitriles, as the most important products. Sulforaphane is one of the investigated isothiocyanates and is particularly abundant in broccoli var. italica in the form of its corresponding glucosinolate glucoraphanin (GRA) (4-methylsulfinylbutyl glucosinolate) [29]. GRA can be converted into sulforaphane and glucobrassicin to indolyl-3-carbinol; both hydrolyzed derivatives are active against carcinogenesis as demonstrated by many in vitro experiments or in vivo studies [28, 30].

The quality and quantity of GLS differ among the plants species, among the different plant organs (tubercle or leaves), and in function of the ontogeny. The profile of these compounds is not only determined by the plant genetic constitution but also influenced by the environmental conditions [31]. Generally, high levels of GLS occur in response to temperature [32], exposition to different wavelengths [33], nutrients availability [34], and signaling molecules as the salicylic acid (SA) [25], jasmonic acid (JA), and methyl jasmonate (MeJA) [31, 35]. Exogenous applications of SA and its analogous acids, damage by herbivory or treatment with JA, induce increases of indole GLS in B. napus [36], B. campestris [37], and B. juncea [35]. Microorganism infection and/or mechanical damages can promote the biosynthesis of indole GLS and aromatic 2-phenylethyl GLS in B. rapa through synthesizing molecules (e.g., methyl jasmonate or jasmonic acid) [31]. In addition, many compounds such as phenolics, terpenoids, and compounds containing sulfur also regulate the biosynthesis [38].

Saline stress (150 mM NaCl) can reduce the total GLS levels, due to the decrease of both aliphatic GLP, as indole (GBS and MBGS), in broccoli sprouts [23]. This decrease is attributed to cell damages induced by Na accumulation [39]. However, studies with broccoli sprouts determined that the decrease in the GLS level in response to excessive contents of NaCl (44% in comparison to the control—0 mM NaCl) can be decreased by the application of MeJA, applied daily from the third day of growth of 10-day-old broccoli sprouts [23].

Brassica foods (e.g., cabbage, cauliflower, broccoli, Brussels sprouts, turnips, and kale) are consumed raw, frozen, or after domestic thermal processing (cooking). Generally, the conventional cooking methods, such as boiling, steaming, pressure cooking, and microwaving, reduce the content of glucosinolates to approximately 30–60%, depending on the method and analyzed compound [40]. Leaves (turning greens) and young-sprouting shoots (turning tops) of B. rapa cooked in steam showed maintenance of GLS content, compared to raw vegetable, by preventing leaching and solubilization of these metabolites. By contrast, conventional boiling and high-pressure cooking methods induced losses of GLS levels (64%), and the degradation of different GLS classes (e.g., aliphatic or indolic) was similar in both cooking methods [40]. However, some compounds with important pharmacologic activities can be formed after the thermal processing by hydrolysis (e.g., 2-aminothiophene and dimeric 1,4-dithiane-2,5-diacetonitrile), increasing the bioactive potential in brassica vegetables [41].

The thermal treatment causes denaturation of enzymes that catalyze the degradation of nutrients and some metabolites. When brassica vegetables are chopped, ground, or chewed, there is a rupture of the tissues, and the GLSs enter in contact with the myrosinase, inducing the conversion to isothiocyanates, nitriles, thiocyanates, epithionitriles, oxazolidine-2-thiones, and epithioalkanes [42]. The hydrolysis products, mainly during the storage and processing, as well as the myrosinase activity of the gut microbiota, can affect the total content and bioavailability of these compounds [26]. In addition, the glucosinolates are water-soluble compounds and are generally lost by leaching, in methods that use water for cooking.

3.2. Polyphenols

The phenolic compounds are a group of secondary metabolites present in the vegetal kingdom. The most disseminated and diversified groups of polyphenols are the flavonoids, which have C6-C3-C6 flavone skeleton. The flavonoids are important phenolic phytochemicals containing a basic structure of two aromatic benzene rings separated by a heterocyclic-oxygenated ring [43]. The flavonoids and the hydroxycinnamic acids are widely distributed in plants and are important bioactive compounds in the human diet. The dietetic flavonoids have antiviral, anti-inflammatory, antihistaminic, and antioxidant properties. Flavonoids and phenolic acids are the most characterized groups of phenolic compounds in brassicas and can protect the plants against UV radiation, microorganisms, and predator insects [44]. In cabbage, many (poly)phenolic compounds were identified, including myricetin, quercetin, kaempferol, luteolin, delphinidin, cyanidin, and pelargonidin [45].

Generally, the phenolic compounds are produced through the phenylpropanoid pathway. Biotic or abiotic stresses, such as elicitors, were reported for inducing alterations in the phenolic compounds contents, as described in broccoli sprouts [23, 46]. In addition, the quality and quantity of the phenols differ among the plant species and among the plant organs. For example, broccoli sprouts have higher phenolic levels (1133.85 mg/100 g FW), when compared with mature broccoli inflorescences (63.4 mg/100 g FW) [45]. Most of the phenolic compounds present in broccoli sprouts are the hydroxycinnamic acids (sinapic acid derivatives), approximately 98% of the total phenolics found [23].

In saline-stress conditions, there is a possibility for a decrease to occur up to 30% in the phenolic compounds’ content in brassicas (e.g., broccoli sprout). It is important to highlight that the increase or decrease of these compounds in this situation depends on the plant sensibility to salt and on the development stage when the plant was submitted to the stress [23]. The exogenous application of elicitors can also induce the phenolic compounds’ biosynthesis, affecting the contents of antioxidant and nutritional compounds in brassicas. This technique can be a viable tool to obtain vegetables with higher levels of these bioactive compounds. Studies with broccoli sprouts showed that the prolonged application of low concentrations of SA and MeJA during the sprouting significantly increased the content of phenolic compounds. Exogenous SA (50 μM) applied for 5 days or 100 μM SA for 7 days achieved flavonoids-enriched broccoli sprouts by 24 and 33%, respectively. A 10 μM MeJA was a highly efficient treatment, promoting increases of 31 and 23% in the concentration of flavonoids and total phenolics, respectively [47].

The culinary process is a source of several alterations, both physical and biochemical, modifying the phytochemical constituents present in the vegetables, resulting in changes in the nutritional values [48] of brassicas. During the cooking process, the phenolic compounds are highly reactive and can be significantly modified, including the release of conjugated compounds (bound forms), oxidation, degradation, and polymerization [49]. Generally, the effect of boiling in brassica vegetables can lead to significant polyphenol losses. During the boiling, because the phenolic compounds are water-soluble, there might be losses by leaching, besides the breaking of these compounds during the thermal processing. The analysis of the water used in experiments with boiled brassica vegetables (e.g., in watercress) shows the presence of total phenols in the water (9.35 ± 0.12 mg GAE/g DW), confirming the loss by leaching (raw watercress – 14.86 ± 2.02 mg GAE/g DW). The quantity of phenols found in the water and in cooked material (residual phenols) is not different from the quantity present in raw watercress [50]. In opposition, in these studies, a minimum deleterious effect was demonstrated when microwaving and steaming were used in the content of phenolic compounds. This minimum effect occurs according to the quantity of water used and to the inactivation of oxidative enzymes, which prevent the rupture and the degradation of the phenolic biosynthesis [50].

3.3. Carotenoids

Carotenoids are a class of phytonutrients that are responsible for the colors red, orange, and light yellow in many vegetables and fruits. Most of the brassica vegetables contain carotenoids, such as β-carotene, lutein, zeaxanthin, neoxanthin, violaxanthin, and folate, which have important antioxidant, anticarcinogenic properties, and provitamin A [51]. Carotenoids have shown to have functions during the photosynthesis and show an important role in defense mechanisms apart from the essential nutrients. These compounds are involved in biotic and abiotic stresses response and development, acting as signaling molecules, and in addition, they are related to processes such as photomorphogenesis, nonphotochemical quenching and lipid peroxidation, and attracting pollinators [52].

Brassica vegetables are rich in carotenoids and, among the varieties of B. oleracea, kale has the highest levels of lutein and beta-carotene [51]. In Chinese cabbage (B. rapa ssp. pekinensis), lutein and β-carotene are mainly distributed in the older leaves and in the flowers, while the zeaxanthin and violaxanthin levels are relatively low [53].

In order for the carotenoid absorption to occur by gut enterocytes, the mechanical and/or enzymatic disruption of the food matrix is necessary. In addition, due to the hydrophobic character of these chemical molecules, the formation of micelles before its absorption is also necessary [54]. Since the carotenoids in fruits and vegetables are present in the chromoplasts, their substructure and the cell wall are the main barriers to the bioavailability of these compounds [55]. Thus, thermal processing as the boiling or the steaming can have positive effects in bioavailability, collaborating to the food matrix disruption, even though negative effects caused by the carotenoids degradation were also reported [56].

The processing methods used in brassica vegetables generally increase the carotenoid content [50]. The thermal processing can cause quantitative and qualitative changes by isomerization processes. An increase in the carotenoid content in brassica vegetables, such as broccoli, Brussels sprouts, cabbage, cauliflower, and watercress, after boiling and steaming, were reported in many studies [33, 50]. The increase in the total carotenoid content after thermal treatments can also be explained by changes in the cell wall, due to the cellulose degradation, improving the extraction of these compounds, as a result of the denaturation of carotenoids/protein complexes caused after the thermal processing [57]. However, high temperatures can lead to isomerization processes, decreasing the food nutritional values. β-carotene and lutein degradation for the formation of cis-isomer (4–40%) during the thermal processing was described in some studies with brassica vegetables (e.g., broccoli and kale) [58]. Thus, a higher retention of cis-isomers was registered in brassica vegetables thermally processed in comparison to trans-isomers, leading to losses in the vitamin A content in these foods.


4. Conclusion

Brassica vegetables have attracted increasing attention due to the presence of phytochemicals with beneficial recognized functions to the human organism, reducing the risk of diseases. Variations in the agronomic conditions (e.g., vegetal species, cultivars, development stages, plant organs, fertilization, soil, and pH) and climatic factors (e.g., luminous intensity and water availability) are known for affecting the content and the profile of compounds from the secondary metabolism.

Many studies show that stress can lead to the accumulation of bioactive compounds in plants, generating the production of foods with more benefits to the human health. In contrast, the growth and development are affected, because there is a reallocation of primary metabolites for the formation of secondary metabolites. This reflects in the biomass production and, certainly, in the species production. However, these metabolites, such as GLS, phenolic compounds, and carotenoids, play a fundamental role in the plants’ defense against microorganisms, possibly leading to a better adaptation of the plants to the environment and, consequently, to the reduction in the use of agrochemicals. The current knowledge of the climatic factors that affect the content and profile of these phytochemicals in Brassicaceae is of scientific and economical interest and can be the base to elaborate strategies for producing plants more resistant to plagues and diseases, reducing the use of agrochemicals and increasing the productivity with a higher nutraceutical potential.



The authors gratefully acknowledge the support by São Paulo Research Foundation (FAPESP—Brazil), process 2016/22665-2 and 2016/00972-0, São Paulo State University, and Conselho Nacional de Pesquisa (CNPq e Brazil), process 305177/2015-0.


Conflict of interest

The authors affirm that there is no conflict of interest.


  1. 1. Rakow GI. 1 Species Origin and Economic Importance of Brassica. In: Berlin HS, editor. Brassica. Berlin, Heidelberg; 2004. p. 3-11
  2. 2. Card SD, Hume DE, Roodi D, McGill CR, Millner JP, Johnson RD. Beneficial endophytic microorganisms of Brassica - A review. Biological Control. 2015;90:102-112
  3. 3. Hafidh RR, Abdulamir AS, Abu Bakar F, Jalilian FA, Jahanshiri F, Abas F, Sekawi Z. Novel anticancer activity and anticancer mechanisms of Brassica oleracea L. Var. capitata f. rubra. European Journal of Integrative Medicine. 2013;5(5):450-464
  4. 4. Park S, Arasu MV, Jiang N, Choi SH, Lim YP, Park JT, Al-Dhabi NA, Kim SJ. Metabolite profiling of phenolics, anthocyanins and flavonols in cabbage (Brassica oleracea var. capitata). Industrial Crops and Products. 2014;60:8-14
  5. 5. Forte MBA, Sanctis R, Leonetti G, Manfredelli S, Urbano V. Dietary chemoprevention of colorectal cancer. Annali Italiani di Chirurgia. 2008;79(4):261-268
  6. 6. Maria B, Klingen I, Birch ANE, Bones AM, Bruce TJA, Johansen TJ, Meadow R, Mølmann J, Seljåsen R, Smart LE, Stewart D. Phytochemicals of Brassicaceae in plant protection and human health - Influences of climate, environment and agronomic practice. Phytochemistry. 2011;72(7):538-556
  7. 7. Maynard DN, Hochmuth GJ. Knott’s Handbook for Vegetable Growers. 5th ed. New York: John Wiley & Sons; 2006
  8. 8. Bjorkman T, Pearson KJ. High temperature arrest of inflorescence development in broccoli (Brassica oleracea var. italica L.). Journal of Experimental Botany. 1998;49(318):101-106
  9. 9. Ruelland E, Zachowski A. How plants sense temperature. Environmental and Experimental Botany. 2010;69(3):225-232
  10. 10. Lalla JG, Laura VA, Rodrigues APDC, Seabra Júnior S, Silveira DSS, Zago VH, Dornas MF. Competição de cultivares de brócolis tipo cabeça única em Campo Grande. Horticultura Brasileira. 2010;28(3):360-363
  11. 11. Thakur BS. Adaptability for yield in some mid-late and late group cauliflower (Brassica oleracea var botrytis) genotypes under the mid-hill conditions of Himachal Pradesh. The Indian Journal of Agriculture Sciences. 2017;76:37-40
  12. 12. Nespoli A, Cochev JS, Neves SMAS SS. Vegetable Production By Family Agriculture in Alta Floresta, Matogrossense Amazon. CAMPO-TERRITÓRIO Rev. Geogr. agrária. 2015;10:159-91
  13. 13. Pereira AA, FMV RE, Fahey KKS, Carvalho R. Influence of temperature and ontogeny on the levels of glucosinolates in broccoli (Brassica oleracea Var. Italica) sprouts and their effect on the induction of mammalian phase 2 enzymes. Journal of Agricultural and Food Chemistry. 2002;50(21):6239-6244
  14. 14. Ciska E, Martyniak-Przybyszewska B, Kozlowska H. Content of glucosinolates in cruciferous vegetables grown at the same site for two years under different climatic conditions. Journal of Agricultural and Food Chemistry. 2000;48(7):2862-2867
  15. 15. Zhang H, Schonhof I, Krumbein A, Gutezeit B, Li L, Stützel H, Schreiner M. Water supply and growing season influence glucosinolate concentration and composition in turnip root (Brassica rapa ssp. rapifera L.). Journal of Plant Nutrition and Soil Science. 2008;171(2):255-265
  16. 16. Pocasap P, Weerapreeyakul N, Tanthanuch W, Thumanu K. Sulforaphene in Raphanus sativus L. var. caudatus Alef increased in late-bolting stage as well as anticancer activity. Asian Pacific Journal of Tropical Biomedicine. 2017;7(11):998-1004
  17. 17. Cardenia V, Vivarelli F, Cirillo S, Paolini M, Rodriguez-Estrada MT, Canistro D. Dietary effects of Raphanus sativus cv Sango on lipid and oxysterols accumulation in rat brain: A lipidomic study on a non-genetic obesity model. Chemistry and Physics of Lipids. 2017;207:206-213
  18. 18. Hubbard TD, Murray IA, Nichols RG, Cassel K, Podolsky M, Kuzu G, Tian Y, Smith P, Kennett MJ, Patterson AD, Perdew GH. Dietary broccoli impacts microbial community structure and attenuates chemically induced colitis in mice in an Ah receptor dependent manner. Journal of Functional Foods. 2017;37:685-698
  19. 19. Tong T, Niu YH, Yue Y, chan Wu S, Ding H. Beneficial effects of anthocyanins from red cabbage (Brassica oleracea L. var. capitata L.) administration to prevent irinotecan-induced mucositis. Journal of Functional Foods. 2017;32:9-17
  20. 20. Jia X, Zhong L, Song Y, Hu Y, Wang G, Sun S. Consumption of citrus and cruciferous vegetables with incident type 2 diabetes mellitus based on a meta-analysis of prospective study. Primary Care Diabetes. 2016;10(4):272-280
  21. 21. Traka MH. Health benefits of glucosinolates. Advances in Botanical Research. 2016;80:247-279
  22. 22. Dekić MS, Radulović NS, Stojanović NM, Randjelović PJ, Stojanović-Radić ZZ, Najman S, Stojanović S. Spasmolytic, antimicrobial and cytotoxic activities of 5-phenylpentyl isothiocyanate, a new glucosinolate autolysis product from horseradish (Armoracia rusticana P. Gaertn., B. Mey. & Scherb., Brassicaceae). Food Chemistry. 2017;232:329-339
  23. 23. Hassini I, Martinez-Ballesta MC, Boughanmi N, Moreno DA, Carvajal M. Improvement of broccoli sprouts (Brassica oleracea L. var. italica) growth and quality by KCl seed priming and methyl jasmonate under salinity stress. Scientia Horticulturae (Amsterdam). 2017;226(May):141-151
  24. 24. Hanschen FS, Lamy E, Schreiner M, Rohn S. Reactivity and stability of glucosinolates and their breakdown products in foods. Angewandte Chemie International Edition. 2014;53(43):11430-11450
  25. 25. Yi GE, Robin AHK, Yang K, Park JI, Hwang BH, Nou IS. Exogenous methyl jasmonate and salicylic acid induce subspecies-specific patterns of glucosinolate accumulation and gene expression in Brassica oleracea L. Molecules. 2016;21(10)
  26. 26. Verkerk R, Schreiner M, Krumbein A, Ciska E, Holst B, Rowland I, de Schrijver R, Hansen M, Gerhäuser C, Mithen R, Dekker M. Glucosinolates in Brassica vegetables: The influence of the food supply chain on intake, bioavailability and human health. Molecular Nutrition & Food Research. 2009;53(Suppl. 2):219-265
  27. 27. Vicas SI, Teusdea AC, Carbunar M, Socaci SA, Socaciu C. Glucosinolates profile and antioxidant capacity of Romanian Brassica vegetables obtained by organic and conventional agricultural practices. Plant Foods for Human Nutrition. 2013;68(3):313-321
  28. 28. Shapiro TA, Fahey JW, Wade KL, Stephenson KK, Talalay P. Chemoprotective glucosinolates and isothiocyanates of broccoli sprouts: Metabolism and excretion in humans. Cancer Epidemiology, Biomarkers & Prevention. 2001;10(5):501-508
  29. 29. Fahey JW, Zhang Y, Talalay P. Broccoli sprouts: An exceptionally rich source of inducers of enzymes that protect against chemical carcinogens. Proceedings of the National Academy of Sciences. 1997;94(19):10367-10372
  30. 30. Zanichelli F, Capasso S, Di Bernardo G, Cipollaro M, Pagnotta E, Cartenì M, Casale F, Iori R, Giordano A, Galderisi U. Low concentrations of isothiocyanates protect mesenchymal stem cells from oxidative injuries, while high concentrations exacerbate DNA damage. Apoptosis. 2012;17(9):964-974
  31. 31. Wiesner M, Hanschen FS, Schreiner M, Glatt H, Zrenner R. Induced production of 1-methoxy-indol-3-ylmethyl glucosinolate by jasmonic acid and methyl jasmonate in sprouts and leaves of pak choi (Brassica rapa ssp. chinensis). International Journal of Molecular Sciences. 2013;14(7):14996-15016
  32. 32. Martínez-Ballesta M d C, Moreno DA, Carvajal M. The physiological importance of glucosinolates on plant response to abiotic stress in Brassica. International Journal of Molecular Sciences. 2013;14(6):11607-11625
  33. 33. Kopsell DA, Sams CE. Increases in shoot tissue pigments, glucosinolates, and mineral elements in sprouting broccoli after exposure to short-duration blue light from light emitting diodes. Journal of the American Society for Horticultural Science. 2013;138(1):31-37
  34. 34. Chun JH, Kim S, Arasu MV, Al-Dhabi NA, Chung DY, Kim SJ. Combined effect of nitrogen, phosphorus and potassium fertilizers on the contents of glucosinolates in rocket salad (Eruca sativa Mill.). Saudi Journal of Biological Sciences. 2017;24(2):436-443
  35. 35. Augustine R, Bisht NC. Biotic elicitors and mechanical damage modulate Glucosinolate accumulation by co-ordinated interplay of glucosinolate biosynthesis regulators in polyploid Brassica juncea. Phytochemistry. 2015;117(1):43-50
  36. 36. Bodnaryk RP. Potent effect of jasmonates on indole glucosinolates in oilseed rape and mustard. Phytochemistry. 1994;35(2):301-305
  37. 37. Ludwig-Müller J, Schubert B, Pieper K, Ihmig S, Hilgenberg W. Glucosinolate content in susceptible and resistant Chinese cabbage varieties during development of clubroot disease. Phytochemistry. 1997;44(3):407-414
  38. 38. Thiruvengadam M, Chung IM. Selenium, putrescine, and cadmium influence health-promoting phytochemicals and molecular-level effects on turnip (Brassica rapa ssp. rapa). Food Chemistry. 2015;173:185-193
  39. 39. Çiçek N, Çakirlar H. The effect of salinity on some physiological. Bulgarian Journal of Plant Physiology. 2002;28(1-2):66-74
  40. 40. Francisco M, Moreno DA, Elena M, Ferreres F, García-viguera C, Velasco P. Simultaneous identification of glucosinolates and phenolic compounds in a representative collection of vegetable Brassica rapa. 2009;1216:6611-6619
  41. 41. Hanschen FS, Kaufmann M, Kupke F, Hackl T, Kroh LW, Rohn S, Schreiner M. Brassica vegetables as sources of epithionitriles: Novel secondary products formed during cooking. Food Chemistry. 2017;245(October):564-569
  42. 42. Grubb CD, Abel S. Glucosinolate metabolism and its control. Trends in Plant Science. 2006;11(2):89-100
  43. 43. Harborne JB, Baxter H, Moss GP. Phytochemical Dictionary: A Handbook of Bioactive Compounds from Plants. 2nd ed. London: Taylor & Francis; 1999
  44. 44. Verma N, Shukla S. Impact of various factors responsible for fluctuation in plant secondary metabolites. Journal of Applied Research on Medicinal and Aromatic Plants. 2015;2:105-113
  45. 45. Singh J, Upadhyay AK, Prasad K, Bahadur A, Rai M. Variability of carotenes, vitamin C, E and phenolics in Brassica vegetables. Journal of Food Composition and Analysis. 2007;20(2):106-112
  46. 46. Pérez-Gregorio MR, Regueiro J, González-Barreiro C, Rial-Otero R, Simal-Gándara J. Changes in antioxidant flavonoids during freeze-drying of red onions and subsequent storage. Food Control. 2011;22(7):1108-1113
  47. 47. Pérez-Balibrea S, Moreno DA, García-Viguera C. Improving the phytochemical composition of broccoli sprouts by elicitation. Food Chemistry. 2011;129(1):35-44
  48. 48. Palermo M, Pellegrini N, Fogliano V. The effect of cooking on the phytochemical content of vegetables. Journal of the Science of Food and Agriculture. 2014;94(6):1057-1070
  49. 49. Gliszczyńska-Swigło A, Ciska E, Pawlak-Lemańska K, Chmielewski J, Borkowski T, Tyrakowska B. Changes in the content of health-promoting compounds and antioxidant activity of broccoli after domestic processing. Food Additives & Contaminants., vol. 23, no. 11, pp. 1088-1098, 2006.
  50. 50. Giallourou N, Oruna-Concha MJ, Harbourne N. Effects of domestic processing methods on the phytochemical content of watercress (Nasturtium officinale). Food Chemistry. 2016;212:411-419
  51. 51. Kopsell DA, Kopsell DE, Lefsrud MG, Curran-Celentano J, Dukach LE. Variation in lutein, β-carotene, and chlorophyll concentrations among Brassica oleracea cultigens and seasons. Hortscience. 2004;39(2):361-364
  52. 52. Farré G, Sanahuja G, Naqvi S, Bai C, Capell T, Zhu C, Christou P. Travel advice on the road to carotenoids in plants. Plant Science. 2010;179(1-2):28-48
  53. 53. Baek KJ, SA JY, Lim SH, Park SU. Metabolic profiling in Chinese cabbage (Brassica rapa L. subsp. pekinensis) cultivars reveals that glucosinolate content is correlated with carotenoid content. Journal of Agricultural and Food Chemistry. 2016;64(21):4426-4434
  54. 54. Sy C, Gleize B, Dangles O, Landrier JF, Veyrat CC, Borel P. Effects of physicochemical properties of carotenoids on their bioaccessibility, intestinal cell uptake, and blood and tissue concentrations. Molecular Nutrition & Food Research. 2012;56(9):1385-1397
  55. 55. Kaulmann A, André CM, Schneider YJ, Hoffmann L, Bohn T. Carotenoid and polyphenol bioaccessibility and cellular uptake from plum and cabbage varieties. Food Chemistry. 2016;197:325-332
  56. 56. Palmero P, Panozzo A, Simatupang D, Hendrickx M, Van Loey A. Lycopene and β-carotene transfer to oil and micellar phases during in vitro digestion of tomato and red carrot based-fractions. Food Research International. 2014;64:831-838
  57. 57. Khachik F, Beecher GR, Goli MB, Lusby WR. Separation, identification, and quantification of carotenoids in fruits, vegetables and human plasma by high performance liquid chromatography. Pure and Applied Chemistry. 1991;63(1):71-80
  58. 58. Colle IJP, Lemmens L, Knockaert G, Van Loey A, Hendrickx M. Carotene degradation and isomerization during thermal processing: A review on the kinetic aspects. Critical Reviews in Food Science and Nutrition. 2016;56(11):1844-1855
  59. 59. Finley. Proposed criteria for assessing the efficacy of cancer reduction by plant foods enriched in carotenoids, glucosinolates, polyphenols and selenocompounds. Annals of Botany. 2005;95:1075-1096
  60. 60. Schonhof I, Blankenburg D, Müller S, Krumbein A. Sulfur and nitrogen supply influence growth, product appearance, and glucosinolate concentration of broccoli. Journal of Soil Science and Plant Nutrition. 2007;170:65-72
  61. 61. Charron CS, Sams CE. Glucosinolate Content and Myrosinase Activity in Rapid-cycling Brassica oleracea Grown in a Controlled Environment. Journal of the American Society for Horticultural Science. [Internet]. 2004;129:321-330
  62. 62. Flaishman MA, Peles Y, Dahan Y, Milo-Cochavi S, Frieman A, Naor A. Differential response of cell-cycle and cell-expansion regulators to heat stress in apple (Malus domestica) fruitlets. Plant Science. [Internet]. Elsevier Ireland Ltd; 2015;233:82-94
  63. 63. Makus DJ, Lester G, N TL, Dec LW, N TL, Long W. Effect of soil type, Light intensity, and Cultivar on leaf nutrients in mustard greens. 2002;23-28
  64. 64. Maggio A, De Pascale S, Ruggiero C, Barbieri G. Physiological response of field-grown cabbage to salinity and drought stress. European Journal of Agronomy. 2005;23:57-67
  65. 65. Pathirana I, Thavarajah P, Siva N, Wickramasinghe ANK, Smith P, Thavarajah D. Moisture deficit effects on kale (Brassica oleracea L. var. acephala) biomass, mineral, and low molecular weight carbohydrate concentrations. Scientia Horticulturae. [Internet]. Amsterdam: Elsevier; 2017;226:216-222
  66. 66. Khan MAM, Ulrichs C, Mewis I. Influence of water stress on the glucosinolate profile of Brassica oleracea var. italica and the performance of Brevicoryne brassicae and Myzus persicae. Entomologia Experimentalis et Applicata. 2010;137:229-336
  67. 67. Issarakraisila M, Ma Q, Turner DW. Photosynthetic and growth responses of juvenile Chinese kale (Brassica oleracea var. alboglabra) and Caisin (Brassica rapa subsp. parachinensis) to waterlogging and water deficit. Scientia Horticulturae. (Amsterdam). 2007;111:107-113
  68. 68. López-Berenguer C, García-Viguera C, Carvajal M. Are root hydraulic conductivity responses to salinity controlled by aquaporins in broccoli plants? Plant Soil. 2006;279:13-23
  69. 69. López-Berenguer C, Martínez-Ballesta MDC, Moreno DA, Carvajal M, García-Viguera C. Growing hardier crops for better health: Salinity tolerance and the nutritional value of broccoli. Journal of Agricultural and Food Chemistry. 2009;57:572-578
  70. 70. Jensen CR, Mogensen VO, Mortensen G, Fieldsend JK, Milford GFJ, Andersen MN, et al. Seed glucosinolate, oil and protein contents of field-grown rape (Brassica napus L.) affected by soil drying and evaporative demand. Field Crops Research. 1996;47:93-105
  71. 71. Anjana SU, Muhammad I. Agronomy for sustainable development. Ital. Journal of Agronomy. 2008;3:77-78
  72. 72. Grattan SR, Grieve CM. Salinity-mineral nutrient relations in horticultural crops. Amsterdam: Journal of Agronomy. 1998;78:127-157
  73. 73. Marino D, Ariz I, Lasa B, Santamaría E, Fernández-Irigoyen J, González-Murua C, et al. Quantitative proteomics reveals the importance of nitrogen source to control glucosinolate metabolism in Arabidopsis thaliana and Brassica oleracea. Journal of Experimental Botany. 2016;67:3313-3323
  74. 74. De Pascale S, Maggio A, Pernice R, Fogliano V, Barbieri G. Sulphur fertilization may improve the nutritional value of Brassica rapa L. subsp. sylvestris. European Journal of Agronomy. 2007;26:418-424
  75. 75. Li S, Schonhof I, Krumbein A, Li L, Stützel H, Schreiner M. Glucosinolate Concentration in Turnip (Brassica rapa ssp. rapifera L.) Roots as Affected by Nitrogen and Sulfur Supply. European Journal of Agronomy. [Internet]. 2007;55:8452-8457

Written By

Cristine Vanz Borges, Santino Seabra Junior, Franciely S. Ponce and Giuseppina Pace Pereira Lima

Submitted: 08 November 2017 Reviewed: 31 January 2018 Published: 24 October 2018