Phytochemical Composition, Sensory Acceptance, and Cultivation Potential of Sanguisorba verrucosa, Eruca vesicaria, and Scorzonera laciniata

Abstract

Three wild edible plant species native to the Mediterranean region, Sanguisorba verrucosa, Eruca vesicaria, and Scorzonera laciniata, were investigated to evaluate their potential for cultivation and integration into human diets. All three species were positively rated in sensory evaluations and exhibited high levels of specific metabolites of nutritional and health-related interest. Moderate concentrations of β-carotene were found across all species. Notably, S. verrucosa and E. vesicaria contained appreciable amounts of vitamin C, and the phenolic content in S. verrucosa exceeded that of many commonly consumed vegetables. Each species also proved to be a rich source of distinct organic acids: S. verrucosa for fumaric acid, E. vesicaria for citric acid, and S. laciniata for quinic acid. Although domestication led to a reduction in several bioactive compounds, the nutritional value of these plants remains significant. The compositional and sensory profiles of these species highlight their promise as leafy vegetables for sustainable diets and as functional food ingredients. Furthermore, their cultivation could support biodiversity conservation efforts and reduce harvesting pressure on wild populations, contributing to more sustainable agricultural practices.

1. Introduction

The modern human diet is often based on the consumption of a few highly cultivated species, and diversifying dietary intake to include a broader array of plant species is thus a promising way to improve human welfare. Despite their global prevalence and cultural significance, wild edible plants (WEPs) are frequently underestimated as significant dietary components in developed countries [1]. However, they are a key component in people’s diets worldwide [2], playing a crucial role in food security, especially in regions with limited agricultural resources or harsh climates [3,4]. The Food and Agriculture Organisation [5] reported that over 100 million people in the EU, 20% of the population, eat wild foods, and 65 million (14%) gather some wild food at least occasionally [6,7]. Moreover, there is currently a growing interdisciplinary interest in the study of wild plants as sources of both bioactive compounds and novel ingredients for haute cuisine. On the one hand, WEPs have rich nutritional profiles, providing essential micronutrients and macronutrients, including vitamins, proteins, minerals, and dietary fibers [8,9,10]. Furthermore, numerous studies have highlighted their therapeutic potential due to their antioxidant, antimicrobial, antiinflammatory, and antitumor properties [10,11,12,13], and they are natural antidiabetics as well [14]. On the other hand, the interest in haute cuisine is evidenced by chefs incorporating traditionally consumed wild species, particularly those from the Mediterranean diet, into their menus. Research by Łuczaj and Pieroni [15] highlighted the successful adoption of diverse local wild plants in avant-garde restaurants worldwide. Additionally, recent studies have explored the potential of wild plants to be marketed as novel functional foods, emphasizing innovative and health-promoting culinary applications [16].

For this study, three different species grown in the Mediterranean area were chosen: Sanguisorba verrucosa (S. verrucosa), Eruca vesicaria (E. vesicaria), and Scorzonera laciniata (S. laciniata). The genus Sanguisorba has proven to be a rich source of phytochemicals, with over 270 compounds identified to date. Among these, Sanguisorba officinalis stands out, with its extracts exhibiting a range of pharmacological activities, including notable anticancer, antiviral, and antiinflammatory effects [17]. More recently, Sanguisorba minor (S. minor), used as an edible plant in the Mediterranean diet, has been reported as a good source of bioactive compounds, particularly polyphenols, with a potential use in food supplements and pharmaceutical formulations [18]. Few studies have been found on S. verrucosa, such as some reports on the consumption of tender leaves and stems in salads in Spain [19]. Rivera et al. [20] documented the traditional medicinal use of S. verrucosa across various applications related to the circulatory, digestive, and integumentary systems. The genus Eruca, part of the Brassicaceae family, includes species commonly known as rocket salads, which hold an important place in the Mediterranean diet due to their rich phytochemical content [21,22]. In addition to their culinary relevance, traditional uses of these plants have been recorded in various ethnobotanical sources [23,24]. While Eruca sativa Mill. has been extensively studied, E. vesicaria has attracted comparatively limited scientific attention, despite its utilization in traditional dishes such as salads, omelets, and gazpachos [20]. Regarding Scorzonera species, some are utilized as vegetables or spices in parts of Asia and Europe [25], and a recent review encompassing 55 species highlighted their significant pharmacological potential [26]. Specifically, the young leaves of S. laciniata are occasionally consumed in salads [20], and evidence of their phytochemical richness and significant antioxidant activity suggests potential applications in dietary supplementation, as well as in microbiological and pharmacological research [27].

WEPs are predominantly harvested from their natural habitats. However, traditional foraging has several limitations, such as inconsistent supply, limited quality control, and the potential for ecological degradation due to overharvesting. In response to these challenges, recent research has increasingly turned to the cultivation of WEPs as a sustainable alternative [28,29,30,31]. Emerging evidence indicates that domestication and cultivation may offer significant advantages, such as enhanced yields and greater consistency in both nutritional and phytochemical profiles [32,33]. These findings underscore the potential of integrating WEP cultivation into rural Mediterranean agriculture, which may also contribute to the preservation of natural ecosystems by alleviating harvesting pressure on wild populations. Given the culinary and pharmaceutical potential of the three selected Mediterranean species and the limited available data on their organoleptic properties and cultivation performance, this study aims to investigate their sensory attributes and chemical composition. Special attention is given to the effects of cultivation on leaf composition, with a focus on evaluating the potential for large-scale commercial production and the corresponding implications for quality parameters.

2. Materials and Methods

The two-year study included both wild and cultivated harvesting.

2.1. Gathering of Wild Plants

Initial plant material was collected from various natural habitats, predominantly within the Region of Murcia. The taxonomic identification was performed by Antonio Robledo Miras (BSc Botany). The collection adhered to established protocols, i.e., recording parameters such as collection date, collector, specific locality, geographical coordinates, altitude, habitat description, land use, geological features, drainage characteristics, number of sampled plants, and estimated population size. Visual documentation, including images of whole plants and detailed leaf morphology, is provided for S. oleraceus and S. tenerrimus in Figure 1. Sampling was conducted over a two-month period at the beginning of the year (January to March). To ensure data representativeness, five distinct batches of each species were independently sampled from diverse locations (Figure 2) that comprised a heterogeneous mosaic of Quaternary, calcareous, siliceous, and mixed sedimentary substrates, including alluvial and colluvial deposits, marls, conglomerates, sandstones, schists, and other lithologies that generate diverse edaphic and hydrological conditions. Each batch was treated as an independent sample and contained at least 500 g of the edible portion from multiple individuals. These individuals were representative of the population at a mature harvest stage, characterized by fully expanded leaves and optimal leaf size. Furthermore, viable whole plants were transferred to IMIDA’s facilities for cultivation, facilitating seed production for subsequent germination and cultivation trials. These seeds were also accessioned into the conservation collection of IMIDA’s germplasm bank (BAGERIM). Seed cleaning was performed manually, with no disinfection procedures applied to preserve natural germination characteristics.

2.2. Germination Assay in Petri Dishes

One hundred seeds from each species were distributed across three Petri dishes (33, 33, and 34 seeds per dish) lined with filter paper. Deionized water was supplied on demand to maintain consistent moisture levels. The Petri dishes were taken to a controlled environment growth chamber with the following conditions: 16/8 h light/night photoperiod, at a 25 °C (day)/20 °C (night) temperature, 75% relative humidity, and 350 μmol m⁻² s⁻¹ flux density. Radicle emergence was established as the criterion for germination, at which point the germinated seeds were removed from the Petri dish. The duration of this germination trial was 15 days.

2.3. Field Germination Assays

During the spring season, an experiment was conducted using the obtained seeds at the IMIDA experimental facilities in La Alberca, Murcia, Spain. Prior to seed sowing, the plot was tilled, and compost was applied at a rate of 1 kg m⁻², and then thoroughly integrated into the soil via a second pass with a tractor. After that, seeds were sown with an optimum plant density established as 400 plants m⁻². The germination rates obtained from controlled Petri dish assays and seed weight (seeds per gram) were used to calculate the appropriate dose of each species.

Four different 1 m² plots (replications) were used for each species with a randomized block design. After the seeds were manually distributed into each plot, they were irrigated using an exudative pipe, and this continued until the end of the experiment. Twenty-four days after sowing, germination monitoring was carried out using a 60 cm × 60 cm template placed in the center of each of the 1 m² plots. Previously, in each plot, five 10 cm × 10 cm cells were randomly chosen.

2.4. Cultivation Trial

This experiment was carried out during the autumn–winter period. Seeds were initially germinated in Petri dishes in order to obtain better homogeneity. After radical emergence, the germinated seeds were transplanted to seedbeds containing commercial seedbed substrate. After that, the trays were taken to a growth chamber under controlled conditions with temperatures of 25 °C during the day and 20 °C during the night, 75% of humidity, and a photoperiod of 16 h day/8 h night. The seedlings were later transplanted into the experimental plots. A planting density of 400 plants m⁻² was implemented, utilizing a 5 cm × 5 cm planting frame. The experimental design followed a randomized block design, with four blocks allocated to each species. Irrigation was delivered through a porous-pipe system and sustained for the duration of the experiment. Applications were timed at regular intervals and adjusted as necessary to maintain optimal soil moisture while preventing oversaturation. This regimen standardized water availability across individuals, resulting in uniform growth patterns. Plants were harvested during April–May when leaf length was between 5 and 10 cm and prior to flowering, corresponding to the mature harvest stage established for wild plants, characterized by fully expanded leaves and optimal leaf size. Samples consisted of the youngest aerial plant parts. Four replicates were collected for each species, with each replicate containing ten individual plants. These replicates were subsequently pooled and used for sensory attribute analysis on the same day.

2.5. Metabolite Analysis

2.5.1. Metabolite Extraction

At least 100 g from each replicate of both wild-gathered and cultivated plants were homogenized, flash-frozen in liquid nitrogen, and stored at −80 °C for subsequent metabolite analysis.

For the analysis of soluble sugars, organic acids, and total phenolic content, an extraction on 3 g of frozen material was carried out as described by López et al. [34], with 10 mL of ultrapure water (Millipore, Molsheim, France) in a Polytron (PT-MR 3100, Malters, Switzerland), and 25 mL of ethyl acetate (J.T. Baker, Deventer, Holland, The Netherlands) to remove pigments. The aqueous fraction was used for analyzing soluble sugars, organic acids, and total phenolic content.

The extraction of carotenoids and chlorophylls was performed according to Hernández et al. [35], by taking 1 g of frozen material. The extraction solvent consisted of 25 mL of a 1:1 (v/v) mixture of methanol and tetrahydrofuran (Scharlau, Sentmenat, Spain), supplemented with MgO (Merck, Darmstadt, Germany) and 0.1% (w/v) butylated hydroxytoluene (BHT) (Sigma-Aldrich, Saint Louis, MO, USA) to inhibit degradation. Post-extraction, the solvent was evaporated to achieve a final concentrated volume of 2 mL. β-apo-8′carotenal (Sigma-Aldrich, Saint Louis, MO, USA) was added to the extract to serve as an internal standard for accurate quantification.

For the extraction of vitamin C (ascorbic and dehydroascorbic), 3 g of fresh material were homogenized with EDTA, 0.05% (w/v), and dithiothreitol (Sigma, Steinheim, Germany).

2.5.2. Determination of Sugar and Organic Acids

The quantification of soluble sugars was performed using a Hewlett–Packard model 1100 High-Performance Liquid Chromatography (HPLC) system (Waldbronn, Germany) equipped with a Refractive Index (RI) detector. Chromatographic separation was achieved using a CARBOSep CHO-682 LEAD column (300 mm × 7.8 mm i.d., Transgenomic, Omaha, NE, USA). The isocratic mobile phase consisted of ultrapure water, delivered at a constant flow rate of 0.4 mL min⁻¹. To determine the linearity of the detector response and to establish the limits of detection for sucrose, glucose, and fructose, a series of standard solutions (≥99.5% purity, Sigma, Steinheim, Germany) was prepared and injected. These standards spanned a concentration range of 1–10 g⋅L⁻¹.

Organic acids were determined using a liquid chromatography system (Agilent 1200; Agilent Technologies, Santa Clara, CA, USA) coupled to a triple quadrupole tandem mass spectrometer (MS/MS). The analytical methodology was adapted from the protocol described by Flores et al. [36]. For calibration and to ascertain the linearity of the detector response, standard solutions were prepared from commercially sourced external standards (Sigma-Aldrich, St. Louis, MO, USA).

2.5.3. Chlorophyll and Carotenoid Analysis

The analysis of carotenoids and chlorophylls was performed according to Hernández et al. [35], utilizing an Agilent 1200 High-Performance Liquid Chromatography (HPLC) system (Agilent Technologies, Santa Clara, CA, USA) equipped with a photodiode array detector (PAD). Chromatographic separation was achieved using a binary solvent system consisting of methanol (solvent A) and methyl tert-butyl ether (solvent B) (Scharlau, Sentmenat, Spain). The mobile phase was delivered at a constant flow rate of 1.0 mL⋅min⁻¹ with the following gradient program: an initial 15% solvent B for 20 min, followed by a linear gradient increasing to 30% solvent B over 20 min. This composition was then maintained for 10 min, culminating in an 80-min linear gradient to 90% solvent B. For quantification, carotenoids and chlorophylls were determined against commercially available external standards obtained from DHI LAB (Hoersholm, Denmark). Luteoxanthin was quantified relative to antheraxanthin, and the cis isomers of β-carotene were quantified with respect to their all-trans isomer.

2.5.4. Ascorbic Acid Analysis

The quantification of vitamin C (ascorbic acid and its derivative, dehydroascorbic acid) was carried out with an HPLC equipped with an MS/MS detector, according to Fenoll et al. [37]. The mobile phase utilized was a 0.2% (v/v) solution of formic acid (Scharlau, Sentmenat, Spain), delivered at a flow rate of 0.4 mL min⁻¹. For calibration, L-ascorbic acid standard solutions (Sigma-Aldrich, St. Louis, MO, USA) were prepared, with the addition of 0.05% (w/v) EDTA to ensure stability.

2.5.5. Total Phenolic Content Analysis

The total phenolic content was assessed using the Folin–Ciocâlteu method. A standard curve was generated from gallic acid (Fluka, Steinheim, Germany) solutions, with concentrations ranging from 50 to 400 mg⋅L⁻¹. Phenolic concentrations in samples were then calculated based on their absorbance at 765 nm, measured with a Shimadzu UV-2401PC spectrophotometer (Kyoto, Japan).

2.6. Sensory Assessment

A panel of fifteen trained assessors was used to assess the sensory profiles of the cultivated plants through the application of a descriptive analysis. Prior to evaluation, plants were prepared by thoroughly washing and then immersed in a solution of water containing food-grade hypochlorite. The leaves were subsequently rinsed with abundant drinking water, and excess moisture was removed using a kitchen centrifuge. The prepared leaves were then refrigerated until the sensory evaluation session. For the tasting session, the samples were presented raw and undressed in appropriately coded, food-grade plastic cups. To mitigate bias, the tasting order for each panelist was randomized. A separate sample, consisting of a single leaf from each species, was prepared for the assessment of leaf color and shape. All samples were coded with unique three-digit random numbers to prevent any potential influence that sequential numbering or alphabetical codes might exert on panelists’ judgments. Sensory perception data were primarily captured numerically, based on an ideal reference pattern, a defined scale, or responses to specific questions, as described in reference [38]. Initially, trained tasters quantitatively evaluated five distinct sensory attributes: color, shape, smell, taste (flavor), and texture. Each attribute was rated using a nine-point hedonic scale, with “9” representing “I like it very much” (maximum value) and “1” signifying “I dislike it very much” (minimum value). Upon completion of the individual attribute evaluations, the panelists provided an overall assessment of general acceptability, integrating their perceptions of the aforementioned characteristics.

A subsequent questionnaire quantitatively assessed specific taste characteristics, including bitterness, sweetness, pungency (hotness), acidity, and texture (further detailed as crispness, stringiness, or fibrousness). These attributes were rated on a unipolar scale of ten values, ranging from “0” (minimum value) to “9” (maximum value). Assessors were also provided an open-ended section at the end of the questionnaire to record additional comments and relevant specific observations. The sensory evaluation sessions were conducted between 12:30 p.m. and 14:00 p.m. in a controlled environment. The tasting area was quiet, well-lit, and free from extraneous odors to prevent interference with the assessment process. Each panelist performed their evaluations individually, with no time constraints imposed. To ensure palate cleansing between samples, filtered water was provided to each assessor.

2.7. Statistical Analysis

The statistical analysis of the data was performed using IBM SPSS Advanced Statistics (version 25.0). Depending on the data distribution, either an Analysis of Variance (one-way ANOVA) or the Kruskal–Wallis test was applied.

3. Results and Discussion

3.1. Germination and Cultivation

The germination rates of wild species are generally lower than those of domesticated leafy vegetables [39,40]. Moreover, differences in germination percentages have been reported between direct sowing and pre-sowing on Petri dishes for wild species [33]. Therefore, assessing seed germination viability is essential for determining the most effective propagation strategy. Marked differences in germination performance were observed between controlled conditions in Petri dishes and open field conditions. In Petri dishes, germination was relatively high for all three species after 15 days, with values of 71.0 ± 11.4% for S. laciniata, 79.7 ± 17.6% for S. verrucosa, and 91.8 ± 7.3% for E. vesicaria. In contrast, field germination was lower, particularly for S. verrucosa (6.7 ± 2.9%) as compared to E. vesicaria (19.5 ± 8.9%) and S. laciniata (26.3 ± 7.2%). Based on the obtained results, and to ensure maximum uniformity in the cultivation trial, seeds were initially germinated in Petri dishes and then transferred to seedbeds.

3.2. Soluble Sugars

The concentrations of the three soluble sugars varied among the three studied species. In E. vesicaria, the sugars were present in decreasing order as glucose, sucrose, and fructose. In L. laciniata, the order was glucose, fructose, and sucrose, whereas in S. verrucosa, glucose was the most abundant, followed by sucrose and fructose (

Table 1. Primary and secondary metabolite concentrations (µg gFW-1) in cultivated S. verrucosa, E. vesicaria, and S. laciniata. Values are mean ± SD (n = 4).

Metabolite	S. verrucosa	E. vesicaria	S. laciniata
Glucose	5275.0 ± 303.7	1877.3 ± 182.5	2527.3 ± 630.7
Sucrose	4699.7 ± 197.6	560.0 ± 29.2	1953.2 ± 217.8
Fructose	3837.1 ± 384.7	406.0 ± 21.1	2324.0 ± 541.5
Citric	762.4 ± 1.3	1235.9 ± 56.9	528.4 ± 80.7
Malic	445.4 ± 8.1	428.6 ± 18.6	1759.5 ± 72.3
Tartaric	2.3 ± 0.3	5.3 ± 0.2	7.8 ± 0.9
Fumaric	802.1 ± 38.7	54.2 ± 1.6	196.8 ± 34.8
Succinic	60.0 ± 4.6	30.8 ± 2.7	79.4 ± 9.0
Quinic	13.9 ± 0.7	9.6 ± 0.5	285.0 ± 5.3
Malonic	12.5 ± 1.3	9.9 ± 0.4	6.5 ± 1.0
Isocitric	21.2 ± 0.7	50.7 ± 3.1	11.7 ± 3.3
Ketoglutaric	2.8 ± 0.2	9.1 ± 0.6	16.7 ± 1.7
Glutamic	7.1 ± 0.3	7.4 ± 0.4	4.2 ± 0.4
Skimic	6.5 ± 0.0	0.9 ± 0.0	3.0 ± 0.3
Chlorophyll a	64.3 ± 1.2	96.4 ± 3.4	108.4 ± 3.4
Chlorophyll b	31.8 ± 0.9	45.4 ± 1.0	45.0 ± 1.5
All-trans-β-carotene	30.0 ± 1.0	16.0 ± 0.4	36.6 ± 0.0
Lutein	10.7 ± 0.3	13.0 ± 0.3	13.8 ± 0.2
All-trans violaxanthin	2.9 ± 0.0	3.6 ± 0.1	5.5 ± 0.2
9-cis-Neoxanthin	1.8 ± 0.1	2.6 ± 0.1	2.3 ± 0.0
9-cis-β-carotene	2.9 ± 0.1	18.3 ± 0.0	3.5 ± 0.0
Luteoxanthin	3.1 ± 0.2	6.4 ± 0.1	2.6 ± 0.1
13-cis-β-carotene	1.0 ± 0.1	1.7 ± 0.0	1.3 ± 0.0
Vitamin C	933.4 ± 204.3	1012.9 ± 128.5	530.8 ± 99.3
Total phenolics	3098.3 ± 93.6	246.7 ± 19.2	286.6 ± 11.8

). This composition differs from the patterns observed in other Sanguisorba species. For instance, in S. minor, Karkanis et al. [41] reported fructose and glucose as the dominant sugars, while Ceccanti et al. [30] found sucrose to be most abundant, followed by glucose and fructose. These differences may be attributed to interspecific variation as well as environmental, genetic, or methodological factors influencing carbohydrate metabolism.

Among the species analysed, S. verrucosa showed the highest concentrations of all three sugars, with particularly elevated levels of glucose and sucrose as compared to L. laciniata and E. vesicaria. Additionally, S. verrucosa and L. laciniata had a markedly higher fructose content than other wild cultivated species, such as Sonchus oleraceus (S. oleraceus) and Sonchus tenerrimus (S. tenerrimus) [33]. When compared with commonly cultivated leafy vegetables, the studied species generally exhibited higher levels of soluble sugars. Glucose content in S. verrucosa was higher than that reported for both lettuce and lamb’s lettuce [42,43]. Similarly, fructose concentrations in S. verrucosa and L. laciniata exceeded those in lamb’s lettuce, though they remained below the levels observed in lettuce. All three species showed higher sucrose levels than lamb’s lettuce and baby lettuce [43,44]. As compared with lettuce, S. verrucosa had higher sucrose concentrations, L. laciniata had similar levels, and E. vesicaria exhibited lower values [42].

3.3. Organic Acids

In the present study, the predominant organic acids identified across all three species were citric, malic, and fumaric acids (Table 1). However, the composition and concentration of the organic acids varied significantly among the species. In S. verrucosa, fumaric acid was the predominant organic acid, with levels exceeding those in E. vesicaria and S. laciniata by 15-fold and 4-fold, respectively. When compared to other wild species within the same genus, distinct profiles emerged. For instance, in S. minor, only a minor concentration of fumaric acid [30] or even trace amounts [8,41] were detected as compared to other organic acids. Regarding other cultivated wild and commonly consumed leafy vegetables, such as lettuce and lamb’s lettuce, fumaric acid was found in higher concentrations in S. verrucosa [33,42,43]. The elevated fumaric acid content in S. verrucosa underscores the potential industrial relevance of this species, as fumaric acid is widely used as a food acidulant in beverages and baking powders [45]. Additionally, fumaric acid serves as a pharmaceutically active compound in the treatment of psoriasis and multiple sclerosis [46], highlighting its dual importance in both the food and pharmaceutical industries.

E. vesicaria was notable for its high content of citric acid and its positional isomer, isocitric acid (Table 1). This species also exhibited higher citric acid levels as compared to other cultivated wild species, such as S. oleraceus and S. tenerrimus [33] and similar concentration to lamb’s lettuce [43]. Citric acid has been shown to protect both the liver and brain from oxidative damage and to support tissue regeneration, owing to its antioxidant properties [47]. Additionally, it contributes to reducing urinary acidity, alleviating stress, and slowing age-related processes. In the food industry, citric acid also plays a role in inhibiting lipid oxidation, thereby extending the shelf life of food products [48].

In S. laciniata, malic acid was the most abundant organic acid, with concentrations approximately four times higher than those observed in S. verrucosa and E. vesicaria (Table 1). This species also presented slightly higher malic acid levels as compared to other wild edible species, such as S. oleraceus and S. tenerrimus [33], but lower than commonly consumed vegetables such as lettuce and lamb’s lettuce [42,43]. Malic acid has been reported to enhance the perception of sweetness when combined with sucrose [49]. In addition to malic acid, S. laciniata was distinguished by its elevated contents of quinic, fumaric, succinic, and ketoglutaric acids. Quinic acid, in particular, was found at levels 21 and 30 times higher than in S. verrucosa and E. vesicaria, respectively, while ketoglutaric acid concentrations were six and two times higher than in the same species. These results suggest that S. laciniata may serve as a valuable natural source of quinic acid, a compound associated with multiple biological activities, including antioxidant, antimicrobial, anticancer, antiviral, antidiabetic, antiageing, antinociceptive, and analgesic properties [50]. Finally, S. laciniata and S. verrucosa may serve as valuable sources of succinic acid, with S. verrucosa also being a notable source of fumaric acid when compared to commonly cultivated vegetables such as lettuce [33,42,43].

3.4. Carotenoids

Among the three species analyzed, S. laciniata showed the highest total carotenoid content (65.6 µg g⁻¹), followed by E. vesicaria (61.6 µg g⁻¹) and S. verrucosa (52.4 µg g⁻¹). The values observed were lower than those reported for other cultivated wild species, including S. tenerrimus and S. oleraceus [33], as well as S. minor [32]. In the case of lettuce, Kim et al. [51] documented comparable or higher concentrations, depending on the cultivar. Furthermore, in comparison to lamb’s lettuce, the carotenoid content in the studied species was consistently lower across all cultivation conditions [35].

Regarding the carotenoid profile, carotenes represented a substantial proportion of the total carotenoid content, ranging from 58% to 65% in the studied species (Table 1). Whereas in S. laciniata and S. verrucosa, all-trans-β-carotene was identified as the predominant carotene, E. vesicaria exhibited higher levels of 9-cis-β-carotene, surpassing the concentration of the all-trans isomer. Interestingly, previous studies on Eruca sativa have identified zeaxanthin as the principal carotenoid, followed by β-carotene, cryptoxanthin, and lutein [52]. This discrepancy with our results may be attributable to interspecific variation within the genus Eruca, differences in environmental conditions, or methodological factors such as extraction and quantification techniques. Although total β-carotene concentrations in the three species were lower than those reported for vegetables such as chicory, dandelion, and both cultivated and wild forms of rocket [53], they may still be regarded as relatively good sources of this provitamin A compound, with levels generally exceeding those typically found in lettuce [51]. β-carotene is the major carotenoid present in the human diet [54], being the most potent precursor to vitamin A, and is naturally found as a mixture of its various cis and trans isomers. It has a potent antioxidant capacity and can function as a lipid scavenger and a singlet oxygen quencher [55]. Moreover, β-carotene has numerous associated health benefits, such as reducing the risk of cardiovascular diseases and specific cancers, enhancing the immune system, and offering protection against age-related macular degeneration, a primary cause of irreversible blindness in adults [56]. In vegetables, this carotenoid is predominantly found in the all-trans configuration, which can be converted into cis isomers during processing and intestinal digestion [57,58]. These cis forms differ in their biological functions, generally exhibiting a lower provitamin A activity and a reduced antioxidant capacity. Additionally, evidence from both in vitro and in vivo studies indicates that the all-trans isomer of β-carotene is more readily absorbed and utilized by the body than its cis counterparts [59,60]. According to our results, S. verrucosa and S. laciniata may represent more effective dietary sources of β-carotene than E. vesicaria, due to the greater proportion of the all-trans isomer present in these species, even though total β-carotene concentrations were similar.

Among the xanthophylls, lutein was the most abundant in all three species, constituting over 50% of the total xanthophyll fraction. This observation aligns with established findings that identify lutein as the predominant xanthophyll in the photosynthetic apparatus of higher plants [61]. Lutein and zeaxanthin are typically abundant in leafy vegetables such as kale, spinach, broccoli, and lettuce [62]. However, in the present analysis, zeaxanthin levels were undetectable in all species evaluated. The highest lutein concentration was observed in S. laciniata and E. vesicaria, and the lowest in S. verrucosa, being in all cases lower than those reported for certain cultivated Sonchus species [33] and for green vegetables such as parsley, spinach, and kale [63]. Luteoxanthin was the second most abundant xanthophyll, representing 11–25% of total xanthophylls. The highest concentration of luteoxanthin was observed in E. vesicaria, followed by S. verrucosa and S. laciniata, with all values falling within the range previously reported for S. oleraceus and S. tenerrimus [33]. Luteoxanthin has been reported to possess biological activities of pharmacological interest, including antimicrobial effects against Helicobacter pylori and anticancer properties. More recently, it has been shown to have an inhibitory activity against the angiotensin-converting enzyme 2 (ACE2) receptor, which is utilized by SARS-CoV-2 for cellular entry, suggesting a potential role in the development of antiviral agents [64,65,66]. According to previous findings, which identified lutein, β-carotene, violaxanthin, and neoxanthin as the predominant carotenoids present in the chloroplasts of terrestrial plants [67,68], violaxanthin and neoxanthin were the third and fourth most abundant xanthophylls in the species analysed. The highest concentrations of violaxanthin were observed in S. laciniata, with neoxanthin levels comparable to those found in S. verrucosa and E. vesicaria, and other cultivated and wild leafy vegetables [33,53]. Similar to the other carotenoids, neoxanthin and violaxanthin are recognized for their strong antioxidant potential [69].

3.5. Chlorophylls

Chlorophyll concentrations in all three species (Table 1) fell within the range previously reported for S. minor [32] and were slightly lower than those observed in cultivated wild species such as S. oleraceus and S. tenerrimus [33]. These values were considerably lower than those reported for lettuce [42], as well as for other cultivated leafy vegetables, including garden rocket and wild rocket (Diplotaxis spp.) [53], but were comparable to or higher than those found in lamb’s lettuce [35]. Although less extensively studied than other plant-derived phytochemicals, chlorophylls and their derivatives have been suggested to exhibit various therapeutic properties. Notably, while the pharmacological effects of medicinal plants are typically attributed to their secondary metabolites (e.g., alkaloids, flavonoids, terpenes) [70,71], emerging evidence indicates that photosynthetic pigments, particularly chlorophylls, may also contribute significantly to their health-promoting properties [72,73]. In particular, these pigments may function as antioxidants, antimutagenic agents, and anticancer agents [74].

3.6. Vitamin C

The vitamin C content varied markedly among the species studied (Table 1). S. verrucosa and E. vesicaria were particularly rich in vitamin C, with levels sufficient to meet the recommended daily intake (RDI) for adult women (75 mg) with the consumption of just 80 g of S. verrucosa or 74 g of E. vesicaria. These findings indicate that both species may serve as effective natural sources of dietary vitamin C. When compared to spinach, which has been reported to contain up to 1500 µg g⁻¹ of vitamin C in certain studies [75,76], E. vesicaria and S. verrucosa supplied approximately 64% and 59% of that amount, respectively. Additionally, all three species examined exhibited significantly higher vitamin C concentrations than those reported for lamb’s lettuce grown under varying fertilisation conditions [35]. Relative to lettuce, S. verrucosa and E. vesicaria contained approximately five times more vitamin C, while S. laciniata contained around three times more [42]. Moreover, in comparison to other cultivated wild leafy vegetables, E. vesicaria and S. verrucosa showed vitamin C levels approximately twice as high as those found in S. oleraceus and S. tenerrimus [33], further highlighting their nutritional potential. These results are particularly relevant given the essential role of vitamin C in human health. As a key antioxidant, vitamin C supports numerous metabolic functions [77] and plays a central role in immune system function [78]. Since humans are incapable of synthesizing vitamin C endogenously, a regular intake from dietary sources is necessary to maintain adequate physiological levels [79]. In this context, the high concentrations found in S. verrucosa and E. vesicaria underscore their value as important contributors to dietary vitamin C intake.

3.7. Total Phenolic Content

The total phenolic content in the three species analyzed was notably high, with S. verrucosa exhibiting a particularly elevated concentration (Table 1). Leafy vegetables are generally rich in phenolic compounds, which contribute significantly to their antioxidant properties. S. verrucosa stands out as an excellent source of phenolic compounds, with a total phenolic content comparable to that reported for curry leaves (Murraya koenigii) (3469–5085 µg/100 g FW), which is considered among the vegetables with the highest phenolic content [80]. The content of S. verrucosa is generally much higher than the values reported for spinach, a leafy vegetable that is considered to be rich in phenolic compounds, with values ranging from 800 to 2088 µg/100 g FW [81,82,83,84,85]. Other vegetables, such as Chinese cabbage, broccoli, and lettuce, also show a high phenolic content, often above 1000 µg/g FW [86,87], values which were always lower than in S. verrucosa. When compared with cultivated S. tenerrimus and S. oleraceus, S. verrucosa showed markedly higher phenolic concentrations (up to 16 times more) [33]. This remarkable phenolic concentration is particularly relevant given the importance of phenolic compounds in Sanguisorba species due to their bioactive properties [41]. From a nutritional perspective, the inclusion of these wild species in the human diet could be highly beneficial.

3.8. Sensory Attributes

Median values were used to summarize the sensory evaluation parameters, as they provide a more reliable measure of central tendency in the presence of asymmetrical distributions or outliers. The qualitative satisfaction evaluation of the cultivated species showed a relatively high general acceptance in most parameters (Figure 3). In terms of color, all three species showed similar interquartile ranges, suggesting a generally consistent perception. However, S. laciniata had a slightly lower median, and E. vesicaria showed a greater variability, indicating a less favorable and more inconsistent evaluation, respectively. For shape, S. verrucosa and E. vesicaria received high median scores (7–8), suggesting a positive perception of leaf form. In contrast, S. laciniata had a lower median (5), indicating a less favorable assessment. Odor was the lowest-rated attribute overall, with a median score of 5 across all species. In E. vesicaria, despite a limited central dispersion, there was notable variability with extreme values, reflecting inconsistent evaluations. Regarding taste, S. verrucosa and S. laciniata received relatively high medians (7 and 6), suggesting good acceptance. E. vesicaria had a lower median (4), indicating a less favorable taste. Texture was generally rated positively, with medians of 7 for S. verrucosa and E. vesicaria, and 6 for S. laciniata. The overall scores varied: S. verrucosa received the highest median (7), followed by S. lacinata (5) and E. vesicaria (4). E. vesicaria also exhibited the greatest variability and most outliers. In summary, S. verrucosa was rated most favorably across attributes, with higher median scores and more consistent evaluations, indicating a strong overall preference among panelists.

The sensory evaluation revealed notable differences among the three species across all attributes (Figure 4). E. vesicaria stood out with the highest median bitterness and the widest range, whereas S. verrucosa and S. laciniata were markedly less bitter, with median values of 2 and 1, respectively, and no values exceeding 5. For crunchiness, E. vesicaria also had the highest median (7), indicating that it was perceived as the crunchiest, although its lower whisker extended considerably, and thereby suggesting variability. The other two species had moderate median values (4) and broader distributions. All species showed a similar median in fibrosity (4–5), indicative of a moderate fibrous texture, with E. vesicaria showing the greatest variability within the interquartile range. Sweetness was generally low across species; E. vesicaria had the lowest median (2), while S. verrucosa and S. laciniata had higher, though still modest, median values (4). E. vesicaria also exhibited a much higher pungency than the other two species, which had very low median values (0–1) and narrow ranges, indicating a general lack of pungency. Lastly, acidity was perceived as low in all species (medians 1–2), although E. vesicaria and S. laciniata showed broader ranges, suggesting occasional samples with perceptible acidity, unlike the more uniformly mild S. verrucosa. The results highlight distinct sensory differences among the three species evaluated. E. vesicaria was characterized by higher levels of bitterness, crunchiness, and pungency, as well as a greater variability in several attributes. In contrast, S. verrucosa and S. laciniata showed milder profiles, with consistently lower scores for bitterness and pungency and slightly higher sweetness. Overall, these differences suggest that each species offers a unique sensory profile, which may inform on their potential applications in culinary use or selection in breeding programs aimed at specific flavor qualities.

3.9. Wild-Gathered vs. Cultivated Plants

The changes in the metabolite profiles associated with the organoleptic and functional quality following cultivation are shown in Figure 5, expressed as percentages relative to their levels in wild-collected plants. Supplementary Table S1 provides the detailed compositions of primary and secondary metabolites in wild specimens. The domestication of the three studied species resulted in a consistent decline across all analyzed metabolites. Concerning specific metabolite families, E. vesicaria exhibited the greatest reductions in total sugar and organic acid contents after cultivation. The most substantial decrease in carotenoids was observed in S. verrucosa, while the largest reductions in chlorophyll concentrations occurred in both S. verrucosa and E. vesicaria. The standard deviations associated with vitamin C levels reflect the high variability among wild populations from different geographical locations. Total phenolic compounds decreased by approximately 60% across all species following domestication.

Numerous studies have consistently demonstrated that wild edible plants tend to contain higher concentrations of phytochemicals as compared to their domesticated counterparts [88,89]. In the case of Sanguisorba minor, domestication has not been shown to significantly alter free sugar content, although wild populations exhibited slightly higher levels of fructose and glucose [30]. Conversely, an increase in organic acid concentrations has been observed in cultivated plants [30,41]. Regarding pigment content, in a previous study evaluating the effect of open-field, pot, and soilless cultivation as compared to wild plants, carotenoid and chlorophyll contents showed strong species- and system-dependent variation, with open-field cultivation generally resulting in higher carotenoid levels than other cultivation methods [90]. A general decline in total phenolic compounds following domestication has been reported in several species, including S. minor [90], Portulaca oleracea, and Porophyllum ruderale [91]. However, it has also been shown that the phenolic content, and consequently the antimicrobial and antitumor potential, of S. minor can be enhanced under specific substrate and fertilization regimes [41]. Closely related species, such as S. oleraceus and S. tenerrimus, exhibit distinct responses to cultivation. Notably, S. tenerrimus showed significant increases in total soluble sugars, organic acids, β-carotene, and vitamin C, whereas S. oleraceus experienced a decrease in phenolic content [33]. These interspecific differences are likely attributable to both species-specific physiological responses and the substantial influence of environmental factors. Abiotic conditions, including light intensity, temperature, water availability, and soil nutrient levels, significantly affect the biochemical composition of both wild and cultivated plants [92,93]. Cultivated plants were grown with organic amendments and regular irrigation, whereas wild plants were collected from habitats mostly characterized by low rainfall and poor soil fertility. Elevated levels of sugars, organic acids, and bioactive compounds in wild populations may reflect adaptive responses to environmental stressors such as drought, nutrient limitations, and poor soil conditions. Under such conditions, wild plants accumulate soluble sugars to maintain osmotic balance and cellular integrity [94,95], while organic acids such as malate, citrate, and succinate contribute to stress tolerance by enhancing resistance to water deficits, metal toxicity, and hypoxic conditions [96]. Additionally, exposure to environmental stress can induce the synthesis of secondary metabolites, particularly phenolic compounds, as part of the plant’s defense mechanisms [97]. Similarly, vitamin C functions as an antioxidant, and its production is enhanced under stress to counteract reactive oxygen species [98]. Consequently, wild-collected plants tend to have higher vitamin C levels, reflecting their exposure to greater environmental stress compared to cultivated plants.

4. Conclusions

The integration of WEPs into both agricultural systems and human diets offers numerous advantages. WEPs are well adapted to Mediterranean climates, and their cultivation contributes to biodiversity conservation efforts by alleviating harvesting pressure on wild populations. From a nutritional perspective, WEPs contribute to dietary diversification and are rich in bioactive compounds, offering potential health benefits while also opening venues for rural development and local economic growth. Although the domestication process led to some decline in phytochemical content in the three species studied, they continue to exhibit a significant nutritional value. After cultivation, all three species retained sugar levels higher than those found in many commonly consumed vegetables and remained valuable sources of specific organic acids: S. verrucosa for fumaric acid, E. vesicaria for citric acid, and S. laciniata for quinic acid. Additionally, moderate levels of β-carotene were observed, exceeding those in lamb’s lettuce, and both S. verrucosa and E. vesicaria contained appreciable amounts of vitamin C. Notably, S. verrucosa demonstrated phenolic concentrations surpassing those of most traditional vegetables. These findings indicate that targeted fertilization and cultivation strategies could further enhance the nutritional quality of these species, underscoring their relevance in the development of sustainable and health-oriented food systems.