Next Article in Journal
Establishment and Optimization of Flavonoid Extraction and Detection System for Hemerocallis
Next Article in Special Issue
Phytochemical Analysis and Specific Activities of Bark and Flower Extracts from Four Magnolia Plant Species
Previous Article in Journal
Influence of Selenium, Titanium, and Silicon Nanoparticles on the Growth, Yield, and Fruit Quality of Mango under Drought Conditions
Previous Article in Special Issue
Synergistic Effect of a Plant-Derived Protein Hydrolysate and Arbuscular Mycorrhizal Fungi on Eggplant Grown in Open Fields: A Two-Year Study
Font Type:
Arial Georgia Verdana
Font Size:
Aa Aa Aa
Line Spacing:
Column Width:

Variability in Chemical Profile and Bioactivities of the Flesh of Greek Pumpkin Landraces

Maria G. Leichtweis
Adriana K. Molina
Maria Inês Dias
Ricardo C. Calhelha
Tânia C. S. P. Pires
Ourania Pavli
M. Beatriz P. P. Oliveira
Spyridon A. Petropoulos
Lillian Barros
1,2,* and
Carla Pereira
Centro de Investigaçaão de Montanha (CIMO), Instituto Politeécnico de Bragança, Campus de Santa Apoloónia, 5300-253 Bragança, Portugal
Laboratório Associado para a Sustentabilidade e Tecnologia em Regiões de Montanha (SusTEC), Instituto Politécnico de Bragança, Campus de Santa Apolónia, 5300-253 Bragança, Portugal
Laboratory of Genetics and Plant Breeding, Department of Agriculture, Crop Production and Rural Environment, University of Thessaly, Fytokou Street, 38446 Volos, Greece
REQUIMTE—Science Chemical Department, Faculty of Pharmacy, University of Porto, Rua Jorge Viterbo Ferreira no. 228, 4050-313 Porto, Portugal
Laboratory of Vegetable Production, Department of Agriculture, Crop Production and Rural Environment, University of Thessaly, Fytokou Street, 38446 Volos, Greece
Authors to whom correspondence should be addressed.
Horticulturae 2023, 9(11), 1232;
Submission received: 3 September 2023 / Revised: 7 November 2023 / Accepted: 13 November 2023 / Published: 15 November 2023


The aim of this study was to evaluate the chemical profile of the flesh and yield parameters of Greek pumpkin genotypes, including nine local landraces and two commercially available cultivars, focusing on valorizing the genetic pool of Cucurbita sp. with high added value products. Yield parameters (mean fruit weight and total fruit yield) recorded high variability with genotypes V8 and V2 showing the highest fruit yield. Moreover, genotype V11 was the most abundant in glucose and total sugars and scored the highest sweetness index suggesting good taste and promising marketing attributes. The highest antioxidant activity (OxHLIA assay) was assessed in the V8 genotype, while the V2 genotype showed the highest α-, β- and total tocopherols content. Oxalic acid was the main organic acid, followed by malic and citric acids, while organic acid composition varied among the tested genotypes. Moreover, the flesh extracts showed varied antimicrobial activity against several bacteria and fungi, while no toxicity against non-tumor cells was recorded. In conclusion, our results make evident the presence of high innate variability in terms of crop performance, chemical composition and bioactive properties not only between the different genotypes but also at the intra-populational level. This finding is of high importance for the valorization of the local genetic pool of Cucurbita species through the selection of elite genotypes with high yield and quality of fruit, contributing to the conservation of valuable genetic material and limitation of the risk of genetic erosion due to neglect of local landraces.

1. Introduction

Among cucurbits, Cucurbita is considered a genus with considerable diversity in terms of the plant growth habits, fruit morphology and disease resistance, as well as the nutritional and phytochemical profile of fruit, which is mainly attributed to its complex domestication trajectories [1]. The genus presents a pre-Holocene distribution, originally in the form of bitter fruits [2], while its domestication took place in northern South and Central America ca. 10,000 B.P. [3], being included among the earliest crop domestication records. Although the relationships between wild and domesticated species are not well elucidated [4], it is known that domestication involved traits related to fruit quality, primarily focusing on the loss of bitterness and fruit size [1]. Indeed, among all plant species cultivated for their fruit, squashes and pumpkins produce the most profoundly sizable fruit in comparison to their non-domesticated wild forms [5].
Pumpkin is the fruit of Cucurbita species that belongs to the Cucurbitaceae family and it is traditionally consumed all over the world. Pumpkins belong to the genus Cucurbita L., which encompasses 9 species—C. maxima, C. pepo, C. moschata, C. argyrosperma, C. digitata, C. ficifolia, C. foetidissima, C. okeechobeensis and C. palmata—the first three being the most common around the world [6]. Although these three species have common requirements for optimum growth and development, due to their wide adaptation to soil and climatic conditions, they vary remarkably in fruit morphology (e.g., fruit size, shape, skin and flesh color and flesh structure), as well as in nutritional value [7]. Moreover, interspecies hybridization results in considerable heterogeneity at the intrapopulational level with a large number of subspecies and cultivars that vary in size, shapes, colours and taste [8]. The vernacular term of pumpkin is commonly used to describe the fruit of Cucurbita pepo L., C. moschata Duchesne ex Poir, C. maxima Duchesne, C. fificolia Bouché and C. argyrosperma K.Koch species, while it is interchangeably used with squash or winter squash terms [9]. In addition to the pleasant flavour and versatility, the known nutritional value and functional properties of pumpkin have gathered significant interest in scientific studies and health-conscious consumers [10]. As global consumer trends increasingly lean towards healthier and sustainable food choices, the diverse nutraceutical properties of pumpkin fruit have earned it a place of importance in modern diets [11,12,13].
The flesh (pulp) is the most appreciated part of the fruit, which represents more than 70% of the whole fruit [14]; it is a good source of carbohydrates, dietary fibres (especially pectins) and proteins, while it contains significant amounts of fat and minerals, varying among species and genotypes [15,16,17]. In nineteen accessions of Cucurbita ssp evaluated by Nwofia et al. [17], the crude fibres varied from 0.55% to 1.04% in fresh weight (moisture from 78.46 to 91.97%). Considering an adequate intake of 25 g/day of fibre by ESFA [2], this fruit contributes to a balanced diet. The high mineral content of iron, potassium, phosphorus, magnesium, selenium, calcium and copper has been well documented so far in different pumpkin varieties [18,19]. In the pulp of Cucurbita maxima, 184.34 ± 1.24 mg/100 g dw of potassium and 53.67 ± 0.19 mg/100 g dw of iron was found [17], while the “Waïgoré dollugo” dish, which consists of boiled pumpkin (78.2% of pumpkin fruit and 21.8% of water), presented 2753.0 ± 3.2 mg/100 g dw of potassium and 4.3 ± 0.2 mg/100 g dw of iron [20], which adequate intake and average requirement is, according to EFSA (European Food Safety Authority), about 3500 and 6 mg/day, respectively [21]. In addition to macronutrients, pumpkin is also a rich source of essential micronutrients such as vitamins, antioxidant compounds and other phytochemicals. Several studies have shown that pumpkins are a notable source of vitamins A, C and E, which play crucial roles in supporting immune function and acting as potent antioxidants to combat oxidative stress, in addition to the presence of alkaloids, flavonoids, polyphenols, tannins, phytosterols and cucurbitacins [18,19,22,23]. Kaur et al. [19] also highlighted the great amounts of pro-vitamin A carotenoids in pumpkin and their importance in developing important antioxidant effects, reducing the risk of certain diseases including cancer. These compounds are known as antioxidant and antimicrobial agents for their capacity to neutralize harmful free radicals in the body, preventing oxidative damage in cells and tissues, and inhibiting the growth or killing of microorganisms, such as bacteria or fungi, combating infections or preventing their spread, respectively. Contents of 171.9 μg/g of carotenoids, 2–10 mg/100 g of vitamin C, and 9–10 mg/100 g of vitamin E are reported in pumpkin flesh, which also contributes to a healthy diet (average requirement of 9 g/day of carotenoids with provitamin A activity, and 110 mg/day of vitamin C, of which adequate intake is 13 mg/day) [21].
Beyond its nutrients, pumpkins possess functional properties that are of particular interest to the food industry. Sharma et al. [24] recently reviewed the health-promoting potential of pumpkin in terms of antioxidant, anticancer, anti-inflammatory, anti-obesity, anti-diabetes, antimicrobial and other bioactivities, as well as its application in the food industry, as a cooked, powdered or pureed ingredient in different food products. Their attractive taste and vibrant color make them suitable for the formulation of health-promoting food products, including soups and snacks [25,26,27].
Pumpkin cultivation is widely practiced across the globe, being generally a warm-season crop that thrives in well-drained, nutrient-rich soils with good sunlight exposure. Its cultivation and appreciation may vary according to the wide variety of pumpkin species and cross-pollinations between cultivated species and/or wide relatives, which results in high genetic diversity and inherent polymorphism [28,29,30]. According to data from FAO (Food and Agriculture Organization of the United Nations), in 2021 the pumpkin production (the aggregate of pumpkins, squash and gourds) was approximately 24 million tonnes harvested from 1.5 million hectares, while the world’s leading producers were China (approximately 7.4 million tonnes), followed by Ukraine (approximately 1.43 million tonnes), Russian Federation (approximately 1.2 million tonnes) and USA (approximately 1.1 million tonnes) [31]. Within the European Union, pumpkin is cultivated on 84,270 hectares with a total production of 2.7 million tonnes, while the main producers are Spain (789,780 tonnes), Italy (601,660 tonnes), France (410,360 tonnes) and Poland (316,300 tonnes) [31]. In Greece, although pumpkin does not occupy many hectares of agricultural land (2170 hectares with annual production of 61,120 tonnes [31]), there is wide genetic diversity with several local landraces being cultivated all through the country. The annual world production over the last ten years fluctuated between 21.9 and 26.6 million tonnes, whereas the harvested area showed decreasing trends dropping from almost 2.0 million hectares in 2015 to 1.5 million hectares in 2021 [31]. On the other hand, in Greece, there has been a great increase in harvested area and annual production over the last ten years, since harvests have increased from 536 ha to 2170 ha and annual production from 673 tonnes to 61120 tonnes [31]. Fruit yield may also vary with values that range between 37 to 98 tonnes/ha [32], 25.2 to 55.7 tonnes/ha [33] or 7.3 to 41.8 tonnes/ha [34], depending on fertilization rate and irrigation level.
However, despite the rising pumpkin production during the last few years, the cultivation of this valuable crop faces mounting challenges due to the intensifying impacts of abiotic stress factors. The climate changes observed in the world have changed patterns of temperature and rainfall, thus increasing the occurrence of extreme weather events, affecting the yield of several crop species, including pumpkin [35,36]. Moreover, the intense exploitation of the soil without proper management also has negative effects through land degradation, drought and salinization of irrigation water and agricultural land [37]. These effects have challenged the academic community in search of alternatives that aim to provide high-yield and quality crops, through the understanding of the effects of these adversities on traditional crops as well in the search and promotion of new, more promising crops [37,38,39].
Considering the high fruit polymorphism, more and more studies of the characterization in terms of composition and functional properties of different pumpkin genotypes are being disseminated [40,41,42]. However, the genetic pool of Greek pumpkin germplasm is not adequately characterized so far, especially when considering the high genetic variability of the species which could be a valuable tool for the adaptation of modern horticulture in the changing climate conditions. Therefore, the present study aimed to provide useful information regarding the biochemical and nutritional diversity in pumpkin genotypes cultivated in Greece. For this purpose, eleven local landraces and commercially available genotypes were cultivated under the same conditions and nutritional and chemical profiles, and the functional properties of fruit flesh were investigated.

2. Materials and Methods

2.1. Plant Material and Growing Conditions

The experiment was conducted at the experimental farm of the University of Thessaly, Greece during the growing period of 2021 (e.g., May–November, 2021). Seeds from eleven genotypes (V1–V11) were directly sown in the soil on 19 May 2021. The description of genotypes is presented in Table 1. The genotypes included 9 local landraces collected from the respective regions, while two commercial genotypes were also included (e.g., F1 Fytro FS-243, Fytro seeds S.A., Athens, Greece and Big Max, Geniki Fytotechniki S.A., Athens, Greece). Seeds were sown in rows at distances of 1.6 m within each row and 2.0 m between the rows. Each row consisted of 32 plants, with 3 rows for each genotype being used (96 plants for each genotype in total), while rows were laid out according to the completely randomized design (n = 3). Before sowing, a complex fertilizer (18-9-18, N-P-K; Complex Haifa Turbo K + Mg + S + Fe and Zn; Haifa Group, Israel) was applied as a base dressing at a rate of 250 kg/ha and was evenly distributed with a rototiller. Irrigation was applied via a drip irrigation system (one emitter per plant with a water supply of 6 L/h) at regular intervals based on the environmental conditions during the experimental period. Irrigation was scheduled based on the recordings of soil moisture content taken at regular intervals aiming to retain 100% of field capacity. Soil moisture content was recorded with PR2 Profile Probe (Delta T PR2/4 +HH2; Delta-T devices Ltd., Burwell, UK) using access tubes 40 cm long, while measurements were taken at soil depths of 10, 20, 30 and 40 cm. Access tubes were established with Del-ta-T augering and extraction kit (PR-ASK1-S, Delta-T Devices Ltd., Cambridge, UK), using one access tube per genotype (15 tubes in total).
After crop establishment, fertilization was applied via the drip irrigation system (fertigation). In particular, on 16 July 2023 plants received ammonium nitrate (34.5% nitrogen; 40 kg/ha), Solusop (0-0-52, N-P-K; Agri.fe.m Ltd., Athens, Greece; 20 kg/ha) and Disper bloom (Disper Bloom; Agrofarm S.A., Athens, Greece; 1.5 kg/ha). On 26 July 2023, plants were foliar sprayed with Disper Bloom (100 g/100 L of water) and Root and Leaf (20-202-20, N-P-K +TE; 200 g/100 L of water). On 28 July 2023, plants were fertigated with 40 kg of ammonium nitrate and 20 kg of Solusop. On 12 August 2023, plants were treated with 25 kg of ammonium nitrate, 25 kg/ha of Solusop and 75 kg/ha of Mannitol 3 GR (Agrofarm S.A., Greece). Weeds were controlled chemically with pre-emergence herbicides, as well as manually with a hand hoe during the growing period and until crop establishment. Pests and pathogens were chemically controlled based on the recommended practices of the crop.
Harvest took place when fruit reached maturity from 25 August 2021 to 22 November 2021, depending on the genotype. Due to polymorphism that some genotypes showed (e.g., V2, V4, V5, V7 and V9), two types of fruit that differed in their form were harvested, as described in Table 1. After harvest, fruit yield was calculated as the sum of the weight of all the fruit from each cultivar extrapolated to the harvested area of one hectare.

2.2. Samples Preparation

After collection, fruits were weighted and 15 randomly selected fruits from each genotype and fruit form were cut in half; after removing seeds and pericarp flesh samples were collected, put in an air-sealed bag and stored under freezing conditions (−19 °C). Then, the frozen samples were lyophilized, ground to powder and stored at deep-freezing conditions until further analysis. To obtain the flesh samples, the fruits were peeled and we removed the seeds and fibrous strands. The fleshes were then lyophilized (Sublimator model EKS, manufactured by Christian Zirbus Co. in Osterode am Harz, Germany), crushed (~20 mesh) and stored until subsequent analysis. The applied lyophilization program for drying the plant biomass was the following:
  • Step 1: −35 °C for 2 h at atmospheric pressure (1000 mbar);
  • Step 2: from −35 °C to −20 °C in 6 h under vacuum 0.150 mbar;
  • Step 3: from −20 °C to 0 °C in 12 h under vacuum 0.150 mbar;
  • Step 4: from 0 °C to 10 °C in 12 h under vacuum 0150 mbar;
  • Step 5: from 10 °C to 25 °C in 12 h under vacuum 0.150 mbar.
For the bioactivities, the hydroethanolic (80 ethanol: 20 water) flesh extracts were obtained by maceration for 1 h, twice, with a solid/liquid ratio of 1 g/30 mL, following the standard methodology of the group, previously described in Leichtweis et al. [42]. The extracts were then lyophilized as described above and stored until subsequent analysis.

2.3. Nutritional Characterization

The AOAC procedures [43] were followed to obtain the nutritional composition of flesh samples, including the total fat, crude protein, carbohydrates, dietary fibre content, ash and total energy. The fat content was extracted with petroleum ether in a Soxhlet apparatus; for the crude protein, we used the macro-Kjeldahl method (equipment Pro-Nitro A, Selecta, Barcelona, Spain) and a nitrogen-to-protein conversion factor of 6.25; the total dietary fibre content was determined using a combination of enzymatic and gravimetric method; and, the ash content was assessed by incinerating the samples at 550 ± 10 °C.
Total carbohydrates were calculated by difference, and the total energy was calculated using the following equation:
energy (kcal) = 4 × (g protein + g carbohydrates) + 9 × (g fat) + 2 × (g total dietary fibre)
The results were expressed in g/100 dry weight (dw).

2.4. HPLC Assays

Tocopherol content was determined by the HPLC system described above coupled to a fluorescence detector (FP-2020; Jasco, Easton, MD, USA), following the methodology described by Pereira et al. [44]. The detector was configured to excite at 290 nm and measure emission at 330 nm and the chromatographic separation was achieved with a Polyamide II normal-phase column (250 × 4.6 mm; YMC Waters) operating at 30 °C. Tocol was used as an internal standard (IS) and the compound isoforms (α-, β-, γ-, and δ) were identified by comparisons with authentic standards. The quantities were obtained using the following calibrations curves: α-tocopherol, y = 1.295x, R2 = 0.991, LOD = 18.06 ng/mL, LOQ = 60.20 ng/mL); β-tocopherol, y = 0.396x, R2 = 0.992, LOD = 25.82 ng/mL, LOQ = 86.07 ng/mL); and γ-tocopherol, y = 0.567x, R2 = 0.991, LOD = 14.79 ng/mL, LOQ = 49.32 ng/mL. The results were expressed in mg/100 g dw.
The organic acid characterization was performed by ultra-fast liquid chromatography (UFLC, Shimadzu Corporation, Kyoto, Japan) coupled to a photodiode array detector (PDA), using 215 and 245 nm (for ascorbic acid) as preferred wavelengths [45]. Separation was achieved on a SphereClone (Phenomenex, Torrance, CA, USA) reverse phase C18 column (5 μm, 250 mm × 4.6 mm i.d.) thermostatted at 35 °C. Identification and quantification were achieved by comparison with commercial standards. The calibration curves were as follows: oxalic acid, y = 8 × 106x + 331,789; R2 = 0.9912; malic acid, y = 942,562x + 38,506, R2 = 0.9987; ascorbic acid, y = 5 × 107x + 449,262; R2 = 0.9813; shikimic acid, y = 8 × 107x + 567,119; R2 = 0.9903; citric acid, y = 968,367x − 12,295; R² = 0.9974; and fumaric acid, y = 9 × 107x − 100,894, R2 = 0.9986. The results were expressed in g/100 g dw, except for fumaric acid (mg/100g dw).
The analysis of free sugars was carried out using HPLC coupled to a refractive index detector, more specifically an integrated system with a pump (Knauer, Smartline system 1000, Berlin, Germany), degasser system (Smartline manager 5000), auto-sampler (AS-2057 Jasco, Easton, MD, USA) and an RI detector (Knauer Smartline 2300), following a previously described method [46]. A Eurospher 100-5 NH2 column (4.6 × 250 mm, 5 mm, Knauer) operating at 30 °C (7971 R Grace oven) was used. For this analysis, melezitose was used as IS for the quantification method and the compounds were identified by chromatographic comparisons with authentic standards D(-)-fructose, D(+)-sucrose, D(+)-glucose, D(+)-trehalose and D(+)-raffinose pentahydrate. The results were expressed in g/100 g dw. The content of individual free sugars was used to calculate the sweetness index (SI) of fruit through the formula [45]:
S I = f r u c t o s e   c o n t e n t × 157.5 + g l u c o s e   c o n t e n t × 67.5 + s u c r o s e   c o n t e n t × 100
Fatty acids were determined by gas–liquid chromatography with flame ionization detection (GC-FID), a DANI model GC 1000 instrument and a Macherey–Nagel column (30 m × 0.32 mm ID × 0.25 μm df), following the methodology published by Pereira et al. [44]. The identification and quantification were performed by comparing the retention times of peaks detected to the ones of FAME peaks of commercial standards (FAME reference standard mixture, standard 47885-U, Sigma-Aldrich, St. Louis, MO, USA). The contents were presented in relative percentages.

2.5. Bioactivities

The antioxidant capacity of the flesh extracts was evaluated using two in vitro cell-based assays. The first screening was obtained using porcine (Sus scrofa) brain homogenates, to observe the ability of the extract to inhibit lipid peroxidation by the thiobarbituric acid reactive substances inhibition (TBARS) assay, as described by Pereira et al. [47], and the second one using a red cell suspension of healthy sheep blood, investigating the extracts capacity to inhibit the oxidative hemolysis (OxHLIA), as described by Backes et al. [48]. The results were expressed in IC50 values (μg/mL), meaning the extract concentration that inhibits 50% of oxidation and that delays 50% of erythrocyte hemolysis for 60 min, respectively. Trolox was used as a positive control in both assays.
For the antimicrobial activity, the extracts were tested against eight bacteria and two fungi related to food contamination, more specifically, Bacillus cereus (ATCC 11778), Staphylococcus aureus (ATCC 25923), Listeria monocytogenes (ATCC 19111), Escherichia coli (ATCC 25922), Enterobacter cloacae (ATCC 49741), Pseudomonas aeruginosa (ATCC 9027), Salmonella enterica subsp (ATCC 13076), Yersinia enterocolitica (ATCC 8610), Aspergillus fumigatus (ATCC 204305) and Aspergillus brasiliensis (ATCC 16404). The microdilution and p-iodonitrotetrazolium chloride (INT) methods were used [49]. Streptomycin and ampicillin were used as positive controls for bacteria and ketoconazole and bifonazole for fungi. The results were expressed in mg/mL.
Additionally, the cytotoxic effects were assessed in a primary culture of non-tumor porcine liver cells (PLP2) derived from porcine liver obtained from a local slaughterhouse. The evaluation was conducted using the Sulforhodamine B (SRB) colorimetric assay [50]. The IC50 values (inhibiting 50% of the net cell growth) for extract concentration, presented in μg/mL, were compared to the positive control, Ellipticine.

2.6. Statistical Analysis

For chemical analysis, fruit samples were analysed in triplicate, and the results were expressed as mean ± standard deviation. For comparing two groups of data, a Student’s t-test was used, while for three or more groups, the one-way analysis of variance (ANOVA) was applied. Prior to analysis, normal distribution and homogeneity of variance were assessed using the Shapiro–Wilk and Levene tests, respectively. For homoscedastic data with p > 0.05, Tukey’s honestly significant difference (HSD) test was employed and, for heteroscedastic data, Tamhane’s T2 multiple comparison test was used. All analyses were conducted at a 5% significance level using IBM SPSS Statistics software (Version 22.0, IBM Corp, Armonk, NY, USA).
In addition, principal component analysis (PCA) was performed to identify the contribution of each variable to the total genetic diversity and classify the studied pumpkin landraces based on their chemical composition and antioxidant activities of fruit flesh. All statistical analyses were carried out using the StatGraphics Centurion-XVII statistical package (StatPoint Technologies Inc., Warrenton, VA, USA).

3. Results and Discussion

3.1. Crop Performance

The results of crop performance (e.g., total fresh yield of fruit, number of fruit per plant and mean fruit weight) are presented in Table 2, where significant differences among the studied genotypes were recorded. In particular, the highest fruit yield was recorded for the V8 genotype (34,069 kg/ha), followed by V2 and V4 genotypes (30,695 kg/ha and 27,997 kg/ha, respectively), whereas the lowest overall yield was recorded for V5 genotype with a production of only 7618 kg/ha. Moreover, significant differences were recorded in the number of fruit per plant where V1 recorded the highest mean value (6.1 fruit per plant) with an average fruit weight of 1.0 kg per fruit. On the other hand, the lowest number of fruits per plant was recorded in V5 with an average weight of 3.6 kg. Significant differences were also recorded in mean fruit weight where V3 recorded the highest mean value (8.1 kg) followed by V2, V8 and V11 (6.3, 6.1 and 5.6 kg, respectively). These results indicate that the high number of fruit is accompanied by smaller fruit weight, as in the case of V2, V3, V8 and V11. The high variability recorded in the tested genotypes is mainly attributed to the differences in their genetic background, especially regarding the local landraces which are acclimatized in specific edaphoclimatic conditions through the cultivation in specific regions with particular microclimates. According to the literature, genotypic diversity in Cucurbita species is expected to result in differences in crop performance [51] and quality features [52,53], while differences observed in multi-site experiments can justify the acclimatization of pumpkin genotypes to specific environmental conditions and agronomic practices [54,55]. Moreover, the differences in plant morphology and phenology (e.g., vine length, fruit maturation, number of branches, etc.) may influence crop performance [56], especially when plants are grown under the same conditions and agronomic practices as those in our study.

3.2. Nutritional Profile

The proximate composition of the studied genotypes is presented in Table 3, including two samples of the same genotype that varied in fruit morphology as indicated in Table 1 (e.g., V2, V4, V5, V7 and V9). A variable content was recorded with significant differences not only among the studied genotypes but also between the different fruit types of the same genotype. In particular, fat content ranged between 0.38 and 1.17 g/100 g dw with the highest content being recorded in V2 T, V6 and V11, whereas the lowest content was found in the flesh of V4 R genotype. Protein and ash content ranged between 8.0–21.4 g/100 g dw and 3.5–10.95 g/100 g dw, respectively, with V9 C genotype having the highest content (21.4 g/100 g and 10.95 g/100 g dw, respectively), whereas the lowest overall content of protein and ash was recorded in V4 C and V5 R genotypes (8.6 g/100 g dw and 3.5 g/100 g dw), respectively. V1 showed the highest energy content (360.1 Kcal/100 g dw), V2 had the highest content in carbohydrates (72.2 g/100 g dw), while V9 C was the richest in fibres (27.4 g/100 g dw). The detected amounts of macronutrients were within the same range of literature reports regarding the flesh of Cucurbita species [40,57,58,59], although protein and fat content was higher and lower than other studies, respectively [60]. The high variability in macronutrients presented in our study indicates the genetic inter- and intra-population diversity of the studied germplasm, which could be due to breeding through the farmer’s seed system which facilitates heterogeneity, especially in the case of local landraces as in our study and can justify the recorded differences with other reports in the literature [61,62].
The main free sugars detected in the flesh of the studied genotypes were glucose, sucrose and fructose, followed by trehalose and raffinose (Table 4), while typical chromatographs of samples rich in fructose and sucrose are presented in Figure 1. The highest amounts of the main sugars were recorded in V11 (26.9 g/100 g dw) and in both fruit types of V2 (9.7 and 10.0 g/100 g dw for T and C fruit types, respectively) and V7 (14.0 g/100 g dw for both fruit types), while trehalose was the highest in V2 C (0.67 g/100 g dw) and raffinose in V2 T, V6 and V8 genotypes (and 0.29, 0.27 and 0.25 g/100 g dw, respectively). The highest total free sugar content was recorded for the local landrace V11 due to the increased content of glucose. Moreover, the same genotype (V11) presented the highest sweetness index (3367) due to the overall composition of individual and total free sugars which indicates a sweeter taste and is highly associated with better acceptability from consumers [63]. Similarly to our study, Seroczyńska et al. [64] evaluated twelve forms of C. maxima fruit and identified sucrose, fructose and glucose in variable amounts depending on the genotype while they detected mannose in specific genotypes. The same sugars were detected by Dhenge et al. [65] in C. moschata fruit although a high fraction of total free sugars was not tentatively identified. Moreover, Kostecka-Gugała et al. [66] highlighted the intra-species variability between different cultivars of C. pepo, C. maxima, C. moschata and C. ficifolia in terms of soluble sugar content which explains the differences recorded in our study, while similar results were reported for indigenous and hybrid varieties of (Cucurbita maxima Linn) by Amin et al. [59].
The fatty acids composition of the flesh of the studied genotypes is presented in Table 5. The main fatty acids were palmitic (19.6–63.4%), oleic (1.98–14.1%), linoleic (5.1–35.8%) and γ-linolenic acids (0.5–38.8%). So far, most of the reports in the literature focused on the fatty acids composition of seeds and seed oils where palmitic, oleic and linoleic acids are highlighted as the main compounds [59,60], while the edible flowers contain mostly myristic, oleic, stearic and heneicosanoic acid [67] and the leaves palmitic, linoleic and linolenic acids [68]. To the best of our knowledge, there is a lack of scientific literature regarding the composition of fatty acids in the flesh of pumpkin fruit. The majority of fatty acids were classified as saturated and polyunsaturated fatty acids which accounted for approximately 85 to 95% of total fatty acids, while the rest of the identified compounds were monounsaturated fatty acids SFA and PUFA, respectively. The only exceptions were genotypes V2 C and V4 C where SFA was the predominant class of fatty acids accounting for approximately 85% due to the high content of palmitic acid, whereas V6 and V11 genotypes had the highest content of PUFA (approximately 65%). Moreover, the PUFA/SFA and n-6/n-3 values were higher than 0.45 and lower than 4.0, respectively, for all the tested genotypes indicating high nutritional value of pumpkin pulp [69], except for genotypes V2 C and V4 C where high amounts of PUFA and very low amounts of n-3 fatty acids were recorded.

3.3. Bioactivities

3.3.1. Antioxidant Capacity

Varied lipid peroxidation and oxidative haemolysis inhibition capacity were recorded among the tested genotypes (Table 6). In particular, V4 C and V7 P genotypes recorded the highest antioxidant activity for the TBARS assay (lowest IC50 values), while the V8 genotype was the most effective in the case of the OxHLIA assay. In any case, none of the tested extracts was more effective than Trolox which was used as the positive control for both assays. Recently, Leichtweis et al. [42,70] determined the antioxidant activity in different parts of Cucurbita fruit cultivated in Portugal, Algeria and Greece suggested a variable response to TBARS and OxHLIA assays depending on the fruit type and the growing location, while seeds recorded the highest antioxidant activity compared to peels and endocarp. On the other hand, Jahan et al. [71] suggested that flowers showed the highest antioxidant activity, followed by flesh, leaves, seeds and peels, which indicates the effect of the implemented assay on the obtained results. Moreover, the addition of powder from pumpkin fruit and flowers improved the capacity of lipid peroxidation inhibition in cooked sausages and chicken [72,73]. In the studies of Rolnik et al. [71,74], where the inhibition of lipid peroxidation was determined via the TBARS assay, it was reported a significant variability among cucurbit vegetables, including pumpkin, zucchini and squash which highlights the importance of the genotypic impact on the antioxidant capacity of Cucurbita fruit. In contrast, Tarwadi and Angte [75] who compared fruit and root vegetables classified pumpkin fruit among the species with poor performance in terms of the TBARS assay.

3.3.2. Antimicrobial Activity

The antimicrobial properties of the extracts obtained from the fruit of the tested genotypes are presented in Table 7. In general, despite some slight differences among the tested extracts in terms of MIC. MBC and MFC values, none of them showed higher activity against various Gram− and Gram+ bacterial strains and two species of Aspergillus compared to streptomycin, ampicillin and ketoconazole. However, almost all the tested extracts showed the ability to inhibit the growth of at least one bacterial strain (MIC values up to 10 mg/mL), except for the case of V9 R where no effects were recorded. Moreover, the lowest overall MIC values were recorded for V2 T against Salmonella enterocolitica (5 mg/mL); V6 against and Salmonella enterocolitica (5 mg/mL for both bacteria); V7 F against Yersinia enterocolitica (2.5 mg/mL); V8 against Escherichia coli (5 mg/mL); V9 C against Enterobacter cloacae (5 mg/mL); and V10 against Enterobacter cloacae and Yersinia enterocolitica (5 mg/mL). Similarly, most of the tested extracts showed efficacy against at least one of the studied Aspergillus species, except for V1, V2, V4 C and V5 genotypes. Similar to our study, Leichtweis et al. [42,70] suggested significant antimicrobial properties from fruit parts (peels, endocarp and seeds) of different pumpkin genotypes. Moreover, Mokhtar et al. [76] reported varied antimicrobial properties of C. moschata fruit depending on their ripeness with extracts from mature fruit being more effective against various bacterial strains compared to young and ripe fruit. According to the same authors, this finding was mainly attributed to the highest content of polyphenols detected in this particular stage [76]. Badr et al. [57] also recorded moderate efficacy of rind and flesh extracts against Bacillus subtilis and B cereus. Hussain et al. [77] suggested seed extracts showed higher activity against Candida albicans, Fusarium oxysporum, Mucor miehei and Trichoderma spp. compared to peels and flesh, whereas flesh extracts recorded greater activity against Salmonella typhi, Escherichia coli, Bacillus subtilis and Streptococcus aureus compared to the other two extracts. The beneficial effects of different pumpkin powders, extracts, isolates, purified bioactives and pumpkin-based functional food products are well reported in the review performed by Hussain et al. [78], which corroborates its health promotion properties. In contrast, Saavedra et al. [7] did not record any antimicrobial effects for peel and seed extracts obtained from C. pepo byproducts (shells and seeds).
The reported antimicrobial activity of the pumpkin pulp extracts assessed in the present study, together with the antioxidant activity described above, supports the importance of including this nutritious and bioactive fruit in healthy diets.
None of the tested samples presented cytotoxic activity against the non-tumor porcine liver cells (up to 400 μg/mL), which indicates the high potential of using pumpkin flesh as a natural antioxidant agent in the food industry. This finding is in agreement with the previous reports of Leichtweis et al. [42,70], who did not record any cytotoxic effect on non-tumor cells for the extracts from different parts of pumpkin fruit. Moreover, according to Gaweł-Bęben et al. [79], peel extracts did not show any toxic effects against human keratinocytes up to the concentration of 1000 μg/mL suggesting its safe use in the cosmeceutical industry. Similarly, the extracts of zucchini fruit showed a high potential for use in cosmetic emulsions since they did not show toxic effects against HaCaT cell lines or reconstructed human epidermis [80]. This satisfactory result was expected since pumpkin pulp has been used for thousands of years for animal and human consumption. However, considering mainly local varieties, which are not yet widely cultivated and consumed, these results add new safety insights, since varieties of the same species can considerably vary in terms of chemical composition. In fact, the presence of bioactive compounds with cytotoxic activity against tumour cell lines is widely reported in pumpkins, such as, for example, cucurbitin [13,81], and in the present study we aimed to confirm their safety, testing the obtained extracts against non-tumour cells.

3.4. Chemical Characterization

3.4.1. Organic Acids

The main detected organic acids were oxalic and malic acid, followed by citric and fumaric acid, whereas ascorbic and shikimic acid were either detected in traces or not detected (Table 8). The V9 C genotype recorded the highest content of oxalic and citric acids (6.4 and 3.2 g/100 g dw, respectively), while malic acid was the highest in V4 R and V10 genotypes (5.38 and 5.2 g/100 g dw, respectively) and fumaric acid in V6 genotype (0.0943 mg/100 g dw). According to Priecina and Karklina [82] oxalic acid was the major detected compound in fresh pumpkin fruit followed by malic and citric acid, while the amounts of oxalic acid were considerably higher than those of our study (approximately 15 g/100 g dw). Moreover, Iswaldi et al. [83] tentatively identified eight organic acids and derivatives in three zucchini cultivars, although the authors did not quantify the detected compounds. Abbas et al. [84] suggested that organic acid composition and content in pumpkins may be affected by the growing conditions, the genotype and the developmental stage of fruits. In particular, they indicated a significant decrease in organic acids content throughout the maturation process, while they suggested oxalic and fumaric acids as the prevalent ones at all the developmental stages [85]. In contrast, Nawirska-Olszańska et al. [85] and Zhou et al. [86] reported the presence of only malic, citric and fumaric acid with varied content among the different genotypes of pumpkin species (C. pepo, C. maxima and C. moschata). Therefore, any contradictions among reports from the literature could be associated with differences in the genotype, the growing conditions and the harvesting stage of fruit.

3.4.2. Tocopherols

The main detected vitamin E isoform was α-tocopherol which recorded in amounts that ranged between 0.218 (V7 F) and 4.90 mg/100 g dw (V2 T), followed by β- and γ-tocopherols (Table 9). So far, there is scarce literature regarding the tocopherol composition in the flesh of pumpkin fruit since most of the studies refer to seeds and seed oils. Similarly to our study, Kim et al. [40] suggested α-tocopherol as the main compound in the flesh of three pumpkin species (C. pepo, C. moschata and C. maxima), while γ-tocopherol was detected only in low amounts in C. moschata fruit. Kulczynski and Gramza-Michałowska [9] detected only α- and γ-tocopherols in the pulp of C. pepo and C. moschata fruit in amounts that were towards the highest range of our study, whereas the other two vitamin E isoforms (β- and δ-tocopherols) were not detected. Therefore, the results of our study along with the literature reports suggest that pumpkin fruit flesh can be a rich source of tocopherols (especially α-tocopherol), although a high genotypic variability should be expected at the intra- or inter-species level.

3.5. Principal Component Analysis

The principal component analysis (PCA) was implemented to reveal groups and highlight similarities and differences based on multivariate data. The processing of our data showed that the first eleven principal components (PCs) were associated with Eigen values higher than 1, explaining 95.9% of the cumulative variance, with PC1 accounting for 24.6%, PC2 for 18.4% and PC3 for 13.3%; and PC4 for 8.8%, PC% for 7.9%, PC6 for 6.3%, PC7 for 4.7%, PC8 for 3.9%, PC9 for 3.6%, PC10 for 2.4% and PC11 for 2.1%. In particular, PC1 showed a positive correlation with C18:2n6c and PUFA and a negative correlation with C8:0, C10:0, C15:0, C16:0, C17:0, C20:0, C22:0, C23:0, C24:0 and SFA. On the other hand, PC2 showed a positive correlation with ash, fibres, fructose, oxalic acid and raffinose content, as well as with TBARS, C22:1 and C24:1, whereas a negative correlation with carbohydrates and energy content was recorded. Finally, PC3 showed a positive correlation with fat, fumaric acid, glucose, total sugars and total tocopherols content, whereas a negative correlation was observed for C20:3n6, C22:1, C24:1 and citric acid. Therefore, PCA may help to discriminate the studied landraces as depicted in the following scatterplots and loading plots. The scatterplot in Figure 2 shows four distinct groups of the studied landraces based on their chemical composition and bioactive properties of fruit flesh.
The loading plot of PC1 and PC2 presents the following correlations: the upper left quadrant included fibres, ash, oxalic acid, fructose, glucose, total sugars, total organic acids, malic acid, TBARS, C8:0, C15:0, C16:1, C17:1, C20:0 and C23:0; the lower left quadrant included SFA, C10:0, C12:0, C14:0, C16:0, C17:0, C18:0, C22:0, C22:6n3, C24:0, trehalose, OxHLIA and carbohydrates; the upper right quadrant included C13:0, C18:2n6, C18:3n6, C20:1, C20:3n6, C21:0, C22:1, C24:1, α-tocopherol, β-tocopherol, γ-tocopherol, total tocopherols, raffinose, fumaric acid, protein, fat and PUFA; the lower right quadrant included C18:1n9c, C18:3n3, C20:5n3, C22:2, citric acid, MUFA, sucrose and energy content (Figure 3).
Moreover, the loading plot of PC1 and PC3 also revealed groups of positively correlated variables (Figure 4). The upper left quadrant included glucose, fructose, trehalose, total sugars, malic acid, carbohydrates, fibres, TBARS, OxHLIA, C16:1, C17:0, C17:1, C22:0 and C22:6n3; the lower left quadrant included C8:0, C10:0, C12:0, C14:0, C15:0, C16:0, C18:0, C20:0, C23:0, C24:0, SFA, oxalic acid, total organic acids and ash; the upper right quadrant included fumaric acid, fat, α-tocopherol, β-tocopherol, γ-tocopherol, total tocopherols, fat, raffinose, MUFA, energy content, C13:0, C18:1n9c and C18:3n3; the lower right quadrant included C18:3n6, C18:2n6c, C20:3n6, C20:1, C20:5n3, C21:0, C22:1, C22:2, C24:1, PUFA, sucrose, protein and citric acid.

4. Conclusions

The genetic pool of Greek pumpkin germplasm is not adequately characterized so far, especially when considering the high genetic variability of the species which could be a valuable tool for the adaptation of modern horticulture in the changing climate conditions. The heterogeneity in terms of agronomic performance and chemical profile of fruit in pumpkin germplasm makes it necessary to evaluate local landraces aiming to identify promising genotypes with high yield and quality of fruit. Our results indicate that the studied local landraces and commercial genotypes had a high variability not only in fruit yield but also in all the tested parameters related to the chemical composition and bioactive properties of fruit flesh. The local landraces V2 and V8 (from “Trikala” and “Laconia”, respectively) recorded the highest fruit yield, while the V2 landrace also had the highest tocopherols (α-, β- and total tocopherols) and fructose, trehalose and raffinose content. On the other hand, V8 landrace was the richest in linoleic acid and also presented the highest antioxidant activity for the OxHLIA assay. However, the rest of the tested landraces presented interesting features such as V11 (“Voutirato”) which had the highest sweetness index. Finally, the flesh extracts of most of the tested genotypes showed promising antimicrobial properties, while none of them was toxic against non-tumor cells. Nevertheless, our results support the importance of the conservation and valorization of local genetic material for the selection of elite genotypes with improved nutritional value and quality. However, further research is needed in order to reveal the genetic heterogeneity of the studied material that would help to introduce local landraces in breeding programs.

Author Contributions

Conceptualization, M.G.L., O.P., A.K.M., S.A.P., L.B. and C.P.; data curation, M.G.L., O.P., S.A.P. and C.P.; funding acquisition, S.A.P. and L.B.; investigation, M.G.L., O.P., A.K.M., T.C.S.P.P. and M.I.D.; methodology, M.G.L., O.P., A.K.M., T.C.S.P.P., M.I.D., R.C.C. and C.P.; resources, S.A.P. and L.B.; supervision, M.B.P.P.O., L.B., S.A.P. and C.P.; validation, M.B.P.P.O., S.A.P., L.B. and C.P.; writing—original draft, M.G.L., O.P. and S.A.P.; writing—review and editing, S.A.P., C.P. and L.B. All authors have read and agreed to the published version of the manuscript.


This research was funded by the Foundation for Science and Technology (FCT), P.I. (FCT, Portugal) through national funds FCT/MCTES to CIMO (UIDB/00690/2020 and UIDP/00690/2020), SusTEC (LA/P/0007/2021) and UIDB/50006/2020. National funding by FCT, P.I., through the institutional scientific employment program contract for M.I.D., R.C.C., C.P. and L.B. contracts, and M.G. and A.K.M. PhD grants (2020.06706.BD and 2020.06231.BD, respectively); FCT, P.I., within the scope of the Project PRIMA Section 2—Multi-topic 2019: PulpIng (PRIMA/0007/2019); the MICINN for the Juan de la Cierva Formación contract for T.C.S.P.P. (FJC20120-045405-I); and the General Secretariat for Research and Technology of the Ministry of Development and Investments under the PRIMA Program. PRIMA is an Art.185 initiative supported and co-funded under Horizon 2020, the European Union’s Program for Research and Plants 2022, 11, 800 16 of 18 Innovation (Prima2019-08).

Data Availability Statement

Data are contained within the article.

Conflicts of Interest

The authors declare no conflict of interest.


  1. Chomicki, G.; Schaefer, H.; Renner, S.S. Origin and domestication of Cucurbitaceae crops: Insights from phylogenies, genomics and archaeology. New Phytol. 2020, 226, 1240–1255. [Google Scholar] [CrossRef] [PubMed]
  2. Kistler, L.; Newsom, L.A.; Ryan, T.M.; Clarke, A.C.; Smith, B.D.; Perry, G.H. Gourds and squashes (Cucurbita spp.) adapted to megafaunal extinction and ecological anachronism through domestication. Proc. Natl. Acad. Sci. USA 2015, 112, 15107–15112. [Google Scholar] [CrossRef] [PubMed]
  3. Smith, B.D. The initial domestication of Cucurbita pepo in the Americas 10,000 years ago. Science 1997, 276, 932–934. [Google Scholar] [CrossRef]
  4. Kates, H.R.; Soltis, P.S.; Soltis, D.E. Evolutionary and domestication history of Cucurbita (pumpkin and squash) species inferred from 44 nuclear loci. Mol. Phylogenet. Evol. 2017, 111, 98–109. [Google Scholar] [CrossRef]
  5. Savage, J.A.; Haines, D.F.; Holbrook, N.M. The making of giant pumpkins: How selective breeding changed the phloem of Cucurbita maxima from source to sink. Plant Cell Environ. 2015, 38, 1543–1554. [Google Scholar] [CrossRef]
  6. Integrated Taxonomic Information System—Report. Cucurbita, L. Available online: (accessed on 3 September 2023).
  7. Saavedra, M.J.; Aires, A.; Dias, C.; Almeida, J.A.; De Vasconcelos, M.C.B.M.; Santos, P.; Rosa, E.A. Evaluation of the potential of squash pumpkin by-products (seeds and shell) as sources of antioxidant and bioactive compounds. J. Food Sci. Technol. 2015, 52, 1008–1015. [Google Scholar] [CrossRef] [PubMed]
  8. Sun, H.; Wu, S.; Zhang, G.; Jiao, C.; Guo, S.; Ren, Y.; Zhang, J.; Zhang, H.; Gong, G.; Jia, Z.; et al. Karyotype Stability and Unbiased Fractionation in the Paleo-Allotetraploid Cucurbita Genomes. Mol. Plant 2017, 10, 1293–1306. [Google Scholar] [CrossRef] [PubMed]
  9. Kulczynski, B.; Gramza-Michałowska, A. The profile of secondary metabolites and other bioactive compounds in Cucurbita pepo L. and Cucurbita moschata pumpkin cultivars. Molecules 2019, 24, 2945. [Google Scholar] [CrossRef] [PubMed]
  10. Petropoulos, S.; Ferreira, I.; Barros, L. Phytochemicals in Vegetables: A Valuable Source of Bioactive Compounds; Petropoulos, S.A., Ferreira, I.C.F.R., Barros, L., Eds.; Bentham Science Publishers Ltd.: Sharjah, United Arab Emirates, 2018. [Google Scholar]
  11. Weaver, C.M. Bioactive foods and ingredients for health. Adv. Nutr. 2014, 5, 306S–311S. [Google Scholar] [CrossRef] [PubMed]
  12. Hussain, A.; Kausar, T.; Sehar, S.; Sarwar, A.; Ashraf, A.H.; Jamil, M.A.; Noreen, S.; Rafique, A.; Iftikhar, K.; Quddoos, M.Y.; et al. A Comprehensive review of functional ingredients, especially bioactive compounds present in pumpkin peel, flesh and seeds, and their health benefits. Food Chem. Adv. 2022, 1, 100067. [Google Scholar] [CrossRef]
  13. Montesano, D.; Rocchetti, G.; Putnik, P.; Lucini, L. Bioactive profile of pumpkin: An overview on terpenoids and their health-promoting properties. Curr. Opin. Food Sci. 2018, 22, 81–87. [Google Scholar] [CrossRef]
  14. Rico, X.; Gullón, B.; Alonso, J.L.; Yáñez, R. Recovery of high value-added compounds from pineapple, melon, watermelon and pumpkin processing by-products: An overview. Food Res. Int. 2020, 132, 109086. [Google Scholar] [CrossRef]
  15. Mohaammed, S.; Paiko, Y.; Mann, A.; Ndamitso, M.; Mathew, J.; Maaji, S. Proximate, Mineral and Anti-nutritional Composition of Cucurbita maxima Fruits Parts. J. Chem. Res. 2015, 19, 37–49. [Google Scholar]
  16. Fedha, M.; Mwasaru, M.; Njoroge, C.; Ojijo, N.; Ouma, G. Effect of drying on selected proximate composition of fresh and processed fruits and seeds of two pumpkin species. Agric. Biol. J. N. Am. 2010, 1, 1299–1302. [Google Scholar] [CrossRef]
  17. Nwofia, G.E.; Nwogu Victoria, N.; Nwofia Blessing, K. Nutritional variation in fruits and seeds of pumpkins (Cucurbita spp) accessions from Nigeria. Pak. J. Nutr. 2012, 11, 848–858. [Google Scholar] [CrossRef]
  18. Ratnam, N.; Najibullah, M.; Ibrahim, M. A Review on Cucurbita pepo. Int. J. Pharmacogn. Phytochem. Res. 2017, 9, 1190–1194. [Google Scholar] [CrossRef]
  19. Kaur, S.; Panghal, A.; Garg, M.K.; Mann, S.; Khatkar, S.K.; Sharma, P.; Chhikara, N. Functional and nutraceutical properties of pumpkin—A review. Nutr. Food Sci. 2020, 50, 384–401. [Google Scholar] [CrossRef]
  20. Ponka, R.; Bouba, A.A.; Fokou, E.; Tambe, S.T.; Beaucher, E.; Piot, M.; Leonil, J.; Gaucheron, F. Protein, mineral and amino acid content of some Cameroonian traditional dishes prepared from pumpkin (Cucurbita maxima Duch.). J. Food Compos. Anal. 2015, 43, 169–174. [Google Scholar] [CrossRef]
  21. EFSA. EFSA Dietary Reference Values. Available online: (accessed on 25 October 2023).
  22. Adnan, M.; Gul, S.; Batool, S.; Fatima, B.; Rehman, A.; Yaqoob, S.; Shabir, H.; Yousaf, T.; Mussarat, S.; Ali, N.; et al. A review on the ethnobotany, phytochemistry, pharmacology and nutritional composition of Cucurbita pepo L. J. Phytopharm. 2017, 6, 133–139. [Google Scholar] [CrossRef]
  23. Dhiman, A.K.; Sharma, K.D.; Attri, S. Functional constituents and processing of pumpkin: A review. J. Food Sci. Technol. 2009, 46, 411–417. [Google Scholar]
  24. Sharma, P.; Kaur, G.; Kehinde, B.A.; Chhikara, N.; Panghal, A.; Kaur, H. Pharmacological and biomedical uses of extracts of pumpkin and its relatives and applications in the food industry: A review. Int. J. Veg. Sci. 2020, 26, 79–95. [Google Scholar] [CrossRef]
  25. Caili, F.; Huan, S.; Quanhong, L. A review on pharmacological activities and utilization technologies of pumpkin. Plant Foods Hum. Nutr. 2006, 61, 73–80. [Google Scholar] [CrossRef]
  26. Bochnak, J.; Świeca, M. Potentially bioaccessible phenolics, antioxidant capacities and the colour of carrot, pumpkin and apple powders—Effect of drying temperature and sample structure. Int. J. Food Sci. Technol. 2020, 55, 136–145. [Google Scholar] [CrossRef]
  27. Lemus-Mondaca, R.; Marin, J.; Rivas, J.; Sanhueza, L.; Soto, Y.; Vera, N.; Puente-Díaz, L. Pumpkin seeds (Cucurbita maxima). A review of functional attributes and by-products. Rev. Chil. Nutr. 2019, 46, 783–791. [Google Scholar] [CrossRef]
  28. Abdein, M.A.E.-H. Genetic Diversity between Pumpkin Accessions Growing in the Northern Border Region in Saudi Arabia Based on Biochemical and Molecular Parameters. Egypt. J. Bot. 2018, 58, 463–476. [Google Scholar] [CrossRef]
  29. Kaźmińska, K.; Sobieszek, K.; Targońska-Karasek, M.; Korzeniewska, A.; Niemirowicz-Szczytt, K.; Bartoszewski, G. Genetic diversity assessment of a winter squash and pumpkin (Cucurbita maxima Duchesne) germplasm collection based on genomic Cucurbita-conserved SSR markers. Sci. Hortic. 2017, 219, 37–44. [Google Scholar] [CrossRef]
  30. Hamdi, K.; Ben-Amor, J.; Mokrani, K.; Mezghanni, N.; Tarchoun, N. Assessment of the genetic diversity of some local squash (Cucurbita maxima Duchesne) populations revealed by agro-morphological and chemical traits. J. New Sci. 2017, 42, 2306–2317. [Google Scholar]
  31. FAO. Statistics Division Production and Trade Statistics. Available online: (accessed on 3 September 2023).
  32. Swiader, J.M.; Sipp, S.K.; Brown, R.E. Pumpkin growth, flowering, and fruiting response to nitrogen and potassium sprinkler fertigation in sandy soil. J. Am. Soc. Hortic. Sci. 1994, 119, 414–419. [Google Scholar] [CrossRef]
  33. Biesiada, A.; Nawirska, A.; Kucharska, A.; Sokół-Łȩtowska, A. The effect of nitrogen fertilization methods on yield and chemical composition of pumpkin (Cucurbita maxima) fruits before and after storage. Veg. Crops Res. Bull. 2009, 70, 203–211. [Google Scholar] [CrossRef]
  34. Yavuz, D.; Seymen, M.; Yavuz, N.; Türkmen, Ö. Effects of irrigation interval and quantity on the yield and quality of confectionary pumpkin grown under field conditions. Agric. Water Manag. 2015, 159, 290–298. [Google Scholar] [CrossRef]
  35. Kumari, A.; Lakshmi, G.A.; Krishna, G.K.; Patni, B.; Prakash, S.; Bhattacharyya, M.; Singh, S.K.; Verma, K.K. Climate Change and Its Impact on Crops: A Comprehensive Investigation for Sustainable Agriculture. Agronomy 2022, 12, 3008. [Google Scholar] [CrossRef]
  36. Habib-ur-Rahman, M.; Ahmad, A.; Raza, A.; Hasnain, M.U.; Alharby, H.F.; Alzahrani, Y.M.; Bamagoos, A.A.; Hakeem, K.R.; Ahmad, S.; Nasim, W.; et al. Impact of climate change on agricultural production; Issues, challenges, and opportunities in Asia. Front. Plant Sci. 2022, 13, 925548. [Google Scholar] [CrossRef]
  37. Petropoulos, S.A.; Levizou, E.; Ntatsi, G.; Fernandes, Â.; Petrotos, K.; Akoumianakis, K.; Barros, L.; Ferreira, I.C.F.R. Salinity effect on nutritional value, chemical composition and bioactive compounds content of Cichorium spinosum L. Food Chem. 2017, 214, 129–136. [Google Scholar] [CrossRef] [PubMed]
  38. Petropoulos, S.A.; Fernandes, Â.; Dias, M.I.; Pereira, C.; Calhelha, R.C.; Chrysargyris, A.; Tzortzakis, N.; Ivanov, M.; Sokovic, M.D.; Barros, L.; et al. Chemical composition and plant growth of Centaurea raphanina subsp. mixta plants cultivated under saline conditions. Molecules 2020, 25, 2204. [Google Scholar] [CrossRef]
  39. Polyzos, N.; Paschoalinotto, B.H.; Compocholi, M.; Pinela, J.; Heleno, S.A.; Calhelha, R.C.; Dias, M.I.; Barros, L.; Petropoulos, S.A. Fertilization of pot-grown Cichorium spinosum L.: How it can affect plant growth, chemical profile, and bioactivities of edible parts? Horticulturae 2022, 8, 890. [Google Scholar] [CrossRef]
  40. Kim, M.Y.; Kim, E.J.; Kim, Y.N.; Choi, C.; Lee, B.H. Comparison of the chemical compositions and nutritive values of various pumpkin (Cucurbitaceae) species and parts. Nutr. Res. Pract. 2012, 6, 21–27. [Google Scholar] [CrossRef]
  41. Stryjecka, M.; Krochmal-Marczak, B.; Cebulak, T.; Kiełtyka-Dadasiewicz, A. Assessment of Phenolic Acid Content and Antioxidant Properties of the Pulp of Five Pumpkin Species Cultivated in Southeastern Poland. Int. J. Mol. Sci. 2023, 24, 8621. [Google Scholar] [CrossRef]
  42. Leichtweis, M.G.; Molina, A.K.; Pires, T.C.S.; Dias, M.I.; Calhelha, R.; Bachari, K.; Ziani, B.E.C.; Oliveira, M.B.P.P.; Pereira, C.; Barros, L. Biological activity of pumpkin byproducts: Antimicrobial and antioxidant properties. Molecules 2022, 27, 8366. [Google Scholar] [CrossRef]
  43. AOAC. Official Methods of Analysis of AOAC International, 20th ed.; AOAC: Rockville, MD, USA, 2016. [Google Scholar]
  44. Pereira, C.; Barros, L.; Carvalho, A.M.; Ferreira, I.C.F.R. Nutritional composition and bioactive properties of commonly consumed wild greens: Potential sources for new trends in modern diets. Food Res. Int. 2011, 44, 2634–2640. [Google Scholar] [CrossRef]
  45. Brown, A.C.; Summers, W.L. Carbohydrate Accumulation and Color Development in Watermelon. J. Am. Soc. Hortic. Sci. 2022, 110, 683–687. [Google Scholar] [CrossRef]
  46. Barros, L.; Pereira, E.; Calhelha, R.C.; Dueñas, M.; Carvalho, A.M.; Santos-Buelga, C.; Ferreira, I.C.F.R. Bioactivity and chemical characterization in hydrophilic and lipophilic compounds of Chenopodium ambrosioides L. J. Funct. Foods 2013, 5, 1732–1740. [Google Scholar] [CrossRef]
  47. Pereira, C.; Barros, L.; Ferreira, I.C.F.R. Analytical tools used to distinguish chemical profiles of plants widely consumed as infusions and dietary supplements: Artichoke, milk thistle, and borututu. Food Anal. Methods 2014, 7, 1604–1611. [Google Scholar] [CrossRef]
  48. Backes, E.; Leichtweis, M.G.; Pereira, C.; Carocho, M.; Barreira, J.C.M.; Kamal Genena, A.; José Baraldi, I.; Filomena Barreiro, M.; Barros, L.; Ferreira, I.C.F.R. Ficus carica L. and Prunus spinosa L. extracts as new anthocyanin-based food colorants: A thorough study in confectionery products. Food Chem. 2020, 333, 127457. [Google Scholar] [CrossRef]
  49. Kuete, V.; Ango, P.Y.; Fotso, G.W.; Dwf Kapche, G.; Dzoyem, J.P.; Wouking, A.G.; Ngadjui, B.T.; Abegaz, B.M. Antimicrobial activities of the methanol extract and compounds from Artocarpus communis (Moraceae). BMC Complement. Altern. Med. 2011, 25, 11–42. [Google Scholar] [CrossRef] [PubMed]
  50. Pereira, C.; Calhelha, R.C.; Barros, L.; Ferreira, I.C.F.R. Antioxidant properties, anti-hepatocellular carcinoma activity and hepatotoxicity of artichoke, milk thistle and borututu. Ind. Crops Prod. 2013, 49, 61–65. [Google Scholar] [CrossRef]
  51. Jamal Uddin, A.; Faruq, M.; Roni, M.; Taufique, T.; Mehraj, H. Growth and yield performance of four pumpkin (Cucurbita moschata) lines. Int. J. Bus. Soc. Sci. Res. 2014, 2, 113–115. [Google Scholar]
  52. Gomes, R.S.; Machado, R.; De Almeida, C.F.; Chagas, R.R.; De Oliveira, R.L.; Delazari, F.T.; Da Silva, D.J.H. Brazilian germplasm of winter squash (Cucurbita moschata D.) displays vast genetic variability, allowing identification of promising genotypes for agro-morphological traits. PLoS ONE 2020, 15, e0230546. [Google Scholar] [CrossRef]
  53. Soltani, F.; Karimi, R.; Kashi, A. Estimation of Genetic Diversity in Cucurbita Species Using Morphological and Phytochemical Analysis. Int. J. Veg. Sci. 2017, 23, 42–53. [Google Scholar] [CrossRef]
  54. Ghimire, S.; Wszelaki, A.L.; Moore, J.C.; Inglis, D.A.; Miles, C. The use of biodegradable mulches in pie pumpkin crop production in two diverse climates. HortScience 2018, 53, 288–294. [Google Scholar] [CrossRef]
  55. El-Hamed, K.E.-S.A.; Elwan, M.W.M. Dependence of Pumpkin Yield on Plant Density and Variety. Am. J. Plant Sci. 2011, 2, 636–643. [Google Scholar] [CrossRef]
  56. Ahamed, K.; Akhter, B.; Islam, M.; Ara, N.; Humauan, M. An Assessment of Morphology and Yield Characteristics of Pumpkin (Cucurbita moschata) Genotypes in Northern Bangladesh. Trop. Agric. Res. Ext. 2012, 14, 7. [Google Scholar] [CrossRef]
  57. Badr, S.E.A.; Shaaban, M.; Elkholy, Y.M.; Helal, M.H.; Hamza, A.S.; Masoud, M.S.; El Safty, M.M. Chemical composition and biological activity of ripe pumpkin fruits (Cucurbita pepo L.) cultivated in Egyptian habitats. Nat. Prod. Res. 2011, 25, 1524–1539. [Google Scholar] [CrossRef] [PubMed]
  58. Jacobo-Valenzuela, N.; Zazueta-Morales, J.d.J.; Gallegos-Infante, J.A.; Aguilar-Gutierrez, F.; Camacho-Hernández, I.L.; Rocha-Guzman, N.E.; Gonzalez-Laredo, R.F. Chemical and physicochemical characterization of winter squash (Cucurbita moschata D.). Not. Bot. Horti Agrobot. Cluj-Napoca 2011, 39, 34–40. [Google Scholar] [CrossRef]
  59. Amin, M.Z.; Islam, T.; Uddin, M.R.; Uddin, M.J.; Rahman, M.M.; Satter, M.A. Comparative study on nutrient contents in the different parts of indigenous and hybrid varieties of pumpkin (Cucurbita maxima Linn.). Heliyon 2019, 5, e02462. [Google Scholar] [CrossRef] [PubMed]
  60. Salehi, B.; Sharifi-Rad, J.; Capanoglu, E.; Adrar, N.; Catalkaya, G.; Shaheen, S.; Jaffer, M.; Giri, L.; Suyal, R.; Jugran, A.; et al. Cucurbita Plants: From Farm to Industry. Appl. Sci. 2019, 9, 3387. [Google Scholar] [CrossRef]
  61. Louwaars, N.P. Plant breeding and diversity: A troubled relationship? Euphytica 2018, 214, 114. [Google Scholar] [CrossRef]
  62. Pereira, A.M.; Krumreich, F.D.; Ramos, A.H.; Krolow, A.C.R.; Santos, R.B.; Gularte, M.A. Physicochemical characterization, carotenoid content and protein digestibility of pumpkin access flours for food application. Food Sci. Technol. 2020, 40, 691–698. [Google Scholar] [CrossRef]
  63. Marek, G.; Radzanowska, J.; Danilcenko, H.; Jariene, E.; Cerniauskiene, J. Quality of Pumpkin Cultivars in Relation to Sensory Characteristics. Not. Bot. Horti Agrobot. Cluj-Napoca 2008, 36, 73–79. [Google Scholar]
  64. Seroczyńska, A.; Antczak, A.; Kamińska, K.; Korytowska, M.; Korzeniewska, A.; Niemirowicz-szczytt, K.; Radomski, A.; Zawadzki, J. Evaluation of the selected forms of winter squash (Cucurbita maxima Duch.) for the content of free sugars and polysaccharides. Pol. J. Agron. 2014, 16, 69–73. [Google Scholar]
  65. Dhenge, R.; Rinaldi, M.; Ganino, T.; Santi, S.; Ferrarese, I.; Dall’Acqua, S. Variations of polyphenols, sugars, carotenoids, and volatile constituents in pumpkin (Cucurbita moschata) during high pressure processing: A kinetic study. Innov. Food Sci. Emerg. Technol. 2022, 78, 103005. [Google Scholar] [CrossRef]
  66. Kostecka-Gugała, A.; Kruczek, M.; Ledwożyw-Smoleń, I.; Kaszycki, P. Antioxidants and health-beneficial nutrients in fruits of eighteen Cucurbita cultivars: Analysis of diversity and dietary implications. Molecules 2020, 25, 1792. [Google Scholar] [CrossRef] [PubMed]
  67. Ghosh, P.; Rana, S.S. Physicochemical, nutritional, bioactive compounds and fatty acid profiling of Pumpkin flower (Cucurbita maxima), as a potential functional food. SN Appl. Sci. 2021, 3, 216. [Google Scholar] [CrossRef]
  68. Iosypenko, O.O.; Kyslychenko, V.S.; Omelchenko, Z.I.; Burlaka, I.S. Fatty acid composition of vegetable marrows and zucchini leaves. Pharmacia 2019, 66, 201–207. [Google Scholar] [CrossRef]
  69. Guil, J.L.; Torija, M.E.; Giménez, J.J.; Rodriguez, I. Identification of fatty acids in edible wild plants by gas chromatography. J. Chromatogr. A 1996, 719, 229–235. [Google Scholar] [CrossRef] [PubMed]
  70. Leichtweis, M.G.; Molina, A.K.; Petropoulos, S.A.; Carocho, M.; Pires, T.C.S.P.; Dias, M.I.; Calhelha, R.; Oliveira, M.B.P.P.; Pereira, C.; Barros, L. Valorization of Pumpkin Peel as a Source of Bioactive Compounds: Optimization of Heat- and Ultrasound-Assisted Extraction. Molecules 2023, 28, 3168. [Google Scholar] [CrossRef]
  71. Rolnik, A.; Kowalska, I.; Soluch, A.; Stochmal, A.; Olas, B. Comparative Phytochemical, Antioxidant and Haemostatic Studies of Preparations from Selected Vegetables from Cucurbitaceae Family. Molecules 2020, 25, 4326. [Google Scholar] [CrossRef]
  72. Santos, E.M.; Rodriguez, J.A.; Lorenzo, J.M.; Mondragón, A.C.; Pateiro, M.; Gutiérrez, E.; Ferreira, T.A. Antioxidant Effect of Pumpkin Flower (Cucurbita maxima) in Chicken Patties. Foods 2022, 11, 2258. [Google Scholar] [CrossRef]
  73. Bulambaeva, A.A.; Vlahova-Vangelova, T.B.; Dragoev, S.G.; Balev, D.K.; Uzakov, Y.M. Development of new functional cooked sausages by addition of goji berry and pumpkin powder. Am. J. Food Technol. 2014, 9, 180–189. [Google Scholar] [CrossRef]
  74. Rolnik, A.; Skalski, B.; Stochmal, A.; Olas, B. Preparations from selected cucurbit vegetables as antiplatelet agents. Sci. Rep. 2021, 11, 22694. [Google Scholar] [CrossRef]
  75. Tarwadi, K.; Agte, V. Antioxidant and micronutrient quality of fruit and root vegetables from the Indian subcontinent and their comparative performance with green leafy vegetables and fruits. J. Sci. Food Agric. 2005, 85, 1469–1476. [Google Scholar] [CrossRef]
  76. Mokhtar, M.; Bouamar, S.; Di Lorenzo, A.; Temporini, C.; Daglia, M.; Riazi, A. The influence of ripeness on the phenolic content, antioxidant and antimicrobial activities of pumpkins (Cucurbita moschata Duchesne). Molecules 2021, 26, 3623. [Google Scholar] [CrossRef] [PubMed]
  77. Hussain, A.; Kausar, T.; Din, A.; Murtaza, A.; Jamil, M.A.; Noreen, S.; Iqbal, M.A. Antioxidant and Antimicrobial Properties of Pumpkin (Cucurbita maxima) Peel, Flesh and Seeds Powders. J. Biol. Agric. Healthc. 2021, 6, 42–51. [Google Scholar] [CrossRef]
  78. Hussain, A.; Kausar, T.; Sehar, S.; Sarwar, A.; Ashraf, A.H.; Jamil, M.A.; Noreen, S.; Rafique, A.; Iftikhar, K.; Aslam, J.; et al. Utilization of pumpkin, pumpkin powders, extracts, isolates, purified bioactives and pumpkin based functional food products: A key strategy to improve health in current post COVID 19 period: An updated review. Appl. Food Res. 2022, 2, 100241. [Google Scholar] [CrossRef]
  79. Gaweł-Bęben, K.; Czech, K.; Strzępek-Gomółka, M.; Czop, M.; Szczepanik, M.; Lichtarska, A.; Kukula-Koch, W. Assessment of Cucurbita spp. Peel Extracts as Potential Sources of Active Substances for Skin Care and Dermatology. Molecules 2022, 27, 7618. [Google Scholar] [CrossRef] [PubMed]
  80. Piccolella, S.; Bianco, A.; Crescente, G.; Santillo, A.; Baccari, G.C.; Pacifico, S. Recovering Cucurbita pepo cv. ‘Lungo Fiorentino’ Wastes: UHPLC-HRMS/MS metabolic profile, the basis for establishing their nutra- And cosmeceutical valorisation. Molecules 2019, 24, 1479. [Google Scholar] [CrossRef] [PubMed]
  81. Massironi, A.; Biella, S.; de Moura Pereira, P.F.; Scibona, F.; Feni, L.; Sindaco, M.; Emide, D.; Jiménez-Quero, A.; Bianchi, C.L.M.; Verotta, L.; et al. Valorization of pumpkin seed hulls, cucurbitin extraction strategies and their comparative life cycle assessment. J. Clean. Prod. 2023, 427, 139267. [Google Scholar] [CrossRef]
  82. Priecina, L.; Karklina, D. Composition of Major Organic Acids in Vegetables and Spices. CBU Int. Conf. Proc. 2015, 3, 447–454. [Google Scholar] [CrossRef]
  83. Iswaldi, I.; Gómez-Caravaca, A.M.; Lozano-Sánchez, J.; Arráez-Román, D.; Segura-Carretero, A.; Fernández-Gutiérrez, A. Profiling of phenolic and other polar compounds in zucchini (Cucurbita pepo L.) by reverse-phase high-performance liquid chromatography coupled to quadrupole time-of-flight mass spectrometry. Food Res. Int. 2013, 50, 77–84. [Google Scholar] [CrossRef]
  84. Abbas, H.M.K.; Huang, H.X.; Wang, A.J.; Wu, T.Q.; Xue, S.D.; Ahmad, A.; Xie, D.S.; Li, J.X.; Zhong, Y.J. Metabolic and tran-scriptomic analysis of two Cucurbita moschata germplasms throughout fruit development. BMC Genom. 2020, 21, 365. [Google Scholar] [CrossRef]
  85. Nawirska-Olszańska, A.; Biesiada, A.; Sokół-ŁȨtowska, A.; Kucharska, A.Z. Characteristics of organic acids in the fruit of different pumpkin species. Food Chem. 2014, 148, 415–419. [Google Scholar] [CrossRef]
  86. Zhou, C.L.; Mi, L.; Hu, X.Y.; Zhu, B.H. Evaluation of three pumpkin species: Correlation with physicochemical, antioxidant properties and classification using SPME-GC–MS and E-nose methods. J. Food Sci. Technol. 2017, 54, 3118–3131. [Google Scholar] [CrossRef] [PubMed]
Figure 1. Profile A (from V2 T sample), rich in fructose (I) and glucose (II); Profile B (from V1 sample), rich in sucrose (III).
Figure 1. Profile A (from V2 T sample), rich in fructose (I) and glucose (II); Profile B (from V1 sample), rich in sucrose (III).
Horticulturae 09 01232 g001
Figure 2. Three-dimensional scatterplot of principal components 1, 2 and 3 for pumpkin fruit flesh.
Figure 2. Three-dimensional scatterplot of principal components 1, 2 and 3 for pumpkin fruit flesh.
Horticulturae 09 01232 g002
Figure 3. The loading plot of principal components 1 and 2 for pumpkin fruit flesh.
Figure 3. The loading plot of principal components 1 and 2 for pumpkin fruit flesh.
Horticulturae 09 01232 g003
Figure 4. The loading plot of principal components 1 and 3 for pumpkin fruit flesh.
Figure 4. The loading plot of principal components 1 and 3 for pumpkin fruit flesh.
Horticulturae 09 01232 g004
Table 1. Genotype name of pumpkin cultivated in Greece according to the fruit type.
Table 1. Genotype name of pumpkin cultivated in Greece according to the fruit type.
Pumpkin GenotypesSample CodeHarvest Date
Fytro FS-243V126 October–19 November 2021
Landrace from the region of Trikala (Turbinate)V2 T25 August–19 October 2021
Landrace from the region of Trikala (Cylindrical)V2 C25 August–19 October 2021
Big MaxV325 August–19 October 2021
Local landrace “Nychaki” (Cylindrical)V4 C23 September 2021
Local landrace “Nychaki” (Round)V4 R23 September 2021
Local landrace “Leuka Melitis” (Flattened)V5 F5 October–19 November 2021
Local landrace “Leuka Melitis” (Round)V5 R5 October–19 November 2021
Local landrace from the region of LakoniaV625 August 2021
Local landrace from the region of Lakonia (Pyriform)V7 P24 September–19 November 2021
Local landrace from the region of Lakonia (Flattened)V7 F24 September–19 November 2021
Local landrace from the region of LakoniaV822 November 2021
Local landrace “Makedonika prasina” (Cylindrical)V9 C22 November 2021
Local landrace “Makedonika prasina” (Round)V9 R22 November 2021
Local landrace from the region of LaconiaV1022 November 2021
Local landrace (“Voutirato”)V1122 November 2021
Table 2. Crop performance (total fruit weight in kg/ha) of the tested genotypes.
Table 2. Crop performance (total fruit weight in kg/ha) of the tested genotypes.
GenotypeTotal Fruit Weight (Tonnes/ha)Numbers of Fruit/PlantMean Fruit Weight
V115.5 ± 6.6 g6.1 ± 0.5 a1.0 ± 0.1 g
V230.7 ± 5.6 b0.9 ± 0.1 e6.3 ± 0.6 b
V319.6 ± 7.8 e1.0 ± 0.1 d,e8.1 ± 0.7 a
V428.0 ± 3.6 c3.9 ± 0.4 b2.9 ± 0.3 f
V57.6 ± 2.4 j0.4 ± 0.1 g3.6 ± 0.4 e
V610.9 ± 4.5 i1.2 ± 0.2 d3.8 ± 0.4 e
V724.0 ± 9.6 d2.9 ± 0.3 c1.7 ± 0.2 g
V834.1 ± 5.4 a1.1 ± 0.1 d6.1 ± 0.5 b
V916.7 ± 2.2 f0.7 ± 0.1 f4.8 ± 0.4 d
V1027.7 ± 6.3 c4.1 ± 0.3 b2.7 ± 0.3 f
V1111.8 ± 3.9 h0.8 ± 0.1 e,f5.6 ± 0.5 c
Means of the same column followed by different Latin letters are statistically different according to Tukey’s honestly significant difference (HSD) test at p < 0.05.
Table 3. Proximate composition of the flesh of the tested pumpkin genotypes on a dry weight (dw) basis.
Table 3. Proximate composition of the flesh of the tested pumpkin genotypes on a dry weight (dw) basis.
(g/100 g dw)
(g/100 g dw dw)
(g/100 g dw)
(Kcal/100 g dw)
(g/100 g dw)
(g/100 g dw)
V10.83 ± 0.04 c12.6 ± 0.6 g,h4.511 ± 0.009 l360.1 ± 0.4 a69.1 ± 0.7 b13.0 ± 0.1 j
V2 T1.12 ± 0.06 a13.7 ± 0.4 d,e6.07 ± 0.04 j332.8 ± 0.3 h54.9 ± 0.7 h24.27 ± 0.02 c
V2 C0.645 ± 0.002 e12.9 ± 0.1 f,g7.1 ± 0.1 h334.6 ± 1.2 g59.2 ± 0.8 f,g20.2 ± 0.8 f
V30.45 ± 0.02 h18.07 ± 0.03 c7.7 ± 0.2 e321.85 ± 0.02 j49.0 ± 0.2 j24.7 ± 0.5 c
V4 C0.64 ± 0.03 e,f8.6 ± 0.1 k7.1 ± 0.1 g,h332.5 ± 0.1 h62.6 ± 0.2 e21.1 ± 0.2 e,f
V4 R0.38 ± 0.02 h17.46 ± 0.04 c9.08 ± 0.07 d321.7 ± 0.6 j51.1 ± 0.3 i21.9 ± 0.4 d,e
V5 F0.58 ± 0.03 f,g12.61 ± 0.05 g5.126 ± 0.006 k353.70 ± 0.07 b67.3 ± 0.1 c14.34 ± 0.04 i
V5 R0.55 ± 0.03 g9.9 ± 0.3 j3.5 ± 0.1 m361.0 ± 0.1 a72.2 ± 0.6 a13.8 ± 0.4 i,j
V61.17 ± 0.02 a13.4 ± 0.3 e,f6.98 ± 0.05 h341.5 ± 0.8 e60.27 ± 0.03 f18.2 ± 0.3 g
V7 P0.92 ± 0.04 b14.1 ± 0.1 d7.49 ± 0.03 f343.6 ± 0.6 d61.9 ± 0.6 e15.5 ± 0.5 h
V7 F0.70 ± 0.03 d,e11.9 ± 0.1 h,i7.27 ± 0.06 g348.76 ± 0.05 c67.26 ± 0.6 c12.8 ± 0.2 j
V80.76 ± 0.03 c,d11.3 ± 0.4 i7.29 ± 0.02 g329.4 ± 1.6 i58.0 ± 1.3 g22.7 ± 0.9 d
V9 C0.73 ± 0.03 d21.4 ± 0.6 a10.95 ± 0.05 a315.05 ± 0.96 k44.5 ± 0.2 k22.4 ± 0.5 d
V9 R0.65 ± 0.03 e20.3 ± 0.3 b10.19 ± 0.07 b307.7 ± 1.5 l41.4 ± 0.4 l27.4 ± 0.8 a
V100.73 ± 0.03 d9.9 ± 0.2 j9.3 ± 0.1 c313.8 ± 0.1 k53.7 ± 0.2 h26.3 ± 0.1 b
V111.12 ± 0.02 a8.0 ± 0.3 k6.71 ± 0.04 i338.52 ± 0.34 f64.0 ± 0.4 d20.1 ± 0.2 f
Means of the same column followed by different Latin letters are statistically different according to Tukey’s honestly significant difference (HSD) test at p < 0.05; T: turbinate; C: cylindrical; R: round; F: flattened.
Table 4. Free sugar content of the flesh of the tested pumpkin genotypes (g/100 g dry weight (dw)).
Table 4. Free sugar content of the flesh of the tested pumpkin genotypes (g/100 g dry weight (dw)).
V12.42 ± 0.03 i2.50 ± 0.08 j12.2 ± 0.4 b0.46 ± 0.03 c,d,e0.0579 ± 0.0003 h,i17.7 ± 0.5 g
V2 T9.7 ± 0.9 a21.5 ± 0.3 b,c1.4 ± 0.2 i0.45 ± 0.02 c,d,e0.29 ± 0.03 a33 ± 1 b
V2 C10.0 ± 0.3 a22 ± 1 b0.9 ± 0.2 i0.67 ± 0.02 a0.09 ± 0.02 g,h,i34 ± 1 b
V36.57 ± 0.02 e9.6 ± 0.4 g2.6 ± 0.3 g0.23 ± 0.02 i0.23 ± 0.03 b,c,d19.2 ± 0.8 f,g
V4 C7.854 ± 0.001 c,d18.6 ± 0.8 e6.4 ± 0.7 d0.51 ± 0.06 b,c0.13 ± 0.04 f,g33 ± 2 b
V4 R6.6 ± 0.3 e10.2 ± 0.6 g3.47 ± 0.01 f0.47 ± 0.03 c,d0.24 ± 0.03 b,c20.9 ± 0.9 e,f
V5 F3.790 ± 0.002 g4.8 ± 0.1 h3.6 ± 0.2 f0.408 ± 0.005 d,e,f0.18 ± 0.01 d,e,f12.8 ± 0.4 h
V5 R2.55 ± 0.07 i4.66 ± 0.05 h4.91 ± 0.05 e0.46 ± 0.01 c,d,e0.06 ± 0.05 h,i12.6 ± 0.1 h
V68.18 ± 0.04 c20.33 ± 0.06 c,d1.12 ± 0.06 i0.50 ± 0.03 b,c0.27 ± 0.02 a,b30.41 ± 0.09 c
V7 P2.8 ± 0.2 h,i3.1 ± 0.2 i,j14.0 ± 0.5 a0.56 ± 0.02 b0.05 ± 0.01 i20.4 ± 0.8 f
V7 F3.3 ± 0.2 g,h4.7 ± 0.2 h14.0 ± 0.7 a0.50 ± 0.02 b,c0.057 ± 0.008 h,i23 ± 1 d,e
V87.4 ± 0.4 d19 ± 2 d,e4.0 ± 0.2 f0.37 ± 0.02 f,g0.25 ± 0.02 a,b31 ± 1 c
V9 C8.23 ± 0.04 c12.2 ± 0.5 f2.2 ± 0.1 g,h0.26 ± 0.04 h,i0.16 ± 0.02 e,f23.1 ± 0.3 d
V9 R4.9 ± 0.1 f4.3 ± 0.4 h,i10.4 ± 0.3 c0.40 ± 0.06 e,f0.16 ± 0.03 e,f20.2 ± 0.9 f
V106.7 ± 0.4 e19.3 ± 0.8 d,e3.48 ± 0.08 f0.3273 ± 0.0007 g,h0.108 ± 0.004 g,h30 ± 1 c
V118.9 ± 0.4 b26.9 ± 0.9 a1.50 ± 0.06 h,i0.48 ± 0.04 c0.20 ± 0.03 c,d,e38.0 ± 0.5 a
Means of the same column followed by different Latin letters are statistically different according to Tukey’s honestly significant difference (HSD) test at p < 0.05. T: turbinate; C: cylindrical; R: round; F: flattened.
Table 5. Fatty acids composition of the flesh of the tested pumpkin genotypes (relative %).
Table 5. Fatty acids composition of the flesh of the tested pumpkin genotypes (relative %).
CompoundV1V2 TV2 CV3V4 CV4 RV5 FV5 RV6V7 PV7 FV8V9 CV9 RV10V11
C8:0n.d.n.d.0.264 ± 0.006 an.d.0.24 ± 0.01 bn.d.n.d.n.d.n.d.n.d.n.d.n.d.0.141 ± 0.001 cn.d.n.d.n.d.
C10:00.137 ±
0.001 c
n.d.0.1775 ± 0.0007 a0.0830 ± 0.0000 g0.143 ± 0.002 bn.d.n.d.0.0720 ±
0.0000 h
n.d.0.120 ±
0.001 d
0.070 ± 0.002 i0.0680 ± 0.0000 i0.100 ± 0.001 f0.1125 ±
0.0007 e
C12:01.893 ±
0.004 a
0.219 ± 0.03 j,k0.49 ± 0.02 e0.500 ± 0.004e0.61 ± 0.01 d0.3050 ±
0.0000 g,h
0.25 ±
0.01 i,j,k
0.330 ±
0.001 f,g
0.2135 ± 0.0007 k1.49 ±
0.05 b
1.21 ± 0.04 c0.259 ± 0.003 i,j0.342 ± 0.004 f,g0.36 ± 0.01 f02755 ±
00007 h,i
0.125 ±
0.004 l
C13:0n.d.n.d.n.d.0.1020 ± 0.0000 bn.d.n.d.0.058 ±
0.001 d
0.0935 ±
0.0007 c
0.0940 ± 0.0000 cn.d.n.d.n.d.n.d.0.120 ± 0.006 an.d.n.d.
C14:01.898 ±
0.007 a
1.22 ± 0.03 e1.84 ± 0.03 b1.0505 ± 0.0007 g1.5980 ± 0.0000 c0.7960 ±
0.0000 j,k
0.90 ± 0.01 h1.053 ± 0.006 g0.866 ±
0.001 h,i
1.55 ± 0.04 c1.13 ± 0.05 f0.83 ± 0.01 i,j1.29 ± 0.02 d1.21 ± 0.01 e0.748 ± 0.008 k0.83 ± 0.04 i,j
C15:00.079 ±
0.004 k
0.117 ±
0.004 i
0.53 ± 0.01 b0.1690 ± 0.0000 g0.648 ± 0.007 a0.190 ± 0.001 fn.d.0.0895 ±
0.0007 j
0.1350 ± 0.0000 h0.1095 ± 0.0007 i0.0775 ± 0.0007 k0.290 ± 0.003 e0.445 ± 0.005 c0.1945 ±
0.0007 f
0.3455 ±
0.0007 d
0.133 ±
0.006 h
C16:028.9 ± 0.2 e20.9 ± 0.6 k61.4 ± 0.5 b36.46 ± 0.04 c63.4 ± 0.2 a29.8 ± 0.2 d26.97 ±
0.03 f,g
26.3 ± 0.1 h 20.88 ± 0.03 k25.67 ± 0.01 i22.3 ± 0.2 j30.2 ± 0.1 d26.49 ± 0.05 g,h27.3 ± 0.1 f25.982 ±
0.007 h,i
19.6 ± 0.5 l
C16:10.42 ± 0.02 g0.269 ±
0.006 k
0.586 ± 0.006 b0.295 ±
0.004 j
0.53 ± 0.01 c,d0.50 ± 0.02 e0.45 ± 0.01 f0.388 ± 0.002 h0.61 ± 0.02 a0.283 ±
0.009 j,k
0.36 ± 0.02 i0.50 ± 0.01 e0.520 ± 0.004 d,e0.231 ± 0.008 l0.37 ± 0.01 h,i0.55 ± 0.02 c
C17:00.390 ±
0.002 f,g
0.224 ±
0.002 i,j
6.39 ± 0.05 b1.23 ± 0.04 c7.01 ± 0.04 a0.969 ± 0.007 d0.222 ±
0.001 i,j
0.237 ± 0.001 i,j0.1520 ± 0.0000 k0.602 ± 0.004 e0.349 ± 0.001 h0.93 ± 0.01 d0.350 ± 0.004 g,h0.26 ± 0.01 i0.423 ± 0.001 f0.199 ±
0.009 j
C17:10.182 ±
0.006 d
0.107 ±
0.002 h
0.84 ± 0.01 a0.3770 ± 0.0000 c0.65 ± 0.01 bn.d.0.132 ±
0.001 g
0.1465 ±
0.0007 e,f
0.1250 ± 0.0000 g0.377 ±
0.009 c
n.d.0.1355 ±
0.0007 f,g
0.0945 ±
0.0007 i
n.d.0.1805 ±
0.0007 d
0.151 ±
0.006 e
C18:02.5 ± 0.1 e,f,g1.85 ± 0.05 j4.89 ± 0.07 b2.45 ± 0.08 f,g5.30 ± 0.02 a2.614 ± 0.008 e2.25 ± 0.06 h2.02 ± 0.01 i1.568 ± 0.005 k4.2 ± 0.1 c3.0 ± 0.1 d2.53 ± 0.05 e,f2.3 ± 0.1 h1.64 ± 0.04 k2.37 ±
0.04 g,h
1.54 ± 0.06 k
C18:1n9c12.1 ± 0.3 c14.1 ± 0.1 a8.4 ± 0.2 f6.0 ± 0.2 i4.95 ± 0.05 j7.6 ± 0.2 g8.6 ± 0.2 e,f6.53 ± 0.03 h9.57 ± 0.04 d13.30 ± 0.02 b8.9 ± 0.4 e4.720 ± 0.007 j5.86 ± 0.01 i1.980 ± 0.006 l3.919 ± 0.006 k9.46 ± 0.03 d
C18:2n6c22.9 ± 0.2 k34.0 ± 0.8 c5.1 ± 0.1 m25.8 ± 0.2 i5.9 ± 0.2 l28.71 ± 0.08 g27.0 ± 0.1 h32.2 ± 0.2 e30.12 ± 0.03 f24.8 ± 0.1 j29.8 ± 0.2 f35.78 ± 0.03 a31.97 ± 0.04 e33.4 ± 0.1 d34.65 ± 0.07 b25.09 ± 0.07 j
C18:3n60.162 ±
0.006 c
0.080 ±
0.002 d
n.d.n.d.0.54 ± 0.03 an.d.0.216 ±
0.006 b
C18:3n325.9 ± 0.3 f24.9 ± 0.4 g0.512 ± 0.001 l20.4 ± 0.1 j0.60 ± 0.02 l24.48 ± 0.02 h29.81 ± 0.02 c27.3 ± 0.4 e32.87 ± 0.04 b23.42 ± 0.07 i29.53 ± 0.08 c19.003 ±
0.0007 k
25.73 ± 0.07 f29.59 ± 0.09 c28.5 ± 0.1 d38.8 ± 0.3 a
C20:0n.d.n.d.1.03 ± 0.03 a0.3790 ± 0.0000 c0.75 ± 0.03 bn.d.n.d.0.213 ± 0.001 g0.1840 ± 0.0000 h0.311 ±
0.003 d,e
0.25 ± 0.01 f0.289 ± 0.001 e0.335 ± 0.001 d0.37 ± 0.01 c0.2315 ±
0.0007 f,g
0.235 ±
0.008 f,g
C20:1n.d.n.d.n.d.n.d.n.d.n.d.n.d.n.d.0.2015 ± 0.0007 bn.d.n.d.n.d.0.542 ± 0.002 an.d.n.d.n.d.
C21:0n.d.n.d.n.d.n.d.n.d.n.d.n.d.0.1345 ±
0.0007 f
0.1455 ± 0.0007 e0.184 ± 0.002 cn.d.0.241 ± 0.001 a0.1555 ± 0.0007 d0.2135 ±
0.0007 b
C20:3n6n.d.n.d.n.d.n.d.n.d.n.d.n.d.n.d.n.d.n.d.n.d.n.d.0.1355 ± 0.0007n.d.n.d.n.d.
C22:00.68 ± 0.02 j0.811 ±
0.004 i
3.23 ± 0.01 a1.11 ± 0.03 e,f2.02 ± 0.02 b1.26 ± 0.04 c1.16 ± 0.04 d,e1.089 ± 0.006 f0.876 ± 0.009 h1.16 ± 0.01 d,e1.13 ± 0.05 d,e,f1.18 ± 0.02 d0.838 ± 0.003 h,i0.973 ± 0.005 g0.653 ± 0.002 j1.26 ± 0.05 c
C20:5n30.099 ± 0.003n.d.n.d.n.d.n.d.n.d.n.d.n.d.n.d.n.d.n.d.n.d.n.d.n.d.n.d.n.d.
C22:1n.d.n.d.n.d.0.166 ± 0.006 cn.d.0.602 ± 0.009 an.d.n.d.n.d.n.d.n.d.n.d.0.435 ± 0.002 bn.d.n.d.n.d.
C22:20.264 ±
0.004 d
0.163 ± 0.004 hn.d.0.223 ± 0.002 f0.246 ± 0.008 en.d.0.152 ±
0.006 i
0.28 ± 0.01 c0.141 ± 0.005 j0.536 ± 0.006 a0.240 ± 0.007 e0.334 ± 0.002 b0.225 ± 0.001 f0.1840 ±
0.0000 g
0.1775 ±
0.0007 g
0.150 ±
0.004 i,j
C22:6n3n.dn.dn.dn.d0291 ± 0006n.dn.dn.dn.dn.dn.dn.dn.dn.dn.dn.d
C23:00.368 ±
0.001 g,h
0.33 ± 0.01 i0.98 ± 0.04 a0.519 ± 0.009 d0.87 ± 0.03 b0.427 ± 0.006 f0.385 ±
0.008 g,h
0.37 ± 0.01 g,h0.39 ± 0.01 g0.539 ± 0.006 d0.47 ± 0.02 e0.438 ± 0.002 e,f0.537 ± 0.002 d0.615 ± 0.001 c0.360 ± 0.001 h,i0.598 ±
0.006 c
C24:01.14 ± 0.01 h0.78 ± 0.03 i3.4 ± 0.2 b2.63 ± 0.06 c3.696 ± 0.008 a1.67 ± 0.03 e1.38 ± 0.01 f1.141 ± 0.006 h0.854 ± 0.008 i1.36 ± 0.01 f1.25 ± 0.05 g,h2.3 ± 0.1 d1.136 ± 0.004 h1.1905 ±
0.0007 g,h
0.811 ± 0.003 i1.28 ± 0.03 f,g
C24:1n.d.n.d.n.d.n.d.n.d.n.d.n.d.n.d.n.d.n.d.n.d.n.d.0.0875 ± 0.0007n.d.n.d.n.d.
SFA38.0 ± 0.3 e26.4 ± 0.5 k84.6 ± 0.4 b46.7 ± 0.1 c86.3 ± 0.2 a38.8 ± 0.09 e33.58 ± 0.02 h33.1 ± 0.2 h26.36 ± 0.03 k37.3 ± 0.2 d31.2 ± 0.3 j39.53 ± 0.02 d34.41 ± 0.03 g34.6 ± 0.2 g32.20 ± 0.02 i25.8 ± 0.4 l
MUFA12.7 ± 0.2 c14.5 ± 0.1 a9.8 ± 0.3 f6.9 ± 0.2 j6.13 ± 0.07 k8.7 ± 0.2 h9.2 ± 0.2 g7.07 ± 0.04 j10.51 ± 0.05 d13.96 ± 0.00 b9.2 ± 0.4 g5.36 ± 0.02 l7.533 ± 0.006 i2.211 ± 0.003 n4.46 ± 0.01 m10.157 ± 0.008 e
PUFA49.3 ± 0.5 i59.1 ± 0.4 d5.6 ± 0.1 m46.4 ± 0.3 k7.6 ± 0.3 l53.2 ± 0.1 h57.2 ± 0.2 f59.8 ± 0.2 c63.13 ± 0.02 b48.8 ± 0.2 j59.59 ± 0.07 c,d55.12 ± 0.03 g58.05 ± 0.04 e63.2 ± 0.2 b63.34 ± 0.03 b64.0 ± 0.4 a
Means of the same row followed by different Latin letters are statistically different according to Tukey’s honestly significant difference (HSD) test at p < 0.05; n.d.: not detected. T: turbinate; C: cylindrical; R: round; F: flattened; Caprylic acid (C8:0); Capric acid (C10:0); lauric acid (C12:0); tridecylic acid (C13:0); myristic acid (C14:0); pentadecylic acid (C15:0); palmitic acid (C16:0); palmitoleic acid (C16:1); margaric acid (C17:0); heptadecenoic acid (C17:1); stearic acid (C18:0); oleic acid (C18:1n9); linoleic acid (C18:2n6c); γ-linolenic acid (C18:3n6); α-linolenic acid (C18:3n3); arachidic acid (C20:0); gondoic acid (C20:1); heneicosylic acid (C21:0); arachidonic acid (C20:3n6); behenic acid (C22:0); eicosatrienoic acid (C20:5n3); erucic acid (C22:1); docosadienoic acid (C22:2); (C20:6n3); tricosylic acid (C23:0); lignoceric acid (C24:0); nervonic acid (C24:1); SFA: saturated fatty acids; MUFA: monounsaturated fatty acids; PUFA: polyunsaturated fatty acids.
Table 6. Lipid peroxidation inhibition capacity (TBARS) and oxidative haemolysis inhibition capacity (OxHLIA) of the flesh of the tested pumpkin genotypes (inhibition capacity (IC50), μg/mL).
Table 6. Lipid peroxidation inhibition capacity (TBARS) and oxidative haemolysis inhibition capacity (OxHLIA) of the flesh of the tested pumpkin genotypes (inhibition capacity (IC50), μg/mL).
(IC50 1, μg/mL)
OxHLIA 60 min
(IC50 1, μg/mL)
V12877 ± 79 i674 ± 21 c
V2 T3000 ± 120 g,h302 ± 9 h,i
V2 C5339 ± 151 b980 ± 44 a
V33328 ± 164 f236 ± 11 j,k
V4 C1864 ± 76 m114 ± 7 l
V4 R2310 ± 47 k187 ± 7 k
V5 F2556 ± 121 j350 ± 13 g,h
V5 R2218 ± 111 l599 ± 10 d
V63022 ± 103 g284 ± 8 i,j
V7 P1816 ± 91 m385 ± 30 g
V7 F4035 ± 157 d452 ± 12 f
V82915 ± 133 h,i35 ± 3 m
V9 C4149 ± 192 c254 ± 13 i,j
V9 R2981 ± 84 g,h520 ± 17 e
V105694 ± 129 a255 ± 8 i,j
V113561 ± 161 e792 ± 31 b
Trolox139 ± 5 21.8 ± 0.2
1 IC50: extract concentration that inhibits oxidation by 50%; means of the same column followed by different Latin letters are statistically different according to Tukey’s honestly significant difference (HSD) test at p < 0.05. T: turbinate; C: cylindrical; R: round; F: flattened.
Table 7. Antimicrobial and antifungal activity of the flesh of different pumpkin varieties from Greece (mg/mL).
Table 7. Antimicrobial and antifungal activity of the flesh of different pumpkin varieties from Greece (mg/mL).
V1V2 TV2 CV3V4 CV4 RV5 FV5 RV6V7 P
Gram-negative bacteria
Enterobacter cloacae>10>1010>10>10>10>10>10>10>1010>1010>10>10>1010>10>10>10
Escherichia coli>10>1010>1010>1010>1010>1010>1010>1010>105>10>10>10
Pseudomonas aeruginosa>10>10>10>10>10>10>10>10>10>10>10>10>10>10>10>10>10>10>10>10
Salmonella enterocolitica>10>105>1010>1010>1010>1010>1010>1010>105>10>10>10
Yersinia enterocolitica10>10>10>1010>10>10>1010>10>10>10>10>10>10>10>10>1010>10
Gram-positive bacteria
Bacillus cereus>10>10>10>10>10>10>10>10>10>10>10>10>10>10>10>10>10>10>10>10
Listeria monocytogenes>10>10>10>10>10>10>10>10>10>10>10>10>10>10>10>10>10>10>10>10
Staphylococcus aureus10>1010>1010>10>10>1010>10>10>1010>1010>10>10>1010>10
V7 FV8V9 CV9 RV10V11StreptomycinMethicilinAmpicillin
Gram-negative bacteria
Enterobacter cloacae10>10>10>105>10>10>105>1010>100.0070.007n.t.n.t0.150.15
Escherichia coli>10>105>1010>10>10>10>10>1010>100.010.01n.t.n.t.0.150.15
Pseudomonas aeruginosa>10>10>10>10>10>10>10>1010>10>10>100.060.06n.t.n.t.0.630.63
Salmonella enterocolitica10>1010>10>10>10>10>10>10>1010>100.0070.007n.t.n.t.0.150.15
Yersinia enterocolitica2.5>10>10>10>10>10>10>105>10>10>100.0070.007n.t.n.t.0.150.15
Gram-positive bacteria
Bacillus cereus>10>10>10>10>10>10>10>10>10>10>10>100.0070.007n.t.n.t.n.t.n.t.
Listeria monocytogenes>10>10>10>10>10>10>10>10>10>10>10>100.0070.007n.t.n.t.0.150.15
Staphylococcus aureus10>1010>10>10>10>10>1010>1010>100.0070.0070.0070.0070.150.15
V1V2 TV2 CV3V4 CV4 RV5 FV5 RV 6
Aspergillus brasiliensis>10>10>10>10>10>1010>10>10>1010>10>10>10>10>1010>10
Aspergillus fumigatus>10>10>10>10>10>1010>10>10>1010>10>10>10>10>10>10>10
V7 PV7 FV8V9 CV9 RV10V11Ketoconazole
Aspergillus brasiliensis10>1010>1010>1010>1010>1010>1010>100.060.125
Aspergillus fumigatus>10>10>10>1010>10>10>10>10>1010>10>10>100.51
MIC: minimum inhibitory concentration; MBC: minimal bactericidal concentration; MFC: minimal fungicidal concentration; n.t: not tested; T: turbinate; C: cylindrical; R: round; F: flattened.3.3.3. Cytotoxicity.
Table 8. Quantification of the organic acids in the flesh of the studied pumpkin genotypes.
Table 8. Quantification of the organic acids in the flesh of the studied pumpkin genotypes.
g/100 g dw
g/100 g dw
g/100 g dw
g/100 g dw
g/100 g dw
mg/100 g dw
V13.61 ± 0.04 f2.7 ± 0.1 jtrtr1.08 ± 0.02 d0.0091 ± 0.0001 m
V2 T4.45 ± 0.04 e3.96 ± 0.06 f,gn.d.trn.d.0.02825 ±
0.00007 g
V2 C4.8 ± 0.2 d4.1 ± 0.2 d,e,fn.d.trn.d.0.034 ± 0.001 e
V35.50 ± 0.01 b4.0 ± 0.2 e,f,gn.d.n.d.n.d.0.0169 ± 0.0007 k,l
V4 C4.5 ± 0.1 e4.9 ± 0.2 bn.d.n.d.n.d.0.0255 ± 0.0008 h
V4 R5.17 ± 0.03 c5.38 ± 0.07 an.d.n.d.n.d.0.0193 ± 0.0009 i,j
V5 F5.301 ± 0.006 c4.28 ± 0.03 c,d,en.d.tr3.16 ± 0.09 a0.0155 ± 0.0002 l
V5 R1.80 ± 0.03 h3.14 ± 0.07 itrtrn.d.0.031 ± 0.002 f
V63.641 ± 0.006 f5.16 ± 0.06 a,bn.d.n.d.n.d.0.0943 ± 0.0002 a
V7 P3.75 ± 0.07 f4.2 ± 0.1 c,d,e,ftrtr1.38 ± 0.06 c0.021 ± 0.001 i
V7 F3.64 ± 0.02 f4.3 ± 0.2 c,d,e,ftrtr1.62 ± 0.01 b0.0161 ± 0.0003 l
V84.77 ± 0.01 d3.48 ± 0.05 hn.d.trn.d.0.069 ± 0.003 b
V9 C6.4 ± 0.1 a3.7 ± 0.2 g,hn.d.n.d.3.2 ± 0.1 an.d.
V9 R3.40 ± 0.04 g4.3 ± 0.2 c,dtrn.d.n.d.0.0188 ± 0.0009 j,k
V104.79 ± 0.05 d5.2 ± 0.2 a,bn.d.n.d.n.d.0.0393 ± 0.0004 d
V114.36 ± 0.09 e4.50 ± 0.06 cn.d.n.d.n.d.0.061 ± 0.001 c
n.d.: not detected. tr: traces. Means of the same column followed by different Latin letters are statistically different according to Tukey’s honestly significant difference (HSD) test at p < 0.05. T: turbinate; C: cylindrical; R: round; F: flattened.
Table 9. Tocopherol content of the extracts of the studied pumpkin genotypes of pumpkin flesh (mg/100 g dw).
Table 9. Tocopherol content of the extracts of the studied pumpkin genotypes of pumpkin flesh (mg/100 g dw).
Genotypeα-Tocopherolβ-Tocopherolγ-TocopherolTotal Tocopherols
V11.8 ± 0.1 g0.1496 ± 0.0005 dn.d.1.9 ± 0.1 h
V2 T4.90 ± 0.08 a6.59 ± 0.08 an.d.11.5 ± 0.2 a
V2 C0.81 ± 0.03 k0.238 ± 0.003 cn.d.1.05 ± 0.03 k
V31.57 ± 0.02 hn.d.1.05 ± 0.02 d2.62 ± 0.04 f
V4 C1.15 ± 0.03 i,jn.d.0.1998 ± 0.0002 g,h1.35 ± 0.03 j
V4 R2.68 ± 0.06 cn.d.1.619 ± 0.005 b4.30 ± 0.06 d
VP F2.3 ± 0.1 e0.85 ± 0.01 bn.d.3.19 ± 0.08 e
V5 R1.03 ± 0.02 jn.d.1.02 ± 0.03 d2.05 ± 0.04 h
V62.47 ± 0.01 dn.d.4.5 ± 0.1 a7.0 ± 0.1 b
V7 P0.70 ± 0.04 k0.043 ± 0.001 e0.185 ± 0.004 h0.93 ± 0.04 k
V7 F0.218 ± 0.008 mn.d.0.2685 ± 0.0007 f,g0.486 ± 0.009 l
V81.16 ± 0.04 jn.d.0.58 ± 0.02 e1.73 ± 0.02 i
V9 C1.70 ± 0.09 gn.d.0.572 ± 0.003 e2.3 ± 0.1 g
V9 R2.91 ± 0.06 bn.d.1.24 ± 0.03 c4.15 ± 0.04 d
V101.97 ± 0.09 fn.d.0.28 ± 0.01 f2.3 ± 0.1 g
V110.55 ± 0.01 ln.d.4.52 ± 0.07 a5.07 ± 0.08 c
n.d.: not detected. Means of the same column followed by different Latin letters are statistically different according to Tukey’s honestly significant difference (HSD) test at p < 0.05. T: turbinate; C: cylindrical; R: round; F: flattened.
Disclaimer/Publisher’s Note: The statements, opinions and data contained in all publications are solely those of the individual author(s) and contributor(s) and not of MDPI and/or the editor(s). MDPI and/or the editor(s) disclaim responsibility for any injury to people or property resulting from any ideas, methods, instructions or products referred to in the content.

Share and Cite

MDPI and ACS Style

Leichtweis, M.G.; Molina, A.K.; Dias, M.I.; Calhelha, R.C.; Pires, T.C.S.P.; Pavli, O.; Oliveira, M.B.P.P.; Petropoulos, S.A.; Barros, L.; Pereira, C. Variability in Chemical Profile and Bioactivities of the Flesh of Greek Pumpkin Landraces. Horticulturae 2023, 9, 1232.

AMA Style

Leichtweis MG, Molina AK, Dias MI, Calhelha RC, Pires TCSP, Pavli O, Oliveira MBPP, Petropoulos SA, Barros L, Pereira C. Variability in Chemical Profile and Bioactivities of the Flesh of Greek Pumpkin Landraces. Horticulturae. 2023; 9(11):1232.

Chicago/Turabian Style

Leichtweis, Maria G., Adriana K. Molina, Maria Inês Dias, Ricardo C. Calhelha, Tânia C. S. P. Pires, Ourania Pavli, M. Beatriz P. P. Oliveira, Spyridon A. Petropoulos, Lillian Barros, and Carla Pereira. 2023. "Variability in Chemical Profile and Bioactivities of the Flesh of Greek Pumpkin Landraces" Horticulturae 9, no. 11: 1232.

Note that from the first issue of 2016, this journal uses article numbers instead of page numbers. See further details here.

Article Metrics

Back to TopTop