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Exploring wild edible flowers as a source of bioactive compounds: New perspectives in horticulture


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Figure 1

RACI calculated for the 26 studied flower species. Different lower case letters indicate significant differences between species according to Kruskal–Wallis’ stepwise comparison (p < 0.05). RACI, relative antioxidant capacity index. §Flowers sampled from cultivated plants.
RACI calculated for the 26 studied flower species. Different lower case letters indicate significant differences between species according to Kruskal–Wallis’ stepwise comparison (p < 0.05). RACI, relative antioxidant capacity index. §Flowers sampled from cultivated plants.

Figure 2

Total polyphenols and polyphenol classes (flavonols, flavanols, benzoic acids and cinnamic acids) content (mg · 100 g−1) in the flowers of the 26 studied species. Different lower case letters in a row indicate significant differences between species according to Tukey’s post-hoc test (p < 0.05). -, compound not detected. §Flowers sampled from cultivated plants.
Total polyphenols and polyphenol classes (flavonols, flavanols, benzoic acids and cinnamic acids) content (mg · 100 g−1) in the flowers of the 26 studied species. Different lower case letters in a row indicate significant differences between species according to Tukey’s post-hoc test (p < 0.05). -, compound not detected. §Flowers sampled from cultivated plants.

Figure 3

Hierarchical cluster analysis of the selected species according to their TPC and antioxidant activity (FRAP, DPPH and ABTS). The respective botanical family and vegetation community is provided for every species. §Flowers sampled from cultivated plants. ABTS, 2,2′-azino-bis-3-ethylbenzthiazoline-6-sulfonic acid; DPPH, 2,2-diphenyl-1-picrylhydrazyl; FRAP, ferric reducing antioxidant power; TPC, total polyphenol content.
Hierarchical cluster analysis of the selected species according to their TPC and antioxidant activity (FRAP, DPPH and ABTS). The respective botanical family and vegetation community is provided for every species. §Flowers sampled from cultivated plants. ABTS, 2,2′-azino-bis-3-ethylbenzthiazoline-6-sulfonic acid; DPPH, 2,2-diphenyl-1-picrylhydrazyl; FRAP, ferric reducing antioxidant power; TPC, total polyphenol content.

Figure 4

Hierarchical cluster analysis of the selected species according to their polyphenolic profile and vitamin C content. The respective botanical family and vegetation community is provided for every species. §Flowers sampled from cultivated plants.
Hierarchical cluster analysis of the selected species according to their polyphenolic profile and vitamin C content. The respective botanical family and vegetation community is provided for every species. §Flowers sampled from cultivated plants.

Figure A1

Flavonols (hyperoside, isoquercitrin, quercetin, quercitrin and rutin) content (mg · 100 g−1) in the flowers of the 26 studied species. Different lower case letters in a row indicate significant differences between species according to Tukey’s post-hoc test (p < 0.05). Data are means of three biological replicates. -, compound not detected. §Flowers sampled from cultivated plants.
Flavonols (hyperoside, isoquercitrin, quercetin, quercitrin and rutin) content (mg · 100 g−1) in the flowers of the 26 studied species. Different lower case letters in a row indicate significant differences between species according to Tukey’s post-hoc test (p < 0.05). Data are means of three biological replicates. -, compound not detected. §Flowers sampled from cultivated plants.

Figure A2

Flavanols (catechin and epicatechin) content (mg · 100 g−1) in the flowers of the 26 studied species. Different lower case letters in a row indicate significant differences between species according to Tukey’s post-hoc test (p < 0.05). Data are means of three biological replicates. -, compound not detected. §Flowers sampled from cultivated plants.
Flavanols (catechin and epicatechin) content (mg · 100 g−1) in the flowers of the 26 studied species. Different lower case letters in a row indicate significant differences between species according to Tukey’s post-hoc test (p < 0.05). Data are means of three biological replicates. -, compound not detected. §Flowers sampled from cultivated plants.

Figure A3

Benzoic acids (ellagic and gallic acid) content (mg · 100 g−1) in the flowers of the 26 studied species. Different lower case letters in a row indicate significant differences between species according to Tukey’s post-hoc test (p < 0.05). Data are means of three biological replicates. -, compound not detected. §Flowers sampled from cultivated plants.
Benzoic acids (ellagic and gallic acid) content (mg · 100 g−1) in the flowers of the 26 studied species. Different lower case letters in a row indicate significant differences between species according to Tukey’s post-hoc test (p < 0.05). Data are means of three biological replicates. -, compound not detected. §Flowers sampled from cultivated plants.

Figure A4

Cinnamic acids (caffeic, chlorogenic, coumaric and ferulic acid) content (mg · 100 g−1) in the flowers of the 26 studied species. Different lower case letters in a row indicate significant differences between species according to Tukey’s post-hoc test (p < 0.05). Data are means of three biological replicates. -, compound not detected. §Flowers sampled from cultivated plants.
Cinnamic acids (caffeic, chlorogenic, coumaric and ferulic acid) content (mg · 100 g−1) in the flowers of the 26 studied species. Different lower case letters in a row indicate significant differences between species according to Tukey’s post-hoc test (p < 0.05). Data are means of three biological replicates. -, compound not detected. §Flowers sampled from cultivated plants.

Vitamin C content (mg · 100 g−1) in the flowers of the 26 studied species.

Species Vitamin C
Allium ursinum 0.0 -
Bellis perennis 4.4 fi
Centaurea cyanus 3.3 gi
Cichorium intybus 4.0 gi
Dianthus carthusianorum 5.5 di
Dianthus pavonius 16.4 bc
Erythronium dens-canis 6.7 ch
Geranium sylvaticum 7.9 bg
Lavandula angustifolia 2.8 hi
Leucanthemum vulgare 5.9 di
Mentha aquatica 2.6 i
Paeonia officinalis 11.3 be
Primula veris 44.9 a
Primula vulgaris 3.8 gi
Robinia pseudoacacia 4.0 gi
Rosa canina 12.3 bd
Rosa pendulina 7.2 bg
Salvia pratensis 4.0 gi
Sambucus nigra 11.0 bf
Taraxacum officinale 3.5 gi
Trifolium alpinum 15.5 bc
Viola odorata 4.6 ei
Borago officinalis§ 0.0 -
Calendula officinalis§ 11.8 bd
Tagetes patula§ 7.2 bg
Tropaeolum majus§ 17.7 b

List of the 22 wild and 4 cultivated species collected and analysed in the study, with references on their botanical family, vegetation community, food and/or medicinal uses, month, site (WGS84/32N system) and soil (USDA soil taxonomy; the bedrock type is provided in brackets: C, calcareous; S, siliceous)* of sampling in 2017.

Species Botanical family Vegetation community Reference of flower use
Flower sampling
Soil type
Food Medicinal Month Longitude Latitude
Allium ursinum L. Amaryllidaceae Woodlands Lim (2014a) Sobolewska et al. (2015) April 7.878 45.490 Eutrudept (S)
Bellis perennis L. Asteraceae Nutrient rich grasslands Lim (2014a) Lim (2014a) March 7.592 45.065 Hapludalf (S)
Centaurea cyanus L.** Asteraceae Ruderal communities Lim (2014a) Lim (2014a) May 7.896 45.475 Eutrudept (S)
Cichorium intybus L. Asteraceae Ruderal communities Lim (2014a) Street et al. (2013) June 7.843 45.192 Dystrudept (S)
Dianthus carthusianorum L. Caryophyllaceae Dry grasslands - Palma (1964) June 7.219 45.301 Udifluvent (S)
Dianthus pavonius Tausch Caryophyllaceae Nutrient poor grasslands - - July 7.122 44.391 Eutrocryept (C)
Erythronium dens-canis L. Liliaceae Woodlands - - March 7.371 45.157 Dystrudept (C)
Geranium sylvaticum L. Geraniaceae Edges - - June 7.186 45.300 Dystrudept (S)
Lavandula angustifolia Mill.** Lamiaceae Dry grasslands Lim (2014b) Lim (2014b) June - - -
Leucanthemum vulgare Lam. Asteraceae Nutrient rich grasslands Lim (2014a) Prinsloo et al. (2018) April 7.592 45.065 Hapludalf (S)
Mentha aquatica L. Lamiaceae Wetlands Lim (2014b) Alvarado (2018) September 7.485 45.120 Hapludalf (S)
Paeonia officinalis L.** Paeoniaceae Edges Lim (2014b) - April 7.344 45.097 Udorthent (S)
Primula veris L. Primulaceae Woodlands Lim (2014b) Apel et al. (2017) May 6.802 44.968 Eutrudept (C)
Primula vulgaris Huds. Primulaceae Woodlands Lim (2014b) Tuttolomondo et al. (2014) March 7.379 45.145 Dystrudept (C)
Robinia pseudoacacia L. Fabaceae Woodlands Lim (2014a) Jarić et al. (2015) May 7.593 45.065 Hapludalf (S)
Rosa canina L. ** Rosaceae Shrublands Lim (2014b) Nemati et al. (2015) May 7.677 45.715 Udorthent (S)
Rosa pendulina L. Rosaceae Woodlands - - June 7.192 45.301 Dystrudept (S)
Salvia pratensis L. Lamiaceae Dry grasslands Kucekova et al. (2013) Kucekova et al. (2013) May 7.603 45.036 Udifluvent (S)
Sambucus nigra L. Adoxaceae Shrublands Lim (2014a) Młynarczyk et al. (2018) May 7.593 45.064 Hapludalf (S)
Taraxacum officinale Weber** Asteraceae Nutrient rich grasslands Lim (2014a) Lim (2014a) March 7.593 45.064 Hapludalf (S)
Trifolium alpinum L. Fabaceae Nutrient poor grasslands Abbet et al. (2014) Agelet and Vallès (2001) July 7.122 44.390 Eutrocryept (C)
Viola odorata L. Violaceae Ruderal communities Lim (2014b) Lim (2014b) March 7.591 45.065 Hapludalf (S)

Borago officinalis L. Boraginaceae Cultivated Lim (2014a) Gupta and Singh (2010) May -§ -§ -§
Calendula officinalis L. Asteraceae Cultivated Lim (2014a) Lim (2014a) May -§ § -§
Tagetes patula L. Asteraceae Cultivated Lim (2014a) Lim (2014a) July -§ § -§
Tropaeolum majus L. Tropaeolaceae Cultivated Lim (2014b) Lim (2014b) May -§ § -§

Flavonols and flavanols content (mg · 100 g−1) in the flowers of the 26 studied species.

Species Flavonols Flavanols


Hyperoside Isoquercitrin Quercetin Quercitrin Rutin Catechin Epicatechin
Allium ursinum 38.7 b 0.0 - 0.0 - 0.9 c 19.7 b 0.0 - 20.8 g
Bellis perennis 0.0 - 0.0 - 0.0 - 0.0 - 0.0 - 0.4 d 421.0 a
Centaurea cyanus 0.0 - 0.0 - 0.0 - 0.9 c 0.0 - 0.0 - 65.2 c
Cichorium intybus 23.5 bc 16.1 c 0.0 - 0.0 - 13.7 b 19.1 bc 533.3 a
Dianthus carthusianorum 262.4 a 163.6 b 0.0 - 8.0 c 17.8 b 0.0 - 0.1 h
Dianthus pavonius 0.0 - 2,072.0 a 0.0 - 163.6 c 34.1 b 11.7 c 26.3 dg
Erythronium dens-canis 9.0 cd 0.0 - 0.0 - 108.5 c 0.5 c 0.4 d 29.9 cg
Geranium sylvaticum 0.6 e 12.5 c 189.0 a 34.5 c 17.1 b 20.7 bc 37.7 cg
Lavandula angustifolia 17.3 bd 0.0 - 207.3 a 0.0 - 0.0 - 375.6 a 306.6 ab
Leucanthemum vulgare 0.0 - 0.0 - 328.1 a 0.0 - 0.0 - 0.0 - 0.1 h
Mentha aquatica 0.0 - 12.0 c 227.2 a 0.0 - 16.8 b 24.0 bc 59.2 cd
Paeonia officinalis 0.0 - 0.0 - 216.3 a 16.4 c 0.0 - 28.6 b 30.7 cg
Primula veris 14.7 bd 10.8 c 0.0 - 82.3 c 18.1 b 0.4 d 52.0 ce
Primula vulgaris 9.4 cd 100.1 b 0.0 - 109.6 c 107.7 a 0.0 - 23.9 eg
Robinia pseudoacacia 0.0 - 0.0 - 0.0 - 547.3 b 0.0 - 0.0 - 22.0 fg
Rosa canina 38.5 b 130.2 b 205.7 a 35.9 c 0.0 - 26.0 b 158.1 b
Rosa pendulina 15.3 bd 184.5 b 0.0 - 1,353.4 a 12.9 b 0.4 d 388.4 b
Salvia pratensis 0.0 - 0.0 - 0.0 - 0.0 - 0.0 - 0.0 - 25.0 eg
Sambucus nigra 0.0 - 0.0 - 0.0 - 23.4 c 0.0 - 0.0 - 48.3 cf
Taraxacum officinale 0.6 e 0.0 - 0.7 b 0.0 - 0.5 c 0.0 - 0.1 h
Trifolium alpinum 7.4 d 7.9 c 0.0 - 13.5 c 0.5 c 0.0 - 0.1 h
Viola odorata 16.3 bd 15.7 c 0.0 - 10.8 c 14.4 b 0.0 - 21.9 fg
Borago officinalis§ 0.0 - 0.0 - 0.0 - 0.0 - 0.0 - 0.0 - 0.0 -
Calendula officinalis§ 0.0 - 0.0 - 0.0 - 1.7 c 0.0 - 0.4 d 0.1 h
Tagetes patula§ 0.0 - 0.0 - 0.0 - 0.0 - 0.5 c 0.0 - 29.3 cg
Tropaeolum majus§ 0.0 - 0.0 - 0.0 - 619.6 b 0.0 - 0.0 - 49.1 cf

Mobile phases, elution conditions and wavelength used to detect the five classes of compounds with HPLC analysis.

Class of compounds Mobile phase Elution conditions Wavelength (nm)
Cinnamic acids and flavonols A: 10 mM KH2PO4/H3PO4, pH = 2.8B: CH3CN 5%B to 21%B in 17 min + 21%B in 3 min (2 min conditioning time); flow: 1.5 mL min−1 330

Benzoic acids and flavanols A: H2O/CH3OH/HCOOH(5:95:0.1 v/v/v), pH = 2.5B: CH3OH/HCOOH (100:0.1 v/v) 3%B to 85%B in 22 min + 85%B in 1 min (2 min conditioning time); flow: 0.6 mL min−1 280

Vitamin C A: 5 mM C16H33N(CH3)3Br/50 mMKH2PO4, pH = 2.5B: CH3OH Isocratic, ratio of phase A and B: 95:5 in 10 min (5 min conditioning time); flow: 0.9 mL min−1 261, 348

Dry matter, TPC and antioxidant activity (FRAP, DPPH and ABTS assays) in the 26 edible flowers.

Species Dry matter (%) TPC (mg GAE · 100 g−1) Antioxidant activity

FRAP (mmol Fe2+ · kg−1) DPPH (mmol TE · g−1) ABTS (mmol TE · g−1)
Allium ursinum 10.2 jk 184.4 k 4.2 j 7.6 kl 0.7 m
Bellis perennis 16.9 ef 396.3 gi 81.6 cf 24.3 i 13.4 hj
Centaurea cyanus 26.4 b 378.5 hi 68.3 df 23.6 i 17.8 fh
Cichorium intybus 17.3 ef 618.4 df 138.4 ad 69.2 f 26.9 d
Dianthus carthusianorum 27.8 ab 936.3 bd 222.2 ab 81.1 ef 33.6 c
Dianthus pavonius 21.2 cd 752.8 ce 176.1 ac 106.5 c 24.3 de
Erythronium dens-canis 15.1 fh 364.3 hi 53.5 eg 20.4 ik 14.4 hj
Geranium sylvaticum 12.8 i 1,267.8 ab 267.0 ab 152.9 b 55.2 a
Lavandula angustifolia 31.2 a 396.0 gi 89.5 ce 14.8 il 14.0 hj
Leucanthemum vulgare 17.0 ef 448.8 fi 44.3 eh 20.9 ik 10.8 ij
Mentha aquatica 22.3 c 1,061.7 bc 256.0 ab 86.7 de 42.5 b
Paeonia officinalis 13.9 gi 1,930.5 a 303.8 a 226.2 a 55.3 a
Primula veris 18.8 de 1,044.9 bc 230.1 ab 97.1 cd 38.5 bc
Primula vulgaris 9.8 jk 602.9 dg 127.4 bd 41.7 gh 21.5 df
Robinia pseudoacacia 12.9 i 203.8 jk 15.7 i 4.5 l 2.4 m
Rosa canina 16.7 ef 1,396.6 ab 257.5 ab 146.2 b 55.6 a
Rosa pendulina 21.6 cd 1,773.7 a 253.8 ab 154.3 b 55.7 a
Salvia pratensis 17.7 ef 314.7 ij 38.9 fh 8.9 jl 9.0 jl
Sambucus nigra 16.8 ef 508.7 eh 78.8 cf 28.5 hi 18.3 fh
Taraxacum officinale 16.5 ef 159.4 k 13.0 i 7.7 kl 3.3 lm
Trifolium alpinum 10.0 jk 464.6 fi 91.5 ce 50.3 gh 20.3 eg
Viola odorata 13.0 hi 428.4 fi 66.1 dg 22.6 ij 15.6 gi
Borago officinalis§ 15.3 fg 163.4 k 29.7 gi 22.8 i 3.7 km
Calendula officinalis§ 13.7 gi 189.6 k 22.6 hi 3.6 l 9.2 jk
Tagetes patula§ 10.7 j 470.8 fi 143.9 ad 44.1 gh 23.0 df
Tropaeolum majus§ 8.9 k 355.8 hi 45.3 eh 14.8 il 12.8 hj
p *** *** *** *** ***

Benzoic acids and cinnamic acids content (mg · 100 g−1) in 26 flower species.

Species Benzoic acids Cinnamic acids


Ellagic acid Gallic acid Caffeic acid Chlorogenic acid Coumaric acid Ferulic acid
Allium ursinum 15.2 h 0.1 c 0.0 - 0.0 - 0.5 c 357.3 a
Bellis perennis 212.9 ad 0.0 - 15.6 ab 0.0 - 158.5 a 0.0 -
Centaurea cyanus 180.1 ad 0.0 - 0.1 d 0.0 - 0.0 - 0.0 -
Cichorium intybus 23.7 fh 27.5 b 0.0 - 230.0 b 148.5 a 0.0 -
Dianthus carthusianorum 278.5 ac 27.6 b 14.6 ab 0.0 - 0.0 - 0.0 -
Dianthus pavonius 214.4 ad 0.0 - 0.0 - 0.0 - 0.0 - 0.0 -
Erythronium dens-canis 20.4 gh 0.0 - 16.2 a 275.5 a 110.7 a 20.9 bc
Geranium sylvaticum 121.4 be 0.0 - 0.0 - 244.1 b 16.4 b 0.0 -
Lavandula angustifolia 122.8 be 0.0 - 11.7 c 0.2 c 0.0 - 0.0 -
Leucanthemum vulgare 63.2 dg 0.0 - 0.0 - 0.0 - 0.0 - 0.0 -
Mentha aquatica 153.8 bd 0.0 - 15.9 ab 270.6 a 113.8 a 0.0 -
Paeonia officinalis 589.2 a 244.2 a 13.9 b 0.0 - 0.0 - 32.6 bc
Primula veris 27.9 fh 0.1 c 15.2 ab 0.0 - 0.0 - 43.9 b
Primula vulgaris 16.3 h 0.0 - 16.3 a 0.0 - 0.0 - 29.3 bc
Robinia pseudoacacia 79.5 cf 0.0 - 0.0 - 0.0 - 0.0 - 0.0 -
Rosa canina 72.5 dg 0.1 c 0.0 - 232.0 b 0.0 - 0.0 -
Rosa pendulina 410.7 ab 0.0 - 0.0 - 0.0 - 0.0 - 0.0 -
Salvia pratensis 20.4 gh 0.1 c 0.0 - 0.0 - 0.0 - 0.0 -
Sambucus nigra 27.1 fh 0.1 c 15.3 ab 0.0 - 112.0 a 0.0 -
Taraxacum officinale 28.2 fh 0.0 - 15.7 ab 273.3 a 0.0 - 0.0 -
Trifolium alpinum 34.6 eh 0.0 - 0.0 - 0.0 - 0.5 c 0.2 c
Viola odorata 26.2 fh 0.0 - 1.9 d 0.0 - 113.4 a 0.2 c
Borago officinalis§ 0.1 i 27.9 b 0.0 - 0.0 - 0.0 - 0.0 -
Calendula officinalis§ 15.2 h 0.0 - 0.0 - 0.0 - 0.0 - 0.0 -
Tagetes patula§ 150.1 bd 0.0 - 0.0 - 0.0 - 0.0 - 0.0 -
Tropaeolum majus§ 214.3 ad 0.0 - 14.1 b 241.2 b 109.6 a 0.0 -

Spearman’s correlation indexes between TPC, antioxidant activity (FRAP, DPPH and ABTS assays) and phenolic compounds recorded in the 26 edible flowers.

FRAP DPPH ABTS Caffeic acid Chlorogenic acid Coumaric acid Ferulic acid Hyperoside Isoquercitrin Quercetin Quercitrin Rutin Ellagic Acid Gallic Acid Catechin Epicatechin Vitamin C
TPC 0.94** 0.93** 0.95** 0.07 0.06 0.05 0.09 0.26* 0.68** 0.43** 0.36** 0.43** 0.47** 0.18 0.50** 0.43** 0.44**
FRAP 1 0.93** 0.96** 0.09 0.09 0.06 0.04 0.23* 0.65** 0.40** 0.26* 0.40** 0.49** 0.21 0.56** 0.49** 0.37**
DPPH 1 0.93** 0.01 0.07 0.01 0.09 0.22 0.68** 0.33** 0.30** 0.42** 0.40** 0.31** 0.49** 0.37** 0.41**
ABTS 1 0.07 0.15 0.07 0.05 0.25* 0.67** 0.36** 0.29** 0.38** 0.47** 0.19 0.52** 0.47** 0.44**
Caffeic acid 1 0.30** 0.44** 0.35** 0.04 −0.01 −0.03 0.08 0.12 0.02 −0.03 −0.07 0.11 −0.05
Chlorogenic acid 1 0.49** −0.07 −0.03 0.04 0.22* 0.06 0.01 −0.04 −0.06 0.35** 0.21 −0.01
Coumaric acid 1 −0.12 −0.08 −0.07 −0.04 0.02 0.05 −0.05 −0.06 0.09 0.40** −0.02
Ferulic acid 1 0.30** −0.03 −0.03 0.17 0.32** −0.22* 0.08 −0.08 −0.05 0.00
Hyperoside 1 0.48** −0.06 0.03 0.44** −0.17 0.14 0.11 0.13 −0.05
Isoquercitrin 1 0.03 0.38** 0.69** 0.17 0.01 0.27* 0.14 0.30**
Quercetin 1 −0.09 −0.08 0.29* 0.06 0.70** 0.20 0.02
Quercitrin 1 0.23 0.20 −0.23* 0.01 0.26* 0.50**
Rutin 1 −0.04 0.04 0.14 0.07 −0.05
Ellagic acid 1 0.06 0.27* 0.34** 0.28*
Gallic acid 1 0.23* −0.09 −0.12
Catechin 1 0.49** 0.05
Epicatechin 1 0.04
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