Citation: Andrea Ertani, Ornella Francioso, Serenella Nardi. Mini review: fruit residues as plant biostimulants for bio-based product recovery[J]. AIMS Agriculture and Food, 2017, 2(3): 251-257. doi: 10.3934/agrfood.2017.3.251
[1] | Albert Nugraha, Asadin Briliantama, M Umar Harun, Li Sing-Chung, Chin Xuan Tan, Vuanghao Lim, Amir Husni, Widiastuti Setyaningsih . Ultrasound-assisted extraction of phenolic compounds from ear mushrooms (Auricularia auricula-judae): Assessing composition and antioxidant activity during fruiting body development. AIMS Agriculture and Food, 2024, 9(4): 1134-1150. doi: 10.3934/agrfood.2024059 |
[2] | Quartieri Maurizio, Sorrenti Giovambattista, Ciriani Alessandro, Baldi Elena, Collina Marina and Toselli Moreno . Combining quince (Cydonia oblonga) rootstock with soil-applied calcium chloride solution as a strategy to control brown spot (Stemphylium vesicarium) incidence in Abbé Fétel pear fruits. AIMS Agriculture and Food, 2019, 4(2): 414-428. doi: 10.3934/agrfood.2019.2.414 |
[3] | Evi Mintowati Kuntorini, Laurentius Hartanto Nugroho, Maryani, Tri Rini Nuringtyas . Maturity effect on the antioxidant activity of leaves and fruits of Rhodomyrtus tomentosa (Aiton.) Hassk.. AIMS Agriculture and Food, 2022, 7(2): 282-296. doi: 10.3934/agrfood.2022018 |
[4] | Alexandros Tsoupras, Eirini Panagopoulou, George Z. Kyzas . Olive pomace bioactives for functional foods and cosmetics. AIMS Agriculture and Food, 2024, 9(3): 743-766. doi: 10.3934/agrfood.2024040 |
[5] | Anthony Temitope Idowu, Oluwakemi Osarumwense Igiehon, Ademola Ezekiel Adekoya, Solomon Idowu . Dates palm fruits: A review of their nutritional components, bioactivities and functional food applications. AIMS Agriculture and Food, 2020, 5(4): 734-755. doi: 10.3934/agrfood.2020.4.734 |
[6] | Stefano Puccio, Anna Perrone, Giuseppe Sortino, Giuseppe Gianguzzi, Carla Gentile, Vittorio Farina . Yield, pomological characteristics, bioactive compounds and antioxidant activity of Annona cherimola Mill. grown in mediterranean climate. AIMS Agriculture and Food, 2019, 4(3): 592-603. doi: 10.3934/agrfood.2019.3.592 |
[7] | Thornthan Sawangwan, Chompoonuth Porncharoennop, Harit Nimraksa . Antioxidant compounds from rice bran fermentation by lactic acid bacteria. AIMS Agriculture and Food, 2021, 6(2): 578-587. doi: 10.3934/agrfood.2021034 |
[8] | Thi Thuy Le, Trung Kien Nguyen, Nu Minh Nguyet Ton, Thi Thu Tra Tran, Van Viet Man Le . Quality of cookies supplemented with various levels of turmeric by-product powder. AIMS Agriculture and Food, 2024, 9(1): 209-219. doi: 10.3934/agrfood.2024012 |
[9] | Sana Hayyat, Mehdi Maqbool, Abdul Hamid, Muhammad Shehzad, Raheel Anwar, Sandra Horvitz, Noosheen Zahid, Muhammad Azam Khan . Appropriate harvest maturity for exploitation of wild black raspberry (Rubus sp.) fruits during shelf life period from Rawalakot, Azad Jammu and Kashmir. AIMS Agriculture and Food, 2025, 10(1): 1-16. doi: 10.3934/agrfood.2025001 |
[10] | Enrica Bargiacchi, Margherita Campo, Annalisa Romani, Gilberto Milli, Sergio Miele . Hydrolysable Tannins from Sweet Chestnut (Castanea sativa Mill.) to improve Tobacco and Food/Feed Quality. Note 1: Fraction characterization, and Tobacco biostimulant effect for gall-nematode resistance. AIMS Agriculture and Food, 2017, 2(3): 324-338. doi: 10.3934/agrfood.2017.3.324 |
Fruit juices and derived products have experienced a growing consumption during the last years due to the positive effects of their ingredients on human health. Indeed, consumers are increasingly becoming aware of health problems related to food intake, therefore demanding health-promoting natural products.
Plant food processing derived by-products (agro-industrial residues) represent a disposal problem for the companies, but they are also promising sources of bioactive compounds, including antioxidants [1,3], which may be employed in agriculture by farmers to foster crop performance [4]. Therefore, agro-industrial residues can be referred to as biostimulant compounds.
According to the European Biostimulants Industry Council [5], biostimulants are defined as products containing substances and/or microorganisms whose function when applied to plants or the rhizosphere is to stimulate natural processes, to enhance/benefit nutrient uptake, nutrient use efficiency, tolerance to abiotic stress, and crop quality. Biostimulants are of remarkable importance for sustainable agriculture as they offer an innovative solution to an increased world demand for high crop productivity under low unsustainable inputs [6]. The sale-market in Europe in particular, has been estimated to grow up to $2 billions by 2018 [7,8]. The European Biostimulants Industry Council [9] does not recognize any direct action of biostimulants against pests; therefore, biostimulants do not fall within the regulatory framework of pesticides. Also, they must operate in plants through different mechanisms than fertilizers, regardless of the presence of nutrients in the products [9]. In this respect, the Biostimulant Coalition in North America specifies that biostimulants are not nutrients, but products that in very little amounts are able to promote plant growth and development, plant nutrient uptake and use efficiency, plant metabolic processes, or act as soil amendments to help ameliorating soil structure, function, or performance. However, because they are manufactured starting from different sources and consist of complex mixtures of active substances, the assignment of specific functions in plants to the individual components of these products is often arduous.
In the last few years, several research has proposed the application of products derived from fruit residues in agriculture because of their content in phenols, which are particularly actives in stimulating plant metabolism [10,11]. There are a number of techniques used for the extraction of phenolic compounds from different types of vegetable residues, most of them using organic solvents (e.g. ethanol), high temperatures, and long extraction times to exhaust the materials [12,13]. However, long extraction times can cause artifacts and organic solvents may pose a risk for the environment and human health. Other extraction procedures have been recently proposed, such as ultrasound [14], pressure [15,16] and supercritical fluid extraction [17]. Controlled enzymatic hydrolysis (Figure 1) has been applied for proteins recovery from residues of animal and fish processing industries [18]. This process is advantageous in terms of environmental impact (low temperature, minimum amount of water) and because several end-products may be recovered.
In a study by Ertani et al. [10], maize (Zea mays L.) plants were supplied for the first time with the dry apple and blueberry residues under controlled conditions. These residues were particularly rich in phenol compounds. Plants were grown for 12 d in a nutrient solution in the absence (control) or in the presence of either cellulolytic dry apple hydrolysate (AP) or dry blueberry cool extract (BB) applied at two different doses. Plants supplied with AP and BB displayed a significant increase in root and leaf biomass and a high content in macronutrients and proteins. Conversely, sucrose and glucose concentrations drastically decreased in foliar tissues of plants treated with AP and BB (Table 1). Of particular interest was the positive impact of AP and BB on the secondary metabolism associated with the synthesis of phenolic compounds, evaluated in terms of activation of phenylalanine ammonia-lyase (PAL; EC 4.3.1.5) gene expression. PAL catalyzes the first step of the phenylpropanoid pathway, by converting phenylalanine into trans-cinnamic acid and tyrosine into p-coumaric acid. In Table 2 the content of soluble phenols in the plant tissues in relation to the treatment of plants with AP and BB is shown [10].
Dosage | Proteins | Sugars | |||
mg g−1 fresh weight | Sucrose | Glucose | Fructose | ||
s | es | mg g−1 dry weight | |||
Control | 1.01 ± 0.41 d | 2.90 ± 0.37 c | 116.73 ± 12.34 a | 27.42 ± 1.33 a | 77.43 ± 0.57 d |
Apple 0.1 | 1.36 ± 0.34 c | 3.19 ± 0.17 b | 77.39 ± 10.02 b | 10.96 ± 1.14 c | 226.66 ± 1.21 a |
Apple 1.0 | 1.94 ± 0.16 a | 3.91 ± 0.10 a | 3.55 ± 1.48 d | 4.05 ± 1.26 d | 223.18 ± 0.68 b |
Blue Berry 0.1 | 1.85 ± 0.10 b | 3.57 ± 0.16 b | 22.47 ± 4.42 c | 22.76 ± 1.18 b | 99.79 ± 0.61 c |
Blue Berry 1.0 | 1.99 ± 0.32 a | 3.83 ± 0.09 a | 8.83 ± 3.27 d | 6.83 ± 2.29 d | 110.12 ± 0.44 c |
Control | Apple 0.1 | Apple 1.0 | Blue Berry 0.1 | Blue Berry 1.0 | |
(mg gallic acid g−1 fw) | |||||
Phenols | 1.02 ± 0.04 c | 1.13 ± 0.05 c | 1.30 ± 0.05 b | 1.48 ± 0.04 a | 1.52 ± 0.09 a |
(mg gallic acid g−1 fw) | |||||
Flavonoids | 0.092 ± 0.005 d | 0.123 ± 0.017 c | 0.111 ± 0.007 c | 0.204 ± 0.010 b | 0.232 ± 0.013 a |
(µg g−1d w) | |||||
Gallic acid | 2.37 ± 0.62 c | 13.73 ± 2.84 b | 22.81 ± 1.47 a | 14.52 ± 2.73 b | 24.42 ± 1.12 a |
Protocatechuic acid | 12.80 ± 1.73 d | 30.82 ± 3.23c | 42.38 ± 1.99 b | 50.52 ± 2.85 a | 49.93 ± 2.16 a |
Syringic acid | n.d. | 26.86 ± 2.44 a | 28.32 ± 2.17 a | 26.02 ± 2.81 a | 25.92 ± 2.10 a |
Vanillic acid | n.d. | 41.27 ± 5.61 a | 20.08 ± 3.72 b | n.d. | n.d. |
p-Hydroxibenzoic acid | n.d. | 5.89 ± 0.98 a | 2.68 ± 0.99 b | 6.15 ± 0.43 a | 6.72 ± 0.35 a |
Caffeic acid | n.d. | n.d. | 2.47 ± 0.63 b | 12.13 ± 1.97 a | 10.86 ± 2.09 a |
Chlorogenic acid | n.d. | n.d. | n.d. | n.d. | n.d. |
p-Coumaric acid | 55.01 ± 4.03 c | 15.61 ± 2.28 e | 149.70 ± 4.34 b | 34.71 ± 2.04 d | 172.46 ± 4.12 a |
Ferulic acid | 75.33 ± 5.62 c | 83.41 ± 3.84 bc | 87.18 ± 3.01 ab | 84.65 ± 4.21 bc | 91.32 ± 4.16 a |
(nmol cinnamic acid mg−1 protein min−1) | |||||
PAL act | 9.481 ± 0.012 c | 23.845 ± 0.008 b | 21.425 ± 0.007 b | 23.855 ± 0.009 b | 30.399 ± 0.017 a |
Phenols are important compounds for their ability to act as antioxidants and protectors against reactive oxygen species [19,20,21]. Protocatechuic, hydroxybenzoic, vanillic and p-coumaric acids influence several physiological effects that include improvement of plant-water relationships, stomatal function and rate of photosynthesis and respiration. These phenols also interact with several phytohormones and enzymes determining a different biosynthesis and flow of carbon into metabolites. Nevertheless, the factors regulating and controlling the quality and quantity of phenols in plant tissues still remain partially unknown.
In other study by Ertani et al. [11], vegetal extracts derived from hawthorn (Crataegus monogyna Jacq.) leaves, red grape (Vitis vinifera L.) skin material and blueberry (Vaccinium vitis-idaea L.) fruits were applied in maize plants in order to verify their biostimulant activity. Their content in phenols was evaluated, as well as the hormone-like activity, which was inferred via indoleacetic acid and isopentenyladenosine quantification. The biostimulant effect was demonstrated in terms of root and leaf biomass improvement, high content in chlorophyll and sugars compared to untreated plants. Moreover, hawthorn, red grape skin and blueberry enhanced the accumulation of p-coumaric acid, whilst hawthorn alone increased the amounts of gallic and p-hydroxybenzoic acids. These evidences were likely imputable to the role of phenols in stimulating plant metabolism, and to the promotion of the auxin-mediated signal transduction pathway [22,23].
Selby and coauthors [24] further analyzed aqueous extracts from the forage grass (Lolium perenne) finding an elicitor of plant defense reactions in the whole plant system and in crop situations. They also showed that an elicitor from one species can be active in a completely unrelated species, thus suggesting that the biostimulant activity from L. perenne extracts might have a broad applicability to a wide range of crops, particularly if its mode-of-action consists in inducing plant defenses.
Sánchez-Gómez et al. [25], demonstrated that grapevines treated with aqueous oak extracts via foliar application were able to modulate the phenolic and aroma composition of the grapes and the derived wines products [26,27]. One of the biggest problem in wine-producing regions is to create alternatives for processing the great amount of grape waste generated during the harvest season. Sánchez-Gómez et al. [25], proposed the use of vine-shoot waste as biostimulant for vitis vinifera metabolism. The authors produced Airén waste vine-shoot aqueous extracts using four solid-liquid extraction techniques, such as conventional solid-liquid extraction (CSLE), solid-liquid dynamic extraction (SLDE-Naviglio), microwave extraction (ME), and pressurized solvent extraction (PSE). The results showed that vine-shoot waste aqueous extracts from the V. vinifera Airén variety obtained by CSLE and SLDE-Naviglio had the highest concentration of phenolic, volatile, and mineral valuable compounds, therefore they may have a new potential use as biostimulants or foliar fertilizers in vineyards. Their use could be implemented in the "Sustainable Viticulture" concept, where the cycle of the vine could be closed, such as using vine-shoot extracts to modulate the grape quality, return nutrient to the soil and enhance terroir effect.
In the paper of Bulgari et al. [28], the effectiveness of raw extracts from leaves or flowers of Borago officinalis L., were assessed in Lactuca sativa Longifolia. The extracts were diluted to 1 or 10 mL L−1, sprayed onto lettuce plants at the middle of the growing cycle and 1 day before harvest. The leaf photosynthetic efficiency (chlorophyll a fluorescence and leaf gas exchanges), levels of ethylene, photosynthetic pigments, nitrate, total sugars and total phenols and flavonoids, including the activity and levels of phenylalanine ammonia lyase (PAL), were assessed. The authors reported that borage extracts increased the primary metabolism by inducing leaf pigment accumulation and the photosynthetic activity. Plant fresh weight increased upon treatments with 10 mL L–1 doses, and chlorophyll a fluorescence data showed that flowers were able to increase the number of active reaction centers per cross section. Ethylene was three-fold lower in flower treatments. Nitrate and sugar levels did not change in response to the different treatments. Total flavonoids and phenols, as well as the total protein levels, the in vitro PAL specific activity, and the levels of PAL-like polypeptides were increased by all borage extracts, with particular regard to flowers. Flower extracts also were efficient in preventing degradation and inducing an increase in photosynthetic pigments during storage.
The use of aqueous extracts produced from fruit juices and derived products as biostimulants in agriculture is gaining attention, even if additional studies are needed to develop new extractive techniques from these sources. Nowadays, there is a trend that encourages the use of free environmental techniques to enhance sustainability; it is the so called "green chemistry". Such techniques are developed to reduce and/or eliminate the use of organic solvents, and the use of water as a solvent for extraction should be preferred. The resulting aqueous extracts will have the advantage of being exempt from certification based on their vegetal origin, although the concentration of bioactive compounds in such extracts may be lower than when other techniques are used, as the exhaustion of the vegetable material does not take place.
All authors declare no conflicts of interest in this paper.
[1] |
Schieber A, Ullrich W, Carle R (2000) Characterization of polyphenols in mango puree concentrate by HPLC with diode array and mass spectrometric detection. Innov Food Sci Emerg Technol 1: 161-166. doi: 10.1016/S1466-8564(00)00015-1
![]() |
[2] |
Balasundram N, Sundram K, Samman S (2006) Phenolic compounds in plant and agrindustrial byproducts: antioxidant activity, occurrence, and potential uses. Food Chem 99: 191-203. doi: 10.1016/j.foodchem.2005.07.042
![]() |
[3] | Solange I, Mussatto LF, Ballesteros SM (2012) Use of agro-industrial wastes in solid-state fermentation processes, In: indus trial waste, Kuan-Yeow Show.Editor, In tech. |
[4] |
da Silva MAG, Roque SAT, Muniz AS, et al. (2008) Use of agro-industrial organic compound: Yield and nutrient absorption in wheat. Comm Soil Sci Plant Anal 39: 1311-1320. doi: 10.1080/00103620802003989
![]() |
[5] | European Biostimulants Industry Council. EBIC and biostimulants in brief. 2012, Available from: http://www.biostimulants.eu/ |
[6] |
Nardi S, Pizzeghello D, Schiavon M, et al. (2016) Plant biostimulants: physiological responses induced by protein hydrolyzed-based products and humic substances in plant metabolism. Sci Agric 73: 18-23. doi: 10.1590/0103-9016-2015-0006
![]() |
[7] | European Biostimulants Industry Council. Economic overview of the biostimulants sector in Europe. 2013. Available from: http://www.biostimulants.eu/2013/04/2013-overview-of-the-european-biostimulants-market/ |
[8] |
Calvo P, Nelson L, Kloepper JW (2014) Agricultural uses of plant biostimulants. Plant Soil 383: 3-41. doi: 10.1007/s11104-014-2131-8
![]() |
[9] | European Biostimulants Industry Council. What are biostimulants? 2012, Available from: http://www.biostimulants.eu/about/what-are-biostimulants/ |
[10] |
Ertani A, Schiavon M, Altissimo A, et al. (2011) Phenol-containing organic substances stimulate phenylpropanoid metabolism in Zea mays. J Plant Nutr Soil Sci 174: 496-503. doi: 10.1002/jpln.201000075
![]() |
[11] | Ertani A, Pizzeghello D, Francioso O, et al. (2014) Capsicum chinensis L. growth and nutraceutical properties are enhanced by biostimulants in a long-term period: chemical and metabolomic approaches. Front Plant Sci 5: 375. |
[12] |
Luque-Rodríguez J M, Luque de Castro MD, Pérez-Juan P, et al. (2007) Dynamic superheated liquid extraction of anthocyanins and other phenolics from red grape skins of winemaking residues. Bioresour Technol 98: 2705-2713. doi: 10.1016/j.biortech.2006.09.019
![]() |
[13] |
Spigno G, Tramelli L, De Faveri DM (2007) Effects of extraction time, temperature and solvent on concentration and antioxidant activity of grape marc phenolics. J Food Eng 81: 200-208. doi: 10.1016/j.jfoodeng.2006.10.021
![]() |
[14] |
Vilkhu K, Mawson R, Simons L, et al. (2008) Applications and opportunities for ultrasound assisted extraction in the food industry: A review. Inn Food Sci Emerging Technol 9: 161-169. doi: 10.1016/j.ifset.2007.04.014
![]() |
[15] |
Delgado de la Torre MP, Ferreiro-Vera C, Priego-Capote F, et al. (2012) Comparison of accelerated methods for the extraction of phenolic compounds from different vine-shoot cultivars. J Agric Food Chem 60: 3051-3060. doi: 10.1021/jf205078k
![]() |
[16] |
Mustafa A, Turner C (2011) Pressurized liquid extraction as a green approach in food and herbal plants extraction: A review. Anal Chim Acta 703: 8-18. doi: 10.1016/j.aca.2011.07.018
![]() |
[17] | Pourmortazavi SM, Rahimi-Nasrabadi M, Hajimirsadeghi SS (2014) Supercritical fluid technology in analytical chemistry: A review. Curr Anal Chem 10: 3-28. |
[18] |
Bhaskar N, Modi K, Govindaraju R, et al. (2007) Utilization of meat industry by products: protein hydrolysate from sheep visceral mass. Bioresour Technol 98: 388-394. doi: 10.1016/j.biortech.2005.12.017
![]() |
[19] | Rice EL (1995) Allelopathy, 2Eds., New York: Academic Press. |
[20] | Northup RR, Yu Z, Dahlgren RA, et al. (1998) Polyphenol control of nitrogen release from pine litter. Nature 77: 227-229. |
[21] | Inderjit (1996) Plant phenolics in allelopathy. Bot Rev 62: 182-202. |
[22] | Brebu M, Vasile C (2010) Thermal degradation of wood: A review. Cellul Chem Technol 44: 353-363. |
[23] |
Hoareau W, Trindade WG, Siegmund B, et al. (2004) Sugar cane bagasse and curaua lignins oxidized by chlorine dioxide and reacted with furfuryl alcohol: Characterization and stability. Polym Degrad Stab 86: 567-576. doi: 10.1016/j.polymdegradstab.2004.07.005
![]() |
[24] |
Selby C, Carmichael EHS, Sharma S (2016) Bio-refining of perennial ryegrass (Lolium perenne): evaluation of aqueous extracts for plant defence elicitor activity using French bean cell suspension cultures. Chem Biol Technol Agric 3: 11. doi: 10.1186/s40538-016-0061-9
![]() |
[25] |
Sánchez-Gómez R, Zalacain A, Alonso GL, et al. (2014) Vine-shoot waste aqueous extracts for re-use in agriculture obtained by different extraction techniques: phenolic, volatile, and mineral compounds. J Agric Food Chem 62: 10861-10872. doi: 10.1021/jf503929v
![]() |
[26] |
Martínez-Gil AM, Angenieux M, Pardo-García AI, et al. (2013) Glycosidic aroma precursors of Syrah and Chardonnay grapes after an oak extract application to the grapevines. Food Chem 138: 956-965. doi: 10.1016/j.foodchem.2012.11.032
![]() |
[27] |
Martínez-Gil AM, Garde-Cerdán T, Martínez L, et al. (2011) Effect of oak extract application to Verdejo grapevines on grape and wine aroma. J Agric Food Chem 59: 3253-3263. doi: 10.1021/jf104178c
![]() |
[28] | Bulgari R, Morgutti S, Cocetta G, et al. (2017) Evaluation of borage extracts as potential biostimulant using a phenomic, agronomic, physiological, and biochemical approach. Front Plant Sci 935: 1-8. |
1. | Andrea Ertani, Serenella Nardi, Ornella Francioso, Santiago Sanchez-Cortes, Michele Di Foggia, Michela Schiavon, Effects of Two Protein Hydrolysates Obtained From Chickpea (Cicer arietinum L.) and Spirulina platensis on Zea mays (L.) Plants, 2019, 10, 1664-462X, 10.3389/fpls.2019.00954 | |
2. | Agnieszka Szparaga, Sławomir Kocira, Ireneusz Kapusta, Grzegorz Zaguła, Prototyping extracts from Artemisia absinthium L. for their biostimulating properties yield-enhancing, and farmer income-increasing properties, 2021, 160, 09266690, 113125, 10.1016/j.indcrop.2020.113125 | |
3. | Matteo Perra, Alba Cuena-Lombraña, Gianluigi Bacchetta, Maria Letizia Manca, Maria Manconi, Richard G. Maroun, Aldo Muntoni, Carlo Ignazio Giovanni Tuberoso, Katarzyna A. Gil, Giorgia De Gioannis, Combining Different Approaches for Grape Pomace Valorization: Polyphenols Extraction and Composting of the Exhausted Biomass, 2022, 14, 2071-1050, 10690, 10.3390/su141710690 | |
4. | Deise Munaro, Carolina Herrera Mazo, Cláudia Marlene Bauer, Leticia da Silva Gomes, Emanuelle Bavaresco Teodoro, Letícia Mazzarino, Maria Beatriz da Rocha Veleirinho, Sidnei Moura e Silva, Marcelo Maraschin, A novel biostimulant from chitosan nanoparticles and microalgae-based protein hydrolysate: Improving crop performance in tomato, 2024, 323, 03044238, 112491, 10.1016/j.scienta.2023.112491 | |
5. | José Alejandro Díaz-Elizondo, Azrrael Ayala-Velazco, Adalberto Benavides-Mendoza, Francisco Javier Enriquez-Medrano, Julia Medrano-Macías, Obtaining Lignin from Nutshells under Mild Extraction Conditions and Its Use as a Biostimulant in Tomato Seedlings, 2024, 10, 2311-7524, 1079, 10.3390/horticulturae10101079 |
Dosage | Proteins | Sugars | |||
mg g−1 fresh weight | Sucrose | Glucose | Fructose | ||
s | es | mg g−1 dry weight | |||
Control | 1.01 ± 0.41 d | 2.90 ± 0.37 c | 116.73 ± 12.34 a | 27.42 ± 1.33 a | 77.43 ± 0.57 d |
Apple 0.1 | 1.36 ± 0.34 c | 3.19 ± 0.17 b | 77.39 ± 10.02 b | 10.96 ± 1.14 c | 226.66 ± 1.21 a |
Apple 1.0 | 1.94 ± 0.16 a | 3.91 ± 0.10 a | 3.55 ± 1.48 d | 4.05 ± 1.26 d | 223.18 ± 0.68 b |
Blue Berry 0.1 | 1.85 ± 0.10 b | 3.57 ± 0.16 b | 22.47 ± 4.42 c | 22.76 ± 1.18 b | 99.79 ± 0.61 c |
Blue Berry 1.0 | 1.99 ± 0.32 a | 3.83 ± 0.09 a | 8.83 ± 3.27 d | 6.83 ± 2.29 d | 110.12 ± 0.44 c |
Control | Apple 0.1 | Apple 1.0 | Blue Berry 0.1 | Blue Berry 1.0 | |
(mg gallic acid g−1 fw) | |||||
Phenols | 1.02 ± 0.04 c | 1.13 ± 0.05 c | 1.30 ± 0.05 b | 1.48 ± 0.04 a | 1.52 ± 0.09 a |
(mg gallic acid g−1 fw) | |||||
Flavonoids | 0.092 ± 0.005 d | 0.123 ± 0.017 c | 0.111 ± 0.007 c | 0.204 ± 0.010 b | 0.232 ± 0.013 a |
(µg g−1d w) | |||||
Gallic acid | 2.37 ± 0.62 c | 13.73 ± 2.84 b | 22.81 ± 1.47 a | 14.52 ± 2.73 b | 24.42 ± 1.12 a |
Protocatechuic acid | 12.80 ± 1.73 d | 30.82 ± 3.23c | 42.38 ± 1.99 b | 50.52 ± 2.85 a | 49.93 ± 2.16 a |
Syringic acid | n.d. | 26.86 ± 2.44 a | 28.32 ± 2.17 a | 26.02 ± 2.81 a | 25.92 ± 2.10 a |
Vanillic acid | n.d. | 41.27 ± 5.61 a | 20.08 ± 3.72 b | n.d. | n.d. |
p-Hydroxibenzoic acid | n.d. | 5.89 ± 0.98 a | 2.68 ± 0.99 b | 6.15 ± 0.43 a | 6.72 ± 0.35 a |
Caffeic acid | n.d. | n.d. | 2.47 ± 0.63 b | 12.13 ± 1.97 a | 10.86 ± 2.09 a |
Chlorogenic acid | n.d. | n.d. | n.d. | n.d. | n.d. |
p-Coumaric acid | 55.01 ± 4.03 c | 15.61 ± 2.28 e | 149.70 ± 4.34 b | 34.71 ± 2.04 d | 172.46 ± 4.12 a |
Ferulic acid | 75.33 ± 5.62 c | 83.41 ± 3.84 bc | 87.18 ± 3.01 ab | 84.65 ± 4.21 bc | 91.32 ± 4.16 a |
(nmol cinnamic acid mg−1 protein min−1) | |||||
PAL act | 9.481 ± 0.012 c | 23.845 ± 0.008 b | 21.425 ± 0.007 b | 23.855 ± 0.009 b | 30.399 ± 0.017 a |
Dosage | Proteins | Sugars | |||
mg g−1 fresh weight | Sucrose | Glucose | Fructose | ||
s | es | mg g−1 dry weight | |||
Control | 1.01 ± 0.41 d | 2.90 ± 0.37 c | 116.73 ± 12.34 a | 27.42 ± 1.33 a | 77.43 ± 0.57 d |
Apple 0.1 | 1.36 ± 0.34 c | 3.19 ± 0.17 b | 77.39 ± 10.02 b | 10.96 ± 1.14 c | 226.66 ± 1.21 a |
Apple 1.0 | 1.94 ± 0.16 a | 3.91 ± 0.10 a | 3.55 ± 1.48 d | 4.05 ± 1.26 d | 223.18 ± 0.68 b |
Blue Berry 0.1 | 1.85 ± 0.10 b | 3.57 ± 0.16 b | 22.47 ± 4.42 c | 22.76 ± 1.18 b | 99.79 ± 0.61 c |
Blue Berry 1.0 | 1.99 ± 0.32 a | 3.83 ± 0.09 a | 8.83 ± 3.27 d | 6.83 ± 2.29 d | 110.12 ± 0.44 c |
Control | Apple 0.1 | Apple 1.0 | Blue Berry 0.1 | Blue Berry 1.0 | |
(mg gallic acid g−1 fw) | |||||
Phenols | 1.02 ± 0.04 c | 1.13 ± 0.05 c | 1.30 ± 0.05 b | 1.48 ± 0.04 a | 1.52 ± 0.09 a |
(mg gallic acid g−1 fw) | |||||
Flavonoids | 0.092 ± 0.005 d | 0.123 ± 0.017 c | 0.111 ± 0.007 c | 0.204 ± 0.010 b | 0.232 ± 0.013 a |
(µg g−1d w) | |||||
Gallic acid | 2.37 ± 0.62 c | 13.73 ± 2.84 b | 22.81 ± 1.47 a | 14.52 ± 2.73 b | 24.42 ± 1.12 a |
Protocatechuic acid | 12.80 ± 1.73 d | 30.82 ± 3.23c | 42.38 ± 1.99 b | 50.52 ± 2.85 a | 49.93 ± 2.16 a |
Syringic acid | n.d. | 26.86 ± 2.44 a | 28.32 ± 2.17 a | 26.02 ± 2.81 a | 25.92 ± 2.10 a |
Vanillic acid | n.d. | 41.27 ± 5.61 a | 20.08 ± 3.72 b | n.d. | n.d. |
p-Hydroxibenzoic acid | n.d. | 5.89 ± 0.98 a | 2.68 ± 0.99 b | 6.15 ± 0.43 a | 6.72 ± 0.35 a |
Caffeic acid | n.d. | n.d. | 2.47 ± 0.63 b | 12.13 ± 1.97 a | 10.86 ± 2.09 a |
Chlorogenic acid | n.d. | n.d. | n.d. | n.d. | n.d. |
p-Coumaric acid | 55.01 ± 4.03 c | 15.61 ± 2.28 e | 149.70 ± 4.34 b | 34.71 ± 2.04 d | 172.46 ± 4.12 a |
Ferulic acid | 75.33 ± 5.62 c | 83.41 ± 3.84 bc | 87.18 ± 3.01 ab | 84.65 ± 4.21 bc | 91.32 ± 4.16 a |
(nmol cinnamic acid mg−1 protein min−1) | |||||
PAL act | 9.481 ± 0.012 c | 23.845 ± 0.008 b | 21.425 ± 0.007 b | 23.855 ± 0.009 b | 30.399 ± 0.017 a |