Here are all the ways this water-packed summer staple can revolutionize your dietary health. By Good Housekeeping Magazine.

Nothing tastes like summer more than a crisp, juicy, refreshing slice of watermelon. Watermelon — which can actually be considered a fruit or a vegetable, according to the National Watermelon Promotion Board — is incredibly nutritious, too. Nutritionists say watermelon is low in calories and sugar and chock-full of vitamins, minerals, and antioxidants, making it a great addition to a healthy diet.

According to the U. Department of Agriculture , 1 cup of watermelon g contains:. Different fruits contain different nutrients, so eating a variety will ensure your body gets everything it needs.

People with diabetes or who may need to count their carbohydrate servings should also pay attention to their watermelon intake, she adds. Eating too much fruit could introduce too much sugar to your diet, leading to blood sugar fluctuations , which can be risky for people with diabetes.

At just 46 calories per cup , watermelon packs a punch when it comes to nutrients. Vitamin C strengthens your immune system and helps the body absorb iron, Derocha says, while vitamin A is crucial for skin and eye health.

Watermelon is also rich in potassium, which works to lower blood pressure and supports nerve functioning, and vitamin B6, which helps the body break down the proteins that you eat and also boosts the immune system and nerve function. Lycopene is a natural compound found in watermelon and other fruits and vegetables that has antioxidant properties.

The substance is also what gives watermelon its red color; but beyond its hue, lycopene is also good for you, too. Frusciante L, Carli P, Ercolano MR, Pemice R, Di-Matteo A, Fogliano V, Pellegrini N Antioxidant nutritional quality of tomato. Mol Nutr Food Res 51 5 — Kamiloglu S, Demirci M, Selen S, Toydemir G, Boyacioglu D, Capanoglu E Home processing of tomatoes Solanum lycopersicum : effects on in vitro bioaccessibility of total lycopene, phenolics, flavonoids, and antioxidant capacity.

J Sci Food Agric 94 11 — FAO Food and Agriculture Organization of the United Nations. FAOSTAT, Production, Accessed 18 Jan Bohm F, Edge R, Burke M, Truscott TG Dietary uptake of lycopene protects human cells from singlet oxygen and nitrogen dioxide-ROS components from cigarette smoke. J Photochem Photobiol B: Biol 64 2—3 — CAS Google Scholar.

Nguyen ML, Schwartz SJ Lycopene: chemical and biological properties. Food Technol 53 2 — Singh P, Goyal GK Dietary lycopene: Its properties and anticarcinogenic effects.

Compr Rev Food Sci Food Saf 7 3 — Naz A, Butt MS, Pasha I, Nawaz H Antioxidant indices of watermelon juice and lycopene extract. Pak J Nutr — Tlili I, Hdider C, Lenucci MS, Ilahy R, Jebari H, Dalessandro G Bioactive compounds and antioxidant activities during fruit ripening of watermelon cultivars.

J Food Comps Anal 24 7 — Hong MY, Hartig N, Kaufman K, Hooshmand S, Figueroa A, Kern M Watermelon consumption improves inflammation and antioxidant capacity in rats fed an atherogenic diet.

Nutr Res 35 3 — Ghavipour M, Saedisomeolia A, Djalali M, Sotoudeh G, Eshranghyan MR, Moghada AM, Wood LG Tomato juice consumption reduces systemic inflammation in overweight and obese females. Br J Nutr 11 — Ferruzzi MG, Nguyen ML, Sander LC, Rock CL, Schwartz SJ Analysis of lycopene geometrical isomers in biological microsamples by liquid chromatography with coulometric array detection.

J Chromatography B 2 — Korea Health Statistics Korea National Health and Nutrition Examination Survey KNHANES VII Korea Center for Disease Control and Prevention; Seoul, Korea, pp 3— Meyers KJ, Watkins CB, Pritts MP, Liu RH Antioxidant and antiproliferative activities of strawberries.

J Agric Food Chem 51 23 — Shin Y Correlation between antioxidant concentrations and activities of Yuja Citrus junos Sieb ex Tanaka and other citrus fruits. Food Sci Biotechnol 21 5 — Bicanic D, Fogliano V, Luterotti S, Swarts J, Piani G, Graziani G Quantification of lycopene in tomato products: comparing the performances of a newly proposed direct photothermal method and high-performance liquid chromatography.

J Sci Food Agric 85 7 — Bicanic D, Swarts J, Luterotti S, Helander P, Fogliano V, Anese M Optothermistor as a breakthrough in the quantification of lycopene content of thermally processed tomato-based foods: Verification versus absorption spectrophotometry and high-performance liquid chromatography.

J Agric Food Chem 53 9 — Brand-Williams W, Cuvelier ME, Berset C Use of a free radical method to evaluate antioxidant activity.

LWT-Food Sci Technol — J Food Comp Anal 24 7 — Re R, Pellegrini N, Proteggente A, Pannala A, Yang M, Rice-Evans C Antioxidant activity applying an improved ABTS radical cation decolorization assay. Free Radical Biol Med 26 9—10 — Hallmann E The influence of organic and conventional cultivation systems on the nutritional value and content of bioactive compounds in selected tomato types.

J Sci Food Agric 92 14 — Kamiloglu S, Boyacioglu D, Capanoglu E The effect of food processing on bioavailability of tomato antioxidants. J Berry Res 3 2 — Gerard KA, Robert JS Microwave heating of apple mash to improve juice yield and quality. Food Sci Technol 37 5 — Tlili I, Hdider C, Lenucci MS, Riadh I, Jebari H, Dalessandro G Bioactive compounds and antioxidant activities of different watermelon Citrullus lanatus Thunb.

Mansfeld cultivars as affected by fruit sampling area. J Food Comp Anal 24 3 — Lee BH, Kim SY, Cho CH, Chung DK, Chun OK, Kim DO Estimation of daily per capita intake of total phenolics, total flavonoids, and antioxidant capacities from fruit and vegetable juices in the Korean diet based the Korea National Health and Nutrition Examination survey Korean J Food Sci Technol 43 4 — Gahler S, Otto K, Bohm V Alterations of vitamin C, total phenolics, and antioxidant capacity as affected by processing tomatoes to different products.

J Agric Food Chem 51 27 — Klopotek Y, Otto K, Böhm V Processing strawberries to different products alters contents of vitamin c, total phenolics, total anthocyanins, and antioxidant capacity. J Agric Food Chem 53 14 — Nagal S, Kaur C, Choudhary H, Singh J, Bhushan-Singh B, Singh KN Lycopene content, antioxidant capacity and colour attributes of selected watermelon Citrullus lanatus Thunb.

Mansfeld cultivars grown in India. Int J Food Sci Nutr 63 8 — Martinez-Valverde I, Periago MJ, Provan G, Chesson A Phenolic compounds, lycopene and antioxidant activity in commercial varieties of tomato Lycopersicum esculentum.

J Sci Food Agric 82 3 — Markovic K, Hruskar M, Vahcic N Lycopene content of tomato products and their contribution to the lycopene intake of Croatians. Nutr Res 26 11 — Perkins-Veazie P, Collins JK, Pair SD, Roberts W Lycopene content differs among red-fleshed watermelon cultivars.

J Sci Food Agric 81 10 — Chun OK, Kim DO, Smith N, Schroeder D, Han JT, Lee CY Daily consumption of phenolics and total antioxidant capacity from fruit and vegetables in the American diet.

J Sci Food Agric 85 10 — Jenab M, Ferrari P, Mazuir M, Tjonneland A, Clavel-chapelon F, Linseisen J, Trichopoulou A, Tumino R, Bueno-de-Mesquita HB, Lund E, Gonzalez CA, JohanssonJ-Key T, Riboli E G S Variations in lycopene blood levels and tomato consumption across European countries based on the European Prospective Investigation into Cancer and Nutrition EPIC study.

J Nutr 8 S—S. Goldbohm RA, Brants HA, Hulshof KF, Van Den Brandt PA The contribution of various foods to intake of vitamin A and carotenoids in The Netherlands.

Int J Vitam Nutr Res 68 6 — Pelz R, Schmidt-Faber B, Heseker H Carotenoid intake in the German national food consumption survey. Z Ernahrungswiss 37 4 — British J Nutr 85 4 — Porrini M, Riso P S What are typical lycopene intakes?

Johnson-Down L, Saudny-Unterberger H, Gray-Donald K Food habits of Canadians: lutein and lycopene intake in the Canadian population. J Am Diet Assoc 7 — PubMed Google Scholar.

Lugasi A, Brio L, Hovarie J, Sagi KV, Brandt S, Barna E Lycopene content of foods and lycopene intake in two groups of the Hungarian population.

Nutr Res 23 8 — Wawrzyniak A, Marciniak A, Rajewska J Lycopene content of selected foods available on the Polish market and estimation of its intake. Pol J Food Nutr Sci 14 55 — McNaughton SA, Marks GC, Gaffney P, Williams G, Green A Validation of a food-frequency questionnaire assessment of carotenoid and vitamin E intake using weighed food records and plasma biomarkers: the method of triads model.

Eur J Clin Nutr 59 2 — Vandevijvere S, Cucu T, Vinkx C, Huvaere K, Huybrechts I, Van Loco J Dietary intake of lycopene by the Belgian adult population.

Public Health Nutr 17 2 — Mazeikiene A, Jakaitiene A, Karciauskaite D, Kucinskiene ZA, Abaravicius JA, Kaminskas A, Kucinskas V Dietary lycopene and cardiovascular health in ethnic Lithuanians. Acta Med Litu 22 4 — Download references. Department of Food Engineering, Dankook University, Cheonan, Chungnam, , Republic of Korea.

Department of Food Science and Technology, Seoul National University of Science and Technology, Seoul, , Republic of Korea. You can also search for this author in PubMed Google Scholar. Conceptualization, YK and YS; methodology, HP, YK, and YS; formal analysis, HP, YK, and YS; writing—original draft preparation, HP; writing—review and editing, YK and YS.

All authors read and approved the final manuscript. Correspondence to Young-Jun Kim or Youngjae Shin. Springer Nature remains neutral with regard to jurisdictional claims in published maps and institutional affiliations. Open Access This article is licensed under a Creative Commons Attribution 4.

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Reprints and permissions. Park, H. Estimation of daily intake of lycopene, antioxidant contents and activities from tomatoes, watermelons, and their processed products in Korea. Appl Biol Chem 63 , 50 Download citation. Received : 30 June Accepted : 28 August Published : 04 September Anyone you share the following link with will be able to read this content:.

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Download PDF. Abstract Tomatoes, watermelons, and processed tomato products contain abundant antioxidant compounds, including lycopene. Introduction As nutrition improved at the national level, the problems related to nutrient deficiency decreased; however, the western dietary pattern has caused nutritional imbalance to emerge, along with various chronic diseases, such as hypertension, cancer, and diabetes [ 1 ].

Materials and methods Materials The samples were purchased from local fruit markets and hypermarkets in the city of Cheonan, South Korea. Total flavonoid analysis The total flavonoid content of samples was measured using the colorimetric assay method [ 17 , 18 ]. Total phenolic analysis The total phenolic content of samples was measured using the Folin-Ciocalteu colorimetric assay method [ 17 , 18 ].

DPPH radical scavenging activity analysis The DPPH radical scavenging activity was measured using a modified method by Shin [ 18 ] and Brand-Williams et al. Every cell, tissue and organ in your body needs water to work properly. Even mild dehydration can drain your energy and make you tired.

Every day you lose water through your breath, perspiration, urine and bowel movements. For your body to function properly, you must replenish its water supply by consuming beverages and foods that contain water.

Watermelon provides about USDA Food Composition Database. Some observational studies have shown an association between a diet rich in the antioxidant lycopene, as well as those using supplemental lycopene and reduced risk of some types of cancer, including breast , prostate and lung cancers.

However, thus far, research results are inconsistent in these areas and randomized controlled trials have not demonstrated a benefit to consuming lycopene from food or supplements. However, there is a large and growing body of research Into the mechanistic and dose-relational effects of lycopene consumption.

Although more research and clinical trials are needed, systematic reviews and meta-analyses link the consumption ofcarotenoids, like lycopene, to a role in maintaining healthy skin.

Sign up here to receive the monthly What About Watermelon? enewsletter with seasonal tips, tricks and recipes. The Board provides this link as a convenience to you; the link should not be considered an endorsement by the Board of the third-party website or the company who owns it.

The Board is not responsible for the quality, safety, completeness, accuracy or nature of the content of the linked website. To return to www. org, simply close the new browser window. Scroll for More. Heart Happy.

The amino acid sequences were compared by multiple sequence alignment using trimAI software Capella-Gutierrez et al. A phylogenetic tree was established with the UPGMA function in MEGA X Kumar et al. Based on the transcript expression RPKM of candidate genes Cla and Cla in the above four varieties, we aggregated their data and produced a trend graph using software GraphPad 8.

We analyzed the expression levels of the two candidate genes at different mature stages of different flesh color watermelons. The flesh color of the F 1 generation was canary yellow in Pop.

Five categories of flesh color, red 87 plants , pale yellow 48 plants , canary yellow plants , and two irregular color patterns consisting of red mixed with pale and canary yellow or red in mixed patterns in the heart and placental tissues of the fruit 18 and 26 plants, respectively , were found in the segregating population.

Canary yellow and pale yellow could also be classified into red and nonred groups by visual observation. These results indicated that a single major recessive gene determined red and nonred color in watermelon based on the genetic background of Pop.

High-performance liquid chromatography analysis of lycopene contents in mature fruit showed that LSW had a high lycopene content with an average value of Comparing the lycopene content and flesh color data, we observed that the plants showed a red flesh color when the lycopene content was higher than We also compared these results with our previously published data Liu et al.

The F 2 generations in both years showed the same genetic ratio for flesh color and similarly divergent trends in lycopene content.

The results of the χ 2 goodness-of-fit test of the segregation ratios and the lycopene content separation analysis in the F 2 populations derived from LSW and COS are shown in Table 1 and Figure 2.

Table 1 Flesh color separation proportion of the parental materials and F 2 population. Figure 2 Segregation analysis of lycopene content in the F 2 population from a cross between LSW and COS parental strains in the year of and In Pop.

Four main flesh colors, red mixed with white, red mixed with white and yellow, canary yellow mixed with white, and white, emerged in the F 2 generation.

For the red and white mixed groups, most of the plants accumulated lycopene only around the seed region, while only two plants had fully red flesh color. Two groups were separated based on flesh color, one group with red or red mixed with white flesh and the other group with yellow, white, and mixed yellow and white flesh.

Approximately seven main flesh colors red, red mixed with white, orange, orange mixed with white, canary yellow, canary yellow mixed with white and mixed canary yellow, orange and red segregated in the BC 1 P 1 generation. According to the classification standards of Pop.

Considering that the F 1 plants of Pop. The separate proportions of the three genetic populations are shown in Table 2. Table 2 The separate proportion of red and nonred plants in the three genetic populations. Two linkage maps consisting of markers CAPS, 37 SSR and CAPS markers were constructed from Pop.

Eight new CAPS markers were developed in the initial mapping region for secondary mapping Figure 3. The order of the CAPS marker locations did not tightly correspond to the reference genome. For Pop. Based on the genome resequencing data, the physical distance between WII04E and WII04EBsaHI-6 was 92, bp.

The 11 CAPS markers from WII04E to CAPSSacI on chromosome 4; Figure 3B confirmed the chromosome region, with 12 candidate genes cosegregating with flesh color in most individuals of Pop.

All the mixed-color plants were heterozygous, similar to the F 1 generation. Figure 3 Genetic linkage groups of chromosome 4 for QTL detection and candidate genes in the effective region. A Genetic linkage group of chromosome 4 with the F 2 generation plants derived from a cross between garden female and PI B Genetic linkage group of chromosome 4 with the F 2 generation plants derived from a cross between LSW and COS.

C Linkage analysis of the CAPS markers in the QTL effective region. D Candidate genes in the QTL detected region. Five candidate genes Cla to Cla , which are presumed to encode proteins, Table S1 were detected in the 92,bp region WII04E to WII04EBsaHI-6 by consulting the Cucurbit Genomics Database Zheng et al.

Based on the results of ORF and BLAST Altschul et al. We also used LSW and PI to determine the polymorphism status of the 11 CAPS markers that cosegregated with flesh color in Pop. Based on the results of the linkage analysis, only WII04EBsaHI-6 was assigned to the region between WIII and WIII , and it narrowed the effective QTL region with an R 2 value of CAPSMboI was polymorphic between LSW and PI , but in Pop.

The genotyping data showed that WII04E and CAPSHindII had distorted segregation in Pop. This result may reflect the fact that PI belongs to C.

mucosospermus , which is another subspecies of watermelon. Considering the genome divergence of different watermelons, separate reference genomes are needed.

For the CAPS marker WII04EBsaHI-6 , all plants having red mixed with white flesh color showed the same electrophoretic bands as LSW, and the individuals with mixed flesh colors were heterozygous, similar to the F 1 generation.

By filtering the recombinant plants using CAPS markers WII04E and WII04E in the F 2 population, which was derived from the enlarged population of Pop. Among these, 4 dominant homozygous individuals, 6 recessive homozygous individuals, and 18 recombinant individuals were identified.

Four CAPS markers CAPSNedI-1 , CAPSEcoRI , CAPSRsaI , CAPSSacI and one known marker, WII04EBsaHI-6 , were developed between the WII04E and WII04E markers. The target region between markers WII04EBsaHI-6 and CAPSRsaI was ultimately acquired by using the above markers to fine map the six recessive homozygous individuals red flesh , and the physical distance was Using the Cucurbit Genomics Database Zheng et al.

Figure 4 Fine mapping of the genes that control the color of watermelon red flesh through recombinant recessive individuals.

P 1 black frame represents the parent material LSW, P 2 white frame represents the parent material PI , F 1 was gray frame. F , F , F , F , F , F , F , and F were recessive recombinant individuals, respectively.

Furthermore, according to fine mapping and combination of the results of CDS blasting with the gene LCYB , Cla was speculated to be on the same locus as LCYB in watermelon. Figure 5 Comparison analysis of the DNA sequence and amino acid sequence of the Cla between LSW and COS. A 3-bp mutation resulted in two amino acid mutations: A gene structure of Cla , including only one exon; B coding sequence analysis of three SNP mutations; C two amino acids mutation due to three SNP mutations.

The black boxes indicate the exons. Figure 6 Comparison analysis of the DNA sequence and amino acid sequence of the Cla between LSW and COS. A 1-bp mutation resulted in one amino acid mutation: A gene structure of Cla , including 15 exons; B coding sequence analysis of one SNP mutations; C one amino acid mutation due to one SNP mutation.

According to the genotyping data, the 11 CAPS markers perfectly cosegregated with flesh color and lycopene content in Pop. To further validate the results of the QTL mapping, we also tested the relationships between lycopene content and the CAPS marker allele pairs WII04EBsaHI-6 and WII04E for all individuals in Pop.

For the CAPS marker WII04EBsaHI-6 , the average content of lycopene in plants with homozygous LSW alleles was Heterozygous individuals accumulated an average content of 2. The same trend was also observed for WII04E The genome data of 4 parental materials and 20 other published watermelon accessions Guo et al.

The 24 watermelon samples exhibited four flesh colors: red, yellowish, green, and white. According to the results, a total of 20 SNP loci were detected in the LCYB sequence among these 24 watermelons, with only one SNP distinguishing red-fleshed color G th plants from nonred T th accessions, except in PI , which had white flesh but also had a G base at the th position in the exon region.

This SNP G th to T th changed valine V: red into glycine nonred. It was also the position of the restriction site of the CAPS marker WII04EBsaHI-6 Figure S3.

To verify the applicability of these markers in different genetic populations and watermelon accessions, the 11 CAPS markers in the effective region were used for genotyping in Pop. Among the 11 CAPS markers, only two markers WII04EBsaHI-6 and W04EII showed polymorphisms between garden parent and PI Considering the flesh color—specific markers and the different polymorphisms of white-fleshed plants, WII04EBsaHI-6 , W04EII , WII04EKpnI-1 , and WII04E were selected to construct a MAS approach for red, yellowish, and white flesh color in 81 watermelon accessions.

Based on the genotyping results, the CAPS markers WII04EBsaHI-6 and W04EII cosegregated with red and pink flesh color in these accessions, except in two PI lines PI and PI , which had yellowish flesh but exhibited the same restriction enzyme fragments as the red and pink plants.

The sizes of the PCR fragments for WII04EBsaHI-6 and W04EII were 1, and bp, respectively. For the CAPS markers WII04EKpnI-1 and WII04E , the PCR amplicons were and bp for all 81 accessions.

In the yellowish group, each plant exhibited or bp bands for WII04EKpnI-1 and or bp bands for WII04E The pink, red, and white plants showed and bp bands with these two markers. Three white-fleshed PI lines PI , PI , and PI had the same results as the yellowish-fleshed plants.

Some white-fleshed PI lines were heterozygous at these marker loci. The consistency ratios of WII04EKpnI-1 and WII04E with yellowish color were To distinguish the three types of flesh color red, yellowish, and white , the four markers were combined for further testing.

Each accession was represented by the restriction enzyme results using the four markers in the order of WII04EBsaHI-6 , W04EII , WII04EKpnI-1 , and WII04E Plants with the marker restriction site were scored as 1, those without were scored as 0, and heterozygotes were recorded as h.

With this method, each material could be given a marker code for flesh color. The results showed that the combination of these four markers could clearly distinguish the three flesh colors.

According to the genotyping results Table S3 , all of the red and pink plants had the code 1, 0, 0, 0, whereas the code for the yellowish plants was 0, 1, 1, 1. For white-fleshed plants, the codes were not uniform, and heterozygous loci were present in some plants, but the code of each white-fleshed watermelon was quite different from those of the pink, red, and yellowish accessions.

With these marker codes, we can easily distinguish the three types of flesh color in watermelon. The SSR marker SSR also showed different polymorphisms in the 81 watermelon accessions but did not cosegregate with flesh color.

The transcriptional expression RPKM of two candidate genes, Cla and Cla , in the above species of watermelons was quantified at different periods of development.

In all the accessions, COS, LSW, red flesh , and PI white flesh , the expression of Cla did not explain the lycopene accumulation difference between different watermelon accessions. Cla had a relatively stable expressing trend in COS line as fruits ripen, while in LSW, it increased as time went on.

The function of LCYB is to cyclase the lycopene into β-carotene, but in high-lycopene accumulation accession LSW, the expression level was higher than the low-lycopene materials COS Figure 7C. In red flesh , the expression level of Cla did not show any significant difference compared to PI Figures 7A, B.

Cla was annotated as the kinesin-like protein that seemed to be not related with lycopene accumulation. The expressing quantity in LSW was decreasing as fruits ripened on the whole and was significantly more than that in COS Figures 7D—F.

Figure 7 The expression levels of two candidate genes Cla and Cla in four different watermelon varieties , LSW, COS, and PI at different maturity stages of the flesh. Cla : A BioProject: PRJNA RPKM of , sample DAP10, DAP18, DAP28, DAP34, DAP: days after pollination.

B BioProject: PRJNA RPKM of PI , sample DAP10, DAP18, DAP26, DAP34, DAP42, DAP C BioProject: PRJNA RPKM of COS and LSW , DAP10, DAP18, DAP26, DAP34, DAP D BioProject: PRJNA RPKM of , sample DAP10, DAP18, DAP28, DAP E BioProject: PRJNA RPKM of PI , sample DAP10, DAP18, DAP26, DAP34, DAP42, DAP F BioProject: PRJNA RPKM of COS and LSW , DAP10, DAP18, DAP26, DAP34, DAP After screening for typical lycopene cyclase genes in different species of cucurbitaceous crops cucumber, melon, watermelon, gourd, pumpkin and tomato, 33 homologous genes were filtered, and a phylogenetic tree was constructed using MEGA X Kumar et al.

The results showed that all genes were divided into three categories, which were defined as lycopene β-cyclase LCYB and lycopene epsilon-cyclase LCYE. LCYB clusters I and II contained 13 and 9 genes, respectively. Additionally, the number of exons was relatively consistent and conservative.

Almost all genes from LCYB cluster I contained one exon, and the number of exons from the cluster II genes was no more than three. The remaining 11 genes from cluster III encoded LYCE , which regulates lycopene to generate the synthesis of α-carotenoids, which in turn produces lutein. The focus candidate gene Cla belonged to LCYB cluster I, and the LCYB genes of watermelon for gourd were most closely related, with melon, cucumber, and pumpkin relatively distant.

Tomato species with higher levels of lycopene accumulation were used in the process of constructing the phylogenetic tree, and the LCYB genes of watermelon were further related to that of tomato.

The phylogenetic tree analysis demonstrates that the candidate gene identified from the fine mapping results participated in regulating red flesh formation through the process of carotenoid production Figure 8.

Figure 8 Phylogenetic tree analysis of lycopene homologous genes in different Cucurbitaceae crops and tomato. The phylogenetic tree was constructed by combining the lycopene genes of melon, watermelon, gourd, and pumpkin in Cucurbitaceae crops with one lycopene gene in tomato.

A total of 33 homologous genes were divided into three clusters. Red flesh color and lycopene content are major influences on watermelon quality and consumption. In previous research, the red flesh color trait in watermelon was found to be controlled by a single recessive gene Henderson et al.

To analyze the genetic inheritance patterns of red flesh color in watermelon, two F 2 populations and one BC population were tested in our research. The segregation of these three populations further verified that one major recessive gene controlled red flesh color and lycopene content.

The segregation of flesh color in Pop. The homozygous recessive allele of the py gene could inhibit the formation of canary yellow, yielding a pale yellow flesh color.

The results from the HPLC data of Pop. The skewed distribution of lycopene content in Pop. Both LSW and garden parent have red flesh color in their mature fruit, but their segregation was quite different when they were crossed with the white-fleshed PI The proportion of full red-fleshed plants was very low in the segregating populations with a genetic basis for white and red flesh color.

Only two full red-fleshed plants were detected in Pop. The Wf gene has been reported as a white flesh-related gene in watermelon, and the segregation ratio was 12 white : 3 yellow : 1 red in the F 2 generation 74 fruits with white and red flesh color materials Shimotsuma, In the research of Zhang and colleagues, watermelon lines red-fleshed and PI FR light yellowish green flesh color, which is regarded as a white-fleshed parental line were used to form F 2 and F 9 -RIL populations.

The F 2 individuals had a white and yellow : 7 red segregation fitting for a ratio of , implying a duplicate effect between the genes for white and red flesh color. In the RIL population with the same parental materials, the segregation changed to 75 white and yellow : 28 red fitting for a ratio of Zhang et al.

Another Wf gene investigation was reported by Gusmini and Wehner , but in this study, COS, with pale yellow flesh color, was used as white-fleshed material to cross with a canary yellow flesh line, NC One hundred thirty-five canary yellow and 49 white actually pale yellow plants were detected in the F 2 generation, which fit a ratio of In our research, the segregation ratio was somewhat different from that of the previous study, which could be explained by the use of different parental lines.

PI , with a pure white flesh color, was different from COS and PI FR. According to the genotyping data with the gene marker WII04EBsaHI-6 in Pop. The ratio of red alleles and nonred alleles was and for Pop. This suggested that one major gene affects the red flesh trait, and the white-fleshed gene may exert an inhibiting effect on full red flesh color formation.

The white flesh color trait was incomplete dominant to the red flesh trait based on the fact that the two F 1 generations of Pop. The white-fleshed trait would be more complicated than pale yellow and red, which still needs further investigation.

Orange-fleshed plants were segregated in Pop. The carotenoid biosynthesis pathway in plants has been thoroughly investigated Cazzonelli and Pogson, , but little research was focused on QTL analysis of lycopene accumulation in watermelon. Red tomato is also a high-lycopene-content plant, which reportedly undergoes pigment development similar to that of watermelon Grassi et al.

The genes in the carotenoid biosynthesis pathway perform through different action modes to regulate carotenoid accumulation and flesh color formation.

Some genes affect pigments at the level of the transcriptome, such as the phytoene synthase PSY gene, which showed significantly different expression between red- and white-fleshed watermelon accessions Grassi et al.

On the other hand, some genes did not show significant differences in expression among fruits with different flesh colors but still exhibited carotenoid accumulation variance based on enzyme activity; for example, the CmOr and BoOr genes acted as the main functional genes for β-carotene accumulation in melon and cauliflower Lu et al.

By performing stepwise increases in mapping population sizes, marker numbers, and multiple genetic populations, a major effective QTL and candidate gene related to lycopene content and red flesh color was refined in a narrow region of chromosome 4.

The two trait-related QTLs shared the same region, and the red flesh color gave a high correlation with the lycopene content in the F 2 generation for Pop.

Based on the preliminary mapping information, new CAPS markers were developed to narrow down the region from , bp to 41, bp. To verify the stability of the QTL we identified, Pop. The same QTL region and markers showed a high detection efficacy for red flesh color or lycopene content in the two populations through linkage analysis and MAS.

In this study, Cla seemed to be the best candidate gene for lycopene accumulation and red flesh color formation. A nonsynonymous substitution arose from one SNP variation between COS and LSW at the th position in the LCYB gene, and this variation could also be detected in another 20 watermelon sequences.

We also performed a transcriptome analysis of COS and LSW at different time points throughout the whole growth period with the RNA of pulp, the expression level of Cla was not the main reason for red and nonred flesh color formation. The results of our transcriptome analysis were also supported by other research showing that the expression difference in the LCYB gene was among red and nonred watermelon and pumpkin Kang et al.

This might indicate that the accumulation of lycopene was not dependent on the expression of LCYB. It was also likely that the difference in protein levels or functionality may regulate the color of the fruit Wang et al. This seemed contradictory when we combined the results of QTL mapping and transcriptome analysis.

In liverwort, the results of functional identification also proved that the LCYB gene had a lycopene degradation capability to produce β-carotene at the enzyme activity level Takemura et al. It is reasonable to speculate that LCYB may regulate lycopene metabolism through protein level.

The nonsynonymous SNP locus in Cla in the th coding region Figure 5 and Figure S3 may be the key site causing the change in enzyme activity, which may still need further investigation.

While most people agree watermelon tastes good, a majority are unaware of the many health benefits this wonderful fruit contains.

Did you know watermelon has more lycopene than any other fresh fruit or vegetable? Lycopene is an antioxidant linked to decreased risk of cancer, heart disease and age-related eye disorders. Vitamin A is important for skin and eye health.

Vitamin B6 helps your body break down the protein you eat, and is also important for the immune system and nerve function.

Vitamin C helps strengthen the immune system and aids in the absorption of iron. Potassium is helpful in lowering blood pressure and is important for nerve function as well. No wonder it's the most common melon eaten in America and is perfect for staying refreshed and hydrated on a hot summer day.

Food Technol 53 2 — Singh P, Goyal GK Dietary lycopene: Its properties and anticarcinogenic effects. Compr Rev Food Sci Food Saf 7 3 — Naz A, Butt MS, Pasha I, Nawaz H Antioxidant indices of watermelon juice and lycopene extract.

Pak J Nutr — Tlili I, Hdider C, Lenucci MS, Ilahy R, Jebari H, Dalessandro G Bioactive compounds and antioxidant activities during fruit ripening of watermelon cultivars. J Food Comps Anal 24 7 — Hong MY, Hartig N, Kaufman K, Hooshmand S, Figueroa A, Kern M Watermelon consumption improves inflammation and antioxidant capacity in rats fed an atherogenic diet.

Nutr Res 35 3 — Ghavipour M, Saedisomeolia A, Djalali M, Sotoudeh G, Eshranghyan MR, Moghada AM, Wood LG Tomato juice consumption reduces systemic inflammation in overweight and obese females.

Br J Nutr 11 — Ferruzzi MG, Nguyen ML, Sander LC, Rock CL, Schwartz SJ Analysis of lycopene geometrical isomers in biological microsamples by liquid chromatography with coulometric array detection.

J Chromatography B 2 — Korea Health Statistics Korea National Health and Nutrition Examination Survey KNHANES VII Korea Center for Disease Control and Prevention; Seoul, Korea, pp 3— Meyers KJ, Watkins CB, Pritts MP, Liu RH Antioxidant and antiproliferative activities of strawberries.

J Agric Food Chem 51 23 — Shin Y Correlation between antioxidant concentrations and activities of Yuja Citrus junos Sieb ex Tanaka and other citrus fruits. Food Sci Biotechnol 21 5 — Bicanic D, Fogliano V, Luterotti S, Swarts J, Piani G, Graziani G Quantification of lycopene in tomato products: comparing the performances of a newly proposed direct photothermal method and high-performance liquid chromatography.

J Sci Food Agric 85 7 — Bicanic D, Swarts J, Luterotti S, Helander P, Fogliano V, Anese M Optothermistor as a breakthrough in the quantification of lycopene content of thermally processed tomato-based foods: Verification versus absorption spectrophotometry and high-performance liquid chromatography.

J Agric Food Chem 53 9 — Brand-Williams W, Cuvelier ME, Berset C Use of a free radical method to evaluate antioxidant activity. LWT-Food Sci Technol — J Food Comp Anal 24 7 — Re R, Pellegrini N, Proteggente A, Pannala A, Yang M, Rice-Evans C Antioxidant activity applying an improved ABTS radical cation decolorization assay.

Free Radical Biol Med 26 9—10 — Hallmann E The influence of organic and conventional cultivation systems on the nutritional value and content of bioactive compounds in selected tomato types.

J Sci Food Agric 92 14 — Kamiloglu S, Boyacioglu D, Capanoglu E The effect of food processing on bioavailability of tomato antioxidants. J Berry Res 3 2 — Gerard KA, Robert JS Microwave heating of apple mash to improve juice yield and quality.

Food Sci Technol 37 5 — Tlili I, Hdider C, Lenucci MS, Riadh I, Jebari H, Dalessandro G Bioactive compounds and antioxidant activities of different watermelon Citrullus lanatus Thunb. Mansfeld cultivars as affected by fruit sampling area. J Food Comp Anal 24 3 — Lee BH, Kim SY, Cho CH, Chung DK, Chun OK, Kim DO Estimation of daily per capita intake of total phenolics, total flavonoids, and antioxidant capacities from fruit and vegetable juices in the Korean diet based the Korea National Health and Nutrition Examination survey Korean J Food Sci Technol 43 4 — Gahler S, Otto K, Bohm V Alterations of vitamin C, total phenolics, and antioxidant capacity as affected by processing tomatoes to different products.

J Agric Food Chem 51 27 — Klopotek Y, Otto K, Böhm V Processing strawberries to different products alters contents of vitamin c, total phenolics, total anthocyanins, and antioxidant capacity. J Agric Food Chem 53 14 — Nagal S, Kaur C, Choudhary H, Singh J, Bhushan-Singh B, Singh KN Lycopene content, antioxidant capacity and colour attributes of selected watermelon Citrullus lanatus Thunb.

Mansfeld cultivars grown in India. Int J Food Sci Nutr 63 8 — Martinez-Valverde I, Periago MJ, Provan G, Chesson A Phenolic compounds, lycopene and antioxidant activity in commercial varieties of tomato Lycopersicum esculentum. J Sci Food Agric 82 3 — Markovic K, Hruskar M, Vahcic N Lycopene content of tomato products and their contribution to the lycopene intake of Croatians.

Nutr Res 26 11 — Perkins-Veazie P, Collins JK, Pair SD, Roberts W Lycopene content differs among red-fleshed watermelon cultivars. J Sci Food Agric 81 10 — Chun OK, Kim DO, Smith N, Schroeder D, Han JT, Lee CY Daily consumption of phenolics and total antioxidant capacity from fruit and vegetables in the American diet.

J Sci Food Agric 85 10 — Jenab M, Ferrari P, Mazuir M, Tjonneland A, Clavel-chapelon F, Linseisen J, Trichopoulou A, Tumino R, Bueno-de-Mesquita HB, Lund E, Gonzalez CA, JohanssonJ-Key T, Riboli E G S Variations in lycopene blood levels and tomato consumption across European countries based on the European Prospective Investigation into Cancer and Nutrition EPIC study.

J Nutr 8 S—S. Goldbohm RA, Brants HA, Hulshof KF, Van Den Brandt PA The contribution of various foods to intake of vitamin A and carotenoids in The Netherlands. Int J Vitam Nutr Res 68 6 — Pelz R, Schmidt-Faber B, Heseker H Carotenoid intake in the German national food consumption survey.

Z Ernahrungswiss 37 4 — British J Nutr 85 4 — Porrini M, Riso P S What are typical lycopene intakes? Johnson-Down L, Saudny-Unterberger H, Gray-Donald K Food habits of Canadians: lutein and lycopene intake in the Canadian population.

J Am Diet Assoc 7 — PubMed Google Scholar. Lugasi A, Brio L, Hovarie J, Sagi KV, Brandt S, Barna E Lycopene content of foods and lycopene intake in two groups of the Hungarian population.

Nutr Res 23 8 — Wawrzyniak A, Marciniak A, Rajewska J Lycopene content of selected foods available on the Polish market and estimation of its intake. Pol J Food Nutr Sci 14 55 — McNaughton SA, Marks GC, Gaffney P, Williams G, Green A Validation of a food-frequency questionnaire assessment of carotenoid and vitamin E intake using weighed food records and plasma biomarkers: the method of triads model.

Eur J Clin Nutr 59 2 — Vandevijvere S, Cucu T, Vinkx C, Huvaere K, Huybrechts I, Van Loco J Dietary intake of lycopene by the Belgian adult population. Public Health Nutr 17 2 — Mazeikiene A, Jakaitiene A, Karciauskaite D, Kucinskiene ZA, Abaravicius JA, Kaminskas A, Kucinskas V Dietary lycopene and cardiovascular health in ethnic Lithuanians.

Acta Med Litu 22 4 — Download references. Department of Food Engineering, Dankook University, Cheonan, Chungnam, , Republic of Korea. Department of Food Science and Technology, Seoul National University of Science and Technology, Seoul, , Republic of Korea.

You can also search for this author in PubMed Google Scholar. Conceptualization, YK and YS; methodology, HP, YK, and YS; formal analysis, HP, YK, and YS; writing—original draft preparation, HP; writing—review and editing, YK and YS.

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Appl Biol Chem 63 , 50 Download citation. Received : 30 June Accepted : 28 August Published : 04 September Anyone you share the following link with will be able to read this content:.

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Skip to main content. Search all SpringerOpen articles Search. Download PDF. Abstract Tomatoes, watermelons, and processed tomato products contain abundant antioxidant compounds, including lycopene. Introduction As nutrition improved at the national level, the problems related to nutrient deficiency decreased; however, the western dietary pattern has caused nutritional imbalance to emerge, along with various chronic diseases, such as hypertension, cancer, and diabetes [ 1 ].

Materials and methods Materials The samples were purchased from local fruit markets and hypermarkets in the city of Cheonan, South Korea. Total flavonoid analysis The total flavonoid content of samples was measured using the colorimetric assay method [ 17 , 18 ].

Total phenolic analysis The total phenolic content of samples was measured using the Folin-Ciocalteu colorimetric assay method [ 17 , 18 ]. DPPH radical scavenging activity analysis The DPPH radical scavenging activity was measured using a modified method by Shin [ 18 ] and Brand-Williams et al.

ABTS radical scavenging activity analysis The ABTS radical scavenging activity of the extracted samples was measured using a modified method by Floegel et al. Daily intake of tomatoes, watermelons, and their processed products The daily intake values of tomatoes, watermelons, and their processed products were estimated using the Korea National Health and Nutrition Examination Survey KNHANES VII-2 that was conducted by the Korea Center for Disease Control and Prevention in [ 16 ].

Estimation of daily per capita intake of antioxidant compounds and activities To estimate the daily per capita intake of lycopene and the antioxidant compounds and activities of tomatoes, watermelons, and their processed products, the results of the KNHANES and chemical analysis of the measured contents were used.

Statistical analysis For statistical analysis, the SPSS 20 program SPSS Inc. Results and discussion Total flavonoid content The total flavonoid content of tomatoes, watermelons, and their processed products, is as presented in Table 1.

Table 1 Levels of antioxidant content and activities of tomato, tomato juice, tomato ketchup, and watermelon Full size table. Table 3 Estimated daily intake of antioxidant contents and activities from tomato, tomato processed product, and watermelon in the Korean diet according to gender Full size table.

Full size image. Table 5 Comparison of estimated daily intake of lycopene in some countries Full size table. Availability of data and materials The datasets used and analyzed in this study are available from the corresponding authors on reasonable request.

References Kwon JH, Shim JE, Park MK Paik HY Evaluation of fruit and vegetables intake for prevention of chronic disease in Korean adults aged 30 years and over: Using the third Korea national health and nutrition examination survey KNHANES III.