Cucumber fruits with increased sugar content

Abstract

The present invention relates to cultivated cucumber plants comprising a QTL on chromosome 1 and / or 3 and / or chromosome 4 of the cucumber genome conferring increased sugar content of the fruits and / or mint-green flesh color of the fruits and / or a mint-green peel color, and to methods for generating such plants, and their use.

Description

[0001] Cucumber fruits with increased sugar content

[0002] FIELD

[0003] The present invention relates to the field of cucumber breeding. Provided is a Quantitative Trait Locus (QTL) located on chromosome 1 (QTL1. 1) of the cucumber genome, which can be used to increase sugar content of cucumber fruits, especially glucose and fructose content, in cultivated cucumbers (Cucumis sativus var. sativus), such as pickling cucumbers (e.g. American pickling, European pickling types), slicing cucumbers (e.g. American slicing), long cucumbers, short cucumbers, European greenhouse cucumbers or Beit-Alpha type cucumbers. Also, malic acid content is reduced in the cucumber fruits, which is due to the same QTL (QTL1.1). The fruits, therefore, have a sweeter overall taste as at least glucose and fructose content is increased and / or malic acid content is reduced.

[0004] BACKGROUND

[0005] Cultivated cucumber (Cucumis sativus var. sativus L.) is an important vegetable crop worldwide. It belongs to the family Cucurbitaceae . It is thought to originate from South East Asia from wild ancestors with small, bitter fruits, such as Cucumis sativus var. hardwickii.

[0006] The cultivated cucumber genome has seven pairs of chromosomes (n = 7) and a haploid genome size of about 367 Mb (Megabases) with an estimated total of about 26,682 genes. The cucumber genome was the first vegetable genome to be sequenced (Huang etal. 2009, Nature Genetics, Volume 41, Number 12, pl275-1283).

[0007] Fruits are harvested at an immature stage, as mature cucumbers are not marketable. The bitter taste originally present in wild cucumbers has been bred out of cultivated cucumbers, but besides cucumber fruits being nonbitter, there is little variation in the taste of harvested cucumbers and no QTLs or genes have been identified that influence taste.

[0008] Herein a QTL has been identified in a wild cucumber donor, which influences taste of cucumber fruits at harvest stage by increasing glucose and fructose content of the harvest-stage fruits. The QTL was identified in a donor that produces inferior quality fruits, with a yellow-white mesocarp. The QTL is referred to as QTL1. 1, or as “sugar QTL” or “glucose and fructose QTL”. It highly correlates with an increase in degree Brix (or total soluble solids, TSS) and may, therefore, also be referred to as “Brix QTL” or “TSS QTL”.

[0009] In one aspect cultivated cucumber fruits are provided herein (and plants and producing such fruits) which comprise QTL1.1 in homozygous or heterozygous form and produce an increased amount of glucose and fructose in the fruits compared to cucumber fruits lacking QTL1. 1. In another aspect cultivated cucumber fruits are provided herein (and plants and producing such fruits) which have an attractive green ‘mint-green colored’ fruit flesh (at least the mesocarp). The green mint-colored fruit flesh is conferred by a QTL on chromosome 3, referred herein to QTL3. 1. The green mint colored fruit flesh can be combined with different immature fruit peel (or skin or rind) colors as described in ‘Genetic regulation and molecular mechanism of immature cucumber peel color: A review’ (Cui et al. 2023, Vegetable Research 3:9, pages 1-6) or in Genome-wide Association Analysis Reveals a Novel QTL CsFS 1 for Fruit Skin Color in Cucumber (Geng et al., 2022, / / doi.org / 10.21203 / rs.3.rs-578494 / v2).

[0010] In another aspect cucumber fruits are provided herein (and plants and producing such fruits) which have an attractive green ‘mint-green colored’ peel color. The mint-green peel color is conferred by the presence of the mutant White-peel allele, referred to as w-allele (or also referred to as QTL3. 1) in combination with a QTL on chromosome 4, referred to as QTL4. 1 (or enhanced peel color QTL4. 1).

[0011] In yet a further aspect cucumber fruits are provided comprising QTL 1.1 (in homozygous or preferably in heterozygous form), QTL3.1 (preferably in homozygous form), QTL3.2 / w-allele (in homozygous form) and QTL4. 1 (preferably in homozygous form), resulting in cucumber fruits as shown in Figures 1 and 2, i.e. comprising a mint-green fruit flesh color, a mint-green fruit peel color and an increased sugar content. In one aspect QTL 1.1, QTL3.1 and QTL4. 1 are from a wild donor, optionally from the same wild donor, and are obtainable from seeds of which a representative sample has been deposited under Accession number NCIMB 44295.

[0012] Thus, in one aspect cultivated cucumber plants, plant parts and cells comprising an introgression fragment from a wild or primitive cucumber are provided, which introgression fragment comprises QTL1. 1 on chromosome 1, wherein QTL1. 1 confers an increased amount of glucose and fructose being present in the cucumber fruits. The introgression fragment which comprises QTL 1.1 comprises one or more donor SNP nucleotides for the markers selected from the group of SNP_01 to SNP_14. In one aspect also the amount of malic acid is reduced in the cucumber fruits of plants comprising QTL1. 1.

[0013] Furthermore, cultivated cucumber plants, plant parts and cells comprising an introgression fragment from a wild or primitive cucumber are provided, which introgression fragment comprises QTL3. 1 on chromosome 3, wherein QTL3. 1 confers a mint-green fruit flesh color to the cucumber fruits, at least the mesocarp of the fruits has a mint green color. The introgression fragment which comprises QTL3.1 comprises one or more donor SNP nucleotides for the markers selected from the group of SNP_15 to SNP_24. The mint green fruit-flesh color has, in one aspect, an RHS color chart rating Yellow-Green Group 145A and / or 145B (Fan 3 Blue-Green Green Green-Yellow Groups of the RHS Color Chart, 5thEdition, The Royal Horticultural Society 2007). In one aspect the RHS color chart rating of the fruit flesh is 145B and / or 145C; or 145A and / or 145B and / or 145C.

[0014] Especially nearer to the seed cavity the flesh color is in one aspect 145C. See also Figure 1 and 2.

[0015] In one aspect the cultivated cucumber plants, plant parts, cells, fruits or seeds are provided herein which comprise both an introgression fragment comprising QTL1.1 (conferring an increased glucose and fructose content to the fruits produced by the plant) and an introgression fragment comprising QTL3. 1 (conferring mintgreen colored fruit flesh to the fruits produced by the plant). In one aspect one or both introgression fragments or one or both QTLs are obtainable from seeds deposited under accession number NCIMB 44295. In one aspect the introgression fragment comprising QTL1. 1 is present in heterozygous form or in homozygous form and / or the introgression fragment comprising QTL3. 1 is present in heterozygous form or in homozygous form in the plant, seeds, plant parts, fruits or cells.

[0016] A commercial cucumber variety with a distinct fruit color is the variety Quinton Fl (Rijk Zwaan). It has a pale light-green exterior skin color and a pale cream-colored flesh. Its taste is bland and watery. It does not contain QTL3. 1, as the SNP marker haplotype is different (data not shown). It also does not comprise QTL1. 1, see e.g. Figures 3-5.

[0017] In yet another aspect the cultivated cucumber plants, plant parts, cells, fruits or seeds are provided herein which comprise an introgression fragment comprising QTL1. 1 (conferring an increased glucose and fructose content to the fruits produced by the plant) and the White-peel mutant allele (w / w or QTL3.2) and an introgression fragment comprising QTL4. 1 (conferring mint-green colored fruit peel to the fruits produced by the plant). In one aspect QTL1. 1 and QTL4. 1 are obtainable from seeds deposited under accession number NCIMB 44295. In one aspect the mutant w-allele is obtainable from seeds deposited under accession number NCIMB 44295. In one aspect the introgression fragment comprising QTL1. 1 is present in heterozygous form or in homozygous form and the w / w-allele is present in homozygous form and the introgression fragment comprising QTL4. 1 is present in homozygous form in the plant, seeds, plant parts, fruits or cells.

[0018] FIGURES

[0019] Figure 1 and 2 - fotos of cultivated cucumber fruits (whole fruits and cut fruit) comprising the introgression fragment with QTL1. 1 (conferring increased glucose and fructose content) and the introgression fragment with QTL3. 1 (conferring mint-green fruit flesh color, e.g. RHS Yellow-Group 145A and / or 145B and / or 145C; e.g. 145C is the color near the seed cavity). The fruits also have a mint green colored peel (e.g. RHS Yellow-Group 145A and / or 145B and / or 145C). The peel and mesocarp are therefore mint-green in color. The mint-green peel color is conferred by the w / w-allele and the introgression fragment comprising QTL4. 1 in homozygous form (enhanced peel color QTL). Figure 3 - average amount of glucose in the cultivated cucumber fruits (of deposited line) comprising the introgression of the QTL1.1 in homozygous form, compared to the recurrent parent and compared to variety Quinton Fl (Rijk Zwaan).

[0020] Figure 4 - average amount of fructose in the cultivated cucumber fruits (of deposited line) comprising the introgression of the QTL1.1 in homozygous form, compared to the recurrent parent and compared to variety Quinton Fl (Rijk Zwaan).

[0021] Figure 5 - average amount of malic acid in the cultivated cucumber fruits (of deposited line) comprising the introgression of the QTL1.1 in homozygous form, compared to the recurrent parent and compared to variety Quinton Fl (Rijk Zwaan).

[0022] Figure 6 - schematic diagram of the location of four QTLs on the seven cucumber chromosomes: QTL1.1 (sugar, glucose and fructose), QTL3. 1 (fruit flesh color), QTL3.2 (exterior skin color, White (W) locus, QTL4. 1 (skin / peel color enhancer).

[0023] GENERAL DEFINITIONS

[0024] The indefinite article "a" or "an" does not exclude the possibility that more than one of the elements is present, unless the context clearly requires that there be one and only one of the elements. The indefinite article "a" or "an" thus usually means "at least one".

[0025] When “seeds of a plant” are referred to, these either refer to seeds from which the plant can be grown or to seeds produced on the plant, after self-fertilization or cross-fertilization.

[0026] "Plant variety" is a group of plants within the same botanical taxon of the lowest grade known, which (irrespective of whether the conditions for the recognition of plant breeder’s rights are fulfilled or not) can be defined on the basis of the expression of characteristics that result from a certain genotype or a combination of genotypes, can be distinguished from any other group of plants by the expression of at least one of those characteristics, and can be regarded as an entity, because it can be multiplied without any change. Therefore, the term “plant variety” cannot be used to denote a group of plants, even if they are of the same kind, if they are all characterized by the presence of a few loci or genes (or phenotypic characteristics due to these specific loci or genes), but which can otherwise differ from one another enormously as regards the other loci or genes. Thus, e.g. a plant defined only by the presence of one or more QTLs is not a plant variety, as thousands of other genes which define a plant variety are undefined and a plant defined only by the presence of one or more QTLs is not uniform and stable for these thousands of genes and the characteristics conferred by these genes. QTL1. 1 and / or QTL3. 1 and / or QTL3.2 and / or QTL4. 1 can be used to develop many different plant varieties, e.g. a long cucumber variety which is uniform and stable for all its physiological and morphological characteristics such as leaf size or shape, leaf margins and color, fruit size and color, warts, plant height, etc. and which also comprises QTL1.1 and / or QTL3.1 and / or QTL3.2 and / or QTL4.1.

[0027] “Fl, F2, F3, etc.” refers to the consecutive related generations following a cross between two parent plants or parent lines. The plants grown from the seeds produced by crossing two plants or lines is called the Fl generation. Selfing the Fl plants results in the F2 generation, etc.

[0028] “F 1 hybrid” plant (or F 1 hybrid seed) is the generation obtained from crossing two inbred parent lines. Thus, Fl hybrid seeds are seeds from which Fl hybrid plants grow. Fl hybrids are more vigorous and higher yielding, due to heterosis. Inbred lines are essentially homozygous at most loci in the genome.

[0029] A “plant line” or “breeding line” refers to a plant and its progeny. As used herein, the term "inbred line" refers to a plant line which has been repeatedly selfed and is nearly homozygous. Thus, an “inbred line” or “parent line” refers to a plant which has undergone several generations (e.g. at least 5, 6, 7 or more) of inbreeding, resulting in a plant line with a high uniformity.

[0030] The term “allele(s)” means any of one or more alternative forms of a gene at a particular locus, all of which alleles relate to one trait or characteristic at a specific locus. In a diploid cell of an organism, alleles of a given gene are located at a specific location, or locus (loci plural) on a chromosome. One allele is present on each chromosome of the pair of homologous chromosomes. A diploid plant species may comprise a large number of different alleles at a particular locus. These may be identical alleles of the gene (homozygous) or two different alleles (heterozygous). Thus, for example reference may herein be made to a “sugar allele” or “glucose and fructose allele” of QTL1. 1 or to the “mint-green flesh color allele” of QTL3. 1. In a SNP marker assay one may also refer to ‘alleles’, wherein one ‘allele’ is the donor SNP nucleotide and the other ‘allele’ is the alternative nucleotide (e.g. the recurrent parent SNP nucleotide).

[0031] The term “gene” means a (genomic) DNA sequence comprising a region (transcribed region), which is transcribed into a messenger RNA molecule (mRNA) in a cell, and an operably linked regulatory region (e.g. a promoter). Different alleles of a gene are thus different alternatives form of the gene, which may be in the form of e.g. differences in one or more nucleotides of the genomic DNA sequence (e.g. in the promoter sequence, the exon sequences, intron sequences, etc.), mRNA and / or amino acid sequence of the encoded protein.

[0032] The term “locus” (loci plural) means a specific place or places or a site on a chromosome where for example a QTL, a gene or genetic marker is found. The sugar locus (or ‘glucose and fructose locus’ or QTL1. 1 locus) is, thus, the location in the genome of cucumber, where QTL1. 1 is found. In cultivated cucumber the QTL1. 1 is found on chromosome 1 (using the chromosome assignment of Huang et al. 2009, Nature Genetics, Volume 41, Number 12, pl275-1283) and world wide web at / / cucurbitgenomics.org / , the genome of Cucumber (Chinese Long) V3) i.e. it is introgressed into the cultivated cucumber genome (i.e. onto chromosome 1) from a wild or primitive cucumber donor. Likewise QTL3.1 and QTL4.1 provided herein are introgressed from a wild or primitive donor into cultivated cucumber, making reference to the cultivated cucumber chromosomes as designated by the Cucumber (Chinese Long) V3 genome (referred to herein as the cucumber reference genome).

[0033] A "quantitative trait locus", or "QTL" is a chromosomal locus that encodes for one or more alleles that affect the expressivity of a continuously distributed (quantitative) phenotype. The sugar locus conferring quantitative trait locus is named QTL1. 1. The interior color locus, conferring mint-green color to the cucumber fruit flesh, is named QTL3. 1 herein. The exterior / peel color enhancer locus, enhancing peel color in a w / w background, is named QTL4. 1 herein.

[0034] “Cucumber genome” and “physical position on the cucumber genome” and “chromosome 1 or 3 or 4” refer to the physical genome of cultivated cucumber, world wide web at / / cucurbitgenomics.org / , the genome of Cucumber (Chinese Long) v3), and the physical chromosomes and the physical position on the chromosomes. So, for example SNP_01 is located at the nucleotide (or ‘base’) positioned physically at nucleotide 3530044 of chromosome 1 of the reference genome.

[0035] “Physical distance” between loci (e.g. between molecular markers and / or between phenotypic markers) on the same chromosome is the actually physical distance expressed in bases or base pairs (bp), kilo bases or kilo base pairs (kb) or megabases or mega base pairs (Mb).

[0036] “Genetic distance” between loci (e.g. between molecular markers and / or between phenotypic markers) on the same chromosome is measured by frequency of crossing-over, or recombination frequency (RF) and is indicated in centimorgans (cM). One cM corresponds to a recombination frequency of 1%. If no recombinants can be found, the RF is zero and the loci are either extremely close together physically or they are identical. The further apart two loci are, the higher the RF.

[0037] “Introgression fragment” or “introgression segment” or “introgression region” refers to a chromosome fragment (or chromosome part or region) which has been introduced into another plant of the same or related species by crossing or traditional breeding techniques, such as backcrossing, i.e. the introgressed fragment is the result of breeding methods referred to by the verb “to introgress” (such as backcrossing). In cucumber, wild or primitive cucumber accessions (e.g. landraces) or wild relatives of cultivated cucumber can be used to introgress fragments of the wild genome into the genome of cultivated cucumber, Cucumis sativus var. sativus L. Such a cultivated cucumber plant thus has a “genome of cultivated Cucumis sativus var. sativus”, but comprises in the genome a fragment of a wild or primitive cucumber or of a wild relative of cucumber, e.g. an introgression fragment of a related wild Cucumis sativus genome, such as Cucumis sativus var. hardwickii, C. sativus var. sikkimensis, Cucumis sativus var. xishuangbannesis, or another wild cucumber or wild relative of cucumber. So, for example, a cultivated cucumber is provided herein comprising a genome of cultivated cucumber, and in that genome there is an introgression fragment on chromosome 1 of cultivated cucumber which confers an increased glucose and fructose content compared to the cultivated cucumber genome lacking the introgression fragments (and having a chromosomes 1 of cultivated cucumber, without the introgression fragments). It is understood that the term “introgression fragment” never includes a whole chromosome, but only a part of a chromosome. The introgression fragment can be large, e.g. even three quarter or half of a chromosome, but is preferably smaller, such as about 15 Mb or less, such as about 10 Mb or less, about 9 Mb or less, about 8 Mb or less, about 7 Mb or less, about 6 Mb or less, about 5 Mb or less, about 4 Mb or less, about 3 Mb or less, about 2.8 Mb, 2.5 Mb or 2 Mb or less, about 1 Mb (equals 1,000,000 base pairs) or less, or about 0.5 Mb (equals 500,000 base pairs) or less, such as about 200,000 bp (equals 200 kilo base pairs) or less, about 100,000 bp (100 kb) or less, about 50,000 bp (50 kb) or less, about 25,000 bp (25 kb) or less.

[0038] “Cultivated cucumber” or “domesticated cucumber” refers to plants of Cucumis sativus var. sativus i.e. varieties, breeding lines or cultivars, cultivated by humans and having good agronomic characteristics, especially producing edible and marketable fruits of good size and quality and uniformity; such plants are not “wild cucumber” or “primitive cucumber” plants , i.e. plants which generally have much poorer yields and poorer agronomic characteristics than cultivated plants and are less uniform genetically and in their physiological and / or morphological characteristics. “Wild plants” of “wild cucumber” include for example ecotypes, landraces or wild accessions or wild relatives of a species. Cultivated cucumber plants (lines or varieties) can also be distinguished from wild or primitive cucumber accessions by the significantly lower amount of SNPs (less than 2,000,000 SNPs) and INDELs (insertions / deletions of shorter than 5bp; less than 150,000 INDELs) in the genome and their significantly lower nucleotide diversity (equal to or less than 2.3 x 10'3K), as described in Table 1 of Qi et al, Nature Genetics December 2013, Vol 45, No. 12, pages 1510 - 1518. SNP numbers, INDEL numbers and nucleotide diversity can be determined as described herein, especially in the section ‘Online Methods’.

[0039] “Indian cucumber group” refers to wild or wild relatives of cucumbers from India, having a high amount of SNPs (more than 3,000,000 SNPs) and INDELs (insertions / deletions of shorter than 5bp; more than 200,000 INDELs) in the genome and high nucleotide diversity (more than 3.0 x 10'3it or even more than 4.0 x 10'3it).

[0040] “Eurasian cucumber group” refers to cultivated cucumbers from central or western Asia, Europe and the United States, having a low amount of SNPs (less than 2,000,000 SNPs, or less than 1,500,000 SNPs) and INDELs (insertions / deletions of shorter than 5bp; less than 150,000 INDELs) in the genome and a low nucleotide diversity (equal to or less than 2.3 x 10'3K, preferably less than 2.0 x 10'3it).

[0041] “East Asian cucumber group” refers to cultivated cucumbers from East Asia, such as China, Korea and Japan, having a low amount of SNPs (less than 2,000,000 SNPs, or less than 1,500,000 SNPs) and INDELs (insertions / deletions of shorter than 5bp; less than 150,000 INDELs, preferably less than 100,000) in the genome and a low nucleotide diversity (equal to or less than 2.3 x 10'3it, preferably less than 2.0 x 10'3it or even less than 1.5 x 10'3it).

[0042] “Xishuangbanna cucumber group” refers to cucumbers from the Xishuangbanna region of China, having a low amount of SNPs (less than 2,000,000 SNPs, or less than 1,500,000 SNPs or even less than 100,000 SNPs) and INDELs (insertions / deletions of shorter than 5bp; less than 150,000 INDELs, preferably less than 100,000) in the genome and a low nucleotide diversity (equal to or less than 2.3 x 10'3it, preferably less than 2.0 x 10'3it or even less than 1.5 x 10'3it).

[0043] “Wild cucumber” or “primitive cucumber” refers to C. sativus var. sativus which generally have much poorer yields and poorer agronomic characteristics than cultivated plants and are less uniform genetically and in their physiological and / or morphological characteristics. Wild plants include for example ecotypes, landraces or wild accessions or wild relatives of a species.

[0044] “Wild relatives of cucumber” refer to Cucumis sativus var. hardwickii, C. sativus var. sikkimensis, Cucumis sativus var. xishuangbannesis.

[0045] “Landrace(s)” refers to primitive cultivars of Cucumis sativus var. sativus developed in local geographic regions, which often show a high degree of genetic variation in their genome and exhibit a high degree of morphological and / or physiological variation within the landrace (e.g. large variation in fruit size, etc.), i.e. are significantly less uniform than cultivated cucumber. Landraces are, therefore, herein included in the group “wild cucumber”, which is distinct from “cultivated cucumber”.

[0046] “Uniformity” or “uniform” relates to the genetic and phenotypic characteristics of a plant line or variety. Inbred lines are genetically highly uniform as they are produced by several generations of inbreeding. Likewise, and the F 1 hybrids which are produced from such inbred lines are highly uniform in their genotypic and phenotypic characteristics and performance.

[0047] The term “sugar allele” or “glucose and fructose allele” refers to an allele found at the locus QTL1.1 introgressed into cultivated cucumber (onto cultivated C. sativus var. sativus chromosome 1) from a wild or primitive cucumber. The term “sugar allele”, thus, also encompasses alleles obtainable from the seeds deposited herein or from other Cucumis accessions. When one (heterozygous) or two (homozygous) sugar alleles are present at the locus QTL1. 1, in the genome, the plant line or variety comprises a significantly higher glucose and fructose levels in the fruits than the control or genetic control lacking the QTL. In cultivated cucumber plant lacking the introgression fragments, the C. sativus var. sativus allele found at the same locus on chromosome 1 is herein referred to as “wild type” allele (wt). The haplotype or genotype of the SNP markers provided herein is indicative of the wild type or of the QTL1. 1 in homozygous or heterozygous form. E.g. the genotype of SNP_01 indicative of QTL1.1 is e.g. ‘CA’ (QTLl.l / wt) or ‘CC’ (QTLl. l / QTL1.1) while the genotype indicative of the wild type, i.e. of the cultivated cucumber, is e.g. ‘AA’ (wt / wt). The genotype of SNP_02 indicative of QTLl. l is e.g. ‘AT’ (QTLl. l / wt) or ‘AA’ (QTLl.l / QTLl.l) while the genotype indicative of the wild type, i.e. of the cultivated cucumber, is e.g. ‘TT’ (wt / wt). Likewise the donor SNP haplotype for SNP_01 and SNP_02 is ‘C-A’ and the SNP genotype for SNP_01 and SNP_02 is ‘CC-AA’). See Table 2 and Table 3.

[0048] The term “flesh color allele” or “fruit flesh color allele” or ‘mint-green fruit flesh color-conferring allele’ refers to an allele found at the locus QTL3.1 introgressed into cultivated cucumber (onto cultivated C. sativus var. sativus chromosome 3) from a wild or primitive cucumber. The term “fruit flesh color allele”, thus, also encompasses alleles obtainable from the seeds deposited herein or from other Cucumis accessions. When one (heterozygous) or two (homozygous) alleles are present at the locus QTL3. 1, in the genome, the plant line or variety comprises a mint-green fruit flesh color. In cultivated cucumber plant lacking the introgression fragments, the C. sativus var. sativus allele found at the same locus on chromosome 3 is herein referred to as “wild type” allele (wt). The haplotype or genotype of the SNP markers provided herein is indicative of the wild type or of the QTL3. 1 in homozygous or heterozygous form. See Table 5 and Table 6.

[0049] The term “white allele” or “mutant white allele” or “w-allele” refers to a mutant allele found at the White-peel locus or W-locus (also referred to as QTL3.2 herein) that may be present in or generated in cultivated cucumber (at the W-locus on cultivated C. sativus var. sativus chromosome 3). The term “w-allele” encompasses the mutant w-allele obtainable from the seeds deposited herein or from other cucumbers as described in the literature. See e.g. Example 5 and Table 9.

[0050] The term “peel / skin color enhancer allele” or “exterior color enhancing allele” refers to an allele found at the locus QTL4. 1 introgressed into cultivated cucumber (onto cultivated C. sativus var. sativus chromosome 4) from a wild or primitive cucumber. The term “peel / skin color enhancer allele”, thus, also encompasses alleles obtainable from the seeds deposited herein or from other Cucumis accessions. When two (homozygous) alleles are present at the locus QTL4. 1 in a genome which is homozygous for the white mutant allele (w / w), the plant line or variety comprises a mint-green fruit peel color. In cultivated cucumber plant lacking the introgression fragments, the C. sativus var. sativus allele found at the same locus on chromosome 4 is herein referred to as “wild type” allele (wt). The haplotype or genotype of the SNP markers provided herein is indicative of the wild type or of the QTL4. 1 in homozygous or heterozygous form. See Table 14 and Table 15.

[0051] A genetic element, an introgression fragment, or a gene or allele conferring a trait (such as fruit sugar levels) is said to be “obtainable from” or can be “obtained from” or “derivable from” or can be “derived from” or “as present in” or “as found in” a plant or seed or tissue or cell if it can be transferred from the plant or seed in which it is present into another plant or seed in which it is not present (such as a line or variety) using traditional breeding techniques without resulting in a phenotypic change of the recipient plant apart from the addition of the trait conferred by the genetic element, locus, introgression fragment, gene or allele. The terms are used interchangeably and the genetic element, locus, introgression fragment, gene or allele can thus be transferred into any other genetic background lacking the trait. Not only seeds deposited and comprising the genetic element, locus, introgression fragment, gene or allele can be used, but also progeny / descendants from such seeds which have been selected to retain the genetic element, locus, introgression fragment, gene or allele, can be used and are encompassed herein, such as commercial varieties developed from the deposited seeds or from descendants thereof. Whether a plant (or genomic DNA, cell or tissue of a plant) comprises the same genetic element, locus, introgression fragment, gene or allele as obtainable from the deposited seeds can be determined by the skilled person using one or more techniques known in the art, such as phenotypic assays, whole genome sequencing, molecular marker analysis (e.g. SNP haplotyping or SNP genotyping), trait mapping, chromosome painting, allelism tests and the like, or combinations of techniques.

[0052] “SNP marker” refer herein to single nucleotide polymorphisms of a genomic sequence linked to e.g. QTL1. 1 or QTL3.1 or QTL4.1 whereby a specific nucleotide, which is also referred to as the ‘donor SNP nucleotide’ (e.g. for SNP_01 a Cytosine at nucleotide 102 of SEQ ID NO: 1, or a Cytosine at nucleotide 102 of a sequence comprising at least 95%, 96%, 97%, 98%, 99% sequence identity to SEQ ID NO: 1, or a sequence comprising the specific nucleotide, is linked to the QTL. This nucleotide, or sequence comprising the nucleotide, is also referred to as the ‘SNP genotype’ or ‘SNP nucleotide’ (or SNP haplotype when referring to several markers) of the plant or plant part, and SNP 01 may be ‘C’ (haploid, on one chromosome) or ‘CC’ (diploid, on both chromosomes). For example, the donor SNP nucleotide of one or more of markers SNP_01 to SNP_14 are linked to QTL1. 1 and are e.g. present on the introgression fragment which comprises QTL1. 1. Markers SNP_15 to SNP_24 are linked to QTL3. 1 and are e.g. present on the introgression fragment which comprises QTL3. 1. Markers SNP_25 to SNP_33 are linked to QTL4.1 and are e.g. present on the introgression fragment which comprises QTL4.1. A SNP marker can be referred to as a nucleotide in a specific sequence (e.g. the donor nucleotide Cytosine at nucleotide 102 of SEQ ID NO: 1 or at the equivalent nucleotide in a sequence comprising at least 95%, 96%, 97%, 98% or 99% sequence identity to SEQ ID NO: 1), or alternatively as a donor nucleotide at a specific position on the chromosome of the reference genome, e.g. the donor nucleotide for SNP_01 is a Cytosine at nucleotide position 3530044 of chromosome 1 of the reference genome Chinese Long V3. The position is shown in Table 2 (for QTL1. 1) and Table 5 (for QTL3. 1) and Table 14 (for QTL4. 1). It is noted that herein the donor SNP nucleotides are referred to as the nucleotide on the plus-strand of a sequence with respect to the reference genome. So, a Cytosine for SNP_01 at nucleotide 102 of SEQ ID NO: 1 refers to the Cytosine at nucleotide 102 of SEQ ID NO: 1 because SEQ ID NO: 1 is the plus-strand with respect of the reference genome. All given sequences are herein on the plus strand as indicated in e.g. Tables 2, 4 and 14. All SNP nucleotides and haplotypes are referred to herein are the plus-strand nucleotides of a certain sequence. Thus, reference to a Cytosine (herein a plus-strand nucleotide) at nucleotide 102 of SEQ ID NO: 1 (herein a plus strand sequence, see Table 2) is understood to refer to a Cytosine at nucleotide 102 of SEQ ID NO: 1 or a Guanine at nucleotide 102 of the complement strand of SEQ ID NO: 1.

[0053] The ‘haplotype ’or “haploid genotype” refers to the haploid genotype of several genetic loci in a plant, especially of several SNP markers or several sequences comprising the SNP markers. For QTL 1.1 the SNP haplotype may thus be the haploid genotype of at least 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12 or more (e.g. all 14) SNP markers of SNP_01 to SNP_14 (or of the sequences comprising the SNP markers). For example, the plant comprising QTL1. 1 may comprise a ‘C’ for SNP_01 at nucleotide 102 in SEQ ID NO: 1 (or at nucleotide 102 of a sequence which is at least 95%, 96%, 97%, 98% or 99% identical to SEQ ID NO: 1), a ‘A’ for SNP_02 at nucleotide 102 in SEQ ID NO: 2 (or at nucleotide 102 of a sequence which is at least 95%, 96%, 97%, 98% or 99% identical to SEQ ID NO: 2), a ‘C’ for SNP_03 in SEQ ID NO: 3 (or at nucleotide 102 of a sequence which is at least 95%, 96%, 97%, 98% or 99% identical to SEQ ID NO: 3), it thus has the SNP haplotype C-A-C for SNP_01 to SNP_03, which is the SNP haplotype of SNP_01 to SNP_03 of the wild cucumber donor (also referred to as donor SNP haplotype). A diploid plant homozygous for the QTL would have the SNP genotype CC-AA-CC for SNP_01 to SNP_03.

[0054] A “Variant” or “orthologous” sequence or a “variant QTL 1. 1 or QTL3. 1 or QTL4. 1” or a “variant of QTL1. 1 or QTL3.1 or QTL4.1” refers to a fruit sugar QTL (QTL1.1) or internal mint green color QTL (QTL3.1) or peel color enhancer QTL (QTL4. 1), or an introgression fragment comprising the QTL, which is derived from a different wild or primitive cucumber donor plant than the QTL 1.1 or QTL3.1 or QTL4.1 present in NCIMB44295. Such a variant QTL can e.g. be identified as having the same SNP haplotype as the QTLs present in NCIMB44295 for at least 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13 or 14 markers (preferably consecutive markers) selected from SNP_01 to SNP_14 for a variant of QTL1. 1 or for at least 2, 3, 4, 5, 6, 7, 8, 9 or 10 markers (preferably consecutive markers) selected from SNP_15 to SNP_24 for a variant of QTL3. 1 or for at least 2, 3, 4, 5, 6, 7, 8 or 9 markers (preferably consecutive markers) selected from SNP_25 to SNP_33 for a variant of QTL4.1. In addition, the variant QTL confers (at least in homozygous form) an increase in fruit glucose and fructose content as described herein. An “increased (or enhanced) fruit sugar content” or a “significantly increased (or enhanced) fruit glucose and fructose content” refers to a cultivated cucumber plant, plant line, hybrid or variety comprising QTL1. 1 (or a variant thereof), having (due to the QTL, especially when in homozygous form) a higher average fruit glucose and fructose content at harvest stage compared to the control plant lacking the QTLs, preferably the genetic control plant or recurrent parent. Preferably the average fruit glucose and fructose level of the line or variety is increased by e.g. at least 5%, 6%, 7%, 8%, 9%, 10%, 11% or more when the QTL1. 1 (or a variant thereof) is in heterozygous form and by e.g. at least 5%, 6%, 7%, 8%, 9%, 11%, 12%, 13%, 14%, 15%, 16%, 17%, 18% or 19% or more when the QTL1.1 (or a variant thereof) is in homozygous form. For example, if the recurrent parent plant line lacking the QTLs has an average fruit glucose and fructose level of 22 g / L (as e.g. measured using the method described in the Examples), the introduction of QTL 1.1 into that line, especially in homozygous form, increases the average fruit glucose and fructose level of the line to at least 23 g / L.

[0055] “Control plant” is a cultivated cucumber genotype, breeding line, hybrid or variety lacking the introgression fragments. The control plant is preferably of the same type as the plant comprising the introgression fragment(s), e.g. long cucumber type, pickling type, short cucumber type, slicer type, etc. For example, the original parent line into which the QTLs are / were introgressed (also referred to as the recurrent parent) is a suitable control. For example Nunhems variety Beesan Fl may be used as suitable control e.g. for short cucumbers.

[0056] “Genetic control” is a cultivated cucumber genotype, breeding line, variety or hybrid which has the same or very similar cultivated genome as the cucumber plant comprising the one or more introgression fragments except that it lacks the introgressions, e.g. chromosome 1 and / or 3 and / or 4 are “wild type”, i.e. cultivated cucumber genome. This is for example a backcross line in the backcrossing program which does not contain the introgression fragments.

[0057] The term “marker assay” refers to a molecular marker assay or genotyping assay which can be used to test whether on cultivated C. sativus var. sativus chromosome 1 and / or 3 and / or 4 an introgression from a wild or primitive cucumber is present which introgression fragment comprises the sugar QTL 1.1 and / or the interior mint green color QTL3.1 and / or the enhanced peel color QTL4.1 (or a variant of any of these) by e.g. determining the genotype or haplotype of any one or more markers linked to the QTL1. 1 and / or QTL3. 1 and / or QTL4.1, e.g. the genotype or haplotype of one or more SNP markers selected from SNP_01 to SNP_14 for QTL 1.1 or the genotype or haplotype of one or more SNP markers selected from SNP_15 to SNP_24 for QTL3.1 or the genotype or haplotype of one or more SNP markers selected from SNP_25 to SNP_33 for QTL4.1. “Flanking markers” are markers which are on either side of the QTL, i.e. the QTL is located on the chromosomal region in-between the flanking markers, e.g. the QTL1. 1 (or a variant QTL1. 1) is in one aspect in between SNP_01 at nucleotide 102 of SEQ ID NO: 1, corresponding to nucleotide 3530044 of chromosome 1 of the reference genome, and SNP_14 at nucleotide 102 of SEQ ID NO: 14, corresponding to nucleotide 6316306 of chromosome 1 of the reference genome; QTL3. 1 (or a variant QTL3. 1) is in one aspect in between SNP_15 at nucleotide 102 of SEQ ID NO: 15, corresponding to nucleotide 1128135 of chromosome 3 of the reference genome, and SNP_24 at nucleotide 102 of SEQ ID NO: 24, corresponding to nucleotide 2585865 of chromosome 3 of the reference genome; QTL4. 1 (or a variant QTL4. 1) is in one aspect in between SNP_25 at nucleotide 102 of SEQ ID NO: 25, corresponding to nucleotide 9330448 of chromosome 4 of the reference genome, and SNP_33 at nucleotide 102 of SEQ ID NO: 33, corresponding to nucleotide 10300571 of chromosome 4 of the reference genome, see Table 2, 5 and 14. As the peak marker (which is SNP_15) for QTL3.1 is at the beginning of chromosome 3, in one aspect QTL3.1 is located in the region starting at the beginning of chromosome 3 (nucleotide 1 of chromosome 3) and ending at SNP_24, corresponding to nucleotide 2585865.

[0058] The SNP markers provided herein are located in the given order on the introgression fragment. “Consecutive” markers refer to markers in the same consecutive order, so e.g. two consecutive markers may be SNP_01 and SNP_02; SNP_02 and SNP_03; SNP_03 and SNP_04, etc. and three consecutive markers may be SNP_01 and SNP_02 and SNP_03; SNP_02 and SNP_03 and SNP_04; etc.

[0059] “Average” or “mean” refers herein to the arithmetic mean and both terms are used interchangeably. The term “average” or “mean” thus refers to the arithmetic mean of several measurements. The skilled person understands that the phenotype of a plant line or variety depends to some extent on growing conditions and that, therefore, arithmetic means of at least 5, 6, 7, 8, 9, 10, 15, 20, 30, 40, 50 or more plants (or plant parts) are measured, preferably in randomized experimental designs with several replicates and suitable control plants grown under the same conditions in the same experiment. “Statistically significant” or “statistically significantly” different or “significantly” different refers to a characteristic of a plant line or variety that, when compared to a suitable control (e.g. the genetic control) show a statistically significant difference in that characteristic (e.g. the p-value is less than 0.05, p < 0.05, using ANOVA) from the (mean of the) control.

[0060] A “recombinant chromosome” refers to a chromosome having a new genetic makeup arising through crossing- over between homologous chromosomes, e.g. a “recombinant chromosome 1”, i.e. a chromosome 1 which is not present in either of the parent plants and arose through a rare double crossing-over event between homologous chromosomes of a chromosome 1 pair. Herein, for example, recombinant cucumber chromosome 1 is provided comprising an introgression fragment from a wild or primitive cucumber donor. The same applies for chromosome 3 and chromosome 4. The term “traditional breeding techniques” encompasses herein crossing, backcrossing, selfing, selection, double haploid production, embryo rescue, protoplast fusion, marker assisted selection, mutation breeding etc., all as known to the breeder (i.e. methods other than genetic modification / transformation / transgenic methods), by which, for example, a recombinant chromosome 1 or 3 or 4 can be obtained, identified and / or transferred.

[0061] “Backcrossing” refers to a breeding method by which a (single) trait, such as a sugar QTL, can be transferred from a (generally inferior) genetic background (e.g. a wild or primitive cucumber; also referred to as “donor”) into a (generally superior) genetic background (also referred to as “recurrent parent”), e.g. cultivated cucumber. An offspring of a cross (e.g. an Fl plant obtained by crossing a wild or primitive cucumber with a cultivated cucumber; or an F2 plant or F3 plant, etc., obtained from selfing the Fl) is “backcrossed” to the parent with the superior genetic background, e.g. to the cultivated parent. After repeated backcrossing, the trait of the (generally inferior) genetic background will have been incorporated into the (generally superior) genetic background.

[0062] “Marker assisted selection” or “MAS” is a process of using the presence of molecular markers (such as SNP markers or INDEL markers), which are genetically linked to a particular locus or to a particular chromosome region (e.g. introgression fragment), to select plants for the presence of the specific locus or region (introgression fragment). For example, a molecular marker physically linked to a sugar QTL, can be used to detect and / or select cucumber plants comprising the sugar QTL on chromosome 1. The closer the linkage of the molecular marker to the locus, the less likely it is that the marker is dissociated from the locus through meiotic recombination. Likewise, the closer two markers are linked to each other the less likely it is that the two markers will be separated from one another (and the more likely they will co -segregate as a unit).

[0063] “Complementary strands” refer to two strands of complementary sequence and may be referred to as sense (or plus) and anti-sense (or minus) strands for double stranded DNA. For any of the sequences provided herein only one strand of the sequence is given, but the complementary strand of the given strand is also encompassed herein. The complementary nucleotides of DNA are A complementary to T, and G complementary to C.

[0064] “Oligonucleotides” or “oligos” or “oligonucleotide primers or probes” are short, single-stranded polymers of nucleic acid, e.g. at least 10, 11, 12, 13, 14, 15, 16, 17, 18, 19, 20, 21, 22, 23, 24 or more nucleotides in length. Oligos may be unmodified or modified with a variety of chemistries depending on their intended use, for example, the addition of 5' or 3' phosphate groups to enable ligation or block extension, respectively, labelling with radionuclides or fluorophores and / or quenchers for use as probes, the incorporation of thiol, amino, or other reactive moieties to enable the covalent coupling of functional molecules such as enzymes, and extension with other linkers and spacers of diverse functionality. DNA oligos are the most commonly used, but RNA oligos are also available. The length of an oligo is usually designated by adding the suffix -mer. For example, an oligonucleotide with 19 nucleotides (bases) is called a 19-mer. For most uses, oligonucleotides are designed to base-pair with a strand of DNA or RNA. The most common use for oligonucleotides is as primers for PCR (polymerase chain reaction). Primers are designed with at least part of their sequence complementary to the sequence targeted for amplification. Optimal primer length for a complementary sequence is e.g. 18 to 22 nucleotides. Optimal primer sequences for PCR are usually determined by primer design software.

[0065] “DNA microarrays” are arrays which have many microscopic spots of DNA, usually oligonucleotides, bound on a solid support. Assay targets can be DNA, cDNA, or cRNA. Depending on the system, the hybridization of targets to specific spots is detected by fluorescence, chemiluminescence, or colloidal silver or gold. Microarrays are used for multiple applications such as simultaneous measurement of the expression of large numbers of genes, enabling genome-wide gene expression analysis, as well as genotyping studies using e.g. single-nucleotide polymorphism (SNP) or InDei analysis.

[0066] “LOD-score” (logarithm (base 10) of odds) refers to a statistical test often used for linkage analysis in animal and plant populations. The LOD score compares the likelihood of obtaining the test data if the two loci (molecular marker loci and / or a phenotypic trait locus) are indeed linked, to the likelihood of observing the same data purely by chance. Positive LOD scores favor the presence of linkage and a LOD score greater than 3.0 is considered evidence for linkage. A LOD score of +3 indicates 1000 to 1 odds that the linkage being observed did not occur by chance.

[0067] “Vegetative propagation”, “vegetative reproduction” or “clonal propagation” are used interchangeably herein and mean the method of taking part of a plant and allowing that plant part to form at least roots where plant part is, e.g., defined as or derived from (e.g. by cutting of) leaf, pollen, embryo, cotyledon, hypocotyl, cells, protoplasts, meristematic cell, root, root tip, pistil, anther, flower, shoot tip, shoot, stem, fruit, petiole, etc. When a whole plant is regenerated by vegetative propagation, it is also referred to as a vegetative propagation. In one aspect propagation by grafting, e.g. a scion onto a rootstock, is included herein.

[0068] “Cell culture” or “tissue culture” refers to the in vitro culture of cells or tissues of a plant.

[0069] “Regeneration” refers to the development of a plant from cell culture or tissue culture or vegetative propagation.

[0070] “Non-propagating cell” refers to a cell which cannot be regenerated into a whole plant.

[0071] “Transgene” or “chimeric gene” refers to a genetic locus comprising a DNA sequence, such as a recombinant gene, which has been introduced into the genome of a plant by transformation, such as Agrobacterium mediated transformation. A plant comprising a fransgene stably integrated into its genome is referred to as “transgenic plant”. An “isolated nucleic acid sequence” or “isolated DNA” refers to a nucleic acid sequence which is no longer in the natural environment from which it was isolated, e.g. the nucleic acid sequence in a bacterial host cell or in the plant nuclear or plastid genome. When referring to a “sequence” herein, it is understood that the molecule having such a sequence is referred to, e.g. the nucleic acid molecule.

[0072] A "host cell" or a "recombinant host cell" or “transformed cell” are terms referring to a new individual cell (or organism) arising as a result of at least one nucleic acid molecule, having been introduced into said cell. The host cell is preferably a plant cell or a bacterial cell. The host cell may contain the nucleic acid as an extra- chromosomally (episomal) replicating molecule, or comprises the nucleic acid integrated in the nuclear or plastid genome of the host cell, or as introduced chromosome, e.g. minichromosome.

[0073] “Sequence identity” and “sequence similarity” can be determined by alignment of two peptide or two nucleotide sequences using global or local alignment algorithms. Sequences may then be referred to as "substantially identical” or “essentially similar” when they are optimally aligned by for example the programs GAP or BESTFIT or the Emboss program “Needle” (using default parameters, see below) share at least a certain minimal percentage of sequence identity (as defined further below). These programs use the Needleman and Wunsch global alignment algorithm to align two sequences over their entire length, maximizing the number of matches and minimises the number of gaps. Generally, the default parameters are used, with a gap creation penalty = 10 and gap extension penalty = 0.5 (both for nucleotide and protein alignments). For nucleotides the default scoring matrix used is DNAFULL and for proteins the default scoring matrix is Blosum62 (Henikoff & Henikoff, 1992, PNAS 89, 10915-10919). Sequence alignments and scores for percentage sequence identity may for example be determined using computer programs, such as EMBOSS as available on the world wide web under ebi.ac.uk / Tools / psa / emboss_needle / ). Alternatively sequence similarity or identity may be determined by searching against databases such as FASTA, BLAST, etc., but hits should preferably be retrieved and aligned pairwise to compare sequence identity. Two proteins or two protein domains, or two nucleic acid sequences have “substantial sequence identity” if the percentage sequence identity is at least 95%, 96%, 97%, 98% or 99% or more (as e.g. determined by Emboss “needle” using default parameters, i.e. gap creation penalty = 10, gap extension penalty = 0.5, using scoring matrix DNAFULL for nucleic acids and Blosum62 for proteins). For marker sequences comprising a SNP nucleotide in between flanking sequence regions, a ‘variant sequence’ (or a ‘sequence comprising substantial sequence identity’) is for example a sequence comprising the same SNP nucleotide at the equivalent position (or at the equivalent nucleotide) in a sequence comprising at least 95%, 96%, 97%, 98% or 99% sequence identity to the other sequence. The % identity is measured over sequences of the same length, e.g. two sequences of 203 nucleotides in length with a SNP at nucleotide 102. This can be done e.g. by pairwise alignment using e.g. Needle or by BLAST analysis of the sequence against e.g. the genomic sequence or chromosome sequence. For example a ‘SNP nucleotide at nucleotide 102 of SEQ ID NO: 1 or at nucleotide 102 in a variant sequence / or at the equivalent position / or at the equivalent nucleotide in a variant sequence’, refers to the SNP nucleotide at the same position (e.g. nucleotide 102 or 101) but the flanking nucleotides to the right and to the left of the SNP may not be 100% identical to the flanking nucleotides to the left and to the right of nucleotide 102 in SEQ ID NO: 1. Both sequences of the same length, when aligned, may therefore only have 95%, 96%, 97%, 98% or 99% sequence identity. Table 3 provides a variant sequence for SNP l l, Table 6 provides a variant sequence for SNP_15, 17 and 20, and Table 15 provides a variant sequence for SNP_25 and 29. In another embodiment a nucleotide sequence is considered to have substantially identical to the given nucleotide sequence if it can be identified using stringent hybridisation conditions.

[0074] “Stringent hybridisation conditions” can be used to identify nucleotide sequences, which are substantially identical to a given nucleotide sequence. Stringent conditions are sequence dependent and will be different in different circumstances. Generally, stringent conditions are selected to be about 5°C lower than the thermal melting point (Tm) for the specific sequences at a defined ionic strength and pH. The Tm is the temperature (under defined ionic strength and pH) at which 50% of the target sequence hybridises to a perfectly matched probe. Typically stringent conditions will be chosen in which the salt concentration is about 0.02 molar at pH 7 and the temperature is at least 60°C. Lowering the salt concentration and / or increasing the temperature increases stringency. Stringent conditions for RNA-DNA hybridisations (Northern blots using a probe of e.g. lOOnt) are for example those which include at least one wash in 0.2X SSC at 63°C for 20min, or equivalent conditions. Stringent conditions for DNA-DNA hybridisation (Southern blots using a probe of e.g. lOOnt) are for example those which include at least one wash (usually 2) in 0.2X SSC at a temperature of at least 50°C, usually about 55°C, for 20 min, or equivalent conditions.

[0075] “Fine-mapping” refers to methods by which the position of a QTL can be determined more accurately (narrowed down) and by which the size of the introgression fragment comprising the QTL is reduced. For example Near Isogenic Lines for the QTL (QTL-NILs) can be made, which contain different, overlapping fragments of the introgression fragment within an otherwise uniform genetic background of the recurrent parent. Such lines can then be used to map on which fragment the QTL is located and to identify a line having a shorter introgression fragment comprising the QTL.

[0076] DETAILED DESCRIPTION

[0077] The present invention relates in one aspect to a cultivated Cucumis sativus var. sativus plant comprising a fruit sugar QTL, referred to as QTL 1.1 (on chromosome 1) and / or a QTL conferring mint green fruit flesh color, referred to as QTL3.1 (on chromosome 3) infrogressed from a wild or primitive cucumber. Thus, the sugar QTL is conferred by an introgression fragment on cultivated cucumber chromosome 1 (comprising QTL1. 1 or a variant thereof) and / or the mint-green flesh color QTL is conferred by an introgression fragment on cultivated cucumber chromosome 3 (comprising QTL3.1 or a variant thereof), wherein said introgression fragment is from a wild or primitive cucumber, referred to as the ‘donor’ of the QTL.

[0078] QTL 1. 1 is in the region starting at SNP_01 (corresponding to nucleotide 3530044 of chromosome 1 ) and ending at SNP_14 (corresponding to nucleotide 6316306 of chromosome 1), see e.g. Table 2.

[0079] QTL3. 1 is in the region starting at SNP_15 (corresponding to nucleotide 1128135 of chromosome 3) and ending at SNP_24 (corresponding to nucleotide 2585865 of chromosome 3), see e.g. Table 5.

[0080] In a further aspect a cultivated Cucumis sativus var. sativus plant comprising a fruit sugar QTL, referred to as QTL 1. 1 (on chromosome 1) and a mutant w-allele in homozygous form (genotype w / w) and QTL4.1 conferring an enhanced fruit peel color, referred to as QTL4. 1 (on chromosome 4), wherein QTL 1.1 and QTL4.1 are introgressed from a wild or primitive cucumber. Thus, the fruit sugar QTL is conferred by an introgression fragment on cultivated cucumber chromosome 1 (comprising QTL1. 1 or a variant thereof) and the fruit peel color-enhancing QTL is conferred by an introgression fragment on cultivated cucumber chromosome 4 (comprising QTL4. 1 or a variant thereof), wherein said introgression fragment on chromosome 1 and on chromosome 4 is from a wild or primitive cucumber, referred to as the ‘donor’ of the QTL. QTL 1. 1 and QTL4. 1 are in a cultivated cucumber line which comprises the mutant w-allele in homozygous form. The presence of QTL1. 1 may in cultivated cucumbers have a negative effect on fruit peel color when the w-allele is present in homozygous form, which negative effect is reduced or eliminated when QTL4. 1 is present in homozygous form.

[0081] QTL4. 1 is in the region starting at SNP_25 (corresponding to nucleotide 9330448 of chromosome 4) and ending at SNP_33 (corresponding to nucleotide 10300571 of chromosome 4), see e.g. Table 14.

[0082] The M -allcIc is on chromosome 3, comprising e.g. SEQ ID NO: 40 and / or 41, which is in the region starting at nucleotide 40340728 and ending at nucleotide 40340871 of chromosome 3, see Table 9.

[0083] When reference is made herein to an introgression fragment on chromosome 1 having an ‘increase in glucose and fructose-level-conferring QTL’ or ‘sugar QTL’ this encompasses various sizes of introgression fragments, e.g. the fragment as found in NCIMB 44295 comprising the donor SNP nucleotide of all SNP markers linked to the QTL (for QTL1.1: SNP_01 to SNP_14), but also smaller introgression fragments (comprising less than these 14 SNP markers, such as only e.g. 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12 or 13 of the SNP markers), where however the fragment remains large enough to confer the increase in glucose and fructose content (compared to the control or genetic control) when the introgression fragment is in heterozygous or preferably in homozygous form in the cultivated cucumber genome. In other words, the fragment retains QTL 1. 1 or a variant thereof, i.e. it still confers the increase in at least glucose and fructose levels of the fruits (compared to the control, e.g. the genetic control) when the introgression fragment is in heterozygous or preferably in homozygous form in the cultivated cucumber genome. Preferably at least the peak marker, which is the marker most closely linked to the QTL, is present, and preferably also at least 1, 2 or 3 of the SNP markers preceding the peak marker and / or following the peak marker on the chromosome fragment. See also Table 2. For QTL1. 1 the peak marker is SNP_11, so preferably the donor nucleotide for at least SNP_10 and SNP_11 is present, or the donor nucleotide for at least SNP_11 and SNP_12 is present, or the donor nucleotide for at least SNP_10, SNP_11 and SNP_12 is present, or the donor nucleotide for at least SNP_8 and / or SNP_09 and / or SNP_10 and for SNP_11 is present; or the donor nucleotide for SNP_14 and / or SNP_13 and / or SNP12 and for SNP_11 is present; or the donor nucleotide for SNP_11 and for at least 2, 3 or more of SNP_8, SNP_09, SNP_10, SNP_12, SNP_13 and / or SNP_14 is present.

[0084] When reference is made herein to an introgression fragment on chromosome 3 having a ‘mint-green fruit flesh color-conferring QTL’ or ‘ fruit flesh color QTL’ this encompasses various sizes of introgression fragments, e.g. the fragment as found in NCIMB 44295 comprising the donor SNP nucleotide of all SNP markers linked to the QTL (for QTL3.1: SNP_15 to SNP_24), but also smaller introgression fragments (comprising less than these 10 SNP markers, such as only e.g. 2, 3, 4, 5, 6, 7, 8 or 9 of the SNP markers), where however the fragment remains large enough to confer the mint green flesh color (compared to the control or genetic control) when the introgression fragment is in heterozygous or preferably in homozygous form in the cultivated cucumber genome. In other words, the fragment retains QTL3.1 or a variant thereof, i.e. it still confers the mint green flesh color of the fruits (compared to the control, e.g. the genetic control) when the introgression fragment is in heterozygous or preferably in homozygous form in the cultivated cucumber genome. Preferably at least the peak marker, which is the marker most closely linked to the QTL, is present, and preferably also at least 1, 2 or 3 of the SNP markers preceding the peak marker and / or following the peak marker on the chromosome fragment. See also Table 5. For QTL3. 1 the peak marker is SNP_15, which is located towards the beginning of the chromosome. Therefore, in one aspect the donor nucleotide for at least SNP_15, SNP_16 and SNP_17 is present, or for at least SNP_15, SNP_16, SNP_17 and / or SNP_18, or for at least SNP_15, SNP_16, SNP_17, SNP_18 and / or SNP_19. In one aspect the entire beginning of the chromosome is from the donor, so starting at nucleotide 1 of the chromosome and comprising at least SNP_15, SNP_16, SNP_17, SNP_18 and / or SNP_19 from the donor.

[0085] When reference is made herein to an introgression fragment on chromosome 4 having a ‘exterior colorenhancing QTL’ or ‘ fruit peel / skin color enhancing QTL’ this encompasses various sizes of introgression fragments, e.g. the fragment as found in NCIMB 44295 comprising the donor SNP nucleotide of all SNP markers linked to the QTL (for QTL4.1: SNP_25 to SNP_33), but also smaller introgression fragments (comprising less than these 10 SNP markers, such as only e.g. 2, 3, 4, 5, 6, 7, 8 or 9 of the SNP markers), where however the fragment remains large enough to enhance the peel color (compared to the control or genetic control) when the introgression fragment is in heterozygous or preferably in homozygous form in the cultivated cucumber genome, especially in a cucumber genome which comprises the mutant w-allele in homozygous form. In other words, the fragment retains QTL4. 1 or a variant thereof, i.e. it still confers the enhanced peel color of the fruits (compared to the control, e.g. the genetic control) when the introgression fragment is in heterozygous or preferably in homozygous form in the cultivated cucumber genome comprising the w / w genotype at the White locus (also referred to as White peel locus). Preferably at least the peak marker, which is the marker most closely linked to the QTL, is present, and preferably also at least 1, 2 or 3 of the SNP markers preceding the peak marker and / or following the peak marker on the chromosome fragment. See also Table 14. For QTL4. 1 the peak marker is SNP_29, so preferably the donor nucleotide for at least SNP_28 and SNP_29 is present, or the donor nucleotide for at least SNP_29 and SNP_30 is present, or the donor nucleotide for at least SNP_28, SNP_29 and SNP_30 is present, or the donor nucleotide for at least SNP_26 and / or SNP_27 and / or SNP_28 and for SNP_29 is present; or the donor nucleotide for SNP_32 and / or SNP_31 and / or SNP30 and for SNP_29 is present; or the donor nucleotide for SNP_29 and for at least 2, 3 or more of SNP_26, SNP_27, SNP_28, SNP_30, SNP_31 and / or SNP_32 is present.

[0086] Further, when reference is made herein to an introgression fragment on chromosome 1 or 3 or 4 having or comprising QTL 1.1 or QTL3.1 or QTL4.1 this encompasses introgression fragments from various donors which comprise the same or variant QTL of the QTLs present in e.g. NCIMB 44295 (as e.g. described in Tables 2 and 5 and 14). Such same or variant QTLs have the same SNP haplotype or SNP genotype for at least 2, 3, 4, 5, 6, 7, 8, 9, 10 or more of the SNP makers described e.g. in Table 2 for QTL1. 1 and in Table 5 for QTL3. 1 or Table 14 for QTL4.1. For example, suitable donors may be found in seed banks such as the ARS-GRIN collection in the US (see world wide web at npgsweb.ars-grin.gov / gringlobal / search) or at the Center for Genetic Resources, Wageningen University, / / cgngenis.wur.nl / ) which comprise the same SNP haplotype or genotype for at least 2, 3, 4, 5, 6, 7, 8, 9, 10 or more (or all) SNP markers of QTL1. 1 (as described in Table 2) or of QTL3.1 (as described in Table 5) or of QTL4. 1 (as described in Table 14). Preferably the wild or primitive cucumber donor which comprises the same SNP haplotype or genotype for at least 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12 or more (or all) SNP markers of QTL 1.1, as described in Table 2, further can be used to transfer the QTL1.1 (or variant) region into cultivated cucumber, which cultivated cucumber then produces fruits which have an average glucose and fructose content in the fruits which is significantly higher than the average glucose and fructose content in the fruits of cultivated cucumber lines or varieties which lack the introgression of QTL1. 1 (or variant thereof), e.g. when grown under the same conditions. As mentioned before, the QTL1. 1 region is preferably present in homozygous form in the cultivated cucumber in order to determine whether the QTL1. 1 region confers a significant increase in at least glucose and fructose content of the fruits comprising the introgression fragment. Preferably the wild or primitive cucumber donor which comprises the same SNP haplotype or genotype for at least 2, 3, 4, 5, 6, 7, 8, 9 or more (or all) SNP markers of QTL3. 1, as described in Table 5, further can be used to transfer the QTL3. 1 region (or variant) region into cultivated cucumber, which cultivated cucumber then produces fruits which have a mint green fruit flesh e.g. when grown under the same conditions. Preferably the QTL3. 1 region is present in homozygous form in the cultivated cucumber in order to determine whether the QTL3. 1 region confers the fruits having a mint green fruit flesh color.

[0087] Preferably the wild or primitive cucumber donor which comprises the same SNP haplotype or genotype for at least 2, 3, 4, 5, 6, 7, 8, 9 or more (or all) SNP markers of QTL4. 1, as described in Table 14, further can be used to transfer the QTL4. 1 region (or variant) region into cultivated cucumber comprising the w / w genotype (i.e. the mutant w-allele in homozygous form), which cultivated cucumber then produces fruits which have an improved fruit peel color e.g. when grown under the same conditions. Preferably the QTL4. 1 region is present in homozygous form in the cultivated cucumber in order to determine whether the QTL4. 1 region confers the peel color enhancing effect.

[0088] Herein below aspects are described for each of QTL1. 1 and QTL3. 1 (or variants of any of these) individually, but it is understood that not only plants and plant parts comprising individual QTLs are encompassed herein, but that plants and plant parts comprising both QTLs are encompassed. In one aspect also the genetic determinants which determine exterior mint green color of the fruits may be combined with QTL1.1 and / or with QTL3. 1. Herein below also aspects for the w-allele and QTL4. 1 (or variants of any of these), optionally in combination with QTL1. 1 and / or QTL3. 1 (or variants of any of these) are described and encompassed herein.

[0089] In one aspect a cultivated Cucumis sativus var. sativus plant is provided comprising at least one or two introgression fragments on chromosome 1 and / or 3 from a wild cucumber donor, wherein said at least one fragment comprises QTL1.1 (or a variant thereof) and the other fragment comprises QTL3.1 (or a variant thereof). Thus, in one aspect QTL 1. 1 (or a variant thereof) is present in the plant (or plant part or seed or fruit), in heterozygous form or preferably in homozygous form, and optionally combined with QTL3. 1 (or a variant thereof), in heterozygous form or homozygous form. In another aspect QTL3. 1 (or a variant thereof) is present in the plant (or plant part or seed or fruit), in heterozygous form or homozygous form.

[0090] In another aspect a cultivated Cucumis sativus var. sativus plant is provided comprising at least two introgression fragments, one on chromosome 1 and one on chromosome 3, both from a wild cucumber donor wherein each of said introgression fragments comprises a Quantitative Trait Locus (QTL) selected from the QTLs designated QTL 1.1 (or a variant thereof) and QTL3.1 (or a variant thereof). As was shown in the Examples, QTL1. 1 increases glucose and fructose content of the marketable fruits, especially in homozygous form, and QTL3. 1 makes the fruit flesh mint-green, especially in homozygous form. In one aspect both QTLs are from the same donor accession, e.g. from seeds deposited under NCIMB 44295 or progeny thereof, e.g. obtained by selfing plants grown from seeds deposited under NCIMB 44295. In a different aspect a cultivated Cucumis sativus var. sativus plant is provided comprising at least two introgression fragments, one on chromosome 1 and one on chromosome 4, both from a wild cucumber donor wherein each of said introgression fragments comprises a Quantitative Trait Locus (QTL) selected from the QTLs designated QTL 1.1 (or a variant thereof) and QTL4. 1 (or a variant thereof). As was shown in the Examples, QTL1. 1 increases glucose and fructose content of the marketable fruits, especially in homozygous form. However, QTL 1.1 may have a negative effect on fruit peel color when it is present in plants that are homozygous for the mutant w-allele (genotype w / w). This negative effect can be counteracted by the presence of QTL4. 1 in preferably homozygous form, whereby the peel color is enhanced again, to e.g. mint-green. In one aspect both QTL 1.1 and QTL4.1 are, therefore, present, especially in a genetic background comprising the w / w genotype. In one aspect both QTL1. 1 and QTL4. 1 are introgressed from the same wild donor accession, e.g. from seeds deposited under NCIMB 44295 or progeny thereof, e.g. obtained by selfing plants grown from seeds deposited under NCIMB 44295. Optionally further QTL3. 1 (or a variant thereof) is further present in the plant, seed or fruit or plant part.

[0091] Thus, in one aspect QTL1.1, QTL4.1, QTL3.2 (w / w) (or variants of any of these) are present in a cultivated cucumber fruit, seed, plant, plant part or cell. All QTLs are found in the deposited seeds and are in one aspect obtainable from these seeds.

[0092] In another aspect QTL1.1, QTL4.1, QTL3.2 (w / w) and QTL3.1 (or variants of any of these) are present in a cultivated cucumber fruit, seed, plant, plant part or cell. All QTLs are found in the deposited seeds and are in one aspect obtainable from these seeds.

[0093] In yet another aspect QTL3.2 (w / w) and QTL4.1 (or variants of any of these) are present in a cultivated cucumber fruit, seed, plant, plant part or cell. All QTLs are found in the deposited seeds and are in one aspect obtainable from these seeds. QTL4. 1 (or variants thereof) enhances the fruit peel color in e.g. a plant comprising the mutant White-allele (genotype w / w) in its genome.

[0094] QTLL 1 and variants of QTLL 1 on chromosome 1

[0095] Thus, in one aspect a cultivated cucumber plant (as well as plant parts such as cells, tissues, leaves, stems, flowers, fruits, seeds, etc.) is provided comprising an introgression fragment from a wild or primitive cucumber, wherein the introgression fragment comprises QTL1.1, or a variant thereof, and wherein the introgression fragment comprises all or part of the region starting at nucleotide (or base) 3530044 of chromosome 1 (corresponding to SNP_01) and ending at nucleotide (or base) 6316306 of chromosome 1 (corresponding to SNP_14). In other words, all or part of the region starting at nucleotide 3530044 of chromosome 1 (SNP_01) and ending at nucleotide 6316306 of chromosome 1 (SNP_14) is, in one aspect, from a wild donor cucumber and comprises QTL 1. 1 or a variant thereof. Which sub-region contains QTL 1. 1 can be identified by e.g. fine-mapping. So, for example if QTL1.1 is found to be in-between SNP_09 and SNP_13, then the plant of the invention only needs to comprise the introgression region starting at nucleotide 4557067 of chromosome 1 (SNP_09) and ending at nucleotide 5550581 (SNP_13) of chromosome 1.

[0096] In one aspect QTL 1. 1 (or a variant thereof) is located in-between marker SNP 01 at nucleotide 102 of SEQ ID NO: 1 (or at nucleotide 102 in a variant sequence of SEQ ID NO: 1) and marker SNP_14 at nucleotide 102 of SEQ ID NO: 14 (or at nucleotide 102 in a variant sequence of SEQ ID NO: 14).

[0097] In another aspect QTL 1. 1 (or a variant thereof) is located in-between marker SNP_08 at nucleotide 102 of SEQ ID NO: 8 (or at nucleotide 102 in a variant sequence of SEQ ID NO: 8) and marker SNP_14 at nucleotide 102 of SEQ ID NO: 14 (or at nucleotide 102 in a variant sequence of SEQ ID NO: 14). In a further aspect QTL1. 1 (or a variant thereof) is located in-between marker SNP_08 at nucleotide 102 of SEQ ID NO: 8 (or at nucleotide 102 in a variant sequence of SEQ ID NO: 8) and marker SNP_13 at nucleotide 102 of SEQ ID NO: 13 (or at nucleotide 102 in a variant sequence of SEQ ID NO: 13). In a further aspect QTL1. 1 (or a variant thereof) is located in-between marker SNP_9 at nucleotide 102 of SEQ ID NO: 9 (or at nucleotide 102 in a variant sequence of SEQ ID NO: 9) and marker SNP_12 at nucleotide 102 of SEQ ID NO: 12 (or nucleotide 102 in a variant sequence of SEQ ID NO: 12). In a further aspect QTL1.1 (or a variant thereof) is located in-between marker SNP_10 at nucleotide 102 of SEQ ID NO: 10 (or at nucleotide 102 in a variant sequence of SEQ ID NO: 10) and marker SNP_12 at nucleotide 102 of SEQ ID NO: 12 (or nucleotide 102 in a variant sequence of SEQ ID NO: 12).

[0098] In another aspect the introgression fragment of the invention (comprising QTL 1.1 or a variant thereof) is a fragment comprising a smaller fragment (part) of the region starting at nucleotide (or base) 3530044 of chromosome 1 and ending at nucleotide (or base) 6316306 of chromosome 1, e.g. having a size of e.g. 3.0 Mb, 2.8 Mb, 2.5 Mb, 2 Mb, 1Mb, 0.5Mb, lOOkb, 50kb, 35kb, 30kb, 20kb, or less and comprising the QTL or a variant thereof. In one aspect the part is at least 5kb, lOkb, 20kb in size, or more.

[0099] In one aspect the cultivated cucumber plant of the invention comprises an introgression fragment from a wild or primitive cucumber, which introgression fragment comprises QTL 1.1 or a variant thereof, wherein the introgression fragment comprises all of part of the region starting at 3.530044 Mb and ending at 6.316306 Mb of the physical chromosome 1.

[0100] In one aspect the introgression fragment on chromosome 1 comprising QTL 1.1, or a variant thereof, is obtainable by crossing a plant grown from NCIMB44295 with another cucumber plant, especially a cultivated cucumber plant, in one aspect a long cucumber type or a short cucumber type or a pickling or slicer type. In one aspect the cultivated cucumber plant of the invention comprising QTL 1. 1, or a variant thereof, is a plant wherein said introgression fragment on chromosome 1 is obtainable by crossing a plant grown from seeds deposited under accession number NCIMB44295 with another cucumber plant. Thus, in one aspect the QTL is the QTL present in seeds deposited under accession number NCIMB44295.

[0101] In a further aspect the cultivated cucumber plant of the invention comprising QTL 1. 1 , or a variant thereof, is a plant wherein said introgression fragment on chromosome 1 is obtainable by crossing a plant comprising the same donor SNP haplotype or SNP genotype for at least 2, 3, 4, 5, 6, 7, 8, 9, 10 or more SNP markers linked to the QTL (i.e. SNP_01 to SNP_14 for QTL1.1, as e.g. shown in Table 2) with another cucumber plant, especially with a cultivated cucumber elite breeding line. Thus, in one aspect the QTL is the variant QTL present in wild donor accessions or in cultivated cucumber which comprise the same donor SNP haplotype or genotype for at least 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12 or more of the SNP markers e.g. as present in NCIMB44295. Preferably the donor also comprises a high fruit glucose and fructose content, and optionally a low malic acid content, as does the cultivated cucumber plant comprising the introgression fragment (especially in homozygous form).

[0102] Herein a high glucose and fructose content of a plant comprising QTL1. 1 (especially in homozygous form) is a content (of glucose plus fructose) which is at least 5%, 6%, 7%, 8%, 9%, 10%, 11%, 12%, 13%, 14%, 15% higher than in the fruits of a plant lacking QTL1. 1 (e.g. a control or recurrent parent), when grown under the same conditions and measured in the same way, e.g. as described in the Examples. A low malic acid content of a plant comprising QTL1. 1 (especially in homozygous form) is a content of malic acid which is at least 5%, 6%, 7%, 8%, 9%, 10%, 11%, 12%, 13%, 14%, 15%, 16%, 17%, 18%, 19%, 20%, 21%, 22% or more lower than in the fruits of a plant lacking QTL 1.1 (e.g. a control or recurrent parent), when grown under the same conditions and measured in the same way. Differences between a plant comprising QTL1. 1 and a plant lacking QTL1. 1 are preferably statistically significant, i.e. the average glucose and fructose and / or malic acid content of the fruits is significantly different between the plants.

[0103] When referring to the SNP markers herein, which are indicative of the presence of the introgression fragment (and / or of the sugar QTL present on the introgression fragment), it is understood that the donor SNP genotype or haplotype which is indicative of the introgression fragment is referred to, i.e. the donor SNP genotype or haplotype as provided e.g. in Table 2. It is noted that the SNP marker genotype can distinguish between the introgression fragment being in homozygous or heterozygous form. In homozygous form the nucleotide is identical, while in heterozygous form the nucleotide is not (necessarily) identical. The SNP genotype of the ‘wild type’ chromosome lacking the introgression fragment is e.g. the other haplotype, e.g. the haplotype of the recurrent parent). So, e.g. the genotype of SNP_01 indicative of the introgression fragment comprising QTL1.1 is e.g. ‘CA’ (QTLl. l / wt) or ‘CC’ (QTL1.1 / QTL1.1) while the SNP genotype indicative of the wild type / genetic control (lacking the introgression fragment) is e.g. ‘AA’ (wt / wt). This can also be written as genotype CX’ (QTLl.l / wt) or ‘CC’ (QTL1.1 / QTL1.1) while the SNP genotype indicative of the wild type / genetic control (lacking the introgression fragment) is e.g. ‘XX’ (wt / wt). X may be any nucleotide (A, T, C or G). Thus, when referring to a plant or plant part (e.g. cell) comprising the introgression fragment in homozygous or heterozygous form, it is understood that the SNP markers linked to the introgression fragment have the corresponding SNP genotype or haplotype.

[0104] So, in one aspect, a cultivated Cucumis sativus var. sativus plant is provided comprising an introgression fragment on chromosome 1 in homozygous or heterozygous form, wherein said introgression fragment confers an increase in fruit glucose and fructose content compared to the fruits produced by the cucumber plant lacking the introgression fragment on chromosome 1, e.g. the genetic control or control variety, when grown under the same conditions.

[0105] The increase in glucose and fructose content is phenotypically expressed as a (statistically significant) higher average glucose plus fructose content of the fruits produced by the cultivated cucumber plant line or variety comprising the introgression fragment on chromosome 1 in homozygous or heterozygous form compared to the genetic control line or variety lacking the introgression fragment on chromosome 1 when grown under the same environment. The average glucose and fructose level increases by at least 5%, 6%, 7%, 8%, 9%, 10%, 11%, 12%, 13%, 14%, 15% or more, especially when the QTL1. 1 is in homozygous form. Also, the individual content of fruit glucose or fruit fructose increases by at least 5%, 6%, 7%, 8%, 9%, 10%, 11%, 12%, 13%, 14%, 15% or more, especially when the QTL1. 1 is in homozygous form. See for example Figure 3 and Figure 4.

[0106] So, for example if QTL1. 1 is introduced into a cucumber line or variety having an average glucose plus fructose content of about 21g / L or about 22 g / L, the line or variety with QTL1.1 will produce at least about 23g / L, 24g / L or more of glucose plus fructose due to the QTL1. 1.

[0107] The effect of QTL1. 1 can, thus, be determined by introducing the QTL into a cucumber plant, in heterozygous or preferably in homozygous form. For example NCIMB44295 can be crossed with a cucumber plant lacking QTL1. 1 and QTL1. 1 can thus be transferred into the background by e.g. backcrossing.

[0108] The plants of the invention therefore comprise a genome of cultivated cucumber, with at least one or two recombinant chromosomes, namely one or two recombinant chromosomes 1 (i.e. heterozygous or homozygous). The recombinant chromosomes comprise a fragment of a wild donor cucumber, which is easily distinguishable from the cultivated cucumber genome by molecular marker analysis, whole genome sequencing, chromosome painting and similar techniques. In one aspect the introgression fragment on chromosome 1 is from a wild or primitive cucumber, comprises the sugar QTL1. 1, or a variant thereof, and comprises all or part of the region starting at nucleotide SNP 01 and ending at SNP_14. Thus, the introgression fragment comprises the QTL1. 1 or a variant thereof and one or more or all (e.g. 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14) SNP markers of the wild donor selected from SNP_01 to SNP 14 as shown in Table 2.

[0109] In one aspect the introgression fragment comprises QTL1. 1 and one or more or all of the donor SNP nucleotides for SNP_01 to SNP_14 (as shown in Table 2), whereby SNP_01 to SNP_14 are at nucleotide 102 of SEQ ID NO: 1 to SEQ ID NO: 14, respectively, or are present at the equivalent nucleotide (e.g. nucleotide 102) of a variant sequence comprising at least 95%, 96%, 97%, 98% or 99% sequence identity to SEQ ID NO: 1 to 14, respectively. In one aspect Table 3 provides such a variant sequence. The donor used in the Examples was sequenced and it was found that SNP_11 was present in the donor in a sequence comprising 99% sequence identity to SEQ ID NO: 11, this sequence is provided as SEQ ID NO: 34 herein.

[0110] In one aspect the presence of the introgression fragment on chromosomes 1 comprising QTL 1. 1 in the genome of the plant or plant cell or plant tissue (or in the DNA extracted therefrom) is detectable by a molecular marker assay which detects one or more molecular markers indicative of the introgression fragment comprising QTL1.1, especially the donor SNP haplotype or genotype for at least 2, 3, 4, 5, 6, 7, 8, 9, 10 or more of SNP_01 to SNP 14, at nucleotide 102 of SEQ ID NO: 1 to 14 (or the complement sequence of SEQ ID NO: 1 to 14, or a sequence comprising at least 95%, 96%, 97%, 98% or 99% sequence identity to SEQ ID NO: 1 to 14 or to the complement of SEQ ID NO: 1 to 14), respectively. However, as mentioned, other techniques may be used, e.g. the SNP genotype of the markers may also be determined by sequencing or by using alternative markers located in between the SNP markers provided herein or within 7cM, or within 5cM, of a marker provided herein; or within 3 Mb, 2.8Mb, 2.5 Mb, 2 Mb, 1 Mb, 0.5 Mb, 0.4Mb, 0.3Mb, 0.2Mb, 0. 1 Mb, 50kb, 20kb, lOkb, 5kb, 2kb, Ikb or less of a marker provided herein.

[0111] In one aspect the presence of the introgression fragment on chromosomes 1 comprising QTL1. 1 in the genome of the plant or plant cell or plant tissue (or in the DNA extracted therefrom) is detectable by detecting the presence of one or more or all of SEQ ID NO: 1 to SEQ ID NO: 14. In another aspect the presence of the introgression fragment on chromosomes 1 comprising QTL1. 1 in the genome of the plant or plant cell or plant tissue (or in the DNA extracted therefrom) is detectable by detecting the presence of one or more or all of SEQ ID NO: 1 to SEQ ID NO: 10, SEQ ID NO: 34 (comprising SNP_11), and SEQ ID NO: 12 to 14.

[0112] It is noted that when referring herein to an introgression fragment or QTL being detectable or being present in a plant ort plant part, all embodiments encompass also methods for detecting the introgression fragment or QTL in the plant or plant part, e.g. by marker assisted selection, SNP haplotyping, genotyping, sequencing etc., and / or methods for selecting a plant or plant part comprising the introgression fragment or QTL.

[0113] When reference is made herein to one or more molecular markers or sequences being “detectable” by e.g. a molecular marker assay, this means of course that the plant or plant part comprises the one or more markers or sequences in its genome, as the marker or sequence would otherwise not be detectable. Further, when referring to a plant or plant part in which the SNP marker or SNP haplotype of the donor or introgression fragment comprising QTL 1.1 or QTL3.1 or QTL4.1 is present or detectable, this also encompasses SNP assays and methods for detecting the SNP marker and the SNP haplotype or genotype present and / or for selecting a plant or plant part comprising the donor SNP haplotype or genotype.

[0114] Furthermore, when a plant is referred to, this also encompasses plant parts, such as cells, tissues, leaves, stems, fruits, flowers, pollen, roots, etc. and seeds from which the plant can be grown, seeds produced by the plant and retaining the QTL.

[0115] Cucumber plants comprising an introgression fragment on chromosome 1 (QTL 1.1 or a variant thereof)

[0116] In one aspect a cultivated Cucumis sativus var. sativus plant comprising an introgression fragment from a wild or primitive cucumber on chromosome 1 in homozygous or heterozygous form is provided, wherein said introgression fragment comprises a Quantitative Trait Locus (QTL) located between the Single Nucleotide Polymorphism marker SNP 01 at nucleotide 102 of SEQ ID NO: 1 (or at nucleotide 102 of a variant of SEQ ID NO: 1) and the Single Nucleotide Polymorphism marker SNP_14 at nucleotide 102 of SEQ ID NO: 14 (or at nucleotide 102 of a variant of SEQ ID NO: 14), which QTL confers an increase in glucose and fructose content to the fruits. In one aspect the QTL is located between base 3530044 (SNP_01) and base 6316306 (SNP_14) of chromosome 1.

[0117] Thus, in one aspect QTL1. 1 (or a variant thereof) is located in the region between SNP_01 in SEQ ID NO: 1 (or in a variant thereof) and SNP_14 in SEQ ID NO: 14 (or in a variant thereof).

[0118] Provided is in one aspect a cultivated Cucumis sativus var. sativus plant comprising an introgression fragment on chromosome 1 from a wild cucumber donor, wherein said introgression fragments comprises a Quantitative Trait Locus (QTL) referred to as QTL1. 1 conferring an increase in glucose and fructose of the cucumber fruit, wherein QTL1. 1 is located on chromosome 1 between the Single Nucleotide Polymorphism marker SNP_01 at nucleotide 102 of SEQ ID NO: 1 and SNP 14 at nucleotide 102 of SEQ ID NO: 14, and wherein said introgression fragment on chromosome 1 comprising QTL1. 1 comprises a SNP haplotype of at least 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13 or 14 markers selected from the group consisting of: a) a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_01 at nucleotide 102 of SEQ ID NO: 1 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 1; b) an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_02 at nucleotide 102 of SEQ ID NO: 2 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 2; c) a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_03 at nucleotide 102 of SEQ ID NO: 3 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 3; d) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_04 at nucleotide 102 of SEQ ID NO: 4 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 4; e) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_05 at nucleotide 102 of SEQ ID NO: 5 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 5; f) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_06 at nucleotide 102 of SEQ ID NO: 6 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 6; g) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_07 at nucleotide 102 of SEQ ID NO: 7 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 7; h) an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_08 at nucleotide 102 of SEQ ID NO: 8 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 8; i) a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_09 at nucleotide 102 of SEQ ID NO: 9 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 9; j) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_10 at nucleotide 102 of SEQ ID NO: 10 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 10; k) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_11 at nucleotide 102 of SEQ ID NO: 11 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 11, such as a Guanine for SNP l 1 at nucleotide 102 of SEQ ID NO: 34; l) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_12 at nucleotide 102 of SEQ ID NO: 12 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 12; m) an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_13 at nucleotide 102 of SEQ ID NO: 13 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 13; n) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_14 at nucleotide 102 of SEQ ID NO: 14 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 14.

[0119] Instead of referring to the SNP haplotype, which refers to only one chromosome of the chromosome pair, one can also refer to the SNP genotype, whereby reference is made to both chromosomes of a pair of chromosomes in a diploid organism. Therefore, a Cytosine (C) for SNP_01 is then referred to as genotype CX or CC, where the first C is for one chromosome 1 of the pair and the X or the second C is for the other chromosome of the pair. The genotype CC for SNP_01 and one or more of the other markers thus has the introgression fragment on both chromosomes and is homozygous for the introgression fragment.

[0120] In one aspect, therefore, a cultivated Cucumis sativus var. sativus plant is provided comprising an introgression fragment on chromosome 1 in homozygous or heterozygous form, wherein said introgression fragment confers an increase in (at least) fruit glucose and fructose content (compared to the plant lacking the introgression fragment, e.g. the genetic control) and wherein said introgression fragment comprises the SNP marker haplotype or genotype of at least 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13 or 14 of the SNP markers selected from the group consisting of: a) the CX or CC genotype for the Single Nucleotide Polymorphism marker SNP_01 at nucleotide 102 of SEQ ID NO: 1 (or at nucleotide 102 in a variant thereof); b) the AX or AA genotype for the Single Nucleotide Polymorphism marker SNP_02 at nucleotide 102 of SEQ ID NO: 2 (or at nucleotide 102 in a variant thereof); c) the CX or CC genotype for the Single Nucleotide Polymorphism marker SNP_03 at nucleotide 102 of SEQ ID NO: 3 (or at nucleotide 102 in a variant thereof); d) the GX or GG genotype for the Single Nucleotide Polymorphism marker SNP_04 at nucleotide 102 of SEQ ID NO: 4 (or at nucleotide 102 in a variant thereof); e) the TX or TT genotype for the Single Nucleotide Polymorphism marker SNP_05 at nucleotide 102 of SEQ ID NO: 5 (or at nucleotide 102 in a variant thereof); f) the TX or TT genotype for the Single Nucleotide Polymorphism marker SNP_06 at nucleotide 102 of SEQ ID NO: 6 (or at nucleotide 102 in a variant thereof); g) the TX or TT genotype for the Single Nucleotide Polymorphism marker SNP_07 at nucleotide 102 of SEQ ID NO: 7 (or at nucleotide 102 in a variant thereof); h) the AX or AA genotype for the Single Nucleotide Polymorphism marker SNP_08 at nucleotide 102 of SEQ ID NO: 8 (or at nucleotide 102 in a variant thereof); i) the CX or CC genotype for the Single Nucleotide Polymorphism marker SNP_09 at nucleotide 102 of SEQ ID NO: 9 (or at nucleotide 102 in a variant thereof); j) the TX or TT genotype for the Single Nucleotide Polymorphism marker SNP_10 at nucleotide 102 of SEQ ID NO: 10 (or at nucleotide 102 in a variant thereof); k) the GX or GG genotype for the Single Nucleotide Polymorphism marker SNP_11 at nucleotide 102 of SEQ ID NO: 11 (or at nucleotide 102 in a variant thereof, such as a Guanine for SNP l 1 at nucleotide 102 of SEQ ID NO: 34); l) the GX or GG genotype for the Single Nucleotide Polymorphism marker SNP_12 at nucleotide 102 of SEQ ID NO: 12 (or at nucleotide 102 in a variant thereof); m) the AX or AA genotype for the Single Nucleotide Polymorphism marker SNP_13 at nucleotide 102 of SEQ ID NO: 13 (or at nucleotide 102 in a variant thereof); n) the GX or GG genotype for the Single Nucleotide Polymorphism marker SNP_14 at nucleotide 102 of SEQ ID NO: 14 (or at nucleotide 102 in a variant thereof). When referring to a SNP in a variant sequence, that variant sequence comprises at least 95%, 96%, 97%, 98% or 99% sequence identity with the mentioned sequence. For example, the wild donor used herein to introgress the QTL1. 1 comprises variation regarding SNP_11. The introgression fragment with QTL1. 1 on it comprises a sequence (SEQ ID NO: 34) which has 99% sequence identity to SEQ ID NO: 11, i.e. SNP_11 is a Guanine at nucleotide 102 in a sequence comprising at least 99% sequence identity to SEQ ID NO: 11, see also the Examples, especially Table 3. In one aspect when referring to a Guanine for SNP_11, reference is made to a Guanine in SEQ ID NO: 34, which is the sequence of the wild donor. For the other SNPs of QTL1. 1 the donor nucleotide is present in the sequence provided herein, i.e. there is no variant sequence in the donor used. X refers to any nucleotide for the sequence on the other chromosome 1 of the pair of chromosomes. In one aspect X may be the nucleotide of the recurrent parent as described in Table 2.

[0121] In one aspect said at least 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13, 14 or 15 markers are consecutive markers.

[0122] In one aspect the plant or plant part comprises a donor SNP haplotype for at least 5 of the SNP markers of SNP_01 to SNP_14 or of SNP_02 to SNP_13 or of SNP_07 to SNP_14, or for at least 10 of the SNP markers of SNP_01 to SNP_14 or of SNP_02 to SNP_13 or of SNP_07 to SNP_14.

[0123] The fragment comprising the QTL1.1 may, thus, be large (comprising the donor nucleotide for SNP_01 to SNP 14), or may be smaller and lack markers having the genotype or haplotype of the wild cucumber donor (i.e. the markers have the cultivated cucumber genotype or haplotype instead, see also Table 2; SNP haplotype of recurrent parent), but it may still confer a significant increase in fruit glucose and fructose levels on the cultivated cucumber plant, i.e. it can still comprise the QTL1.1 or a variant. Such smaller introgression fragments are an embodiment of the invention. Plants having smaller introgression fragments which still confer an increase in average glucose and fructose levels of the fruits (i.e. contain the sugar allele) can be generated using known techniques, such as fine-mapping or similar techniques. For example by starting with a plant comprising the introgression fragment as found in seeds deposited under accession number NCIMB 44295 and crossing such a plant with another cultivated cucumber plant and selfing the progeny of said cross, and / or backcrossing the progeny, to generate a population of plants which may contain recombinants having a smaller introgression fragment on chromosome 1, which fragments still confer a significant increase in at least glucose and fructose content of the fruits in relation to a plant lacking the introgression fragment (such as the genetic control), e.g. a fragment comprising donor SNP nucleotide for markers SNP_08 to SNP_14, or SNP_08 to SNP_13 or SNP_08 to SNP_12 or SNP_09 to SNP_14 or SNP_09 to SNP_13 or SNP_09 to SNP_12, or SNP_10 to SNP_14 or SNP_10 to SNP_13 or SNP_10 to SNP_12. Marker assays can be used to determine the size of the smaller introgression fragment. One or more of the SNP markers with the genotype or haplotype of the wild donor cucumber may be missing. The cultivated cucumber genotype or haplotype is then detected for these SNP markers. The fruit glucose and fructose levels of plants comprising such a smaller introgression fragment can then be compared in an assay as described herein, i.e. growing a plurality of plants comprising the smaller introgression fragment in experiments together with suitable control plants, lacking the introgression fragments. If the average glucose and fructose level remains significantly higher than in the control, then the smaller introgression fragment has retained the QTL1. 1.

[0124] Alternatively, the same or variant QTL (QTL1. 1 or variant QTL1. 1) may be introgressed from a different wild donor accessions, whereby optionally not all donor SNP markers disclosed herein may be present, i.e. the SNP haplotype of the donor accession may only be identical to the SNP haplotype of the QTL1. 1 present in seeds of NCIMB44295 for at least 2, 3, 4, 5, 6, 7, 8, 9, 10 or more SNPs. Such alternative wild cucumber sources can be identified using the SNP markers provided herein, by screening germplasm (i.e. accessions of) wild or primitive cucumber using a marker assay to detect the genotype or haplotype of one or more of markers SNP_01 to SNP_14, or of markers SNP_08 to SNP_14, or SNP_08 to SNP_13 or SNP_08 to SNP_12 or SNP_09 to SNP_14 or SNP_09 to SNP_13 or SNP_09 to SNP_12, or SNP_10 to SNP_14 or SNP_10 to SNP_13 or SNP_10 to SNP_12, or even only a smaller subgroup of these markers (e.g. 2, 3 or 4). Plants comprising the same or variant QTL 1.1 from these donors or from other sources are also an embodiment of the invention. Thus, as long as at least 2, 3, 4, 5, 6, 7, 8, 9, 10 or more (or all) of the donor SNPs of SNP_01 to SNP_14, or of the SNPs of SNP_08 to SNP_14, or SNP_08 to SNP_13 or SNP_08 to SNP_12 or SNP_09 to SNP_14 or SNP_09 to SNP_13 or SNP_09 to SNP_12, or SNP_10 to SNP_14 or SNP_10 to SNP_13 or SNP_10 to SNP_12 are present, the donor may contain QTL 1.1 (or a variant thereof) and is encompassed herein. The skilled person can then introgress the QTL 1. 1 (or a variant thereof) into cultivated cucumber in order to increase fruit sugar content as described herein and in order to confirm that the QTL increases fruit sugar content when present in cultivated cucumber.

[0125] As described above, in one embodiment the cultivated cucumber plant of the invention comprises an introgression fragment comprising at least a subset of SNP markers with the genotype (or haplotype) of the wild donor cucumber, i.e. at least 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12 or more markers of SNP_01 to SNP_14, or at least 3 markers of SNP_08 to SNP_14, or of SNP_08 to SNP_13 or of SNP_08 to SNP_12 or of SNP_09 to SNP_14 or of SNP_09 to SNP_13 or of SNP_09 to SNP_12, or of SNP_10 to SNP_14 or of SNP_10 to SNP_13 or of SNP_10 to SNP_12. In one aspect the cultivated cucumber plant comprises all, or all except 1 or 2 markers of SNP_01 to SNP_14, or of SNP_08 to SNP_14, or SNP_08 to SNP_13 or SNP_08 to SNP_12 or SNP_09 to SNP_14 or SNP_09 to SNP_13 or SNP_09 to SNP_12, or SNP_10 to SNP_14 or SNP_10 to SNP_13 or SNP_10 to SNP_12.

[0126] When referring herein to “all except 1 or 2” of a number of consecutive markers, the 1 or 2 markers lacking are in one aspect the first 1 or 2 markers of the consecutive markers and / or the last one or two of the consecutive markers. So, e.g. all except 1 or 2 markers of SNP_01 to SNP_14 are in one aspect e.g. SNP_02 to SNP_14 or SNP_02 to SNP_13. Preferably at least the peak marker of a QTL is present and preferably also at least 1, 2, 3 or 4 consecutive markers preceding the peak marker and / or following the peak marker are present on the introgression fragment.

[0127] Thus, the introgression fragment (and a cultivated cucumber plant or plant part, e.g., a cell, comprising the introgression fragment) can be detected in a marker assay by detecting the donor SNP genotype or haplotype of the introgression fragment (i.e. of the wild donor cucumber germplasm) of one or more or all of the markers above, preferably at least 2, 3, 4, 5, 6, 7, 8, 9, 10 or more.

[0128] Thus, in one aspect, a Quantitative Trait Locus (QTL1. 1) was found to be present on chromosome 1 of a wild cucumber donor which, when transferred (introgressed) into a cultivated cucumber variety or breeding line, and when present in heterozygous or homozygous form, confers significantly increased average fruit glucose and fructose levels onto the cultivated cucumber plant. The QTL, or the introgression fragment comprising the QTL (comprising the sugar allele), is thus in one aspect additive, i.e. it is sufficient to have the introgression fragment on one of the chromosomes 1 (one recombinant chromosome 1), while the homologous chromosome 1 of the pair may be a (non-recombinant) chromosome 1 of cultivated C. sativus var. sativus lacking the introgression fragment.

[0129] In a specific embodiment, the introgression fragment comprising the QTL1. 1 (or a variant thereof) is derivable from (or derived from) or obtainable from (or obtained from; or as present in) seeds, a representative sample of which has been deposited under accession number NCIMB 44295 or from progeny thereof. The progeny may be any progeny which retain the SNP markers or haplotype indicative of (and linked to) the QTL, e.g. as described in Table 2. Thus, progeny are not limited to selfings or Fl or F2 progeny of the deposit or accession, but can be any progeny, whether obtained by selfing and / or crossing with another cucumber plant.

[0130] In one embodiment the introgression fragment comprising QTL1. 1 (or a variant) is identifiable by one or more of the donor SNP markers linked to the QTL 1.1 and / or indicative of QTL 1.1 described elsewhere herein, especially markers SNP_01 to SNP_14 for the introgression fragment on chromosome 1, or a subset of markers, such as one or more of the markers selected from SNP markers SNP_08 to SNP_14, or SNP_08 to SNP_13 or SNP_08 to SNP_12 or SNP_09 to SNP_14 or SNP_09 to SNP_13 or SNP_09 to SNP_12, or SNP_10 to SNP_14 or SNP_10 to SNP_13 or SNP_10 to SNP_12. In one aspect the invention provides a cultivated cucumber plant, having a genome of cultivated (domesticated) cucumber which produces cucumber fruits with a significantly higher average amount of fructose and glucose in the harvest ready fruits and optionally a significantly reduced average amount of malic acid, wherein the higher glucose and fructose content (and optionally the lower malic acid content) is conferred by an introgression fragment on the cultivated cucumber chromosome 1, wherein said introgression fragment is e.g. obtained by (or obtainable by) crossing a cultivated plant grown from seeds deposited under NCIMB 44295 or progeny of this plant (which comprises one or more the markers disclosed herein linked to the QTL) with a cultivated cucumber plant. Thus in one aspect the cultivated cucumber plant of the invention comprises the same introgression fragment and the same recombinant chromosome 1 as present in NCIMB 44295 (comprising all of the wild donor haplotype for SNP markers SNP 01 to SNP 14 or comprising SEQ ID NO: 1 to 14 or comprising SEQ ID NO: 1 to 10, SEQ ID NO: 34, and SEQ ID NO: 12 to 14), or it comprises a shorter fragment of that introgression fragment, whereby the shorter fragment retains the genetic element conferring increased fruit glucose and fructose levels (QTL 1.1).

[0131] Thus in one aspect the invention relates to a cultivated Cucumis sativus var. sativus plant comprising an introgression fragment comprising QTL 1.1 (or a variant) from a wild cucumber on chromosome 1 in homozygous or heterozygous form and wherein said introgression fragment is the introgression fragment “as in” / is “identical to” / is “the same as in” the seeds deposited under number NCIMB 44295, or is a shorter fragment thereof, but which still confers an increased fruit glucose and fructose level due to the presence of QTL1. 1 (or a variant).

[0132] In yet another embodiment the invention relates to a plant of the invention i.e. a cultivated Cucumis sativus var. sativus plant comprising an introgression fragment comprising QTL1. 1 (or a variant) from a wild cucumber on chromosome 1 in homozygous or heterozygous form and wherein said introgression fragment is the introgression fragment or is a variant of the introgression fragment present in seeds deposited under number NCIMB 44295, i.e. it comprises the QTL 1.1 (or a variant), but the genomic sequence may be different in the variant. As wild accessions will be genetically divergent, the genomic sequence of an introgression fragment comprising QTL1. 1 from other wild or primitive cucumbers will most likely not be identical to the genomic sequence as introgressed into NCIMB 44295. For example certain donor SNP markers linked to QTL1. 1 may be commonly found in various accessions, while other donor SNP markers may only be found in specific accessions. So, for example not all of SNP_01 to SNP_14 may be found in other wild cucumber donors. However, QTL1. 1 (comprising e.g. a variant or ortholog of the sugar allele) may still be present in such wild accessions. The skilled person is capable of identifying and introgressing the QTL1. 1 comprising region found in other wild cucumber donors into cultivated cucumber, e.g. detecting wild accessions comprising the donor SNP markers, or a subset thereof, and transferring these donor SNP markers (or subset) into a cultivated cucumber line or variety and assessing the fruit glucose and fructose content of the cultivated line or variety compared to the line or variety lacking the donor SNP markers (or subset), i.e. lacking the introgression fragment. Even in cases where the donor SNP haplotype for SNP_01 to SNP_14 is identical to the SNP haplotype of QTL1.1 found in seeds of NCIMB 44295, the actual nucleotide sequences flanking the SNP at nucleotide 102 of SEQ ID NO: 1 to 14 may be different in other donors. So other donors may comprise the same SNP nucleotide at nucleotide 102 (or at a position equivalent to position 102), but in a sequence comprising at least 95%, 96%, 97%, 98% or 99% sequence identity to SEQ ID NO: 1 to 14 when e.g. aligned pairwise. This variation can be seen by sequencing the donors and aligning sequences of SEQ ID NO: 1 to SEQ ID NO: 14 with that sequence. An example herein is SNP_11, where the flanking sequence to the left and to the right of the SNP is not 100% identical to SEQ ID NO: 11 but is 99% identical to SEQ ID NO: 11. This sequence is provided in SEQ ID NO: 34.

[0133] In one embodiment a cultivated cucumber plant or plant part is provided wherein the presence of the introgression fragment comprising QTL1.1, or the chromosome 1 region (or variant or orthologous chromosome 1 region), comprising QTL1. 1, is detectable by a molecular marker assay which detects at least 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13 or 14 Single Nucleotide Polymorphism (SNP) markers selected from the group consisting of: a) a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_01 at nucleotide 102 of SEQ ID NO: 1 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 1; b) an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_02 at nucleotide 102 of SEQ ID NO: 2 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 2; c) a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_03 at nucleotide 102 of SEQ ID NO: 3 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 3; d) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_04 at nucleotide 102 of SEQ ID NO: 4 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 4; e) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_05 at nucleotide 102 of SEQ ID NO: 5 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 5; f) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_06 at nucleotide 102 of SEQ ID NO: 6 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 6; g) a Thymine (T) nucleotide for the Sing le Nucleotide Polymorphism marker SNP_07 at nucleotide 102 of SEQ ID NO: 7 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 7; h) an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_08 at nucleotide 102 of SEQ ID NO: 8 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 8; i) a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_09 at nucleotide 102 of SEQ ID NO: 9 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 9; j) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_10 at nucleotide 102 of SEQ ID NO: 10 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 10; k) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_11 at nucleotide 102 of SEQ ID NO: 11 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 11, such as a Guanine for SNP l 1 at nucleotide 102 of SEQ ID NO: 34; l) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_12 at nucleotide 102 of SEQ ID NO: 12 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 12; m) an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_13 at nucleotide 102 of SEQ ID NO: 13 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 13; n) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_14 at nucleotide 102 of SEQ ID NO: 14 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 14.

[0134] In one aspect said at least 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13 or 14 markers which are detected are consecutive markers, e.g. markers from SNP_2 to SNP_13, or from SNP_06 to SNP_13.

[0135] When referring herein to markers ‘from . . . to” this encompasses in one aspect the starting and end marker or, alternatively, only the markers located in between the start and end marker. Likewise, when referring to a region in-between two markers (e.g. a QTL being present in-between two flanking markers), this may either include the start and end marker, or it may include only the region and markers located in-between the starting and end marker. In one aspect a method for screening genomic DNA of wild or cultivated cucumber plants or plant parts by carrying out the above molecular marker assay is encompassed herein. Optionally the method further comprises selecting a plant or plant part comprising the above nucleotide for at least 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13 or all 14 of the SNP markers SNP_01 to SNP_14.

[0136] Thus, in one embodiment the plant (or plant part) according to the invention comprises: at least a Cytosine (C) (i.e. the CC or CX genotype) at nucleotide 102 of SEQ ID NO: 1 (referred to as SNP_01) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO: 1 (in other words there is a Cytosine at the physical position of chromosome 1 shown in Table 2); and / or at least an Adenine (A) (i.e. the AA or AX genotype) at nucleotide 102 of SEQ ID NO: 2 (referred to as SNP_02) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO:2 (in other words there is an Adenine at the physical position of chromosome 1 shown in Table 2); and / or at least a Cytosine (C) (i.e. the CC or CX genotype) at nucleotide 102 of SEQ ID NO: 3 (referred to as SNP_03) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO:3 (in other words there is a Cytosine at the physical position of chromosome 1 shown in Table 2); and / or at least a Guanine (G) (i.e. the GG or GX genotype) at nucleotide 102 of SEQ ID NO: 4 (referred to as SNP_04) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO:4 (in other words there is a Guanine at the physical position of chromosome 1 shown in Table 2); and / or at least a Thymine (T) (i.e. the TT or TX genotype) at nucleotide 102 of SEQ ID NO: 5 (referred to as SNP_05) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO:5 (in other words there is a Thymine at the physical position of chromosome 1 shown in Table 2); and / or at least a Thymine (T) (i.e. the TT or TX genotype) at nucleotide 102 of SEQ ID NO: 6 (referred to as SNP_06) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO:6 (in other words there is a Thymine at the physical position of chromosome 1 shown in Table 2); and / or at least a Thymine (T) (i.e. the TT or TX genotype) at nucleotide 102 of SEQ ID NO: 7 (referred to as SNP_07) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO:7 (in other words there is a Thymine at the physical position of chromosome 1 shown in Table 2); and / or at least an Adenine (A) (i.e. the AA or AX genotype) at nucleotide 102 of SEQ ID NO: 8 (referred to as SNP_08) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO: 8 (in other words there is an Adenine at the physical position of chromosome 1 shown in Table 2); and / or at least a Cytosine (C) (i.e. the CC or CX genotype) at nucleotide 102 of SEQ ID NO: 9 (referred to as SNP_09) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO:9 (in other words there is a Cytosine at the physical position of chromosome 1 shown in Table 2); and / or at least a Thymine (T) (i.e. the TT or TX genotype) at nucleotide 102 of SEQ ID NO: 10 (referred to as SNP_10) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NOTO (in other words there is a Thymine at the physical position of chromosome 1 shown in Table 2); and / or at least a Guanine (G) (i.e. the GG or GX genotype) at nucleotide 102 of SEQ ID NO: 11 or at nucleotide 102 of SEQ ID NO: 34 (referred to as SNP_11) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO: 11, such as a Guanine for SNP l 1 at nucleotide 102 of SEQ ID NO: 34 (in other words there is a Guanine at the physical position of chromosome 1 shown in Table 2); and / or at least a Guanine (G) (i.e. the GG or GX genotype) at nucleotide 102 of SEQ ID NO: 12 (referred to as SNP_12) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO: 12 (in other words there is a Guanine at the physical position of chromosome 1 shown in Table 2); and / or at least an Adenine (A) (i.e. the AA or AX genotype) at nucleotide 102 of SEQ ID NO: 13 (referred to as SNP_13) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO: 13 (in other words there is an Adenine at the physical position of chromosome 1 shown in Table 2); and / or at least a Guanine (G) (i.e. the GG or GX genotype) at nucleotide 102 of SEQ ID NO: 14 (referred to as SNP_14) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO: 14 (in other words there is a Guanine at the physical position of chromosome 1 shown in Table 2).

[0137] In a further one embodiment the presence of the introgression fragment, or the chromosome 1 region (or variant or orthologous chromosome 1 region), comprising QTL1.1, is detectable by a molecular marker assay which detects the donor SNP nucleotide for at least 2, 3, 4, 5, 6, 7, 8, 9 or 10 Single Nucleotide Polymorphism (SNP) markers of the sub-groups consisting of: SNP_08 to SNP_14, or SNP_08 to SNP_13 or SNP_08 to SNP_12 or SNP_09 to SNP_14 or SNP_09 to SNP_13 or SNP_09 to SNP_12, or SNP_10 to SNP_14 or SNP_10 to SNP_13 or SNP_10 to SNP_12.

[0138] The SNP genotype refers to two nucleotides, and genomic sequences comprising one of these two nucleotides, one on each chromosome 1. So a plant having a CC genotype for SNP_01 has an identical nucleotide (C) on both chromosomes (i.e. is homozygous), while a plant having an CX genotype for SNP_01 has one chromosome with an C at nucleotide 102 of SEQ ID NO: 1 (or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO: 1) and one chromosome with a X at nucleotide 102 of SEQ ID NO: 1 (or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO: 1) and is heterozygous, whereby X may be any nucleotide. As the genomic sequences around the SNP markers provided herein may vary slightly in introgression fragments from other wild cucumber donors (i.e. variants or orthologous chromosome 1 regions) it is clear that the nucleotide sequences before and after the SNP may not be 100% identical to the sequences (or complement sequences) provided herein. Therefore, sequences having substantial sequence identity (i.e. at least 95% identity) to the sequences provided herein, but which comprise the same donor SNP nucleotide, are encompassed herein. For example the donor nucleotide G for SNP_11 may be present in a sequence comprising at least 99% sequence identity to SEQ ID NO: 11, such as SEQ ID NO: 34. The ‘complement’ sequence or ‘complementary’ sequence is the other strand of the double stranded DNA molecule. It may also be referred to as ‘reverse complement strand’ when taking the 5 prime (5’) to 3 prime (3’) orientation of the strands into consideration.

[0139] In one aspect, the introgression fragment comprising QTL 1.1, or the chromosome 1 region (or variant or orthologous chromosome 1 region) comprising the QTL (QTL1. 1 or variant), which is detectable by the above one or more markers is from a wild or primitive cucumber. In one aspect it is the same introgression fragment as found on chromosome 1 in seeds deposited under accession number NCIMB 44295, or a smaller fragment retaining the QTL. SNP markers SNP_01 to SNP_14 span a region of about 2.79 Mb. In one aspect the introgression fragment on chromosome 1 is equal to or less than 3 Mb in size, preferably equal to or less than 2.79 Mb in size, more preferably equal to or less than 2.5 Mb, 2.0Mb or 1.5 Mb in size, e.g. equal to or less than 1 Mb. In one aspect the introgression fragment is at least 0.2 Mb, 0.5 Mb, 1.0 Mb, 1.5 Mb, 1.9 Mb, 2.0 Mb, 2.5 Mb, 2.7Mb, 2.8Mb or 3 Mb in size. Thus, various ranges of introgression sizes are encompassed herein, such as fragments less than 3 Mb but more than 0.2 Mb, which retain the QTL1. 1 and one or more of the SNP markers of the donor markers for SNP_01 to SNP_14, or of the subgroups of SNP_08 to SNP_14, or SNP_08 to SNP_13 or SNP_08 to SNP_12 or SNP_09 to SNP_14 or SNP_09 to SNP_13 or SNP_09 to SNP_12, or SNP_10 to SNP_14 or SNP_10 to SNP_13 or SNP_10 to SNP_12. As mentioned before, the location of the QTL 1.1 in the region spanning SNP_01 to SNP_14 can be determined by fmemapping and recombinants comprising QTL1. 1 on a smaller introgression fragment can be generated. The size of an introgression fragment can be easily determined by e.g. whole genome sequencing or Next Generation Sequencing, e.g. as described in Qi et al. 2013 (supra) or in Huang et al. 2009 (supra). Especially introgression regions can be easily distinguished from cultivated genomic regions due to the larger amount of genetic variation (SNPs, INDELs, etc.) in the introgression region.

[0140] To obtain the introgression fragment present on chromosome 1 (comprising QTL1. 1) from the deposited seeds (NCIMB 44295), i.e. to transfer the introgression fragments comprising the QTL to another cultivated cucumber plant, a plant is grown from the seed and the plant is crossed with a cultivated cucumber plant to obtain Fl seeds. As NCIMB 44295 contains two recombinant chromosomes 1 (comprising the introgression fragment) all of the F 1 seed and plants grown therefrom, contain one recombinant chromosome 1 from the NCIMB 44295 parent and one non-recombinant chromosome 1 from the other cultivated parent. Thus, by traditional breeding one can transfer the recombinant chromosome 1 from NCIMB44295 into other cultivated cucumber lines or varieties. Plants which comprise the QTL1. 1 (or a variant) can be screened for, and selected for, by the presence of one or more of the above donor SNP markers in order to identify plants comprising a recombinant chromosome 1.

[0141] To generate shorter introgression fragments (comprising QTL 1.1) meiosis needs to take place and plants comprising the recombinant chromosomes 1, and especially new meiotic recombination events within the introgression fragment, need to be identified. For example, seeds of NCIMB44295 can be selfed one or more times to produce F 1, F2 or F3 plants (or further selfing generations), and / or F 1, F2 or F3 plants (etc.) comprising a recombinant chromosome 1 can be backcrossed to a cultivated parent. Plants which comprise the recombinant chromosome 1 can be screened for, and selected for, by the presence of one or more of the above SNP markers in order to identify plants comprising a smaller introgression fragment. Such new recombinants can then be tested for the presence of the QTL1. 1 on the smaller introgression fragment by determining the average fruit glucose and fructose levels compared to the (genetic) control lacking the introgression fragment.

[0142] Similarly, cultivated cucumber plants comprising QTL 1.1 (or a variant thereof) can be generated and / or identified using different methods. For example, to obtain a cultivated cucumber plant comprising an introgression fragment from a wild donor, a wild donor is identified which comprises one or more of the donor SNP markers linked to QTL1. 1 disclosed herein, e.g. any one, or more, or all of the markers described herein above. This has for example been done for various wild accessions. The identified plant is crossed with a cultivated cucumber plant to obtain Fl seeds. The Fl can be selfed to produce F2, F3, etc. plants, and / or F2 plants or F3 plants, etc., can be backcrossed to the cultivated cucumber parent. Plants which are comprising QTL1. 1 (or a variant thereof) can be screened for, and / or selected for, by the presence of one or more of the above donor SNP markers and / or screened for, and / or selected for, an increased glucose and fructose fruit level compared to the initial cultivated parent (lacking the introgressions). Alternatively, or in addition, QTL mapping can be carried out in order to identify further molecular markers linked to the QTL1. 1 (or a variant thereof) and / or to generate cultivated cucumber plants comprising an introgression fragment on chromosome 1 which confers significantly higher average fruit glucose and fructose levels.

[0143] In one embodiment a method is provided for detecting the presence of the introgression fragment comprising QTL1. 1 or a variant thereof in a (cultivated or wild) cucumber plant or plant part (cell, fruit, fruit part, leaf, stem, etc.), or the chromosome 1 region comprising QTL1.1 or a variant thereof, said method comprises analyzing the genomic DNA of the plant or plant part using a molecular marker assay which detects at least 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13 or all 14 of the markers selected from the group consisting of: a) a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_01 at nucleotide 102 of SEQ ID NO: 1 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 1; b) an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_02 at nucleotide 102 of SEQ ID NO: 2 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 2; c) a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_03 at nucleotide 102 of SEQ ID NO: 3 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 3; d) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_04 at nucleotide 102 of SEQ ID NO: 4 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 4; e) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_05 at nucleotide 102 of SEQ ID NO: 5 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 5; f) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_06 at nucleotide 102 of SEQ ID NO: 6 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 6; g) a Thymine (T) nucleotide for the Sing le Nucleotide Polymorphism marker SNP_07 at nucleotide 102 of SEQ ID NO: 7 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 7; h) an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_08 at nucleotide 102 of SEQ ID NO: 8 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 8; i) a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_09 at nucleotide 102 of SEQ ID NO: 9 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 9; j) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_10 at nucleotide 102 of SEQ ID NO: 10 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 10; k) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_11 at nucleotide 102 of SEQ ID NO: 11 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 11, such as a Guanine for SNP l 1 at nucleotide 102 of SEQ ID NO: 34; l) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_12 at nucleotide 102 of SEQ ID NO: 12 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 12; m) an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_13 at nucleotide 102 of SEQ ID NO: 13 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 13; n) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_14 at nucleotide 102 of SEQ ID NO: 14 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 14.

[0144] The method optionally further comprises selecting the plant or plant part comprising at least 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13 or all 14 of the marker nucleotides of a) to n). The SNP haplotype indicative of the QTL, i.e. the donor SNP haplotype for one or more of SNP_01 to SNP_14 as described elsewhere herein and in e.g. Table 2 or Table 3 may be detected in the plant or plant part. The method may involve screening a population of plants or plant parts (or genomic DNA thereof) using the above SNP markers. Obviously, a SNP assay can be designed for the above SNP markers using the plus strand or the minus strand of the DNA. The method may further comprise generating plants that are homozygous for the selected donor nucleotides of the SNP markers and / or allowing the plants to develop fruits and optionally analyzing the fruit glucose and fructose content. Further aspects of the above methods are described elsewhere herein. The introgression fragment in the plants of the invention is in one aspect a fragment of chromosome 1 (comprising QTL1. 1) which is present in seeds deposited under accession number NCIMB44295 or a smaller version of that fragment retaining the QTL (generated by e.g. recombination within the introgression fragment). The peak QTL1. 1 marker, SNP l 1, is a Guanine for nucleotide 102 of SEQ ID NO: 11 or it is a Guanine for the equivalent nucleotide in a sequence comprising at least 95%, 96%, 97%, 98% or 99% sequence identity to SEQ ID NO: 11, e.g. at nucleotide 102 of SEQ ID NO: 34, which is the genomic sequence present in the deposited seeds.

[0145] QTL1. 1 and the introgression fragment comprising QTL1. 1 is in one aspect in homozygous form in the plant or plant part, such as the fruits. The SNP marker genotype for one or more of SNP_01 to SNP_14 in this aspect comprises the donor SNP nucleotide in homozygous form.

[0146] QTL1. 1 and the introgression fragment comprising QTL1. 1 is in one aspect in heterozygous form in the plant or plant part, such as the fruits. The SNP marker genotype for one or more of SNP_01 to SNP_14 in this aspect comprises the donor SNP nucleotide in heterozygous form. Heterozygous form is especially preferred in a w / w background and / or in combination with QTL4. 1, as described elsewhere herein.

[0147] Also provided are seeds from which a plant of the invention can be grown, as are cucumber fruits harvested from a plant of the invention and comprising the recombinant chromosome 1 in their genome (comprising QTL1. 1 or a variant). Likewise, a plant cell, tissue or plant part of a plant or of a seed is provided comprising at least one recombinant chromosome 1 (comprising QTL 1.1 or a variant), wherein said recombinant chromosome 1 comprises an introgression fragment from a wild or primitive cucumber donor and wherein said introgression fragment comprises QTL1. 1 or a variant thereof.

[0148] The molecular markers described herein may be detected according to standard method. For example, SNP markers can easily be detected using a KASP-assay (see www.kpbioscience.co.uk) or other SNP genotyping assays. For developing a KASP-assay, for example 50 or 70 base pairs upstream and 50 or 70 base pairs downstream of the SNP can be selected and two allele-specific forward primers and one allele specific reverse primer can be designed. See e.g. Allen et al. 2011, Plant Biotechnology J. 9, 1086-1099, especially p097-1098 for KASP assay method.

[0149] Thus, in one aspect, the SNP markers and the presence / absence of the marker associated with QTL 1.1 is determined using a KASP assay, but equally other SNP genotyping assays can be used. For example, a TaqMan SNP genotyping assay, a High Resolution Melting (HRM) assay, SNP- genotyping arrays (e.g. Fluidigm, Illumina, etc.) or DNA sequencing may equally be used. The physical size of an introgression fragment can be determined by various methods, such as physical mapping, sequencing or by visualization of the introgression using Fluorescent in situ hybridization (FISH) images (Verlaan et al. 2011, Plant Journal 68: 1093-1103).

[0150] Cultivated cucumber plants with smaller introgression fragments on chromosome 1 (comprising QTL1. 1 or a variant) can be generated by generating new recombinant plants from a population of plants derived from a cross between a cultivated cucumber plant (lacking the introgressions) and a plant of the invention and selecting recombinant progeny having smaller introgression sizes. Such plants are thus in one aspect derived from (progeny or descendants of) the recombinant chromosome 1 present in plants of which seeds have been deposited under NCIMB44295. Such progeny or descendants which retain the QTL1. 1, and whereof the fruits produce at least a higher (average) amount of glucose and fructose compared to the fruits of plants lacking an introgression as described herein, are encompassed herein.

[0151] QTL3, 1 or variants of QTL3, 1 on chromosome 3

[0152] In another aspect a cultivated cucumber plant (as well as plant parts such as cells, tissues, leaves, stems, flowers, fruits, seeds, etc.) is provided comprising an introgression fragment from a wild or primitive cucumber, wherein the introgression fragment comprises QTL3.1, or a variant thereof, and wherein the introgression fragment comprises all or part of the region starting at nucleotide (or base) 1128135 of chromosome 3 (corresponding to SNP_15) and ending at nucleotide (or base) 2585865 of chromosome 3 (corresponding to SNP_24). In other words, all or part of the region starting at nucleotide 1128135 of chromosome 3 (SNP_15) and ending at nucleotide 2585865 of chromosome 3 (SNP_24) is, in one aspect, from a wild donor cucumber and comprises QTL3. 1 or a variant thereof. Which sub-region contains QTL3. 1 can be identified by e.g. fine- mapping. So, for example if QTL3.1 is found to be in between SNP 16 and SNP 19, then the plant of the invention only needs to comprise the introgression region starting at nucleotide 1135483 of chromosome 3 (SNP_16) and ending at nucleotide 1689700 (SNP_19) of chromosome 3.

[0153] In one aspect QTL3. 1 (or a variant thereof) is located in-between marker SNP_15 at nucleotide 102 of SEQ ID NO: 15 (or at nucleotide 102 in a variant sequence of SEQ ID NO: 15, such as SEQ ID NO: 35) and marker SNP_19 at nucleotide 102 of SEQ ID NO: 19 (or at nucleotide 102 in a variant sequence of SEQ ID NO: 19).

[0154] In another aspect QTL3. 1 (or a variant thereof) is located in-between marker SNP_15 at nucleotide 102 of SEQ ID NO: 15 (or at nucleotide 102 in a variant sequence of SEQ ID NO: 15, such as SEQ ID NO: 35) and marker SNP_18 at nucleotide 102 of SEQ ID NO: 18 (or at nucleotide 102 in a variant sequence of SEQ ID NO: 18). In a further aspect QTL3. 1 (or a variant thereof) is located in-between the beginning of the chromosome 3 (i.e. nucleotide 1 of chromosome 3) and SNP_16, or SNP_17 or SNP_18 or SNP_19. In another aspect the introgression fragment of the invention (comprising QTL3.1 or a variant thereof) is a fragment comprising a smaller fragment (part) of the region starting at nucleotide (or base) 1128135 of chromosome 3 and ending at nucleotide (or base) 2585865 of chromosome 3, e.g. having a size of e.g. 1.5 Mb, 1.46 Mb, 1.3 Mb, 1.0 Mb, 0.5Mb, lOOkb, 50kb, 35kb, 30kb, 20kb, or less and comprising the QTL or a variant thereof. In one aspect the part is at least 5kb, lOkb, 20kb in size, or more.

[0155] In one aspect the cultivated cucumber plant of the invention comprises an introgression fragment from a wild or primitive cucumber, which introgression fragment comprises QTL3.1 or a variant thereof, wherein the introgression fragment comprises all of part of the region starting at 1.457730 Mb and ending at 2.585865 Mb of the physical chromosome 3 or all or part of the region starting at the beginning of chromosome 3 and ending at 2.585865 Mb of the physical chromosome 3.

[0156] In one aspect the introgression fragment on chromosome 3 comprising QTL3.1, or a variant thereof, is obtainable by crossing a plant grown from NCIMB44295 with another cucumber plant, especially a cultivated cucumber plant, in one aspect a long cucumber type or a short cucumber type or a pickling or slicer type.

[0157] In one aspect the cultivated cucumber plant of the invention comprising QTL 3. 1, or a variant thereof, is a plant wherein said introgression fragment on chromosome 3 is obtainable by crossing a plant grown from seeds deposited under accession number NCIMB44295 with another cucumber plant. Thus, in one aspect the QTL is the QTL present in seeds deposited under accession number NCIMB44295.

[0158] In a further aspect the cultivated cucumber plant of the invention comprising QTL3. 1, or a variant thereof, is a plant wherein said introgression fragment on chromosome 3 is obtainable by crossing a plant comprising the same donor SNP haplotype or SNP genotype for at least 2, 3, 4, 5, 6, 7, 8, 9, 10 or more SNP markers linked to the QTL (i.e. SNP_15 to SNP_24 for QTL3.1, as e.g. shown in Table 5) with another cucumber plant, especially with a cultivated cucumber elite breeding line. Thus, in one aspect the QTL is the variant QTL present in wild donor accessions or in cultivated cucumber which comprise the same donor SNP haplotype or genotype for at least 2, 3, 4, 5, 6, 7, 8, 9, 10 or more of the SNP markers e.g. as present in NCIMB44295. Preferably the donor also comprises an internal mint-green fruit flesh color, as does the cultivated cucumber plant comprising the introgression fragment (at least when the introgression fragment and / or QTL3.1 is in homozygous form). Herein a mint-green fruit flesh color of a plant comprising QTL3. 1 (at least when QTL3. 1 is in homozygous form) is in one aspect a fruit flesh color having a RHS color chart rating of Yellow-Green Groupl45A and / or 145B and / or optionally 145C, especially near the center around the seed cavity (Fan 3 Blue-Green Green Green- Yellow Groups of the RHS Color Chart, 5thEdition, The Royal Horticultural Society 2007), see also Figures 1 and 2. In one aspect the fruit flesh color at e.g. harvest stage is significantly more mint green or more greenyellow than in the fruits of a plant lacking QTL3. 1 (e.g. a control or recurrent parent), when grown under the same conditions and measured in the same way, e.g. as described in the Examples. There are different ways to measure color and it is understood that QTL3.1 (especially when present in homozygous form) changes the fruit flesh color significantly compared to the recurrent parent or control plant (e.g. genetic control) lacking QTL3.1.

[0159] When referring to the SNP markers herein, which are indicative of the presence of the introgression fragment (and / or of the fruit flesh color QTL present on the introgression fragment), it is understood that the donor SNP genotype or haplotype which is indicative of the introgression fragment is referred to, i.e. the donor SNP genotype or haplotype as provided e.g. in Table 5 and Table 6. It is noted that the SNP marker genotype can distinguish between the introgression fragment being in homozygous or heterozygous form. In homozygous form the nucleotide is identical, while in heterozygous form the nucleotide is not (necessarily) identical. The SNP genotype of the ‘wild type’ (wt) chromosome lacking the introgression fragment is e.g. the other haplotype, e.g. the haplotype of the recurrent parent). So, e.g. the genotype of SNP_15 indicative of the introgression fragment comprising QTL3.1 is e.g. ‘AC’ (QTL3.1 / wt) or ‘AA’ (QTL3.1 / QTL3.1) while the SNP genotype indicative of the wild type / genetic control (lacking the introgression fragment) is e.g. ‘CC’ (wt / wt). This can also be written as genotype AX’ (QTL3.1 / wt) or ‘AA’ (QTL3.1 QTL3.1) while the SNP genotype indicative of the wild type / genetic control (lacking the introgression fragment) is e.g. ‘XX’ (wt / wt). X may be any nucleotide (A, T, C or G). Thus, when referring to a plant or plant part (e.g. cell) comprising the introgression fragment in homozygous or heterozygous form, it is understood that the SNP markers linked to the introgression fragment have the corresponding SNP genotype or haplotype.

[0160] So, in one aspect, a cultivated Cucumis sativus var. sativus plant is provided comprising an introgression fragment on chromosome 3 in homozygous or heterozygous form, wherein said introgression fragment confers a mint-green fruit flesh color (at least when the introgression fragment is in homozygous form) compared to the fruits produced by the cucumber plant lacking the introgression fragment on chromosome 3, e.g. the genetic control or control variety, when grown under the same conditions.

[0161] The mint-green fruit flesh color is visually determinable or can be measured using color measurement methods known in the art, such as using a spectrophotometer or a color chart. See for example Figure 1 and 2 of fruits with mint green flesh color and mint green peel color. These fruits are homozygous for QTL3. 1. The fruit flesh color of fruits produced by the cultivated cucumber plant line or variety comprising the introgression fragment on chromosome 3 in homozygous or heterozygous form can be compared to the genetic control line or variety lacking the introgression fragment on chromosome 3 when grown under the same environment. Fruit flesh color can also be measured on a scale of 1 to 9, as described in the Examples (see Example 1.2). In one aspect the average fruit flesh color of fruits produced by plants comprising QTL3.1 in homozygous form has an average score of at least 5.0 (yellow-green) or at least 6.0 (green-yellow) but the average score may, in one aspect, be below 7.0 (green) on the scale from 1 to 9 of the Examples. So, an average score of 5.0 or above 5.0 (such as at least 5. 1, 5.2, 5.3, 5.39, 5.4, 5.5, 5.6, 5.7, 5.8, 5.9), or of 6.0 or above 6.0 (such as at least 6.1, 6.2, 6.3, 6.39, 6.4, 6.5, 6.6, 6.7, 6.8, 6.9), but in one aspect below 7.0, is encompassed herein. When QTL3.1 is heterozygous form the average fruit flesh color has in one aspect an average score of at least 5.0 (yellow-green), but may, in one aspect, be below 6.0 (green-yellow). So an average score of 5.0 or above 5.0 (such as at least 5.1, 5.2, 5.25, 5.75, 5.3, 5.39, 5.4, 6.5, 5.6, 5.7, 5.8, 5.9), but (in one aspect) below 6.0, is encompassed herein. In one aspect the fruit flesh color is changed from e.g. light-yellow (average score of 3.0, or above 3.0 but optionally below 4.5 or below 5.0) to at least yellow-green (at least 5.0) (e.g. heterozygous form) or to at least green-yellow (at least 6.0) (e.g. homozygous form). In one aspect the presence of QTL3. 1 increases the average fruit flesh color score by at least 1.0 compared to a plant / fruits lacking QTL3. 1, preferably by at least 1.5, 2.0, 2.5 or 3.0 score points on a scale of score 1 to 9 as described in Example 1.2, where 5.0 is yellow-green and 6.0 is green-yellow and 7.0 is green. So, an average score of e.g. 3.0 in a plant lacking QTL3. 1 will increase to an average score of at least 4.0, preferably at least 4.5, 5.0, 5.5 or 6.0 due to the presence of QTL3. 1 (at least when in homozygous form). It is understood that the increase in fruit flesh color score is higher when QTL3. 1 is in homozygous form than when it is in heterozygous form. Yellow-green and green-yellow may also be referred to as ‘light mint-green’ and ‘mint-green’, respectively, and green may also be referred to as ‘darker mint-green’.

[0162] The effect of QTL3. 1 can, thus, be determined by introducing the QTL into a cucumber plant, in heterozygous or preferably in homozygous form. For example NCIMB44295 can be crossed with a cucumber plant lacking QTL3. 1 and QTL3. 1 can, thus, be transferred into the background by e.g. backcrossing.

[0163] The plants provided herein, therefore, in one aspect comprise a genome of cultivated cucumber, with at least one or two recombinant chromosomes, namely one or two recombinant chromosomes 3 (i.e. heterozygous or homozygous). The recombinant chromosomes comprise a fragment of a wild donor cucumber, which is easily distinguishable from the cultivated cucumber genome by molecular marker analysis, whole genome sequencing, chromosome painting and similar techniques.

[0164] In one aspect the introgression fragment on chromosome 3 is from a wild or primitive cucumber, comprises the fruit-flesh color QTL3.1, or a variant thereof, and comprises all or part of the region starting at nucleotide SNP_15 and ending at SNP_24. Thus, the introgression fragment comprises the QTL3.1 or a variant thereof and one or more or all (e.g. 3, 4, 5, 6, 7, 8, 9, 10) SNP markers of the wild donor selected from SNP_15 to SNP_24 as shown in Table 5 and 6. In one aspect the introgression fragment comprises QTL3. 1 and one or more or all of SEQ ID NO: 15 to SEQ ID NO: 24. In another aspect the introgression fragment comprises QTL3. 1 and one or more or all of SEQ ID NO: 35 (comprising SNP_15), SEQ ID NO: 16 (comprising SNP_16), SEQ ID NO: 36 (comprising SNP_17), SEQ ID NO: 18 (comprising SNP_18), SEQ ID NO: 19 (comprising SNP_19), SEQ ID NO: 37 (comprising SNP_20), SEQ ID NO: 21 to 24 (comprising SNP_21 to SNP_24 respectively).

[0165] In one aspect the presence of the introgression fragment on chromosomes 3 comprising QTL3. 1 in the genome of the plant or plant cell or plant tissue (or in the DNA extracted therefrom) is detectable by a molecular marker assay which detects one or more molecular markers indicative of the introgression fragment comprising QTL3. 1, especially the donor SNP haplotype or genotype for at least 2, 3, 4, 5, 6, 7, 8, 9, 10 or more of SNP_15 to SNP_24, at nucleotide 102 of SEQ ID NO: 15 to 24 (or the complement sequence of SEQ ID NO: 15 to 24, or a sequence comprising at least 95%, 96%, 97%, 98% or 99% sequence identity to SEQ ID NO: 15 to 24 or to the complement of SEQ ID NO: 15 to 24), respectively. However, as mentioned, other techniques may be used, e.g. the SNP genotype of the markers may also be determined by sequencing or by using alternative markers located in between the SNP markers provided herein or within 7cM, or within 5cM, of a marker provided herein; or within 2 Mb, 1.5Mb, 1 Mb, 0.5 Mb, 0.4Mb, 0.3Mb, 0.2Mb, 0. 1 Mb, 50kb, 20kb, lOkb, 5kb, 2kb, Ikb or less of a marker provided herein.

[0166] In one aspect the presence of the introgression fragment on chromosomes 3 comprising QTL3. 1 in the genome of the plant or plant cell or plant tissue (or in the DNA extracted therefrom) is detectable by detecting the presence of one or more or all of SEQ ID NO: 15 to SEQ ID NO: 24. In another aspect the presence of the introgression fragment on chromosomes 3 comprising QTL3. 1 in the genome of the plant or plant cell or plant tissue (or in the DNA extracted therefrom) is detectable by detecting the presence of one or more or all of SEQ ID NO: 35 (comprising SNP_15), SEQ ID NO: 16, SEQ ID NO: 36 (comprising SNP_17), SEQ ID NO: 18, SEQ ID NO: 19, SEQ ID NO: 37 (comprising SNP_20), SEQ ID NO: 21 to 24.

[0167] It is noted that when referring herein to an introgression fragment or QTL being detectable or being present in a plant ort plant part, all embodiments encompass also methods for detecting the introgression fragment or QTL in the plant or plant part, e.g. by marker assisted selection, SNP haplotyping, genotyping, sequencing etc., and / or methods for selecting a plant or plant part comprising the introgression fragment or QTL.

[0168] When reference is made herein to one or more molecular markers or sequences being “detectable” by e.g. a molecular marker assay, this means of course that the plant or plant part comprises the one or more markers or sequences in its genome, as the marker or sequence would otherwise not be detectable. Further, when referring to a plant or plant part in which the SNP marker or SNP haplotype of the donor or introgression fragment comprising QTL3. 1 is present or detectable, this also encompasses SNP assays and methods for detecting the SNP marker and the SNP haplotype or genotype present and / or for selecting a plant or plant part comprising the donor SNP haplotype or genotype.

[0169] Furthermore, when a plant is referred to, this also encompasses plant parts, such as cells, tissues, leaves, stems, fruits, flowers, pollen, roots, etc. and seeds from which the plant can be grown, seeds produced by the plant and retaining the QTL.

[0170] Cucumber plants comprising an introgression fragment on chromosome 3 (QTL 3.1 or a variant thereof)

[0171] In one aspect a cultivated Cucumis sativus var. sativus plant comprising an introgression fragment from a wild or primitive cucumber on chromosome 3 in homozygous or heterozygous form is provided, wherein said introgression fragment comprises a Quantitative Trait Locus (herein referred to as QTL3. 1) located between the Single Nucleotide Polymorphism marker SNP_15 at nucleotide 102 of SEQ ID NO: 15 (or at nucleotide 102 of a variant of SEQ ID NO: 15, such as SEQ ID NO: 35) and the Single Nucleotide Polymorphism marker SNP_24 at nucleotide 102 of SEQ ID NO: 24 (or at nucleotide 102 of a variant of SEQ ID NO: 24), which QTL confers a mint-green fruit flesh color to the fruits, at least when the QTL is in homozygous form. In one aspect the QTL is located between base 1128135 (SNP_15) and base 2585865 (SNP_24) of chromosome 3.

[0172] Thus, in one aspect QTL3. 1 (or a variant thereof) is located in the region between SNP_15 in SEQ ID NO: 15 (or in a variant thereof, such as SEQ ID NO: 35) and SNP_24 in SEQ ID NO: 24 (or in a variant thereof).

[0173] Provided is in one aspect a cultivated Cucumis sativus var. sativus plant comprising an introgression fragments on chromosome 3 from a wild cucumber donor, wherein said introgression fragment comprises a Quantitative Trait Locus (QTL) referred to as QTL3. 1 conferring a mint-green fruit flesh color to the cucumber fruit (at least when the QTL is in homozygous form), wherein QTL3.1 is located on chromosome 3 between the Single Nucleotide Polymorphism marker SNP_15 at nucleotide 102 of SEQ ID NO: 15 (or SNP_15 at nucleotide 102 of SEQ ID NO: 35) and SNP_24 at nucleotide 102 of SEQ ID NO: 24, and wherein said introgression fragment on chromosome 3 comprising QTL3. 1 comprises a SNP haplotype of at least 2, 3, 4, 5, 6, 7, 8, 9 or 10 markers selected from the group consisting of: a) an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_15 at nucleotide 102 of SEQ ID NO: 15 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 15, such as an Adenine for SNP_15 at nucleotide 102 of SEQ ID NO: 35; b) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_16 at nucleotide 102 of SEQ ID NO: 16 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 16; c) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_17 at nucleotide 102 of SEQ ID NO: 17 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 17, such as a Guanine for SNP 17 at nucleotide 102 of SEQ ID NO: 36; d) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_18 at nucleotide 102 of SEQ ID NO: 18 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 18; e) an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_19 at nucleotide 102 of SEQ ID NO: 19 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 19; f) a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_20 at nucleotide 102 of SEQ ID NO: 20 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 20, such as a Cytosine for SNP_20 at nucleotide 101 of SEQ ID NO: 37; g) a Cytosine (C) nucleotide for the Sing le Nucleotide Polymorphism marker SNP_21 at nucleotide 102 of SEQ ID NO: 21 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 21; h) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_22 at nucleotide 102 of SEQ ID NO: 22 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 22; i) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_23 at nucleotide 102 of SEQ ID NO: 23 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 23; j) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_24 at nucleotide 102 of SEQ ID NO: 24 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 24.

[0174] Instead of referring to the SNP haplotype, which refers to only one chromosome of the chromosome pair, one can also refer to the SNP genotype, whereby reference is made to both chromosomes of a pair of chromosomes in a diploid organism. Therefore, an Adenine (A) for SNP_15 is then referred to as genotype AX or AA, where the first A is for one chromosome 3 of the pair and the X or the second A is for the other chromosome of the pair. The genotype AA for SNP_15 and one or more of the other markers thus has the introgression fragment on both chromosomes and is homozygous for the introgression fragment.

[0175] In one aspect, therefore, a cultivated Cucumis sativus var. sativus plant is provided comprising an introgression fragment on chromosome 3 in homozygous or heterozygous form, wherein said introgression fragment confers a mint-green color of the fruit flesh, at least when the QTL is in homozygous form, (especially a different fruit flesh color, especially more green-yellow or more green, than the plant lacking the introgression fragment, e.g. the genetic control) and wherein said introgression fragment comprises the SNP marker haplotype or genotype of at least 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13 or 14 of the SNP markers selected from the group consisting of: a) the AX or AA genotype for the Single Nucleotide Polymorphism marker SNP_15 at nucleotide 102 of SEQ ID NO: 15 (or at nucleotide 102 in a variant thereof, such as an Adenine for SNP_15 at nucleotide 102 of SEQ ID NO: 35); b) the TX or TT genotype for the Single Nucleotide Polymorphism marker SNP_16 at nucleotide 102 of SEQ ID NO: 16 (or at nucleotide 102 in a variant thereof); c) the GX or GG genotype for the Single Nucleotide Polymorphism marker SNP_17 at nucleotide 102 of SEQ ID NO: 17 (or at nucleotide 102 in a variant thereof, such as a Guanine for SNP_17 at nucleotide 102 of SEQ ID NO: 36); d) the TX or TT genotype for the Single Nucleotide Polymorphism marker SNP_18 at nucleotide 102 of SEQ ID NO: 18 (or at nucleotide 102 in a variant thereof); e) the AX or AA genotype for the Single Nucleotide Polymorphism marker SNP_19 at nucleotide 102 of SEQ ID NO: 19 (or at nucleotide 102 in a variant thereof); f) the CX or CC genotype for the Single Nucleotide Polymorphism marker SNP_20 at nucleotide 102 of SEQ ID NO: 20 (or at nucleotide 102 in a variant thereof, e.g. a Cytosine for SNP_20 at nucleotide 101 of SEQ ID NO: 37); g) the CX or CC genotype for the Single Nucleotide Polymorphism marker SNP_21 at nucleotide 102 of SEQ ID NO: 21 (or at nucleotide 102 in a variant thereof); h) the TX or TT genotype for the Single Nucleotide Polymorphism marker SNP_22 at nucleotide 102 of SEQ ID NO: 22 (or at nucleotide 102 in a variant thereof); i) the GX or GG genotype for the Single Nucleotide Polymorphism marker SNP_23 at nucleotide 102 of SEQ ID NO: 23 (or at nucleotide 102 in a variant thereof); j) the TX or TT genotype for the Single Nucleotide Polymorphism marker SNP_24 at nucleotide 102 of SEQ ID NO: 24 (or at nucleotide 102 in a variant thereof).

[0176] When referring to a SNP in a variant sequence, that variant sequence comprises at least 95%, 96%, 97%, 98% or 99% sequence identity with the mentioned sequence. For example, the wild donor used herein to introgress the QTL3.1 comprises variation regarding SNP_15, SNP_17 and SNP_20. The introgression fragment with QTL3. 1 on it comprises a sequence (SEQ ID NO: 35) which has 99% sequence identity to SEQ ID NO: 15, i.e. SNP_15 is a Adenine at nucleotide 102 in a sequence comprising at least 99% sequence identity to SEQ ID NO: 15, see also the Examples, especially Table 6; and it comprises a sequence (SEQ ID NO: 36) which has 98% sequence identity to SEQ ID NO: 17, i.e. SNP_17 is a Guanine at nucleotide 102 in a sequence comprising at least 98% sequence identity to SEQ ID NO: 17, see also the Examples, especially Table 6; and it comprises a sequence (SEQ ID NO: 37) which has 97% sequence identity to SEQ ID NO: 20, i.e. SNP_20 is a Cytosine at nucleotide 101 (i.e. position 101 is at the equivalent nucleotide to nucleotide position 102 in SEQ ID NO: 20) in a sequence comprising at least 97% sequence identity to SEQ ID NO: 20, see also the Examples, especially Table 6. In one aspect when referring to an Adenine for SNP_15, reference is made to an Adenine (at the equivalent position) in SEQ ID NO: 35, which is the sequence of the wild donor. In one aspect when referring to a Guanine for SNP_17, reference is made to a Guanine (at the equivalent position) in SEQ ID NO: 36, which is the sequence of the wild donor. In one aspect when referring to a Cytosine for SNP_20, reference is made to a Cytosine (at the equivalent position) in SEQ ID NO: 37, which is the sequence of the wild donor. For the other SNPs of QTL3. 1 (i.e. SNP_16, SNP_18, SNP_19, SNP_21-24) the donor nucleotide is present in the sequence provided herein, i.e. there is no variant sequence in the donor used. X refers to any nucleotide for the sequence on the other chromosome 3 of the pair of chromosomes. In one aspect X may be the nucleotide of the recurrent parent as described in Table 5.

[0177] In one aspect said at least 2, 3, 4, 5, 6, 7, 8, 9 or 10 markers are consecutive markers.

[0178] The fragment comprising the QTL3.1 may, thus, be large (comprising the donor nucleotide for SNP_15 to SNP_24), or may be smaller and lack markers having the genotype or haplotype of the wild cucumber (i.e. the markers have e.g. the cultivated cucumber genotype or haplotype instead, see also Table 5 (SNP haplotype of recurrent parent), but it may still confer a mint green fruit flesh color onto the cultivated cucumber plant (especially when in homozygous form), i.e. it can still comprise the QTL3.1 or a variant. Such smaller introgression fragments are an embodiment of the invention. Plants having smaller introgression fragments which still confer a mint green fruit flesh color onto the fruits (i.e. contain the fruit flesh color allele) can be generated using known techniques, such as fine-mapping or similar techniques. For example by starting with a plant comprising the introgression fragment as found in seeds deposited under accession number NCIMB 44295 and crossing such a plant with another cultivated cucumber plant and selfing the progeny of said cross, and / or backcrossing the progeny, to generate a population of plants which may contain recombinants having a smaller introgression fragment on chromosome 3, which fragments still confer a mint green fruit flesh color onto the fruits (especially when the fragment is in homozygous form) in relation to a plant lacking the introgression fragment (such as the genetic control), e.g. a fragment comprising donor SNP nucleotide for markers SNP_15 to SNP_20, or SNP_15 to SNP_19 or SNP_15 to SNP_18 or SNP_15 to SNP_17 or beginning of the chromosome to SNP_20 or SNP_19 to SNP_18, or SNP_17. Marker assays can be used to determine the size of the smaller introgression fragment. One or more of the SNP markers with the genotype or haplotype of the wild donor cucumber may be missing. The cultivated cucumber genotype or haplotype is then detected for these SNP markers. The fruit flesh color of plants comprising such a smaller introgression fragment can then be compared in an assay as described herein, i.e. growing a plurality of plants comprising the smaller introgression fragment in experiments together with suitable control plants, lacking the introgression fragments. If the fruit flesh color is changed compared to the control, then the smaller introgression fragment has retained the QTL3.1.

[0179] Alternatively, the same or variant QTL (QTL3. 1 or variant QTL3. 1) may be infrogressed from a different wild donor accessions, whereby optionally not all donor SNP markers disclosed herein may be present, i.e. the SNP haplotype of the donor accession may only be identical to the SNP haplotype of the QTL3. 1 present in seeds of NCIMB44295 for at least 2, 3, 4, 5, 6, 7, 8, 9, 10 or more SNPs. Such alternative wild cucumber sources can be identified using the SNP markers provided herein, by screening germplasm (i.e. accessions of) wild or primitive cucumber using a marker assay to detect the genotype or haplotype of one or more of markers SNP_ 15 to SNP_24, or of markers SNP_15 to SNP_20, or SNP_15 to SNP_19 or SNP_15 to SNP_18 or SNP_15 to SNP_17 or beginning of the chromosome to SNP_20 or SNP_19 to SNP_18 or SNP_17, or even only a smaller subgroup of these markers (e.g. 2, 3 or 4). Plants comprising the same or variant QTL3. 1 from these donors or from other sources are also an embodiment of the invention. Thus, as long as at least 2, 3, 4, 5, 6, 7, 8, 9, 10 or more (or all) of the SNPs of SNP_15 to SNP_24, or of markers SNP_15 to SNP_20, or SNP_15 to SNP_19 or SNP_15 to SNP_18 or SNP_15 to SNP_17 or beginning of the chromosome to SNP_20 or SNP_19 to SNP_18 or SNP_17, are present, the donor may contain QTL3. 1 (or a variant thereof) and is encompassed herein. The skilled person can then infrogress the QTL3. 1 (or a variant thereof) into cultivated cucumber in order to change the fruit flesh color as described herein and in order to confirm that the QTL changes fruit flesh color to mintgreen when present in cultivated cucumber, at least when it is in homozygous form.

[0180] As described above, in one embodiment the cultivated cucumber plant of the invention comprises an introgression fragment comprising at least a subset of SNP markers with the genotype (or haplotype) of the wild donor cucumber, i.e. at least 2, 3, 4, 5, 6, 7, 8, 9, 10 or more markers of SNP_15 to SNP_24, or at least 3 markers of SNP_15 to SNP_20, or of SNP_15 to SNP 19 or of SNP_15 to SNP_18 or of SNP_15 to SNP_17 or of SNP_16 to SNP_20 or of SNP_16 to SNP_19, or of SNP_16 to SNP_18 or of SNP_17 to SNP_20 or of SNP_17 to SNP_19. In one aspect the cultivated cucumber plant comprises all, or all except 1 or 2 markers of SNP_15 to SNP_24, or of SNP_15 to SNP_20, or of SNP_15 to SNP_19 or of SNP_15 to SNP_18 or of SNP_15 to SNP_17 or of SNP 16 to SNP_20 or of SNP_16 to SNP_19, or of SNP 16 to SNP_18 or of SNP_17 to SNP_20 or of SNP_17 to SNP_19.

[0181] Thus, the introgression fragment (and a cultivated cucumber plant or plant part, e.g., a cell, comprising the introgression fragment) can be detected in a marker assay by detecting the donor SNP genotype or haplotype of the introgression fragment (i.e. of the wild donor cucumber germplasm) of one or more or all of the markers above, preferably at least 2, 3, 4, 5, 6, 7, 8, 9 or 10.

[0182] Thus, in one aspect, a Quantitative Trait Locus (QTL3. 1) was found to be present on chromosome 3 of a wild cucumber donor which, when transferred (introgressed) into a cultivated cucumber variety or breeding line, and when present in heterozygous or homozygous form, confers a change of the fruit flesh color to mint-green (at least when QTL3. 1 is in homozygous form). A change in fruit flesh color is also seen when the QTL3. 1 is in heterozygous form, see Examples. In heterozygous form the fruit flesh color changes from e.g. light-yellow to at least yellow-green, see Examples. Thus, in one aspect it is encompassed herein to have the introgression fragment on one of the chromosomes 3 (one recombinant chromosome 3), while the homologous chromosome 3 of the pair may be a (non-recombinant) chromosome 3 of cultivated C. sativus var. sativus lacking the introgression fragment.

[0183] In a specific embodiment, the introgression fragment comprising the QTL3. 1 (or a variant thereof) is derivable from (or derived from) or obtainable from (or obtained from; or as present in) seeds, a representative sample of which has been deposited under accession number NCIMB 44295 or from progeny thereof. The progeny may be any progeny which retain the SNP markers or haplotype indicative of (and linked to) the QTL, e.g. as described in Table 5. Thus, progeny are not limited to selfing progeny or to Fl or F2 progeny of the deposit or accession, but can be any progeny, whether obtained by selfing and / or crossing with another cucumber plant.

[0184] In one embodiment the introgression fragment comprising QTL3. 1 (or a variant) is identifiable by one or more of the donor SNP markers linked to the QTL3.1 and / or indicative of QTL3.1 described elsewhere herein, especially markers SNP_15 to SNP_24 for the introgression fragment on chromosome 3, or a subset of markers, such as one or more of the markers selected from SNP markers SNP_15 to SNP_20, or of SNP_15 to SNP_19 or of SNP_15 to SNP_18 or of SNP_15 to SNP_17 or of SNP_16 to SNP_20 or of SNP_16 to SNP_19, or of SNP_16 to SNP_18 or of SNP_17 to SNP_20 or of SNP_17 to SNP_19. In one aspect the invention provides a cultivated cucumber plant, having a genome of cultivated (domesticated) cucumber which produces cucumber fruits with a mint-green fruit flesh color which is conferred by an introgression fragment on the cultivated cucumber chromosome 3, wherein said introgression fragment is e.g. obtained by (or obtainable by) crossing a cultivated plant grown from seeds deposited under NCIMB 44295 or progeny of this plant (which comprises one or more the markers disclosed herein linked to the QTL) with a cultivated cucumber plant. Thus in one aspect the cultivated cucumber plant of the invention comprises the same introgression fragment and the same recombinant chromosome 3 as present in NCIMB 44295 (comprising all of the wild donor haplotype for SNP markers SNP_15 to SNP_24 or comprising SEQ ID NO: 15 to 24 or comprising SEQ ID NO: 35, SEQ ID NO: 16, SEQ ID NO: 36, SEQ ID NO: 18 and 19, SEQ ID NO: 37 and SEQ ID NO: 21 to 24) or it comprises a shorter fragment of that introgression fragment, whereby the shorter fragment retains the genetic element conferring mint-green fruit flesh color (QTL3. 1).

[0185] Thus in one aspect a cultivated Cucumis sativus var. sativus plant is provided herein comprising an introgression fragment comprising QTL3.1 from a wild cucumber on chromosome 3 in homozygous or heterozygous form and wherein said introgression fragment is the introgression fragment “as in” / is “identical to” / is “the same as in” the seeds deposited under number NCIMB 44295, or is a shorter fragment thereof, but which still confers a mint green fruit flesh color due to the presence of QTL3. 1, at least when QTL3. 1 is in homozygous form.

[0186] In yet another embodiment a cultivated Cucumis sativus var. sativus plant is provided comprising an introgression fragment comprising QTL3.1 (or a variant) from a wild cucumber on chromosome 3 in homozygous or heterozygous form and wherein said introgression fragment is the introgression fragment or is a variant of the introgression fragment present in seeds deposited under number NCIMB 44295, i.e. it comprises the QTL 3.1 (or a variant), but the genomic sequence may be different in the variant. As wild accessions will be genetically divergent, the genomic sequence of an introgression fragment comprising QTL3.1 from other wild or primitive cucumbers will most likely not be identical to the genomic sequence as infrogressed into NCIMB 44295. For example, certain donor SNP markers linked to QTL3. 1 may be commonly found in various accessions, while other donor SNP markers may only be found in specific accessions. So, for example not all of SNP_15 to SNP_24 may be found in other wild cucumber donors. However, QTL3.1 (comprising e.g. a variant or ortholog of the fruit flesh color allele) may still be present in such wild accessions. The skilled person is capable of identifying and infrogressing the QTL3. 1 comprising region found in other wild cucumber donors into cultivated cucumber, e.g. detecting wild accessions comprising the donor SNP markers or a subset thereof and transferring these donor SNP markers (or subset) into a cultivated cucumber line or variety and assessing the fruit flesh color of the cultivated line or variety compared to the line or variety lacking the donor SNP markers (or subset), i.e. lacking the introgression fragment. Even in cases where the donor SNP haplotype for SNP_15 to SNP_24 is identical to the SNP haplotype of QTL3.1 found in seeds of NCIMB 44295, the actual nucleotide sequences flanking the SNP at nucleotide 102 of SEQ ID NO: 15 to 24 may be different in other donors. So other donors may comprise the same SNP nucleotide at nucleotide 102 or at an equivalent position, but in a sequence comprising at least 95%, 96%, 97%, 98% or 99% sequence identity to SEQ ID NO: 15 to 24 when e.g. aligned pairwise. This variation can be seen by sequencing the donors and aligning sequences of SEQ ID NO: 15 to SEQ ID NO: 24 with that sequence. For example, the donor used herein was found to comprise variation in the flanking sequences of 3 of the SNPs, namely for SNP 15, SNP 17 and SNP_20, see Table 6. In the donor SNP_15 is found in a sequence which comprises 99% identity to SEQ ID NO: 15, namely at nucleotide 102 in SEQ ID NO: 35. In one aspect there is, therefore, an Adenine for SNP_15 at nucleotide 102 of SEQ ID NO: 35. SNP_17 is found in a sequence which comprises 98% identity to SEQ ID NO: 17, namely at nucleotide 102 in SEQ ID NO: 36. In one aspect there is, therefore, an Guanine for SNP_17 at nucleotide 102 of SEQ ID NO: 36. SNP_20 is found in a sequence which comprises 97% identity to SEQ ID NO: 20, namely at nucleotide 101 in SEQ ID NO: 37. In one aspect there is, therefore, a Cytosine for SNP_20 at nucleotide 101 of SEQ ID NO: 37.

[0187] In one embodiment a cultivated cucumber plant or plant part is provided wherein the presence of the introgression fragment comprising QTL3.1, or the chromosome 3 region (or variant or orthologous chromosome 3 region), comprising QTL3.1, is detectable by a molecular marker assay which detects at least 2, 3, 4, 5, 6, 7, 8, 9 or 10 Single Nucleotide Polymorphism (SNP) markers selected from the group consisting of: a) an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_15 at nucleotide 102 of SEQ ID NO: 15 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 15, such as an Adenine for SNP_15 at nucleotide 102 of SEQ ID NO: 35; b) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_16 at nucleotide 102 of SEQ ID NO: 16 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 16; c) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_17 at nucleotide 102 of SEQ ID NO: 17 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 17, such as a Guanine for SNP 17 at nucleotide 102 of SEQ ID NO: 36; d) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_18 at nucleotide 102 of SEQ ID NO: 18 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 18; e) an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_19 at nucleotide 102 of SEQ ID NO: 19 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 19; f) a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_20 at nucleotide 102 of SEQ ID NO: 20 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 20, e.g. a Cytosine for SNP_20 at nucleotide 101 of SEQ ID NO: 37; g) a Cytosine (C) nucleotide for the Sing le Nucleotide Polymorphism marker SNP_21 at nucleotide 102 of SEQ ID NO: 21 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 21; h) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_22 at nucleotide 102 of SEQ ID NO: 22 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 22; i) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_23 at nucleotide 102 of SEQ ID NO: 23 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 23; j) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_24 at nucleotide 102 of SEQ ID NO: 24 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 24.

[0188] In one aspect said at least 2, 3, 4, 5, 6, 7, 8, 9 or 10 markers which are detected are consecutive markers.

[0189] In one aspect a method for screening genomic DNA of wild or cultivated cucumber plants or plant parts by carrying out the above molecular marker assay is encompassed herein. Optionally the method further comprises selecting a plant or plant part comprising the above nucleotide for at least 2, 3, 4, 5, 6, 7, 8, 9 or all 10 of the SNP markers SNP_15 to SNP_24.

[0190] Thus, in one embodiment the plant (or plant part) according to the invention comprises: at least an Adenine (A) (i.e. the AA or AX genotype) at nucleotide 102 of SEQ ID NO: 15 (referred to as SNP_15) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO: 15, such as an Adenine for SNP_15 at nucleotide 102 of SEQ ID NO: 35; (in other words there is an Adenine at the physical position of chromosome 3 shown in Table 5); and / or at least a Thymine (T) (i.e. the TT or TX genotype) at nucleotide 102 of SEQ ID NO: 16 (referred to as SNP_16) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO: 16 (in other words there is a Thymine at the physical position of chromosome 3 shown in Table 5); and / or at least a Guanine (G) (i.e. the GG or GX genotype) at nucleotide 102 of SEQ ID NO: 17 (referred to as SNP_17) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO: 17, such as a Guanine for SNP_17 at nucleotide 102 of SEQ ID NO: 36 (in other words there is a Guanine at the physical position of chromosome 3 shown in Table 5); and / or at least a Thymine (T) (i.e. the TT or TX genotype) at nucleotide 102 of SEQ ID NO: 18 (referred to as SNP_18) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO: 18 (in other words there is a Thymine at the physical position of chromosome 3 shown in Table 5); and / or at least an Adenine (A) (i.e. the AA or AX genotype) at nucleotide 102 of SEQ ID NO: 19 (referred to as SNP_19) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO: 19 (in other words there is an Adenine at the physical position of chromosome 3 shown in Table 5); and / or at least a Cytosine (C) (i.e. the CC or CX genotype) at nucleotide 102 of SEQ ID NO: 20 (referred to as SNP_20) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO:20, such as a Cytosine for SNP_20 at nucleotide 101 of SEQ ID NO: 37 (in other words there is a Cytosine at the physical position of chromosome 3 shown in Table 5); and / or at least a Cytosine (C) (i.e. the CC or CX genotype) at nucleotide 102 of SEQ ID NO: 21 (referred to as SNP_21) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO:21 (in other words there is a Cytosine at the physical position of chromosome 3 shown in Table 5); and / or at least a Thymine (T) (i.e. the TT or TX genotype) at nucleotide 102 of SEQ ID NO: 22 (referred to as SNP_22) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO:22 (in other words there is a Thymine at the physical position of chromosome 3 shown in Table 5); and / or at least a Guanine (G) (i.e. the GG or GX genotype) at nucleotide 102 of SEQ ID NO: 23 (referred to as SNP_23) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO:23 (in other words there is a Guanine at the physical position of chromosome 3 shown in Table 5); and / or at least a Thymine (T) (i.e. the TT or TX genotype) at nucleotide 102 of SEQ ID NO: 24 (referred to as SNP_24) or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO:24 (in other words there is a Thymine at the physical position of chromosome 3 shown in Table 5);

[0191] In a further one embodiment the presence of the introgression fragment, or the chromosome 3 region (or variant or orthologous chromosome 3 region), comprising QTL3.1, is detectable by a molecular marker assay which detects the donor SNP nucleotide for at least 2, 3, 4, 5, 6, 7, 8, 9 or 10 Single Nucleotide Polymorphism (SNP) markers of the sub-groups consisting of: SNP_15 to SNP_20, or SNP_15 to SNP_19 or SNP_15 to SNP_18 or SNP_16 to SNP_20 or SNP_16 to SNP_19 or SNP 16 to SNP_18, or SNP_17 to SNP_20 or SNP_17 to SNP_19 or SNP_17 to SNP_18.

[0192] The SNP genotype refers to two nucleotides, and genomic sequences comprising one of these two nucleotides, one on each chromosome 3. So a plant having a AA genotype for SNP_15 has an identical nucleotide (A) on both chromosomes (i.e. is homozygous), while a plant having an AX genotype for SNP_15 has one chromosome with an A at nucleotide 102 of SEQ ID NO: 15 (or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO: 15, such as an Adenine for SNP_15 at nucleotide 102 of SEQ ID NO: 35) and one chromosome with a X at nucleotide 102 of SEQ ID NO: 15 (or at the equivalent nucleotide of a genomic sequence comprising substantial sequence identity to SEQ ID NO: 15, such as an Adenine for SNP_15 at nucleotide 102 of SEQ ID NO: 35) and is heterozygous, whereby X may be any nucleotide. As the genomic sequences around the SNP markers provided herein may vary slightly in introgression fragments from other wild cucumber donors (i.e. variants or orthologous chromosome 3 regions) it is clear that the nucleotide sequences before and after the SNP may not be 100% identical to the sequences (or complement sequences) provided herein. Therefore, sequences having substantial sequence identity (i.e. at least 95% identity) to the sequences provided herein, but which comprise the same donor SNP nucleotide, are encompassed herein, such as for example an Adenine for SNP_15 at nucleotide 102 of SEQ ID NO: 35, a Guanine for SNP_17 at nucleotide 102 of SEQ ID NO: 36 and a Cytosine for SNP_20 at nucleotide 101 of SEQ ID NO: 37. The ‘complement’ sequence or ‘complementary’ sequence is the other strand of the double stranded DNA molecule. It may also be referred to as ‘reverse complement strand’ when taking the 5 prime (5’) to 3 prime (3’) orientation of the strands into consideration.

[0193] In one aspect, the introgression fragment comprising QTL3.1, or the chromosome 3 region (or variant or orthologous chromosome 3 region) comprising the QTL (QTL3. 1 or variant), which is detectable by the above one or more markers is from a wild or primitive cucumber. In one aspect it is the same introgression fragment as found on chromosome 3 in seeds deposited under accession number NCIMB 44295, or a smaller fragment retaining the QTL. SNP markers SNP_15 to SNP_24 span a region of about 1.46 Mb. In one aspect the introgression fragment on chromosome 3 is equal to or less than 1.5 Mb in size, preferably equal to or less than 1.46 Mb in size, more preferably equal to or less than 1.3 Mb, 1.2 Mb or 1. 1 Mb in size, e.g. equal to or less than 1 Mb. In one aspect the introgression fragment is at least 0.2 Mb, 0.5 Mb, 1.0 Mb, 1.45 Mb, 1.46 Mb, 1.5 Mb or 1.6 Mb in size. Thus, various ranges of introgression sizes are encompassed herein, such as fragments less than 1.46 Mb but more than 0.2 Mb, which retain the QTL3. 1 and one or more of the SNP markers of the donor markers for SNP_15 to SNP_24, or of the subgroups of SNP_15 to SNP_20, or SNP_15 to SNP_19, or SNP_15 to SNP_18, or SNP_16 to SNP_20, or SNP_16 to SNP_19, or SNP_16 to SNP_18, or SNP_17 to SNP_20, or SNP_17 to SNP_19, or SNP_17 to SNP_18, or from the beginning of the chromosome (i.e. nucleotide 1) to SNP_16 or to SNP_17 or to SNP_18 or to SNP_19 or to SNP_20, SNP_21, SNP_22, SNP_23 or SNP_24. As mentioned before, the location of the QTL3. 1 in the region spanning SNP_15 to SNP_24 can be determined by fine-mapping and recombinants comprising QTL3. 1 on a smaller introgression fragment can be generated. The size of an introgression fragment can be easily determined by e.g. whole genome sequencing or Next Generation Sequencing, e.g. as described in Qi et al. 2013 {supra) or in Huang et al. 2009 {supra). Especially introgression regions can be easily distinguished from cultivated genomic regions due to the larger amount of genetic variation (SNPs, INDELs, etc.) in the introgression region.

[0194] To obtain the introgression fragment present on chromosome 3 (comprising QTL3. 1) from the deposited seeds (NCIMB 44295), i.e. to transfer the introgression fragments comprising the QTL to another cultivated cucumber plant, a plant is grown from the seed and the plant is crossed with a cultivated cucumber plant to obtain Fl seeds. As NCIMB 44295 contains two recombinant chromosomes 3 (comprising the introgression fragment) all of the Fl seed and plants grown therefrom, contain one recombinant chromosome 3 from the NCIMB 44295 parent and one non-recombinant chromosome 3 from the other cultivated parent. Thus, by traditional breeding one can transfer the recombinant chromosome 3 from NCIMB44295 into other cultivated cucumber lines or varieties. Plants which comprise the QTL3.1 can be screened for, and selected for, by the presence of one or more of the above donor SNP markers in order to identify plants comprising a recombinant chromosome 3.

[0195] To generate shorter introgression fragments (comprising QTL3.1) meiosis needs to take place and plants comprising the recombinant chromosomes 3, and especially new meiotic recombination events within the introgression fragment, need to be identified. For example, seeds of NCIMB44295 can be selfed one or more times to produce F 1, F2 or F3 plants (or further selfing generations), and / or F 1, F2 or F3 plants (etc.) comprising a recombinant chromosome 3 can be backcrossed to a cultivated parent. Plants which comprise the recombinant chromosome 3 can be screened for, and selected for, by the presence of one or more of the above SNP markers in order to identify plants comprising a smaller introgression fragment. Such new recombinants can then be tested for the presence of the QTL3. 1 on the smaller introgression fragment by determining the fruit flesh color compared to the (genetic) control lacking the introgression fragment.

[0196] Similarly, cultivated cucumber plants comprising QTL3.1 (or a variant thereof) can be generated and / or identified using different methods. For example, to obtain a cultivated cucumber plant comprising an introgression fragment from a wild donor, a wild donor is identified which comprises one or more of the donor SNP markers linked to QTL3. 1 disclosed herein, e.g. any one, or more, or all of the markers described herein above. This has for example been done for various wild accessions. The identified plant is crossed with a cultivated cucumber plant to obtain Fl seeds. The Fl can be selfed to produce F2, F3, etc. plants, and / or F2 plants or F3 plants, etc., can be backcrossed to the cultivated cucumber parent. Plants which are comprising QTL3. 1 (or a variant thereof) can be screened for, and / or selected for, by the presence of one or more of the above donor SNP markers and / or screened for, and / or selected for, a mint-green fruit flesh color or at least a different, greener fruit flesh color compared to the initial cultivated parent (lacking the introgressions). Alternatively, or in addition, QTL mapping can be carried out in order to identify further molecular markers linked to the QTL3.1 (or a variant thereof) and / or to generate cultivated cucumber plants comprising an introgression fragment on chromosome 3 which confers mint-green fruit flesh color.

[0197] In one embodiment a method is provided for detecting the presence of the introgression fragment comprising QTL3.1 or a variant thereof in a (cultivated or wild) cucumber plant or plant part (cell, fruit, fruit part, leaf, stem, etc.), or the chromosome 3 region comprising QTL3.1 or a variant thereof, said method comprises analyzing the genomic DNA of the plant or plant part using a molecular marker assay which detects at least 2, 3, 4, 5, 6, 7, 8, 9 or all 10 of the markers selected from the group consisting of: a) an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_15 at nucleotide 102 of SEQ ID NO: 15 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 15, such as an Adenine for SNP_15 at nucleotide 102 of SEQ ID NO: 35; b) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_16 at nucleotide 102 of SEQ ID NO: 16 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 16; c) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_17 at nucleotide 102 of SEQ ID NO: 17 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 17, such as a Guanine for SNP 17 at nucleotide 102 of SEQ ID NO: 36; d) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_18 at nucleotide 102 of SEQ ID NO: 18 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 18; e) an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_19 at nucleotide 102 of SEQ ID NO: 19 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 19; f) a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_20 at nucleotide 102 of SEQ ID NO: 20 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 20, such as a Cytosine for SNP_20 at nucleotide 101 of SEQ ID NO: 37; g) a Cytosine (C) nucleotide for the Sing le Nucleotide Polymorphism marker SNP_21 at nucleotide 102 of SEQ ID NO: 21 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 21; h) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_22 at nucleotide 102 of SEQ ID NO: 22 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 22; i) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_23 at nucleotide 102 of SEQ ID NO: 23 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 23; j) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_24 at nucleotide 102 of SEQ ID NO: 24 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 24.

[0198] The method optionally further comprises selecting the plant or plant part comprising at least 2, 3, 4, 5, 6, 7, 8, 9 or all 10 of the marker nucleotides of a) to j). The SNP haplotype indicative of the QTL, i.e. the donor SNP haplotype for one or more of SNP_15 to SNP_24 as described elsewhere herein and in e.g. Table 5 or 6 may be detected in the plant or plant part. The method may involve screening a population of plants or plant parts using the above SNP markers. Obviously, a SNP assay can be designed for the above SNP markers using the plus strand or the minus strand of the DNA. The method may further comprise generating plants that are homozygous for the selected donor nucleotides of the SNP markers and / or allowing the plants to develop fruits and optionally analyzing the fruit flesh color. Further aspects of the above methods are described elsewhere herein. The introgression fragment in the plants provided herein is in one aspect a fragment of chromosome 3 (comprising QTL3. 1) which is present in seeds deposited under accession number NCIMB44295 or a smaller version of that fragment retaining the QTL (generated by e.g. recombination within the introgression fragment).

[0199] QTL3. 1 and the introgression fragment comprising QTL3. 1 is in one aspect in homozygous form in the plant or plant part, such as the fruits. The SNP marker genotype for one or more of SNP_15 to SNP_24 in this aspect comprises the donor SNP nucleotide in homozygous form.

[0200] Also provided are seeds from which a plant of the invention can be grown, as are cucumber fruits harvested from a plant of the invention and comprising the recombinant chromosome 3 in their genome (comprising QTL3. 1 or a variant). Likewise, a plant cell, tissue or plant part of a plant or of a seed is provided comprising at least one recombinant chromosome 3 (comprising QTL3.1 or a variant), wherein said recombinant chromosome 3 comprises an introgression fragment from a wild or primitive cucumber and wherein said introgression fragment comprises QTL3. 1 or a variant thereof.

[0201] The molecular markers described herein may be detected according to standard method. For example, SNP markers can easily be detected using a KASP-assay (see www.kpbioscience.co.uk) or other SNP genotyping assays. For developing a KASP-assay, for example 50 or 70 base pairs upstream and 50 or 70 base pairs downstream of the SNP can be selected and two allele-specific forward primers and one allele specific reverse primer can be designed. See e.g. Allen et al. 2011, Plant Biotechnology J. 9, 1086-1099, especially p097-1098 for KASP assay method.

[0202] Thus, in one aspect, the SNP markers and the presence / absence of the marker associated with QTL3.1 is determined using a KASP assay, but equally other SNP genotyping assays can be used. For example, a TaqMan SNP genotyping assay, a High Resolution Melting (HRM) assay, SNP- genotyping arrays (e.g. Fluidigm, Illumina, etc.) or DNA sequencing may equally be used.

[0203] The physical size of an introgression fragment can be determined by various methods, such as physical mapping, sequencing or by visualization of the introgression using Fluorescent in situ hybridization (FISH) images (Verlaan et al. 2011, Plant Journal 68: 1093-1103).

[0204] Cultivated cucumber plants with smaller introgression fragments on chromosome 3 (comprising QTL3. 1 or a variant) can be generated by generating new recombinant plants from a population of plants derived from a cross between a cultivated cucumber plant (lacking the introgressions) and a plant of the invention and selecting recombinant progeny having smaller introgression sizes. Such plants are, thus, in one aspect derived from (progeny or descendants of) the recombinant chromosome 3 present in plants of which seeds have been deposited under NCIMB44295. Such progeny or descendants which retain the QTL3. 1, and whereof the fruits produce a mint-green fruit flesh color (at least when QTL3. 1 is in homozygous form) compared to the fruits of plants lacking an introgression as described herein, are encompassed herein.

[0205] As mentioned, plants and plant parts may either comprise individual QTLs in homozygous or heterozygous form, i.e. QTL1.1 or QTL3.1 or both QTLs in homozygous and / or heterozygous form. In one aspect both QTL1. 1 and QTL3. 1 are in homozygous form and the fruits are e.g. sweet with a mint-green fruit flesh.

[0206] In one aspect QTL1.1 and / or QTL3.1, especially with either one or both in homozygous form, are combined with light-green fruit skin (as described e.g. in Geng et al. 2022, supra) or with Light Green Peel or with Yellow Green Peel as described in Cui et al. (2023, supra). The QTL CsFS 1 of Geng et al. is found on chromosome 3 and the candidate gene for light green fruit skin color is Csa3G912920. This gene is expressed at a higher level in fruits with light green skin color. It is located at nucleotides 40678424 to 40673049 of the Chinese Long genome V3, i.e. at the other end of chromosome 3 than QTL3. 1. In one aspect, therefore, plants comprise fruits with interior mint green fruit flesh due to QTL3.1 (preferably in homozygous form) and with exterior light green fruit skin due to QTL CsFSl. In another aspect QTL CsFSl is not present in any of the plants or plant parts described herein.

[0207] In another aspect QTL1. 1 (in homozygous or preferably in heterozygous from) is combined with QTL3.2 (the mutant allele of the White locus, i.e. genotype w / w) and preferably further with QTL4. 1 for enhanced peel color, as described further below. It was found that QTL1. 1 has a negative effect on fruit peel color when the w / w genotype is present in the plant and this negative effect can be compensated again by further adding QTL4. 1. The in-gene marker for the mutant w-allele is provided herein in SEQ ID NO: 40 (plus strand) and 41 (minus strand) and SEQ ID NO: 40 is in the region starting at nucleotide 40340728 and ending at nucleotide 40340871 of chromosome 3. See also Examples, e.g. Table 9.

[0208] In one aspect QTL1. 1 is present in heterozygous form in a cultivated cucumber plant, QTL3.2 is present in the genome in homozygous form (w / w genotype), QTL4.1 is present in the genome in preferably homozygous form and further optionally QTL3.1 is present in the genome in preferably homozygous form in a cultivated cucumber plant, plant part, fruit or seed. The fruits produced by such a plant have an increased glucose and fructose content due to QTL1. 1, they have a mint-green fruit flesh due to QTL3. 1 and they have a mint-green peel color due to the combination of the w / w-genotype and QTL4. 1. Such fruits are e.g. shown in Figure 1 and 2.

[0209] A cultivated cucumber plant, plant part, seed or fruit may, thus, comprise in its genome 1, 2, 3 or 4 QTLs selected from QTL1. 1 (or a variant) and / or QTL3. 1 (or a variant) and / or QTL3.2 (or a variant) and / or QTL4. 1 (or a variant), see e.g. Figure 6. Various combinations are encompassed herein, such as but not limited to: QTL1. 1 in heterozygous or homozygous form;

[0210] QTL1. 1 in heterozygous or homozygous form and QTL3. 1 in preferably homozygous form;

[0211] QTL1.1 in heterozygous or homozygous form and QTL3.2 (w / w) in homozygous form and QTL4.1 in preferably homozygous form;

[0212] QTL1. 1 in heterozygous form and QTL3.2 (w / w) in homozygous form and QTL4. 1 in homozygous form;

[0213] QTL3.2 (w / w) in homozygous form and QTL4. 1 in preferably homozygous form.

[0214] QTL3.1 in preferably homozygous form and QTL3.2 (w / w) in homozygous form and QTL4.1 in preferably homozygous form.

[0215] QTL1. 1 in heterozygous or homozygous form and QTL3. 1 in preferably homozygous form and QTL3.2 (w / w) in homozygous form and QTL4. 1 in preferably homozygous form.

[0216] QTL1. 1 in heterozygous form and QTL3. 1 in homozygous form and QTL3.2 (w / w) in homozygous form and QTL4. 1 in homozygous form.

[0217] The cultivated cucumber plant, plant part or seed provided herein may be an inbred line, an OP (open pollinated variety) or an Fl hybrid. The cultivated cucumber plant may be of any type. Preferably it has good agronomic and good fruit quality characteristics. The cultivated cucumber plant is in one aspect uniform, both genetically and phenotypically. Especially fruit characteristics are uniform, e.g. regarding shape, skin color, skin thickness, skin ribs, skin toughness, spines (spine color, spine density, etc.), presence / absence of warts, length and diameter at edible and marketable maturity, flavour, etc. Likewise seed characteristics (i.e. characteristics of the seeds from which the plant is grown) are uniform, e.g. seed size, seed color, etc. Thus, plants of the line or variety comprising the one or more QTLs, preferably in homozygous form, produce uniform fruits, meaning that there is little variation between fruits of plants grown under the same environmental conditions and when fruits are at the same developmental stage (e.g. for qualitative characteristics at least 98%, 99% or preferably 100% of all plants or plant parts, fruits or seed are identical for the characteristics; for quantitative characteristics at least 90%, 95%, 98% of all plants or plant parts, fruits or seed are identical for the characteristics).

[0218] The cultivated cucumber plant comprising the one or more QTLs (or variants thereof) according to the invention may be of any type, e.g. it may be of one of the following cucumber types: pickling cucumbers (e.g. American pickling, European pickling type), slicing cucumbers (e.g. American slicing), long cucumbers, short cucumbers, European greenhouse cucumbers, Beit-Alpha type cucumbers, oriental trellis type cucumbers, Asian cucumbers (e.g. selected from Indian Mottled cucumber, Chinese Long cucumber, Korean cucumber and Japanese cucumber type). In one aspect the cultivated cucumber according to the invention is an inbred line or a F 1 hybrid of a pickling cucumber type, slicing cucumber type, long cucumber type, short cucumber type, European greenhouse cucumbers, Beit-Alpha type cucumbers, oriental trellis type cucumbers, Chinese long cucumber type, Korean cucumber type or Japanese cucumber type. In a specific embodiment the cucumber is an inbred line or an Fl hybrid of a European greenhouse cucumber or a slicer type cucumber or a short cucumber.

[0219] The plant may be a single cross F 1 hybrid or an inbred line, comprising the one or more QTLs in homozygous and / or heterozygous form. In one aspect it is an Fl hybrid produced by crossing an (inbred) parent plant comprising one or more of the QTLs (or variant) in homozygous form with an (inbred) parent plant lacking the QTL / s (i.e. lacking introgression fragments comprising the QTLs). Thus, in one aspect the Fl hybrid is heterozygous for e.g. QTL1.1. In one aspect the Fl hybrid is homozygous for QTL1.1. In another aspect the Fl hybrid is heterozygous for QTL1. 1 and is homozygous for QTL3. 1, for QTL3.2 (w / w) and for QTL4. 1.

[0220] In another aspect it is an Fl hybrid produced by crossing an (inbred) parent plant comprising one or more of the QTLs (or variants thereof) in homozygous form with an (inbred) parent plant that also comprises one or more of the QTLs (or variants thereof) in homozygous form. Thus, in one aspect the F 1 hybrid is homozygous for e.g. QTL1.1 and / or QTL3.1.

[0221] See the various preferred combinations of QTLs further above. Any of these combinations may be in e.g. an F 1 hybrid seed or plant or fruit.

[0222] In one aspect the F 1 hybrid is a European greenhouse cucumber type or a short cucumber (like variety Beesan Fl, Nunhems), suitable for the traditional glasshouse cultivation or for high- wire cultivation. In the traditional glasshouse cultivation method the main stem of the plant is led up to a horizontal iron wire that is suspended at a height of about two meters above the ground. When the plant reaches this height and attaches to the wire, it is “topped” by removing its growth point in order to terminate further proliferation, whereupon lateral shoots start to develop. These lateral shoots are allowed to grow downward to a height of about 1 meter above the ground, and the growth points are then removed from them. This is followed by flowering and the development of the fruits both on the stem and on the lateral shoots or tendrils, but the fruits on the tendrils develop later than those on the stem. The fruits are harvested about 6 weeks after sowing. In the high-wire cultivation no lateral tendrils are allowed to grow and all the harvest comes from the stem. Specific varieties have been developed by Nunhems which are highly suitable for high-wire cultivation, as they provide a gene called “compact”, see. W02009 / 059777, for example varieties High-Jack, Hi-Power, Hi-Lisa. Thus, in one aspect of the invention the cultivated cucumber plant comprises additionally the compact gene described in W02009 / 059777. The compact gene is preferably present in heterozygous form.

[0223] In one aspect the cucumber is the plant of which seeds were deposited under accession number NCIMB 44295, or progeny thereof, whereby the progeny retain one or more of QTL1. 1 and / or QTL3. 1 and / or QTL4. 1 and / or QTL3.2 (as detectable by the presence of one or more markers as described elsewhere).

[0224] In another aspect the plant according to the invention is not a wild cucumber plant or a wild relative of cucumber or a landrace.

[0225] In yet another aspect the plant according to the invention is a cultivated cucumber of the Eurasian cucumber group, the East Asian cucumber group or the Xishuangbanna cucumber group. In another aspect the plant according to the invention is not a cucumber of the Indian cucumber group.

[0226] In one embodiment the cultivated cucumber plant comprising one or more of QTL1.1 (or a variant) and / or QTL3.1 (or a variant) and / or QTL3.2 (or a variant) and / or QTL4. 1 (or a variant), see the various preferred combinations of QTLs further above, produces seedless fruits without pollination, i.e. is parthenocarpic. Most European greenhouse cucumbers are parthenocarpic, i.e. the female flowers produce fruits without pollination, whereby the fruits remain seedless. Parthenocarpy is genetically controlled and it is known to breeders how to introduce the parthenocarpy trait into a cucumber line or variety (see e.g. Chapter 13 entitled “Cucumber” by T. tatlioglu, page 207-209 in the book Genetic Improvement of Vegetable Crops, Editors G. Kalloo and BO Bergh, Pergamon Press, 2012, ISBN0080408265).

[0227] In a further embodiment of the invention the cultivated cucumber plant comprising one or more of QTL1. 1 (or a variant) and / or QTL3.1 (or a variant) and / or QTL3.2 (or a variant) and / or QTL4.1 (or a variant), see the various preferred combinations of QTLs further above, is primarily gynoecious or entirely gynoecious (producing 100% female flowers). This means that mostly or only female flowers are produced. This trait is also genetically controlled and it is known to breeders how to introduce the gynoecious trait into a cucumber line or variety (see e.g. Chapter 13 entitled “Cucumber” by T. tatlioglu, page 207-209 in the book Genetic Improvement of Vegetable Crops, Editors G. Kalloo and BO Bergh, Pergamon Press, 2012, ISBN0080408265).

[0228] In one aspect the cucumber plant of the invention is both parthenocarpic and gynoecious. Thus, the plant produces primarily or only female flowers, which produce seedless fruits without pollination. In gynoecious cucumbers male flowers can be induced by treatment with silver nitrate. This method is used to produce pollen and to self-pollinate an inbred gynoecious cucumber line. In a different aspect the cucumber plant of the invention is monoecious (produces both male and female flowers), optionally parthenocarpic and monoecious.

[0229] In a further embodiment of the invention the cultivated cucumber plant comprising one or more of QTL1. 1 (or a variant) and / or QTL3.1 (or a variant) and / or QTL3.2 (or a variant) and / or QTL4.1 (or a variant), see the various preferred combinations of QTLs further above, is uniform and genetically stable regarding the morphological characteristics of the fruits produced by said plant, e.g. regarding fruit shape, fruit color, skin thickness, warts, etc.

[0230] Fruit characteristics, such as average fruit length, average fruit diameter, skin thickness, presence / absence of warts, spininess, skin toughness, fruit neck shape, fruit tapering, shape of medial cross section, presence or absence of seeds (parthenocarpy), etc. depend on the cucumber type, i.e. the cultivated genetic background (gene pool) into which the QTL(s) is / are introgressed. Thus, depending on the cucumber type, various fruit shapes, sizes and fruit types are included herein. In one aspect the fruits are seedless.

[0231] The two main types of cucumber fruit grown commercially today in the United States are fresh market (slicing) type and the processing (pickling) type. Varieties and production methods are typically adapted to the end use. Slicing cucumbers are often longer, larger and have darker and thicker skin, whereas pickling / processing cucumbers have a shorter fruit, thinner skin with interior flesh that make them more amenable to pickling. Seedless varieties are generally preferable for both fresh market and for pickling as developing and large seeds are not palatable.

[0232] In one aspect the plant of the invention is a pickling type (processing type) and produces fruits which at edible maturity and / or marketable size have an average fruit length of at least 10 cm, or at least 11 cm, or at least 12 cm, or at least 13 cm and / or a fruit length to diameter ratio of at least 2, at least 2.5, at least 3, or more.

[0233] In a different aspect the plant of the invention is a fresh market type, e.g. a long cucumber type or slicing type, and produces fruits have an average fruit length at edible maturity and / or marketable size which is longer than the pickling type, e.g. at least 15 cm, 16 cm, 17 cm, 18 cm, 19 cm, 20 cm, 25 cm, 26 cm, 'll cm, 28 cm, 29 cm, 30 cm, 32 cm, 40 cm, or more.

[0234] In one aspect the plant of the invention is a long cucumber type producing fruits of marketable size, especially seedless fruits.

[0235] Provided herein are also fruits comprising QTL1.1 and / or QTL3.1, or fruits comprising the various preferred combinations of QTLs further above, as are parts thereof, and food or feed products containing these. In one embodiment the donor SNP markers linked to one or more of QTL1. 1 and / or QTL3. 1 and / or QTL3.2 and / or QTL4. 1 (see the various preferred combinations of QTLs further above) are detectable in the fruits, fruit parts or food or feed products comprising these.

[0236] In one aspect the plant is an indeterminate cucumber. In another aspect the cucumber is determinate.

[0237] Also seeds from which a plant according to the invention can be grown is provided herein, as are cucumber fruits harvested from a plant according to the invention. These comprise the one or more of QTL1.1 and / or QTL3. 1 and / or QTL3.2 and / or QTL4. 1 (see the various preferred combinations of QTLs further above), in their genome and can, therefore, be distinguished from other fruits by the presence of one or more of the donor SNP markers provided herein.

[0238] In one aspect the fruits are bitter free (selected from the groups bitter and bitterfree) at edible maturity and / or at marketable size of the fruits. As already mentioned, the cultivated cucumber plants provided herein are ‘domesticated cucumber’ plants or ‘cultivated cucumber’ plants (and not ‘wild’ plants) and the fruits produced are bitter-free, a trait conferred by the Bt gene on chromosome 5 (Qi et al., Nature Genetics December 2013, Vol 45, No. 12, pages 1510 - 1518).

[0239] In a further aspect the fruit has a thin skin (selected from the groups thick and thin) at edible maturity and / or at marketable size of the fruits.

[0240] In a different embodiment the one or more of QTL1. 1 and / or QTL3. 1 and / or QTL3.2 and / or QTL4. 1 (see the various preferred combinations of QTLs further above) are introgressed into a cucumber type called ‘Compact’, as described in US8710303B2. Thus, the cucumber plants according to the invention comprise the compact gene as described in US8710303B2 in homozygous or heterozygous form, e.g. as present in varieties Hi-Jack and Hi-Lisa (both Nunhems varieties).

[0241] A further embodiment of the invention is a plant cell, tissue or plant part of a plant or of a seed according to the invention comprising at least one recombinant chromosome 1 and / or 3 and / or 4, wherein said recombinant chromosome 1 and / or 3 and / or 4 comprises an introgression fragment from a wild cucumber and wherein said introgression fragment on chromosome 1 comprises a QTL (QTL1. 1 or a variant) that confers higher glucose and fructose levels of the fruits, and wherein the QTL on chromosome 3 (QTL3. 1 or a variant) confers a mintgreen fruit flesh color, wherein said introgression fragment on chromosome 4 comprises a QTL (QTL4. 1 or a variant) conferring an enhanced mint-green skin / peel color in a genome comprising the w / w genotype (QTL3.2).

[0242] Also, the use of a recombinant chromosome 1 and / or 3 and / or 4 comprising an introgression fragment from a wild donor cucumber for breeding cucumber varieties having at least increased fruit glucose and fructose levels and / or comprising a mint-green fruit flesh color and / or comprising mint-green peel color is encompassed herein. In one aspect said recombinant chromosomes 1 and / or 3 and / or 4 is the recombinant chromosome 1 and / or 3 and / or 4 as found in seeds deposited under accession number NCIMB 44295 or is derived from said recombinant chromosome 1 and / or 3 and / or 4 (e.g. is a smaller fragment of the introgression fragment found in said seeds).

[0243] Likewise, the use of a chromosome 1 and / or 3 and / or 4, or the use of QTL1. 1 and / or QTL3. 1 and / or QTL4. 1, as found in seeds deposited under accession number NCIMB 44295, or in progeny thereof, or as found in a wild donor which comprises the same SNP haplotype or SNP genotype for at least 2, 3, 4, 5, 6, 7, 8, 9, 10 or more of the donor SNP markers linked to each of the QTLs, for generating a cultivated cucumber plant comprising an introgression fragment on chromosome 1 and / or 3 and / or 4 is encompassed herein.

[0244] Similarly, the use of plants grown from seeds deposited under accession number NCIMB 44295 or progeny thereof, for generating a cultivated cucumber plant comprising a higher amount of fruit glucose and fructose levels and / or a mint green fruit flesh color and / or mint green peel color is encompassed herein, wherein said traits are conferred by an introgression fragment obtained from chromosome 1 and / or 3 and / or 4 of said plants or progeny thereof.

[0245] Also provided is the use of plants grown from seeds deposited under accession number NCIMB44295 or progeny thereof, or from another wild donor which comprises the same donor SNP haplotype or donor SNP genotype for at least 2, 3, 4, 5, 6, 7, 8, 9, 10 or more of the SNP markers linked to each of the QTLs, for transferring one or more of QTLs QTL1.1 and / or QTL3.1 and / or QTL4. 1 or the introgression fragment or a sub-fragment thereof comprising said QTL to another cucumber plant is provided.

[0246] QTL3.2 (w / w) is also present in the deposited seeds, but can also be derived from other sources or generated de novo.

[0247] In a further aspect methods for identifying or detecting and / or selecting wild or cultivated cucumbers or cucumber parts comprising QTL 1.1 or QTL3.1 or QTL4.1 using one or more of the donor SNP markers disclosed herein are provided. It is noted that embodiments described above or elsewhere herein for products (plants and plant parts) in which the QTL / s or introgression fragments is / are described to be present or ‘detectable’ are understood to apply equally to any of the methods described herein.

[0248] A method is provided for identifying (or detecting and / or selecting) a cucumber plant or plant part comprising an introgression fragments on chromosome 1 from a wild cucumber donor, wherein said introgression fragments comprises a Quantitative Trait Locus (QTL) conferring an increase in glucose and fructose of the cucumber fruit, wherein QTL 1.1 is located on chromosome 1 between the Single Nucleotide Polymorphism marker SNP_01 at nucleotide 102 of SEQ ID NO: 1 and SNP_14 at nucleotide 102 of SEQ ID NO: 14, comprising: a) providing a cucumber plant or plant part or DNA of such plant or plant part, b) screening said plant, plant part or DNA for the presence of the SNP haplotype or genotype of at least 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13 or 14 markers selected from the group consisting of: a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_01 at nucleotide 102 of SEQ ID NO: 1 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 1; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_02 at nucleotide 102 of SEQ ID NO: 2 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 2; a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_03 at nucleotide 102 of SEQ ID NO: 3 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 3; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_04 at nucleotide 102 of SEQ ID NO: 4 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 4; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_05 at nucleotide 102 of SEQ ID NO: 5 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 5; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_06 at nucleotide 102 of SEQ ID NO: 6 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 6; a Thymine (T) nucleotide for the Sing le Nucleotide Polymorphism marker SNP_07 at nucleotide 102 of SEQ ID NO: 7 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 7; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_08 at nucleotide 102 of SEQ ID NO: 8 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 8; a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_09 at nucleotide 102 of SEQ ID NO: 9 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 9; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_10 at nucleotide 102 of SEQ ID NO: 10 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 10; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_11 at nucleotide 102 of SEQ ID NO: 11 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 11, such as a Guanine for SNP l 1 at nucleotide 102 of SEQ ID NO: 34; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_12 at nucleotide 102 of SEQ ID NO: 12 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 12; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_13 at nucleotide 102 of SEQ ID NO: 13 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 13; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_14 at nucleotide 102 of SEQ ID NO: 14 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 14 and c) identifying and / or selecting a plant or plant part comprising the SNP haplotype for at least 1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13 or all 14 of the SNP markers of step b).

[0249] In one aspect the plants or plant parts comprising at least 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13 or 14 markers of SNP_1 to SNP_14 comprise at least the donor nucleotide for the peak marker, which is SNP_11. In one aspect they comprise at least the donor nucleotide for SNP_9, SNP_10, SNP ll, SNP_12 and SNP_13. In another aspect they comprise at least the donor nucleotide for SNP_08, SNP_9, SNP_10, SNP_11, SNP_12, SNP_13 and SNP_14. Therefore, in one aspect in step b) at least these SNP markers are screened and / or in step c) plants or plant parts comprising at least these SNP markers are identified and / or selected.

[0250] In step c) plants or plant parts comprising the introgression fragment in homozygous form or in heterozygous form may be identified and / or selected. It is understood that in step b) the genotype for the markers may be assayed, i.e. the SNP nucleotide on both chromosomes of the pair of chromosome 1, as e.g. disclosed in Table 2 or 3 and elsewhere herein.

[0251] The method may further comprise optionally selfing the selected plant in order to generate a plant homozygous for the introgression fragment.

[0252] The method may also comprise growing the selected plant (or the optionally selfed plants) and allowing fruits to develop and analyzing the fruits for at least the glucose and fructose content, optionally also for malic acid content. This allows phenotypic determination if QTL1. 1 is present in the plant.

[0253] The SNP haplotype or SNP genotype of SNP_1 to SNP_14 are e.g. disclosed in Table 2 or 3 and elsewhere herein. In the above method, thus, also the SNP genotype may be screened and / or selected for. In step c) plants or plant parts comprising the introgression fragment in homozygous form or in heterozygous form may be identified and / or selected.

[0254] The cucumber plant may (in any of the methods herein) be a cultivated cucumber plant or plant part may be any plant or plant part, such as commercial varieties, mapping populations, etc. The cucumber plant may also or alternatively be a wild cucumber plant or plant part, in which case not an ‘introgression fragment’ is detected, but the endogenous chromosome 1 region is analyzed for the presence of the SNP markers linked to QTL1. 1 or a variant thereof. This applies also to the other methods disclosed herein.

[0255] The plant may also be a seed from which a plant can be grown.

[0256] The plant part may be a tissue or tissue sample, a cell, a seed or seed sample, a fruit or fruit sample, a leaf or leaf sample, etc.

[0257] For analyzing the genomic DNA at least crude genomic DNA extraction may be necessary. The presence of a SNP marker allele in the genomic DNA can be detected directly or indirectly. Directly may for example be by nucleic acid hybridization of e.g. oligonucleotide probes. Indirectly may for example be by nucleic acid amplification using e.g. PCR primers which comprise e.g. a tail sequence attached to the primer and during PCR the allele-specific primer binds to the template DNA and elongates, thereby attaching the tail sequence to the newly synthesized strand and in subsequent PCR rounds a FRET cassette (fluorescent resonant energy transfer cassette) binds to the tail and emits fluorescence. The fluorescent signal can then be detected. This is used e.g. in the KASP-assay.

[0258] The SNP nucleotide and haplotype or genotype for SNP_01 to SNP_14 can be detected in the genome of the plant or plant part using known methods, such as known genotyping methods. Various genotyping assays can be used, as long as they can detect SNPs. Genotyping assays are generally based on specific primers used in PCR or thermal cycling reactions (polymerase chain reaction) to amplify either allele and detect the amplification product or on allele-specific oligonucleotide probes, which hybridize to either allele or both. For example genotyping with BHQplus probes uses two allele specific probes and two primers that flank the region of the polymorphism, and during thermal cycling the polymerase encounters the allele-specific probes bound to the DNA and releases a fluorescent signal. Allele discrimination involves competitive binding of the two allele-specific BHQPlus probes (see also biosearchtech.com).

[0259] Examples of genotyping assays are the KASP -assay (by LGC, see www at LGCgenomics.com and also www at biosearchtech.com / products / pcr-kits-and-reagents / genotyping-assays / kasp-genotyping-chemistry), based on competitive allele-specific PCR and end-point fluorescent detection, the TaqMan-assay (Applied Biosytstems), which is also PCR based, HRM assays (High Resolution Melting Assay), wherein allele -specific probes are detected using real time PCR, or the rhAmp assay, based on Rnase H2-dependent PCR, BHQplus genotyping, BHQplex CoPrimer genotyping and many others.

[0260] The KASP-assay is also described in He C, Holme J, Anthony J. ‘SNP genotyping: the KASP assay. Methods Mol Biol. 2014;1145:75-86’ and EP1726664B1 or US7615620 B2, incorporated by reference. The KASP genotyping assay utilizes a unique form of competitive allele-specific PCR combined with a novel, homogeneous, fluorescence-based reporting system for the identification and measurement of genetic variation occurring at the nucleotide level to detect single nucleotide polymorphisms (SNPs) or inserts and deletions (InDeis). The KASP technology is suitable for use on a variety of equipment platforms and provides flexibility in terms of the number of SNPs and the number of samples able to be analyzed. The KASP chemistry functions equally well in 96-, 384-, and 1,536-well microtiter plate formats and has been utilized over many years in large and small laboratories by users across the fields of human, animal, and plant genetics.

[0261] The TaqMan genotyping assays is also described in Woodward J. ‘Bi-allelic SNP genotyping using the TaqMan® assay.’ Methods Mol Biol. 2014; 1145:67-74, US5210015 and US5487972, incorporated herein by reference. With TaqMan(®) technology allele-specific probes are utilized for quick and reliable genotyping of known polymorphic sites. TaqMan assays are robust in genotyping multiple variant types, including single nucleotide polymorphisms, insertions / deletions, and presence / absence variants. To query a single bi-allelic polymorphism, two TaqMan probes labeled with distinct fluorophores are designed such that they hybridize to different alleles during PCR-based amplification of a surrounding target region. During the primer extension phase of PCR, the 5'-3' exonuclease activity of Taq polymerase cleaves and releases the fluorophores from bound probes. At the end of PCR, the emission intensify of each fluorophore is measured and allele determination at the queried site can be made.

[0262] Various genotyping assays can, therefore, be used, which can differentiate between the SNP markers. Provided is, in one aspect, a method for detecting, and optionally selecting, a cucumber plant, seed or plant part comprising at least one copy of an introgression fragment on chromosome 1 comprising QTL1. 1, comprising the steps of: a) providing one or more genomic DNA samples of one or more cucumber plants, seeds or plant parts, b) carrying out a genotyping assay for at least 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13 or 14 markers selected from SNP_1 to SNP_14, using the DNA samples of a) as template, wherein the genotyping assay discriminates between or detects the nucleotide present for the SNP marker at nucleotide 102 of SEQ ID NO: 1 to SEQ ID NO: 14 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 1 to SEQ ID NO: 14, wherein said genotyping assay is based on nucleic acid amplification making use of oligonucleotide primers and / or wherein said genotyping assay is based on nucleic acid hybridization making use of oligonucleotide probes, and optionally c) identifying or selecting a cucumber plant, seed or plant part comprising a SNP haplotype or genotype of at least 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13 or 14 markers selected from the group consisting of: a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_01 at nucleotide 102 of SEQ ID NO: 1 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 1; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_02 at nucleotide 102 of SEQ ID NO: 2 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 2; a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_03 at nucleotide 102 of SEQ ID NO: 3 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 3; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_04 at nucleotide 102 of SEQ ID NO: 4 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 4; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_05 at nucleotide 102 of SEQ ID NO: 5 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 5; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_06 at nucleotide 102 of SEQ ID NO: 6 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 6; a Thymine (T) nucleotide for the Sing le Nucleotide Polymorphism marker SNP_07 at nucleotide 102 of SEQ ID NO: 7 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 7; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_08 at nucleotide 102 of SEQ ID NO: 8 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 8; a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_09 at nucleotide 102 of SEQ ID NO: 9 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 9; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_10 at nucleotide 102 of SEQ ID NO: 10 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 10; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_11 at nucleotide 102 of SEQ ID NO: 11 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 11, such as a Guanine for SNP l 1 at nucleotide 102 of SEQ ID NO: 34; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_12 at nucleotide 102 of SEQ ID NO: 12 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 12; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_13 at nucleotide 102 of SEQ ID NO: 13 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 13; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_14 at nucleotide 102 of SEQ ID NO: 14 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 14.

[0263] In step c) plants or plant parts comprising the introgression fragment in homozygous form or in heterozygous form may be selected. In one aspect the plants or plant parts comprising at least 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13 or 14 markers of SNP_1 to SNP_14 comprise at least the donor nucleotide for the peak marker, which is SNP_11. In one aspect they comprise at least the donor nucleotide for SNP_9, SNP_10, SNP ll, SNP_12 and SNP_13. In another aspect they comprise at least the donor nucleotide for SNP_08, SNP_9, SNP_10, SNP_11, SNP_12, SNP_13 and SNP_14. Therefore, in one aspect in step b) at least these SNP markers are assayed and / or in step c) plants, seeds or plant parts comprising at least these SNP markers are identified and / or selected.

[0264] The method may further comprise optionally selfing the selected plant in order to generate a plant homozygous for the introgression fragment.

[0265] The method may also comprise growing the selected plant (or the optionally selfed plants) and allowing fruits to develop and analyzing the fruits for at least the glucose and fructose content, optionally also for malic acid content. This allows phenotypic determination if QTL1.1 is present in the plant. This applies herein for all methods that comprise a step to detect and / or select one or more of the SNP markers.

[0266] In another aspect a method for generating a PCR amplification product and / or a oligonucleotide hybridization product of (a part of the) genomic DNA of cucumber plants, seeds or plant parts is provided comprising the steps of: a) providing a sample (or a plurality of samples) of genomic DNA of a cucumber plant or of a plurality of plants (e.g. a F2 population, inbred lines, a backcross population, a breeding population, hybrid plants, etc.), b) providing at least a pair of PCR primers or at least one oligonucleotide probe, which primers or (oligonucleotide) probe comprise at least 10, 11, 12, 13, 14, 15, 16, 17, 18, 19, 20, 21, 22 or more consecutive nucleotides of the SNP marker sequences of SEQ ID NO: 1 to SEQ ID NO: 14 (or the complement sequence of any of these) or of SEQ ID NO: 1 to 10, SEQ ID NO: 34 and SEQ ID NO: 12 to 14 (or the complement sequence of any of these) and can hybridize to the genomic SNP marker region and / or amplify part of the genomic SNP marker region in a PCR assay, and c) carrying out a PCR assay using the primer pair or a hybridization assay using the probe of step b) on the sample(s) of step a) to generate a PCR amplification product and / or an oligonucleotide hybridization product, and optionally d) selecting a cucumber plant, seed or plant part comprising a SNP haplotype or genotype of at least 2, 3,

[0267] 4, 5, 6, 7, 8, 9, 10, 11, 12, 13 or 14 markers selected from the group consisting of: a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_01 at nucleotide 102 of SEQ ID NO: 1 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 1; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_02 at nucleotide 102 of SEQ ID NO: 2 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 2; a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_03 at nucleotide 102 of SEQ ID NO: 3 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 3; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_04 at nucleotide 102 of SEQ ID NO: 4 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 4; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_05 at nucleotide 102 of SEQ ID NO: 5 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 5; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_06 at nucleotide 102 of SEQ ID NO: 6 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 6; a Thymine (T) nucleotide for the Sing le Nucleotide Polymorphism marker SNP_07 at nucleotide 102 of SEQ ID NO: 7 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 7; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_08 at nucleotide 102 of SEQ ID NO: 8 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 8; a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_09 at nucleotide 102 of SEQ ID NO: 9 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 9; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_10 at nucleotide 102 of SEQ ID NO: 10 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 10; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_11 at nucleotide 102 of SEQ ID NO: 11 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 11, such as a Guanine for SNP l 1 at nucleotide 102 of SEQ ID NO: 34; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_12 at nucleotide 102 of SEQ ID NO: 12 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 12; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_13 at nucleotide 102 of SEQ ID NO: 13 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 13; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_14 at nucleotide 102 of SEQ ID NO: 14 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 14.

[0268] Further a method of producing C. sativus Fl hybrid plants comprising an introgression fragment on chromosome 1, wherein said fragment comprises QTL1. 1 (or a variant thereof) comprising: a) providing a first inbred cucumber plant comprising a recombinant chromosome 1 in homozygous form having an introgression fragment comprising QTL1. 1 (or a variant thereof) and comprising the donor SNP nucleotide (the donor SNP haplotype or donor SNP genotype) for at least 1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13 or all 14 SNP markers linked to the QTL (as e.g. indicated in Table 2 or 3 and further above herein), optionally wherein said introgression fragment is derivable from (or derived from) NCIMB 44295 or progeny thereof, b) providing a second inbred cucumber plant, c) crossing said cucumber plant of a) with said cucumber plant of b), d) collecting Fl hybrid seeds from said cross.

[0269] The F 1 hybrid seeds collected are also an embodiment of the invention.

[0270] In another aspect a method for generating progeny of NCIMB 44295 is provided, said method comprising: a) growing a plant from seeds deposited under accession number NCIMB 44295; b) selfing said plant one or more times and / or crossing said plant one or more times with another cucumber plant to generate progeny seeds; c) screening said progeny seeds or plants grown from said seeds or parts of the seeds or plants using a molecular marker assay which detects at least 1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13 or all 14 SNP markers selected from the group consisting of: a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_01 at nucleotide 102 of SEQ ID NO: 1 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 1; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_02 at nucleotide 102 of SEQ ID NO: 2 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 2; a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_03 at nucleotide 102 of SEQ ID NO: 3 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 3; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_04 at nucleotide 102 of SEQ ID NO: 4 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 4; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_05 at nucleotide 102 of SEQ ID NO: 5 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 5; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_06 at nucleotide 102 of SEQ ID NO: 6 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 6; a Thymine (T) nucleotide for the Sing le Nucleotide Polymorphism marker SNP_07 at nucleotide 102 of SEQ ID NO: 7 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 7; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_08 at nucleotide 102 of SEQ ID NO: 8 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 8; a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_09 at nucleotide 102 of SEQ ID NO: 9 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 9; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_10 at nucleotide 102 of SEQ ID NO: 10 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 10; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_11 at nucleotide 102 of SEQ ID NO: 11 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 11, such as a Guanine for SNP l 1 at nucleotide 102 of SEQ ID NO: 34; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_12 at nucleotide 102 of SEQ ID NO: 12 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 12; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_13 at nucleotide 102 of SEQ ID NO: 13 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 13; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_14 at nucleotide 102 of SEQ ID NO: 14 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 14; d) identifying and / or selecting a progeny plant comprising the donor SNP nucleotide (the donor SNP haplotype or donor SNP genotype) for at least 1, 2, 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13 or all 14 of the SNP markers of step c) above.

[0271] The donor SNP nucleotide (or haplotype or genotype) is described above and in e.g. Table 2.

[0272] Further provided is a method for identifying and / or selecting a wild or cultivated cucumber plant for the presence in the genome of QTL1. 1 or a variant thereof, comprising: a) screening seeds or parts of the seeds or plants or parts of plants or DNA of such plants or plant parts of one or more wild cucumber accessions or cultivated cucumber plants using a molecular marker assay which detects at least 5, 6, 7, 8, 9, 10, 11, 12, 13 or 14 SNP markers of SNP_01 to SNP_14 for detecting the introgression fragment on chromosome 1 comprising QTL1. 1 (or a variant); b) identifying and / or selecting a wild cucumber accession or cultivated cucumber plant comprising the donor SNP nucleotide (the donor SNP haplotype or donor SNP genotype) for at least 5, 6, 7, 8, 9, 10, 11, 12, 13 or 14 of SNP_01 to SNP_14 for detecting the introgression fragment or region on chromosome 1 comprising QTL1.1 (or a variant); wherein the donor SNP nucleotide is: a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_01 at nucleotide 102 of SEQ ID NO: 1 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 1; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_02 at nucleotide 102 of SEQ ID NO: 2 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 2; a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_03 at nucleotide 102 of SEQ ID NO: 3 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 3; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_04 at nucleotide 102 of SEQ ID NO: 4 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 4; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_05 at nucleotide 102 of SEQ ID NO: 5 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 5; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_06 at nucleotide 102 of SEQ ID NO: 6 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 6; a Thymine (T) nucleotide for the Sing le Nucleotide Polymorphism marker SNP_07 at nucleotide 102 of SEQ ID NO: 7 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 7; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_08 at nucleotide 102 of SEQ ID NO: 8 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 8; a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_09 at nucleotide 102 of SEQ ID NO: 9 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 9; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_10 at nucleotide 102 of SEQ ID NO: 10 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 10; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_11 at nucleotide 102 of SEQ ID NO: 11 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 11, such as a Guanine for SNP l 1 at nucleotide 102 of SEQ ID NO: 34; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_12 at nucleotide 102 of SEQ ID NO: 12 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 12; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_13 at nucleotide 102 of SEQ ID NO: 13 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 13; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_14 at nucleotide 102 of SEQ ID NO: 14 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 14;

[0273] The above method may optionally also comprise selfing the wild accessions or cultivated plants one or more times, e.g. prior to step a).

[0274] Wild accessions may be accessions of wild or primitive cucumber or relatives of cucumber obtained e.g. from seed depositories, such as USDA ARS-GRIN collections, CGN collections, and others.

[0275] As mentioned previously, the above methods may also comprise phenotypically testing the cucumber fruits produced by plants for at least the fruit glucose and fructose levels in order to confirm the presence of QTL1. 1 (or a variant thereof). This may be done e.g. prior to a SNP marker assay and / or after a SNP marker haplotype or genotype has been identified and / or selected, e.g. after selection or identification of one or more plants comprising a SNP haplotype or SNP genotype (for one or more or all of the SNP markers) identical to the one described in Table 2.

[0276] Also a method for transferring QTL1.1 from a wild donor into a cultivated cucumber plant is provided comprising: a) providing a wild cucumber accession (or a progeny thereof obtained by selfing one or more times) comprising the donor SNP nucleotide (the donor SNP haplotype or donor SNP genotype) for at least 5, 6, 7, 8, 9, 10, 11, 12, 13 or 14 of SNP_01 to SNP_14 for detecting the introgression fragment or region on chromosome 1 comprising QTL1. 1 (or a variant) wherein the donor SNP nucleotide is: a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_01 at nucleotide 102 of SEQ ID NO: 1 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 1; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_02 at nucleotide 102 of SEQ ID NO: 2 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 2; a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_03 at nucleotide 102 of SEQ ID NO: 3 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 3; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_04 at nucleotide 102 of SEQ ID NO: 4 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 4; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_05 at nucleotide 102 of SEQ ID NO: 5 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 5; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_06 at nucleotide 102 of SEQ ID NO: 6 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 6; a Thymine (T) nucleotide for the Sing le Nucleotide Polymorphism marker SNP_07 at nucleotide 102 of SEQ ID NO: 7 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 7; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_08 at nucleotide 102 of SEQ ID NO: 8 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 8; a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_09 at nucleotide 102 of SEQ ID NO: 9 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 9; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_10 at nucleotide 102 of SEQ ID NO: 10 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 10; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_11 at nucleotide 102 of SEQ ID NO: 11 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 11, such as a Guanine for SNP_11 at nucleotide 102 of SEQ ID NO: 34; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_12 at nucleotide 102 of SEQ ID NO: 12 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 12; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_13 at nucleotide 102 of SEQ ID NO: 13 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 13; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_14 at nucleotide 102 of SEQ ID NO: 14 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 14; b) crossing said accession with a cultivated cucumber plant to obtain progenies of the Fl, F2, F3 or further selfing generations or BC1, BC2, BC3 or further backcross generations, and optionally c) selecting a progeny plant comprising QTL1. 1 or a variant thereof.

[0277] The cucumber plant in step b is preferably a cultivated cucumber, such as a European greenhouse cucumber or long cucumber type or a slicer or a short cucumber.

[0278] A progeny plant generated by the above method is also an aspect of the invention.

[0279] Also containers and packages containing or comprising seeds from which plants of the invention can be grown are provided herein.

[0280] Also containers and packages containing or comprising cucumbers of the invention are provided herein. These may be labelled as containing sweeter cucumber fruits. Also progeny seeds and progeny plants of plants of the invention are provided, which retain the introgression on chromosome 1 (QTL1. 1 or a variant), or which comprise a smaller introgression of QTL1. 1 (e.g. derivable from the fragment as is present in NCIMB 44295) which still confers the increase in fruit glucose and fructose levels, i.e. which still contains QTL1. 1. Progeny may be any generation obtained by selfing a cucumber plant according to the invention and / or crossing a cucumber plant according to the invention with another cucumber plant one or more times. Progeny are, therefore, either the generation (seeds) produced from the first cross (Fl) or selfing (SI), or any further generation produced by crossing and / or selfing (F2, F3, etc.) and / or backcrossing (BC1, BC2, etc.) one or more selected plants of the Fl and / or S I and / or BC1 generation (or plants of any further generation, e.g. the F2) with another cucumber plant (and / or with a wild cucumber). Progeny are preferably selected to retain introgression fragments from a wild cucumber comprising QTL1. 1. The presence of (or retention of) the introgression fragments comprising the QTL can be determined phenotypically and / or using the molecular marker assay(s) described herein.

[0281] In a further aspect parts of the cucumber plants according to the invention are provided. Parts include for example cells and cell-cultures, tissue cultures, vegetative plant tissues (leaves, roots, etc.), flowers, pollen, embryos, fruits, parts of fruits, etc. The plant parts comprise the introgression fragment on chromosome 1, as described, and as can be detected using one or more of the markers described. Also, when whole plants are regenerated from such cucumber parts, such as cells, cell- or tissue cultures, the regenerated plants comprise the recombinant chromosome 1.

[0282] Thus, also provided is a plant cell, tissue or plant part of a plant or of a seed according the invention comprising at least one recombinant chromosome 1, wherein said recombinant chromosome 1 comprises an introgression fragment from a wild donor cucumber plant and wherein said introgression fragment comprises QTL1. 1 (or a variants).

[0283] Also in vitro cell cultures and in vitro tissue cultures are encompassed herein, of cells or tissues comprising a recombinant chromosome 1 as described. Preferably the cells or tissues can be regenerated into a whole cucumber plant, i.e. the cells are regenerable cells and the tissues comprise regenerable cells. Thus, also vegetative propagations of the plants according to the invention are an embodiment herein. Thus, a vegetatively propagated cultivated cucumber plant is provided which comprises a recombinant chromosome 1 as described herein. In a different aspect non-propagating cells comprising QTL 1.1 (or a variant thereof), is encompassed herein, as are tissues comprising such cells.

[0284] In a specific aspect a cucumber fruit harvested from a plant according to the invention is provided. Marketable cucumber fruits, especially for the fresh market (slicing), are generally graded according to fruit size and quality characteristics after harvest. See e.g. the United States Standards for Grades of Cucumbers, US Department of Agriculture, Effective March 1, 1985 and reprinted January 1997. Herein different grades of cucumbers are distinguished. Thus, in one aspect harvested fruits are provided of U.S. Fancy grade, U.S. Extra No. 1 grade, U.S. No. 1 grade, U.S. No. 1 Small grade, U.S. No. 1 Large grade, U.S. No. 2 grade. Also containers or packages comprising or consisting of harvested cucumber fruits are provided. Again, the cells of the fruits are distinguishable from other cucumber fruits by the presence of QTL1.1 (or a variant thereof) (as determinable in one or more of the molecular marker assays).

[0285] In another aspect the cucumber is a long cucumber type or a short cucumber type or a slicer cucumber type and fruits harvested and optionally processed (e.g. sliced or diced) are provided.

[0286] In another aspect the cucumber is a pickling type and fruits harvested and optionally pickled are provided.

[0287] The invention also provides for a food or feed product comprising or consisting of a plant part described herein preferably a cucumber fruit or part thereof and / or an extract from a plant part described herein. The food or feed product may be fresh or processed, e.g., pickled, canned, steamed, boiled, fried, blanched and / or frozen, etc. For example, containers such as cans, boxes, crates, bags, cartons, Modified Atmosphere Packaging, films (e.g. biodegradable films), etc. comprising plant parts such as fruits or fruit parts (fresh and / or processed) described herein are also provided herein.

[0288] Also encompassed herein is a method for producing a cultivated cucumber plant comprising an introgression fragments on chromosome 1, wherein said introgression fragment comprises QTL1.1 (or a variant thereof), comprising: a) providing a first cultivated cucumber plant, preferably lacking QTL 1.1, b) providing a second cultivated cucumber plant selected from plants grown from seeds deposited under accession number NCIMB44295 or progeny thereof or providing a second cultivated or wild cucumber plant comprising QTL1. 1 (or a variant therof) and comprising the donor SNP haplotype or genotype for at least 5, 6, 7, 8, 9, 10, 11, 12, 13 or all 14 SNP markers linked to the QTL, c) crossing said plant of a) with said plant of b), d) collecting F 1 seeds from said cross and optionally selfing said F 1 plants one or more times to produce an F2 or F3 or further selfing population, e) optionally backcrossing the F 1 plant or an F2 or F3 or further selfing plant to the plant of a) to produce a backcross population, f) optionally selfing the backcross population one or more times, g) identifying a Fl, F2, F3, further selfing or backcross plant which comprises the donor SNP haplotype or genotype for at least 5, 6, 7, 8, 9, 10, 11, 12, 13 or all 14 SNP markers linked to the QTL.

[0289] The donor SNP haplotype or genotype is provided e.g. in Table 2 and further above.

[0290] In a further aspect a method of producing F 1 hybrid plants is provided comprising: a) providing a first inbred cucumber plant comprising an introgression fragment comprising QTL 1.1, wherein said introgression fragment is the fragment as found in NCIMB44295, or a shorter fragment of that introgression fragment and / or wherein the QTL is the QTL as found in NCIMB44295, or wherein the QTL comprising a donor SNP haplotype or genotype for at least 5, 6, 7, 8, 9, 10, 11, 12, 13 or all 14 SNP markers linked to the QTL, b) providing a second inbred cucumber plant (optionally comprising an introgression fragment comprising QTL1. 1, wherein said introgression fragment is the fragment as found in NCIMB44295, or a shorter fragment of that introgression fragment and / or wherein the QTL is the QTL as found in NCIMB44295, or wherein the QTL comprising a donor SNP haplotype or genotype for at least 5, 6, 7, 8, 9, 10, 11, 12, 13 or all 14 SNP markers linked to the QTL), c) crossing said plant of a) with said plant of b), d) collecting Fl hybrid seeds from said cross.

[0291] Also provided is the use of plants grown from seeds deposited under accession number NCIMB 44295, or progeny thereof, for generating a cultivated cucumber plant which produces fruits comprising an increased glucose and fructose level, wherein said increased fruit sugar level is conferred by an introgression fragment obtained from chromosome 1 of said plants or progeny.

[0292] Also, the molecular marker sequences (and isolated nucleic acid molecules comprising the sequence) disclosed herein and their use in detecting and / or generating cucumber plants comprising said QTLs described herein are encompassed herein.

[0293] Further a method of growing a plant comprising QTL1. 1 (or a variant thereof) in the field, or in glasshouses or tunnels, whereby the fruits comprise higher average glucose and fructose levels compared to the fruits of plants lacking the QTL.

[0294] A method is provided for identifying (or detecting and / or selecting) a cucumber plant or plant part comprising an introgression fragments on chromosome 3 from a wild cucumber donor, wherein said introgression fragments comprises a Quantitative Trait Locus (QTL) conferring a mint-green fruit flesh color of the cucumber fruit, wherein QTL3.1 is located on chromosome 3 between the Single Nucleotide Polymorphism marker SNP_15 at nucleotide 102 of SEQ ID NO: 15 (or SNP_15 at nucleotide 102 of SEQ ID NO: 35) and SNP_24 at nucleotide 102 of SEQ ID NO: 24, comprising: a) providing a cucumber plant or plant part or DNA of such plant or plant part, b) screening said plant, plant part or DNA for the presence of the SNP haplotype or genotype of at least 2, 3, 4, 5, 6, 7, 8, 9 or 10 markers selected from the group consisting of: an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_15 at nucleotide 102 of SEQ ID NO: 15 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 15, such as an Adenine for SNP_15 at nucleotide 102 of SEQ ID NO: 35; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_16 at nucleotide 102 of SEQ ID NO: 16 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 16; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_17 at nucleotide 102 of SEQ ID NO: 17 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 17, such as a Guanine for SNP 17 at nucleotide 102 of SEQ ID NO: 36; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_18 at nucleotide 102 of SEQ ID NO: 18 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 18; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_19 at nucleotide 102 of SEQ ID NO: 19 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 19; a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_20 at nucleotide 102 of SEQ ID NO: 20 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 20, such as a Cytosine for SNP_20 at nucleotide 101 of SEQ ID NO: 37; a Cytosine (C) nucleotide for the Sing le Nucleotide Polymorphism marker SNP_21 at nucleotide 102 of SEQ ID NO: 21 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 21; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_22 at nucleotide 102 of SEQ ID NO: 22 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 22; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_23 at nucleotide 102 of SEQ ID NO: 23 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 23; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_24 at nucleotide 102 of SEQ ID NO: 24 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 24 and c) identifying and / or selecting a plant or plant part comprising the SNP haplotype for at least 1, 2, 3, 4, 5, 6, 7, 8, 9 or all 10 of the SNP markers of step b).

[0295] In one aspect the plants or plant parts comprising at least 3, 4, 5, 6, 7, 8, 9 or 10 markers of SNP_15 to SNP_24 comprise at least the donor nucleotide for the peak marker, which is SNP 15. In one aspect they comprise at least the donor nucleotide for SNP_15, SNP_16, SNP_17, SNP_18 and SNP_19. In another aspect they comprise at least the donor nucleotide for SNP_15, SNP_16, SNP_17 and SNP_18. Therefore, in one aspect in step b) at least these SNP markers are screened and / or in step c) plants or plant parts comprising at least these SNP markers are identified and / or selected.

[0296] In step c) plants or plant parts comprising the introgression fragment in homozygous form or in heterozygous form may be identified and / or selected. It is understood that in step b) the genotype for the markers may be assayed, i.e. the SNP nucleotide on both chromosomes of the pair of chromosome 3, as e.g. disclosed in Table 5 or 6 and elsewhere herein.

[0297] The method may further comprise optionally selfing the selected plant in order to generate a plant homozygous for the introgression fragment.

[0298] The method may also comprise growing the selected plant (or the optionally selfed plants) and allowing fruits to develop and analyzing the fruits for their fruit flesh color, especially the mesocarp color. This allows phenotypic determination if QTL3. 1 is present in the plant.

[0299] The SNP haplotype or SNP genotype of SNP_15 to SNP_24 are e.g. disclosed in Table 5 or 6 and elsewhere herein. In the above method, thus, also the SNP genotype may be screened and / or selected for. In step c) plants or plant parts comprising the introgression fragment in homozygous form or in heterozygous form may be identified and / or selected.

[0300] The cucumber plant may (in any of the methods herein) be a cultivated cucumber plant or plant part may be any plant or plant part, such as commercial varieties, mapping populations, etc. The cucumber plant may also or alternatively be a wild cucumber plant or plant part, in which case not an ‘introgression fragment’ is detected, but the endogenous chromosome 3 region is analyzed for the presence of the SNP markers linked to QTL3. 1 or a variant thereof. This applies also to the other methods disclosed herein.

[0301] The plant may also be a seed from which a plant can be grown.

[0302] The plant part may be a tissue or tissue sample, a cell, a seed or seed sample, a fruit or fruit sample, a leaf or leaf sample, etc.

[0303] For analyzing the genomic DNA at least crude genomic DNA extraction may be necessary. The presence of a SNP marker allele in the genomic DNA can be detected directly or indirectly. Directly may for example be by nucleic acid hybridization of e.g. oligonucleotide probes. Indirectly may for example be by nucleic acid amplification using e.g. PCR primers which comprise e.g. a tail sequence attached to the primer and during PCR the allele-specific primer binds to the template DNA and elongates, thereby attaching the tail sequence to the newly synthesized strand and in subsequent PCR rounds a FRET cassette (fluorescent resonant energy transfer cassette) binds to the tail and emits fluorescence. The fluorescent signal can then be detected. This is used e.g. in the KASP-assay.

[0304] The SNP nucleotide and haplotype or genotype for SNP_15 to SNP_24 can be detected in the genome of the plant or plant part using known methods, such as known genotyping methods. Various genotyping assays can be used, as long as they can detect SNPs. Genotyping assays are generally based on specific primers used in PCR or thermal cycling reactions (polymerase chain reaction) to amplify either allele and detect the amplification product or on allele-specific oligonucleotide probes, which hybridize to either allele or both. For example genotyping with BHQplus probes uses two allele specific probes and two primers that flank the region of the polymorphism, and during thermal cycling the polymerase encounters the allele-specific probes bound to the DNA and releases a fluorescent signal. Allele discrimination involves competitive binding of the two allele-specific BHQPlus probes (see also biosearchtech.com).

[0305] Examples of genotyping assays are the KASP-assay (by LGC, see www at LGCgenomics.com and also www at biosearchtech.com / products / pcr-kits-and-reagents / genotyping-assays / kasp-genotyping-chemistry), based on competitive allele-specific PCR and end-point fluorescent detection, the TaqMan-assay (Applied Biosytstems), which is also PCR based, HRM assays (High Resolution Melting Assay), wherein allele -specific probes are detected using real time PCR, or the rhAmp assay, based on Rnase H2-dependent PCR, BHQplus genotyping, BHQplex CoPrimer genotyping and many others.

[0306] The KASP-assay is also described in He C, Holme J, Anthony J. ‘SNP genotyping: the KASP assay. Methods Mol Biol. 2014;1145:75-86’ and EP1726664B1 or US7615620 B2, incorporated by reference. The KASP genotyping assay utilizes a unique form of competitive allele-specific PCR combined with a novel, homogeneous, fluorescence-based reporting system for the identification and measurement of genetic variation occurring at the nucleotide level to detect single nucleotide polymorphisms (SNPs) or inserts and deletions (InDeis). The KASP technology is suitable for use on a variety of equipment platforms and provides flexibility in terms of the number of SNPs and the number of samples able to be analyzed. The KASP chemistry functions equally well in 96-, 384-, and 1,536-well microtiter plate formats and has been utilized over many years in large and small laboratories by users across the fields of human, animal, and plant genetics.

[0307] The TaqMan genotyping assays is also described in Woodward J. ‘Bi-allelic SNP genotyping using the TaqMan® assay.’ Methods Mol Biol. 2014; 1145:67-74, US5210015 and US5487972, incorporated herein by reference. With TaqMan(®) technology allele-specific probes are utilized for quick and reliable genotyping of known polymorphic sites. TaqMan assays are robust in genotyping multiple variant types, including single nucleotide polymorphisms, insertions / deletions, and presence / absence variants. To query a single bi-allelic polymorphism, two TaqMan probes labeled with distinct fluorophores are designed such that they hybridize to different alleles during PCR-based amplification of a surrounding target region. During the primer extension phase of PCR, the 5'-3' exonuclease activity of Taq polymerase cleaves and releases the fluorophores from bound probes. At the end of PCR, the emission intensity of each fluorophore is measured and allele determination at the queried site can be made.

[0308] Various genotyping assays can, therefore, be used, which can differentiate between the SNP markers.

[0309] Provided is, in one aspect, a method for detecting, and optionally selecting, a cucumber plant, seed or plant part comprising at least one copy of an introgression fragment on chromosome 3 comprising QTL3. 1, comprising the steps of: a) providing one or more genomic DNA samples of one or more cucumber plants, seeds or plant parts, b) carrying out a genotyping assay for at least 2, 3, 4, 5, 6, 7, 8, 9 or 10 markers selected from SNP_15 to SNP_24, using the DNA samples of a) as template, wherein the genotyping assay discriminates between or detects the nucleotide present for the SNP marker at nucleotide 102 of SEQ ID NO: 15 to SEQ ID NO: 24 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 15 to SEQ ID NO: 24 (such as SNP_15 at nucleotide 102 of SEQ ID NO: 35, SNP_17 at nucleotide 102 of SEQ ID NO: 36 and SNP_20 at nucleotide 101 of SEQ ID NO: 37), wherein said genotyping assay is based on nucleic acid amplification making use of oligonucleotide primers and / or wherein said genotyping assay is based on nucleic acid hybridization making use of oligonucleotide probes, and optionally c) identifying and / or selecting a cucumber plant, seed or plant part comprising a SNP haplotype or genotype of at least 2, 3, 4, 5, 6, 7, 8, 9 or 10 markers selected from the group consisting of: an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_15 at nucleotide 102 of SEQ ID NO: 15 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 15, such as an Adenine for SNP_15 at nucleotide 102 of SEQ ID NO: 35; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_16 at nucleotide 102 of SEQ ID NO: 16 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 16; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_17 at nucleotide 102 of SEQ ID NO: 17 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 17, such as a Guanine for SNP 17 at nucleotide 102 of SEQ ID NO: 36; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_18 at nucleotide 102 of SEQ ID NO: 18 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 18; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_19 at nucleotide 102 of SEQ ID NO: 19 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 19; a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_20 at nucleotide 102 of SEQ ID NO: 20 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 20, such as a Cytosine for SNP_20 at nucleotide 101 of SEQ ID NO: 37; a Cytosine (C) nucleotide for the Sing le Nucleotide Polymorphism marker SNP_21 at nucleotide 102 of SEQ ID NO: 21 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 21; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_22 at nucleotide 102 of SEQ ID NO: 22 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 22; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_23 at nucleotide 102 of SEQ ID NO: 23 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 23; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_24 at nucleotide 102 of SEQ ID NO: 24 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 24.

[0310] In step c) plants or plant parts comprising the introgression fragment in homozygous form or in heterozygous form may be selected.

[0311] In one aspect the plants or plant parts comprising at least 2, 3, 4, 5, 6, 7, 8, 9 or 10 markers of SNP_15 to SNP_24 comprise at least the donor nucleotide for the peak marker, which is SNP_15. In one aspect they comprise at least the donor nucleotide for SNP_15, SNP_16, SNP_17, SNP_18 and SNP_19. In another aspect they comprise at least the donor nucleotide for SNP_15, SNP_16, SNP_17 and SNP_18. Therefore, in one aspect in step b) at least these SNP markers are assayed and / or in step c) plants, seeds or plant parts comprising at least these SNP markers are identified and / or selected.

[0312] The method may further comprise optionally selfing the selected plant in order to generate a plant homozygous for the introgression fragment.

[0313] The method may also comprise growing the selected plant (or the optionally selfed plants) and allowing fruits to develop and analyzing the fruit flesh color, especially the mesocarp color. This allows phenotypic determination if QTL3.1 is present in the plant. This applies herein for all methods that comprise a step to detect and / or select one or more of the SNP markers.

[0314] In another aspect a method for generating a PCR amplification product and / or a oligonucleotide hybridization product of (a part of the) genomic DNA of cucumber plants, seeds or plant parts is provided comprising the steps of: a) providing a sample (or a plurality of samples) of genomic DNA of a cucumber plant or of a plurality of plants (e.g. a F2 population, inbred lines, a backcross population, a breeding population, hybrid plants, etc.), b) providing at least a pair of PCR primers or at least one oligonucleotide probe, which primers or (oligonucleotide) probe comprise at least 10, 11, 12, 13, 14, 15, 16, 17, 18, 19, 20, 21, 22 or more consecutive nucleotides of the SNP marker sequences of SEQ ID NO: 15 to SEQ ID NO: 24 (or the complement sequence of any of these) or of SEQ ID NO: 35, SEQ ID NO: 16, SEQ ID NO: 36, SEQ ID NO: 18 and 19, SEQ ID NO: 37, SEQ ID NO: 21 to 24 (or the complement of any of these) and can hybridize to the genomic SNP marker region and / or amplify part of the genomic SNP marker region in a PCR assay, and c) carrying out a PCR assay using the primer pair or a hybridization assay using the probe of step b) on the sample(s) of step a) to generate a PCR amplification product and / or an oligonucleotide hybridization product, and optionally d) selecting a cucumber plant, seed or plant part comprising a SNP haplotype or genotype of at least 2, 3, 4, 5, 6, 7, 8, 9 or 10 markers selected from the group consisting of: an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_15 at nucleotide 102 of SEQ ID NO: 15 or at the equivalent position in a sequence comprising at least 95% sequence identify to SEQ ID NO: 15, such as an Adenine for SNP_15 at nucleotide 102 of SEQ ID NO: 35; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_16 at nucleotide 102 of SEQ ID NO: 16 or at the equivalent position in a sequence comprising at least 95% sequence identify to SEQ ID NO: 16; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_17 at nucleotide 102 of SEQ ID NO: 17 or at the equivalent position in a sequence comprising at least 95% sequence identify to SEQ ID NO: 17, such as a Guanine for SNP 17 at nucleotide 102 of SEQ ID NO: 36; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_18 at nucleotide 102 of SEQ ID NO: 18 or at the equivalent position in a sequence comprising at least 95% sequence identify to SEQ ID NO: 18; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_19 at nucleotide 102 of SEQ ID NO: 19 or at the equivalent position in a sequence comprising at least 95% sequence identify to SEQ ID NO: 19; a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_20 at nucleotide 102 of SEQ ID NO: 20 or at the equivalent position in a sequence comprising at least 95% sequence identify to SEQ ID NO: 20, such as a Cytosine for SNP_20 at nucleotide 101 of SEQ ID NO: 37; a Cytosine (C) nucleotide for the Sing le Nucleotide Polymorphism marker SNP_21 at nucleotide 102 of SEQ ID NO: 21 or at the equivalent position in a sequence comprising at least 95% sequence identify to SEQ ID NO: 21; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_22 at nucleotide 102 of SEQ ID NO: 22 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 22; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_23 at nucleotide 102 of SEQ ID NO: 23 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 23; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_24 at nucleotide 102 of SEQ ID NO: 24 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 24.

[0315] Further a method of producing C. sativus Fl hybrid plants comprising an introgression fragment on chromosome 3, wherein said fragment comprises QTL3. 1 (or a variant thereof) comprising: a) providing a first inbred cucumber plant comprising a recombinant chromosome 3 in homozygous form having an introgression fragment comprising QTL3. 1 (or a variant thereof) and comprising the donor SNP nucleotide (the donor SNP haplotype or donor SNP genotype) for at least 1, 2, 3, 4, 5, 6, 7, 8, 9 or all 10 SNP markers linked to the QTL (as e.g. indicated in Table 5 or 6 and further above herein), optionally wherein said introgression fragment is derivable from (or derived from) NCIMB 44295 or progeny thereof, b) providing a second inbred cucumber plant, c) crossing said cucumber plant of a) with said cucumber plant of b), d) collecting Fl hybrid seeds from said cross.

[0316] The F 1 hybrid seeds collected are also an embodiment of the invention.

[0317] In another aspect a method for generating progeny of NCIMB 44295 is provided, said method comprising: a) growing a plant from seeds deposited under accession number NCIMB 44295; b) selfing said plant one or more times and / or crossing said plant one or more times with another cucumber plant to generate progeny seeds; c) screening said progeny seeds or plants grown from said seeds or parts of the seeds or plants using a molecular marker assay which detects at least 1, 2, 3, 4, 5, 6, 7, 8, 9 or all 10 SNP markers selected from the group consisting of: an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_15 at nucleotide 102 of SEQ ID NO: 15 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 15, such as an Adenine for SNP_15 at nucleotide 102 of SEQ ID NO: 35; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_16 at nucleotide 102 of SEQ ID NO: 16 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 16; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_17 at nucleotide 102 of SEQ ID NO: 17 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 17, such as a Guanine for SNP 17 at nucleotide 102 of SEQ ID NO: 36; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_18 at nucleotide 102 of SEQ ID NO: 18 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 18; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_19 at nucleotide 102 of SEQ ID NO: 19 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 19; a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_20 at nucleotide 102 of SEQ ID NO: 20 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 20, such as a Cytosine for SNP_20 at nucleotide 101 of SEQ ID NO: 37; a Cytosine (C) nucleotide for the Sing le Nucleotide Polymorphism marker SNP_21 at nucleotide 102 of SEQ ID NO: 21 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 21; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_22 at nucleotide 102 of SEQ ID NO: 22 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 22; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_23 at nucleotide 102 of SEQ ID NO: 23 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 23; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_24 at nucleotide 102 of SEQ ID NO: 24 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 24. d) identifying and / or selecting a progeny plant comprising the donor SNP nucleotide (the donor SNP haplotype or donor SNP genotype) for at least 1, 2, 3, 4, 5, 6, 7, 8, 9 or all 10 of the SNP markers of step c) above.

[0318] The donor SNP nucleotide (or haplotype or genotype) is described above and in Table 5.

[0319] Further provided is a method for identifying and / or selecting a wild or cultivated cucumber plant for the presence in the genome of QTL3. 1 or a variant thereof, comprising: a) screening seeds or parts of the seeds or plants or parts of plants or DNA of such plants or plant parts of one or more wild cucumber accessions or cultivated cucumber plants using a molecular marker assay which detects at least 5, 6, 7, 8, 9 or 10 SNP markers of SNP_15 to SNP_24 for detecting the introgression fragment on chromosome 3 comprising QTL3. 1 (or a variant); b) identifying and / or selecting a wild cucumber accession or cucumber plant comprising the donor SNP nucleotide (the donor SNP haplotype or donor SNP genotype) for at least 5, 6, 7, 8, 9 or 10 of SNP_15 to SNP_24 for detecting the introgression fragment or region on chromosome 3 comprising QTL3. 1 (or a variant); wherein the donor SNP nucleotide is: an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_15 at nucleotide 102 of SEQ ID NO: 15 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 15, such as an Adenine for SNP_15 at nucleotide 102 of SEQ ID NO: 35; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_16 at nucleotide 102 of SEQ ID NO: 16 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 16; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_17 at nucleotide 102 of SEQ ID NO: 17 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 17, such as a Guanine for SNP 17 at nucleotide 102 of SEQ ID NO: 36; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_18 at nucleotide 102 of SEQ ID NO: 18 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 18; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_19 at nucleotide 102 of SEQ ID NO: 19 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 19; a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_20 at nucleotide 102 of SEQ ID NO: 20 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 20 such as a Cytosine for SNP_20 at nucleotide 101 of SEQ ID NO: 37; a Cytosine (C) nucleotide for the Sing le Nucleotide Polymorphism marker SNP_21 at nucleotide 102 of SEQ ID NO: 21 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 21; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_22 at nucleotide 102 of SEQ ID NO: 22 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 22; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_23 at nucleotide 102 of SEQ ID NO: 23 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 23; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_24 at nucleotide 102 of SEQ ID NO: 24 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 24.

[0320] The above method may optionally also comprise selfing the wild accessions or cultivated plants one or more times, e.g. prior to step a).

[0321] Wild accessions may be accessions of wild or primitive cucumber or relatives of cucumber obtained e.g. from seed depositories, such as USDA ARS-GRIN collections, CGN collections, and others. As mentioned previously, the above methods may also comprise phenotypically analyzing the cucumber fruits produced by plants for their fruit flesh color in order to confirm the presence of QTL3. 1 (or a variant thereof). This may be done e.g. prior to a SNP marker assay and / or after a SNP marker haplotype or genotype has been identified and / or selected, e.g. after selection or identification of one or more plants comprising a SNP haplotype or SNP genotype (for one or more or all of the SNP markers) identical to the one described in Table 5.

[0322] Also, a method for transferring QTL3.1 from a wild donor into a cultivated cucumber plant is provided comprising: a) providing a wild cucumber accession (or a progeny thereof obtained by selfing one or more times) comprising the donor SNP nucleotide (the donor SNP haplotype or donor SNP genotype) for at least 5, 6, 7, 8, 9 or 10 of SNP_15 to SNP_24 for detecting the introgression fragment or region on chromosome 3 comprising QTL3. 1 (or a variant) wherein the donor SNP nucleotide is: an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_15 at nucleotide 102 of SEQ ID NO: 15 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 15, such as an Adenine for SNP_15 at nucleotide 102 of SEQ ID NO: 35; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_16 at nucleotide 102 of SEQ ID NO: 16 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 16; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_17 at nucleotide 102 of SEQ ID NO: 17 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 17, such as a Guanine for SNP 17 at nucleotide 102 of SEQ ID NO: 36; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_18 at nucleotide 102 of SEQ ID NO: 18 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 18; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_19 at nucleotide 102 of SEQ ID NO: 19 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 19; a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_20 at nucleotide 102 of SEQ ID NO: 20 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 20, such as a Cytosine for SNP_20 at nucleotide 101 of SEQ ID NO: 37; a Cytosine (C) nucleotide for the Sing le Nucleotide Polymorphism marker SNP_21 at nucleotide 102 of SEQ ID NO: 21 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 21; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_22 at nucleotide 102 of SEQ ID NO: 22 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 22; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_23 at nucleotide 102 of SEQ ID NO: 23 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 23; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_24 at nucleotide 102 of SEQ ID NO: 24 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 24; b) crossing said accession with a cultivated cucumber plant to obtain progenies of the Fl, F2, F3 or further selfing generations or BC1, BC2, BC3 or further backcross generations, and optionally c) selecting a progeny plant comprising QTL3. 1 or a variant thereof.

[0323] The cucumber plant in step b is preferably a cultivated cucumber, such as a European greenhouse cucumber or long cucumber type or a slicer or a short cucumber.

[0324] A progeny plant generated by the above method is also an aspect of the invention.

[0325] Also containers and packages containing or comprising seeds from which plants of the invention can be grown are provided herein.

[0326] Also containers and packages containing or comprising cucumbers of the invention are provided herein. These may be labelled as containing sweeter cucumber fruits.

[0327] Also progeny seeds and progeny plants of plants of the invention are provided, which retain the introgression on chromosome 3 (QTL3. 1 or a variant), or which comprise a smaller introgression of QTL3. 1 (e.g. derivable from the fragment as is present in NCIMB 44295) which still confers mint-green fruit flesh, i.e. which still contains QTL3.1. Progeny may be any generation obtained by selfing a cucumber plant according to the invention and / or crossing a cucumber plant according to the invention with another cucumber plant one or more times. Progeny are, therefore, either the generation (seeds) produced from the first cross (Fl) or selfing (S I), or any further generation produced by crossing and / or selfing (F2, F3, etc.) and / or backcrossing (BC1, BC2, etc.) one or more selected plants of the F 1 and / or S 1 and / or BC 1 generation (or plants of any further generation, e.g. the F2) with another cucumber plant (and / or with a wild cucumber). Progeny are preferably selected to retain introgression fragments from a wild cucumber comprising QTL3. 1. The presence of (or retention of) the introgression fragments comprising the QTL can be determined phenotypically and / or using the molecular marker assay(s) described herein.

[0328] In a further aspect parts of the cucumber plants according to the invention are provided. Parts include for example cells and cell-cultures, tissue cultures, vegetative plant tissues (leaves, roots, etc.), flowers, pollen, embryos, fruits, parts of fruits, etc. The plant parts comprise the introgression fragment on chromosome 3, as described, and as can be detected using one or more of the markers described. Also, when whole plants are regenerated from such cucumber parts, such as cells, cell- or tissue cultures, the regenerated plants comprise the recombinant chromosome 3.

[0329] Thus, also provided is a plant cell, tissue or plant part of a plant or of a seed according the invention comprising at least one recombinant chromosome 3, wherein said recombinant chromosome 3 comprises an introgression fragment from a wild donor cucumber plant and wherein said introgression fragment comprises QTL3. 1 (or a variants).

[0330] Also in vitro cell cultures and in vitro tissue cultures are encompassed herein, of cells or tissues comprising a recombinant chromosome 3 as described. Preferably the cells or tissues can be regenerated into a whole cucumber plant, i.e. the cells are regenerable cells and the tissues comprise regenerable cells. Thus, also vegetative propagations of the plants according to the invention are an embodiment herein. Thus, a vegetatively propagated cultivated cucumber plant is provided which comprises a recombinant chromosome 3 as described herein. In a different aspect non-propagating cells comprising QTL3. 1 (or a variant thereof), is encompassed herein, as are tissues comprising such cells.

[0331] In a specific aspect a cucumber fruit harvested from a plant according to the invention is provided. Marketable cucumber fruits, especially for the fresh market (slicing), are generally graded according to fruit size and quality characteristics after harvest. See e.g. the United States Standards for Grades of Cucumbers, US Department of Agriculture, Effective March 1, 1985 and reprinted January 1997. Herein different grades of cucumbers are distinguished. Thus, in one aspect harvested fruits are provided of U.S. Fancy grade, U.S. Extra No. 1 grade, U.S. No. 1 grade, U.S. No. 1 Small grade, U.S. No. 1 Large grade, U.S. No. 2 grade. Also containers or packages comprising or consisting of harvested cucumber fruits are provided. Again, the cells of the fruits are distinguishable from other cucumber fruits by the presence of QTL3.1 (or a variant thereof) (as determinable in one or more of the molecular marker assays).

[0332] In another aspect the cucumber is a long cucumber type or a short cucumber type or a slicer cucumber type and fruits harvested and optionally processed (e.g. sliced or diced) are provided.

[0333] In another aspect the cucumber is a pickling type and fruits harvested and optionally pickled are provided.

[0334] The invention also provides for a food or feed product comprising or consisting of a plant part described herein preferably a cucumber fruit or part thereof and / or an extract from a plant part described herein. The food or feed product may be fresh or processed, e.g., pickled, canned, steamed, boiled, fried, blanched and / or frozen, etc. For example, containers such as cans, boxes, crates, bags, cartons, Modified Atmosphere Packaging, films (e.g. biodegradable films), etc. comprising plant parts such as fruits or fruit parts (fresh and / or processed) described herein are also provided herein.

[0335] Also encompassed herein is a method for producing a cultivated cucumber plant comprising an introgression fragments on chromosome 3, wherein said introgression fragment comprises QTL3.1 (or a variant thereof), comprising: a) providing a first cultivated cucumber plant, preferably lacking QTL3. 1, b) providing a second cultivated cucumber plant selected from plants grown from seeds deposited under accession number NCIMB44295 or progeny thereof or providing a second cultivated or wild cucumber plant comprising QTL3. 1 (or a variant therof) and comprising the donor SNP haplotype or genotype for at least 5, 6, 7, 8, 9 or all 10 SNP markers linked to the QTL, c) crossing said plant of a) with said plant of b), d) collecting F 1 seeds from said cross and optionally selfing said F 1 plants one or more times to produce an F2 or F3 or further selfing population, e) optionally backcrossing the F 1 plant or an F2 or F3 or further selfing plant to the plant of a) to produce a backcross population, f) optionally selfing the backcross population one or more times, g) identifying a Fl, F2, F3, further selfing or backcross plant which comprises the donor SNP haplotype or genotype for at least 5, 6, 7, 8, 9 or all 10 SNP markers linked to the QTL. The donor SNP haplotype or genotype is provided e.g. in Table 5 and further above.

[0336] In a further aspect a method of producing F 1 hybrid plants is provided comprising: a) providing a first inbred cucumber plant comprising an introgression fragment comprising QTL3.1, wherein said introgression fragment is the fragment as found in NCIMB44295, or a shorter fragment of that introgression fragment and / or wherein the QTL is the QTL as found in NCIMB44295, or wherein the QTL comprising a donor SNP haplotype or genotype for at least 5, 6, 7, 8, 9 or all 10 SNP markers linked to the QTL, b) providing a second inbred cucumber plant (optionally comprising an introgression fragment comprising QTL3. 1, wherein said introgression fragment is the fragment as found in NCIMB44295, or a shorter fragment of that introgression fragment and / or wherein the QTL is the QTL as found in NCIMB44295, or wherein the QTL comprising a donor SNP haplotype or genotype for at least 5, 6, 7, 8, 9 or all 10 SNP markers linked to the QTL), c) crossing said plant of a) with said plant of b), d) collecting Fl hybrid seeds from said cross.

[0337] Also provided is the use of plants grown from seeds deposited under accession number NCIMB 44295, or progeny thereof, for generating a cultivated cucumber plant which produces fruits comprising a mint-green fruit flesh, wherein said mint-green fruit flesh color is conferred by an introgression fragment obtained from chromosome 3 of said plants or progeny.

[0338] Also, the molecular marker sequences (and isolated nucleic acid molecules comprising the sequence) disclosed herein and their use in detecting and / or generating cucumber plants comprising said QTLs described herein are encompassed herein.

[0339] Further a method of growing a plant comprising QTL3. 1 (or a variant thereof) in the field, or in glasshouses or tunnels, whereby the fruits comprise a mint-green fruit flesh color.

[0340] In a preferred aspect, QTL1. 1 (or a variant), which confers an increase in at least glucose and fructose of the cucumber fruits, is combined in a single plant with QTL3.1, which confers mint-green fruit flesh. Optionally also the peel is mint-green, e.g. yellow-green or green-yellow. Also, QTL 1.1 is in one aspect combined with QTL4. 1, e.g. in a genetic background comprising the w / w gene in its genome.

[0341] Thus, all aspects described herein for the individual QTLs i.e. for plants and plant parts comprising these and for methods of detecting or selecting or using these, may herein be combined and are encompassed herein for the combination of e.g. QTL1.1 (or a variant thereof) and QTL3.1 (or a variant thereof) or any of the other combinations encompassed herein for the four QTLs.

[0342] Overcoming negative effects of QTLL 1 on fruit peel / skin color

[0343] It was found that QTL1. 1 had in one aspect a negative effect on fruit peel color when the QTL was present in a genetic background that contained the white mutant (w allele) at the White-locus on chromosome 3 in homozygous form (genotype w / w). The mutant white allele (w) is also referred to herein as QTL3.2.

[0344] The negative effect on the fruit peel color did not affect the sugar content, i.e. the fruits contained elevated glucose and fructose levels due to QTL1. 1, but the fruits looked very pale, almost white in appearance.

[0345] The exterior fruit color was paler when QTL 1.1 was in homozygous form than when QTL 1.1 was in heterozygous form in the w / w background.

[0346] It was further found that a QTL on chromosome 4, referred herein to QTL4. 1, could restore a mint green peel color when added in homozygous form to QTL 1.1 (in a w / w background). The restoration of the mint green peel color was seen both when QTL4. 1 was added to plants that were heterozygous for QTL1. 1 and for plants that were homozygous for QTL1. 1, but the peel-color enhancement effect was much better when QTL1. 1 was in heterozygous form, see Table 11. Herein QTL4.1 was named ‘peel color enhancer’ or ‘exterior color enhancer’ or ‘peel color restorer’ or ‘exterior color restorer’ as its presence (especially in homozygous form) enhanced or conferred a mint-green color to the immature cucumber fruit peel, probably by counteracting the reduction in chlorophyll content and chloroplast content of the fruit peel caused by the w / w mutant allele (see e.g. Tang et al. 2018, Int. J. Sci. 19, 1493; doi: 10.3390 and Liu et al. 2016, Theor. Appl. Genetics 129: 1247- 1256, supra).

[0347] The peel color enhancement effect of QTL4. 1 in a w / w background was the highest in plants that lacked QTL 1. 1 (see Tables 10 to 13), but then the fruits of these plants did not have elevated sugars. Still, this shows that QTL4. 1 is useful in restoring mint-green peel color to the fruit peel when the w / w mutant allele is present. Therefore, seeds, plants and plant parts, especially fruits, which comprise the w / w mutant allele and QTL4. 1 in (preferably) homozygous form are one embodiment herein.

[0348] Further embodiments are seeds, plants and plant parts, especially fruits, which comprise the w / w mutant allele and QTL4.1 in (preferably) homozygous form and which further comprise QTL 1.1 in homozygous form or preferably in heterozygous form.

[0349] Yet further embodiments provided herein are seeds, plants and plant parts, especially fruits, which comprise the w / w mutant allele (QTL3.2 in homozygous form) and QTL4. 1 in homozygous form, which further comprise QTL1.1 in homozygous form, or preferably in heterozygous form, and which further comprise QTL3.1 (conferring mint-green fruit flesh color) in homozygous form. All four QTLs are present in the seeds deposited under accession number NCIMB 44295 and are obtainable from such seeds.

[0350] In one aspect a cultivated Cucumis sativus var. sativus plant (or seed or plant part) is provided comprising:

[0351] A) the mutant w-allele of the White peel locus in homozygous form, wherein the cucumber genome comprises e.g. SEQ ID NO: 40 (and / or SEQ ID NO: 41) on chromosome 3 in homozygous form;

[0352] B) an introgression fragment on chromosome 4 from a wild cucumber donor, preferably in homozygous form, wherein said introgression fragments comprises a Quantitative Trait Locus (QTL) referred to as QTL4. 1 conferring an enhanced mint-green fruit peel color, wherein QTL4. 1 is located on chromosome 4 between the Single Nucleotide Polymorphism (SNP) markers SNP_25 at nucleotide 102 of SEQ ID NO: 25 and SNP_33 at nucleotide 102 of SEQ ID NO: 33, and wherein said introgression fragment on chromosome 4 comprising QTL4. 1 comprises a SNP haplotype of at least 2, 3, 4, 5, 6, 7, 8 or 9 markers selected from the group consisting of: a) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_25 at nucleotide 102 of SEQ ID NO: 25 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 25, such as a Thymine for SNP_25 at nucleotide 102 of SEQ ID NO: 38; b) a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_26 at nucleotide 102 of SEQ ID NO: 26 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 26; c) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_27 at nucleotide 102 of SEQ ID NO: T1 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: Tl\ d) a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_28 at nucleotide 102 of SEQ ID NO: 28 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 28; e) an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_29 at nucleotide 102 of SEQ ID NO: 29 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 29, such as an Adenine for SNP_29 at nucleotide 102 of SEQ ID NO: 39; f) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_30 at nucleotide 102 of SEQ ID NO: 30 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 30; g) an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_31 at nucleotide 102 of SEQ ID NO: 31 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 31; h) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_32 at nucleotide 102 of SEQ ID NO: 32 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 32; i) a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_33 at nucleotide 102 of SEQ ID NO: 33 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 33.

[0353] In one aspect the above plant (or seed or plant part) further comprises:

[0354] C) an introgression fragment on chromosome 1 from a wild cucumber donor, wherein said introgression fragments comprises a Quantitative Trait Locus (QTL) referred to as QTL1. 1 conferring an increase in glucose and fructose of the cucumber fruit, wherein QTL1. 1 is located on chromosome 1 between the Single Nucleotide Polymorphism marker SNP_01 at nucleotide 102 of SEQ ID NO: 1 and SNP_14 at nucleoti...

Claims

CLAIMS1. A cultivated Cucumis sativus var. sativus plant comprising an introgression fragment on chromosome1 from a wild cucumber donor, wherein said introgression fragment comprises a Quantitative Trait Locus (QTL) referred to as QTL1.1 conferring an increase in glucose and fructose of the cucumber fruit compared to a plant lacking QTL1. 1, wherein QTL1. 1 is located on chromosome 1 between the Single Nucleotide Polymorphism (SNP) markers SNP_01 at nucleotide 102 of SEQ ID NO: 1 and SNP_14 at nucleotide 102 of SEQ ID NO: 14, and wherein said introgression fragment on chromosome 1 comprising QTL1. 1 comprises a SNP haplotype of at least 5, preferably at least 9, 10, 11, 12 or 13 markers selected from the group consisting of: a) a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_01 at nucleotide 102 of SEQ ID NO: 1 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 1; b) an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_02 at nucleotide 102 of SEQ ID NO: 2 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 2; c) a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_03 at nucleotide 102 of SEQ ID NO: 3 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 3; d) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_04 at nucleotide 102 of SEQ ID NO: 4 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 4; e) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_05 at nucleotide 102 of SEQ ID NO: 5 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 5; f) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_06 at nucleotide 102 of SEQ ID NO: 6 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 6;g) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_07 at nucleotide 102 of SEQ ID NO: 7 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 7; h) an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_08 at nucleotide 102 of SEQ ID NO: 8 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 8; i) a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_09 at nucleotide 102 of SEQ ID NO: 9 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 9; j) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_10 at nucleotide 102 of SEQ ID NO: 10 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 10; k) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_11 at nucleotide 102 of SEQ ID NO: 11 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 11, such as a Guanine for SNP l 1 at nucleotide 102 of SEQ ID NO:

34. l) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_12 at nucleotide 102 of SEQ ID NO: 12 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 12; m) an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_13 at nucleotide 102 of SEQ ID NO: 13 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 13; n) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_14 at nucleotide 102 of SEQ ID NO: 14 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 14.

2. The plant according to claim 1, wherein the plant further comprises an introgression fragment on chromosome 3 from a wild cucumber donor, wherein said introgression fragment comprises a Quantitative Trait Locus (QTL) referred to as QTL3. 1 conferring a mint-green fruit flesh color having an RHS color chart rating of Yellow-Green Group 145A and / or 145B, wherein QTL3.1 is located on chromosome 3 between the SNP markers SNP_15 at nucleotide 102 of SEQ ID NO: 15 and SNP_24 at nucleotide 102 of SEQ ID NO: 24, and- 167 - wherein said introgression fragment on chromosome 3 comprising QTL3.1 comprises a SNP haplotype of at least 5, preferably at least 6, 7, 8, 9 or 10 markers selected from the group consisting of: an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_15 at nucleotide 102 of SEQ ID NO: 15 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 15, such as an Adenine for SNP_15 at nucleotide 102 of SEQ ID NO:

35. a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_16 at nucleotide 102 of SEQ ID NO: 16 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 16; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_17 at nucleotide 102 of SEQ ID NO: 17 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 17, such as a Guanine for SNP 17 at nucleotide 102 of SEQ ID NO:

36. a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_18 at nucleotide 102 of SEQ ID NO: 18 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 18; an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_19 at nucleotide 102 of SEQ ID NO: 19 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 19; a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_20 at nucleotide 102 of SEQ ID NO: 20 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 20, such as a Cytosine for SNP_20 at nucleotide 101 of SEQ ID NO:

37. a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_21 at nucleotide 102 of SEQ ID NO: 21 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 21; a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_22 at nucleotide 102 of SEQ ID NO: 22 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 22; a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_23 at nucleotide 102 of SEQ ID NO: 23 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 23;- 168 - a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_24 at nucleotide 102 of SEQ ID NO: 24 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 24.

3. The plant according to claim 1, wherein the introgression fragment comprising QTL1.1 is in homozygous form.

4. The plant according to claim 2 or 3, wherein the introgression fragment comprising QTL3.1 is in homozygous form.

5. The plant according to any one of the preceding claims, wherein the introgression fragment comprising QTL1. 1 and the introgression fragment comprising QTL3. 1 is obtainable from seeds deposited under accession number NCIMB 44295 or from progeny thereof.

6. The plant according to any one of the preceding claims, wherein the QTL1.1 is the QTL present in seeds deposited under accession number NCIMB 44295.

7. The plant according to any one of the preceding claims, further comprises a mutant w-allele at the White-peel locus on chromosome 3 in homozygous form and comprising an introgression fragment on chromosome 4 from a wild cucumber donor, wherein said introgression fragment comprises a Quantitative Trait Locus (QTL) referred to as QTL4. 1 conferring a mint-green fruit peel color, wherein QTL4. 1 is located on chromosome 4 between the SNP markers SNP_25 at nucleotide 102 of SEQ ID NO: 25 and SNP_33 at nucleotide 102 of SEQ ID NO: 33, and wherein said introgression fragment on chromosome 4 comprising QTL4.1 comprises a SNP haplotype of at least 5, preferably at least 6, 7, 8 or 9 markers selected from the group consisting of:- a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_25 at nucleotide 102 of SEQ ID NO: 25 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 25, such as a Thymine for SNP_25 at nucleotide 102 of SEQ ID NO: 38;- a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_26 at nucleotide 102 of SEQ ID NO: 26 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 26;- a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_27 at nucleotide 102 of SEQ ID NO: T1 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 27;- 169 -- a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_28 at nucleotide 102 of SEQ ID NO: 28 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 28;- an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_29 at nucleotide 102 of SEQ ID NO: 29 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 29, such as an Adenine for SNP_29 at nucleotide 102 of SEQ ID NO: 39;- a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_30 at nucleotide 102 of SEQ ID NO: 30 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 30;- an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_31 at nucleotide 102 of SEQ ID NO: 31 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 31;- a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_32 at nucleotide 102 of SEQ ID NO: 32 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 32;- a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_33 at nucleotide 102 of SEQ ID NO: 33 or at the equivalent position in a sequence comprising at least 95% sequence identity to SEQ ID NO: 33.

8. The plant according to claim 7, wherein the mutant w-allele comprises SEQ ID NO: 40 at the Whitepeel locus on chromosome 3.

9. Seeds from which a plant according to any one of the preceding claims can be grown.

10. A cucumber fruit harvested from a plant according to any one of claims 1 to 8.

11. A plant cell, tissue or plant part of a plant according to any one of claims 1 to 8.

12. A method for detecting the presence of the introgression fragment comprising QTL1.1 or a variant thereof in a cucumber plant or plant part, said method comprises analyzing the genomic DNA of the plant or plant part using a molecular marker assay which detects at least 3, 4, 5, 6, 7, 8, 9, 10, 11, 12, 13 or all 14 of the markers selected from the group consisting of:- 170 - a) a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_01 at nucleotide 102 of SEQ ID NO: 1 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 1; b) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_02 at nucleotide 102 of SEQ ID NO: 2 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 2; c) a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_03 at nucleotide 102 of SEQ ID NO: 3 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 3; d) a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_04 at nucleotide 102 of SEQ ID NO: 4 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 4; e) an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_05 at nucleotide 102 of SEQ ID NO: 5 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 5; f) an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_06 at nucleotide 102 of SEQ ID NO: 6 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 6; g) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_07 at nucleotide 102 of SEQ ID NO: 7 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 7; h) an Adenine (A) nucleotide for the Single Nucleotide Polymorphism marker SNP_08 at nucleotide 102 of SEQ ID NO: 8 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 8; i) a Cytosine (C) nucleotide for the Single Nucleotide Polymorphism marker SNP_09 at nucleotide 102 of SEQ ID NO: 9 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 9;j) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_10 at nucleotide 102 of SEQ ID NO: 10 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 10; k) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_11 at nucleotide 102 of SEQ ID NO: 11 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 11; l) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_12 at nucleotide 102 of SEQ ID NO: 12 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 12; m) a Thymine (T) nucleotide for the Single Nucleotide Polymorphism marker SNP_13 at nucleotide 102 of SEQ ID NO: 13 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 13; n) a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_14 at nucleotide 102 of SEQ ID NO: 14 or at the equivalent position in a sequence comprising at least 97% sequence identity to SEQ ID NO: 14.

13. The method according to claim 12, wherein said molecular marker assay is a SNP genotyping assay.

14. The method according to claim 12 or 13 further comprising selecting a plant or plant part comprising 9, 10, 11, 12, 13 or all 14 nucleotides of SNP_01 to SNP_14.

15. The method according to any one of claims 12 to 14, wherein the plant or plant part comprises a Guanine (G) nucleotide for the Single Nucleotide Polymorphism marker SNP_11 at nucleotide 102 of SEQ ID NO: 34.

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