Tomato plants with increased type iv trichome density and glandular trichomes producing an altered ACYL sugar composition

Abstract

The present invention relates to polynucleotide comprising a lncRNA gene according to SEQ ID No: 2, or according to a sequence having at least 50% sequence identity to SEQ ID No: 2, and an AP2c gene according to SEQ ID No: 3, or according to a sequence having at least 50% sequence identity to SEQ ID No: 3, which polynucleotide confers an increased density of type IV trichomes. The invention further relates to a plant comprising said polynucleotide and a SlASAT4 gene according to SEQ ID No. 21, or comprising a homologous sequence having at least 99.6% sequence identity to SEQ ID No. 21 wherein the presence of the polynucleotide leads to increased density of type IV trichomes and wherein the presence of said SlASAT4 gene leads to an increased ratio of S4:17 / S3:15 acyl sugars produced in glandular trichomes present on said plant when compared to a control plant lacking the polynucleotide and the SlASAT4 gene. Such a plant also was found to be resistant to Bemisia tabaci. The invention also discloses markers for the identification of the polynucleotide and the SlASAT4 gene and their use for the identification and development of Solanum lycopersicum plants. Also methods for identifying, producing and selecting of plants and seed are disclosed.

Description

[0001] TOMATO PLANTS WITH INCREASED TYPE IV TRICHOME DENSITY AND GLANDULAR TRICHOMES PRODUCING AN ALTERED ACYL SUGAR COMPOSITION

[0002] FIELD OF THE INVENTION

[0003] The present invention relates to the field of plant biotechnology. Specifically, the present invention relates to a combination of genetic elements which combination confers resistance towards Bemisia tabaci in Solanaceae plants and to a plant comprising such combination of genetic elements, as well as to methods for identifying and selecting such a plant, and to plant parts, progeny, seed and fruit of such a plant. The invention further relates to methods and markers for producing such a plant, and to methods and genetic markers for selection of such a plant. The invention also relates to a polynucleotide, and to the use of said combination of genetic elements to confer resistance towards Bemisia tabaci. The invention is also directed to a marker for identification of said combination of genetic elements in a plant, and to use of said marker.

[0004] BACKGROUND OF THE INVENTION

[0005] Commercial vegetable production is affected by many conditions that influence the growth and development of the crop. The choice of the grower for a certain variety is a determining factor, and genetics of the selected variety forms the basis for the result that can be achieved. In addition, there are many external factors that influence the outcome, for example, the disease pressure. Growing conditions like climate, soil, and the use of inputs like fertilizer play a major role. There are various ways of cultivating other crops, among which, the most common are: open field, greenhouse and shade house production. The presence of pests and diseases also affects the total yield that can be reached.

[0006] Plants of the species Solarium lycopersicum (tomato) belong to the nightshade family, also known as Solanaceae . Within this family it is nowadays grouped in the genus Solanum. which does not only harbor tomato, but also the important food crops potato, (sweet) pepper and eggplant. It is a perennial, herbaceous, flowering plant species which is native to South America.

[0007] Other species that are related to tomato within the Solanum genus are for example Solanum galapagense, Solanum pimpinellifolium, Solanum chilense, Solanum peruvianum and Solanum habrochaites . Although it is known that crossing can be considerably difficult, these species are used to obtain traits that are valuable in growing tomato plants and for fruits obtained from such plants. In the recent history, advancement in tomato breeding has led to tomato varieties having, for example higher yield, higher disease resistance and increased fruit shelflife. More specifically, resistance to insects is an important asset for tomato plants as insects can greatly damage the plant by sucking plant sap from essential tissues, causing damage like leaf stippling, leaf bleaching, leaf curling and growth distortion and leaving sticky honeydew that may serve as substrate for mold growth. On the other hand, insects often serve as vector for viruses. There is thus a need for a reliable resistance towards Bemisia tabaci also known as whitefly. This resistance is preferably active at all plant stages and preferably the resistance is trichome mediated.

[0008] The surface of the various plant parts of tomato (Solanum lycopersicum) is covered with trichomes, both non-glandular and glandular. Non-glandular trichomes are usually regarded as ‘hairs’ and do not produce, store, or secrete specific biochemical compounds. A variety of biochemical compounds in tomato, however, are produced in glandular trichomes. A glandular trichome typically consists of a stalk, made up of one or more cells, and one or more glandular cells at the tip of the stalk that form the glandular head. Four different types of glandular trichomes are identified in tomato and related Solanum species, namely types I, IV, VI, and VII. These types differ in size and length of the stalks, and in number of secretory cells that form the glandular head (McDowell et al., Plant Physiology Vol. 155, 524-539 (2011)). The unicellular glands, comprising one glandular cell that forms the glandular head, are classified as secreting glands, while multicellular glands are classified as storage glands. Luckwill (The Genus Lycopersicon: An historical, biological, and taxonomic survey of the wild and cultivated tomatoes, Aberdeen University Press, 1943) provides a clear overview and description of all trichome types and their phenotypes.

[0009] It is the object of the present invention to obtain plants which show an increased density of type IV trichomes and which are resistant to Bemisia tabaci.

[0010] In the research leading to the present invention, it was found that the presence of a polynucleotide on chromosome 2 of the tomato genome confers increased density of type IV trichomes. It was furthermore found that this increased density in combination with the presence of glandular trichomes producing an increased ratio of S4: 17 / S3: 15 acyl sugars has a remarkable effect on resistance of tomato and related Solanum plants to whitefly (Bemisia tabaci). Plants having an increased density of type IV trichomes on their leaves, petioles and / or stem as a result of the presence in their genome of the polynucleotide in combination with an increased ratio of S4: 17 / S3: 15 acyl sugars produced in glandular trichomes as a result of the presence of the SIASAT4 gene on chromosome 1 in their genome show resistance to infestation with Bemisia tabaci when compared to control plants not comprising the polynucleotide and the SIASAT4 gene.

[0011] This research further revealed that said polynucleotide comprises two genes: the first one is a gene that is transcribed as a long non-coding RNA (IncRNA), and the second gene is the AP2c gene. It is essential that the polynucleotide comprises at least those two genes in order to confer the phenotype of the invention as is illustrated in Example 1. The trait of the invention is transmitted genetically in a monogenic, intermediate fashion. “Intermediate inheritance” is also known as incompletely dominant inheritance, in which the phenotype of the F 1 progeny is intermediate between the two (more contrasting, extreme) phenotypes of both parents.

[0012] Thus the present invention provides a polynucleotide, herein referred to as the “polynucleotide of the invention”, wherein said polynucleotide comprises at least a IncRNA gene according to SEQ ID No: 2, or according to a sequence having at least 50% sequence identity to SEQ ID No: 2 and an AP2c gene according to SEQ ID No: 3, or according to a sequence having at least 50% sequence identity to SEQ ID No: 3, which polynucleotide confers an increased density of type IV trichomes, when expressed in a plant

[0013] In a preferred embodiment, the polynucleotide of the invention is the polynucleotide as present in the genome of a Solanum lycopersicum plant, representative seed of which was deposited with the NCIMB under deposit number NCIMB 44403.

[0014] In another preferred embodiment, the polynucleotide of the invention can also be defined as the genomic fragment that is located between and comprises the markers having SEQ ID no. 6 and SEQ ID no. 20. This genomic fragment is present in the genome of a Solanum lycopersicum plant, representative seed of which was deposited with the NCIMB under deposit number NCIMB 44403.

[0015] In an even more preferred embodiment, said genomic fragment is located between and comprises the markers having SEQ ID no. 8 and SEQ ID no. 20. This genomic fragment is present in the genome of a Solanum lycopersicum plant, representative seed of which was deposited with the NCIMB under deposit number NCIMB 44403.

[0016] This invention also relates to a polynucleotide which comprises a IncRNA gene sequence having at least 60% sequence identity, at least 70% sequence identity, at least 80% sequence identity, at least 90% sequence identity, at least 95% sequence identity, at least 96% sequence identity, at least 97% sequence identity, at least 98% sequence identity, or at least 99% sequence identity to SEQ ID No: 2.

[0017] This invention also relates to a polynucleotide which comprises a AP2c gene sequence having at least 60% sequence identity, at least 70% sequence identity, at least 80% sequence identity, at least 90% sequence identity, at least 95% sequence identity, at least 96% sequence identity, at least 97% sequence identity, at least 98% sequence identity, or at least 99% sequence identity to SEQ ID No: 3.

[0018] The term “at least 50% sequence identity” as used henceforth in this application is always intended to also encompass at least 60% sequence identity, at least 70% sequence identity, at least 80% sequence identity, at least 90% sequence identity, at least 95% sequence identity, at least 96% sequence identity, at least 97% sequence identity, at least 98% sequence identity, or at least 99% sequence identity.

[0019] This invention also relates to a plant comprising a polynucleotide comprising a IncRNA gene according to SEQ ID No: 2, or according to a sequence having at least 50% sequence identity to SEQ ID No: 2 and an AP2c gene according to SEQ ID No: 3, or according to a sequence having at least 50% sequence identity to SEQ ID No: 3, which confers an increased density of type IV trichomes, when expressed in a plant, when compared to a control plant lacking the polynucleotide. The phenotype observed on plants of the invention, i.e. the increased density of type IV trichomes, is also referred to as the “first trait of the invention”.

[0020] In a preferred embodiment, a plant of the invention comprises a IncRNA gene according to SEQ ID No: 2, or according to a sequence having at least 50% sequence identity to SEQ ID No: 2 and an AP2c gene according to SEQ ID No: 3, or according to a sequence having at least 50% sequence identity to SEQ ID No: 3, which genes confer an increased density of type IV trichomes, when expressed in a plant, when compared to a control plant lacking said genes.

[0021] In the research leading to the invention, the inventors have also identified the role of the SIASAT4 gene in combination with the polynucleotide of the invention. Despite the fact that it was already known that the SIASAT4 gene according to SED ID No. 21 was involved in acyl sucrose acetylation (ref. https: / / www.pnas.org / doi / epdf / 10.1073 / pnas.1207906109). the inventors have verified that only plants comprising both a specific allele of the SIASAT4 gene and the polynucleotide of the invention show increased resistance to Bemisia tabaci.

[0022] The inventors found that a plant, which comprises the polynucleotide of the invention conferring the increased density of type IV trichomes phenotype, and a SIASAT4 gene which comprises SEQ ID No. 21, or comprises a homologous sequence having at least 99.6% sequence identity to SEQ ID No. 21, which SIASAT4 gene confers an increased ratio of S4: 17 / S3: 15 acyl sugars produced in glandular trichomes present on said plant is resistant to Bemisia tabaci. This SIASAT4 gene according to SEQ ID No. 21 or having a sequence having at least 99.6% sequence identity to SEQ ID No. 21 is herein defined as the “SASAT4 gene of the invention” and is also referred to as the “favorable SIASAT4 allele”. The definition "SIASA T4 gene of the invention” and the term “favorable SIASAT4 allele” are used interchangeably herein. The phenotype observed on plants comprising the SIASAT4 gene of the invention, i.e. glandular trichomes that produce an increased ratio of S4: 17 / S3: 15 acyl sugars, is also referred to as the “second trait of the invention”. A plant comprising both the polynucleotide of the invention and the SIASAT4 gene of the invention is referred to as a “plant of the invention”.

[0023] In another embodiment, the invention also relates to a plant, which comprises the polynucleotide of the invention and a SgASAT4 gene which comprises SEQ ID No. 25, or comprises a homologous sequence having at least 99.6% sequence identity to SEQ ID No. 25, which SgASAT4 gene confers an increased ratio of S4: 17 / S3: 15 acyl sugars produced in glandular trichomes present on said plant. This SgASAT4 gene is also referred to as the "SgASA T4 gene of the invention”. The phenotype observed on plants comprising the SgASAT4 gene of the invention, i.e. glandular trichomes that produce an increased ratio of S4: 17 / S3: 15 acyl sugars, is also part of the invention. A plant comprising both the polynucleotide of the invention and the SgASAT4 gene of the invention is also part of the invention. In a preferred embodiment, the SIASAT4 gene of the invention and the SgASAT4 gene of the invention can be used interchangeably in plants of the invention.

[0024] An unfavorable allele of the SIASAT4 gene is also disclosed herein. The inventors have determined that if a plant comprises the polynucleotide of the invention and this version of the S1ASAT4 gene, said version of the SIASAT4 gene, further referred to as “unfavorable allele” or “unfavorable SIASAT4 allele”, does not confer an increased ratio of S4: 17 / S3: 15 acyl sugars produced in glandular trichomes present on said plant. The unfavorable SIASAT4 allele comprises SEQ ID No. 23.

[0025] There are 6 SNPs (single nucleotides polymorphisms) identified between the SIASAT4 gene of the invention (SEQ ID No. 21) and the unfavorable SIASAT4 allele (SEQ ID No. 23). This concerns the following positions on the coding sequence of SEQ ID No. 21 : position 67, 94, 447, 887, 944 and 1317. On position 67, the favorable SIASAT4 allele has an A, whereas the unfavorable SIASAT4 allele has a C. On position 94, the favorable SIASAT4 allele has a G, whereas the unfavorable SIASAT4 allele has a C. This SNP is also the SNP indicated in the marker with SEQ ID No. 27 as disclosed herein. On position 447, the favorable SIASAT4 allele has an A, whereas the unfavorable SIASAT4 allele has a C. On position 887, the favorable SIASAT4 allele has a C, whereas the unfavorable SIASAT4 allele has a T. On position 944, the favorable SIASAT4 allele has a T, whereas the unfavorable SIASAT4 allele has a C. On position 1317, the favorable SIASAT4 allele has an A, whereas the unfavorable SIASAT4 allele has a T.

[0026] A plant comprising the polynucleotide of the invention and comprising the SIASAT4 gene of the invention also is resistant to Bemisia tabaci. The inventors have found that there is a direct correlation between plants showing the increased density of type IV trichomes in combination with the presence of glandular trichomes that produce an increased ratio of S4: 17 / S3: 15 acyl sugars and resistance to Bemisia tabaci when plants are subjected to infestation. The increased type IV trichome density means that the level of acyl sugars from type IV trichomes is increased per square millimeter. If this is combined with glandular trichomes producing an increased ratio of S4: 17 / S3: 15 acyl sugars that are present on said plant, this gives rise to resistance to Bemisia tabaci.

[0027] A plant of the invention comprises the polynucleotide of the invention and the

[0028] SIASAT4 gene of the invention homozygously or heterozygously, i.e. the polynucleotide of the invention and the SIASAT4 gene of the invention can be present on both chromosomes of a chromosome pair in the genome of a plant, or on only one chromosome of a chromosome pair. A plant of the invention comprises a plant of an inbred line, a hybrid (Fl) plant or variety, an open pollinated (OP) variety, a doubled haploid or a plant of a segregating population.

[0029] The plant of the invention is a plant from the Solanaceae family. This family notably comprises the genera Capsicum, Solatium. Nicotiana, Petunia. In a further aspect of the invention, the plant of the invention is a member of the Solanum genus, more preferably a plant of the Solanum lycopersicum species.

[0030] In a preferred embodiment, the plant of the invention is a Solanum lycopersicum plant, wherein the polynucleotide of the invention and the SIASAT4 gene of the invention are as present in the genome of a Solanum lycopersicum plant, representative seed of which was deposited with the NCIMB under deposit number NCIMB 44403.

[0031] The resistance to Bemisia tabaci is caused by the increased type IV trichome density phenotype of the invention in combination with glandular trichomes producing an increased ratio of S4: 17 / S3: 15 acyl sugars. The inventors have studied the inheritance phenotype of both the polynucleotide of the invention and the SIASAT4 gene of the invention. For both the polynucleotide and the SIASAT4 gene it was concluded that they have a semi-dominant mode of inheritance which leads to an intermediate inheritance of the resistance in a plant of the invention. As used herein, intermediate means that a higher level of resistance is found when a polynucleotide of the invention and the SIASAT4 gene of the invention are homozygously present. The heterozygous presence of a polynucleotide of the invention and the SIASAT4 gene of the invention, however, already confers improved resistance to Bemisia tabaci. The resistance to Bemisia tabaci of both homozygous and heterozygous plants makes the plants more suitable for cultivation under conditions where Bemisia tabaci is present. Therefore, both heterozygous and homozygous plants have an improved value in use for resistance to Bemisia tabaci. For heterozygous plants however, the improved resistance may only be observed under certain environmental conditions, e.g. under low or lower insect pressure. In addition, heterozygous plants can be used for development of homozygous plants through crossing and selection, which heterozygous plants also for that reason form a part of this invention. In a preferred embodiment, both the polynucleotide of the invention and the SLASAT4 gene of the invention are homozygously present in a plant.

[0032] The term “increased density of type IV trichomes” relates to an increase in the number of type IV trichomes on a certain surface. In general, observation of trichome phenotypes in tomato wild and cultivated species is described in a paper by Luckwill et al., titled “The genus Lycopersicon: an historical, biological, and taxonomic survey of the wild and cultivated tomatoes.” The University Press, Aberdeen, p 44 (1943). In order to examine the trichome phenotype in the context of the current invention, trichomes are observed generally on the abaxial side of the first fully developed leaf counted from the top of the plant (3rd leaf from the top). The scores as indicated herein are observed on leaves, petioles or the combination thereof. Type IV trichome density can also be observed on the main stem. For the trichome observations, a binocular microscope can be used.

[0033] In order to examine or phenotype the increased density of type IV trichomes, the inventors use five scoring scales that are depicted in Table 1.

[0034] Table 1: scoring scales for type IV trichome density on tomato leaves and petioles

[0035] In the context of the current invention the type IV trichome density of plants is observed in both young plants (until about 6 weeks old) as well as in adult plants.

[0036] It was determined that a plant comprising the polynucleotide of the invention shows “an increased density of type IV trichomes” if the phenotype shows at least one average scoring class higher compared to the phenotype of a near-isogenic or control plant not comprising the polynucleotide of the invention. In general, when a population of plants is examined for increased type IV trichome density, multiple plants and if possible, multiple replicates are tested and subsequently scored. This results in average scores for different genotypes, as is also shown in the examples provided below.

[0037] The term “increased ratio of S4: 17 / S3: 15 acyl sugars” relates to an increase in the acetylation of S3: 15 acyl sugars into S4: 17 acyl sugars. S3: 15 as defined herein is an acyl sucrose with three acyl groups of carbon chain lengths 5, 5, and 5 for a total of 15 carbons. S4: 17 as defined herein is an acyl sucrose with four acyl chains of length 2, 5, 5, and 5 for a total of 17 carbons. With “increased ratio” it is meant that a significantly higher amount of S4: 17 acyl sugars is observed in comparison with the amount observed for S3: 15 acyl sugars. The biochemical conversion of S3: 15 into S4: 17 depends on the SIASAT4 gene of the invention. When this enzymatic step works efficiently, plants accumulate more S4: 17 and thus are resistant to Bemisia tabaci. If the amount of S4: 17 and S3: 15 acyl sugars is the same, the ratio is 1. If the amount of S3: 15 acyl sugars produced is higher than the amount of S4: 17 acyl sugars produced, the ratio is smaller than 1. If the amount of S3 : 15 acyl sugars produced is smaller than the amount S4 : 17 acyl sugars produced, the ratio is larger than 1. The ratio observed in plants comprising both the polynucleotide of the invention and SIASAT4 gene of the invention is always larger than at least 1, but can also be larger than 2. The ratio observed in plants comprising both the polynucleotide of the invention and the unfavorable SIASAT4 allele is smaller than 1.

[0038] The term “resistance to Bemisia tabaci” and / or “resistant to Bemisia tabaci” relates to the capacity of a plant to withstand infestation of the plant by the insect. Whitefly resistance can typically be evaluated by two types of assessments, a free-choice (or preference) test and no-choice (non-preference) test. In a free-choice test, whiteflies are given the choice between two or more different hosts of which it is able to choose the most preferred host(s). In a no-choice test, only one host is accessible for the whitefly and whiteflies that cannot feed on it will be hampered in their growth or die. Reliable parameters for whitefly resistance assessments are very important. Parameters used to describe resistance are density and / or survival of a particular developmental stage of whitefly including adults, eggs or nymphs. Those parameters might measure similar or different resistance factors.

[0039] In order to examine or phenotype the resistance to Bemisia tabaci, the inventors use five scoring scales that are depicted in Table 2.

[0040] Table 2: scoring scales for infection of Bemisia tabaci on tomato leaves. According to the scoring scales, scale 1 and 2 are considered as being resistant to Bemisia tabaci. Scale 3 is considered to be intermediate resistant and scale 4 and 5 are considered to be susceptible to infestation with Bemisia tabaci. In general, when a population of plants is examined for resistance to Bemisia tabaci, multiple plants and if possible, multiple replicates are tested and subsequently scored. This results in average scores for different genotypes, as is also shown in the examples provided below. Therefore, in the context of the current invention an average score up to scale 3 based on the scoring scales of Table 2 is regarded as being resistant to Bemisia tabaci. An average score between scales 3 and 4 is regarded as being intermediate resistant to Bemisia tabaci. An average score between scales 4 and 5 is regarded as susceptible to Bemisia tabaci. In Figure 1, indicative pictures are provided of the description provided for the five classes as comprised in Table 2.

[0041] A "control plant" is intended to mean a plant that has the same genetic background as the plant of the present invention, wherein the control plant does not comprise the polynucleotide of the invention that confers increased density of type IV trichomes and does not comprise the SIASAT4 gene of the invention that confers glandular trichomes to produce an increased ratio of S4: 17 / S3: 15 acyl sugars. Not having the polynucleotide of the invention and the SIASAT4 gene of the invention is intended to mean that the polynucleotide and the SIASAT4 gene are not present in the genome of the control plant. When used in comparison with the plant of the invention, the control plant is grown for the same length of time and under the same conditions as the plant of the invention. Thus, a control plant may be a near-isogenic line, an inbred line or a hybrid, provided that they have the same genetic background as the plant of the present invention, except that it does not have the polynucleotide of the invention that confers an increased density of type IV trichomes and the SIASAT4 gene of the invention that causes glandular trichomes to produce an increased ratio of S4: 17 / S3: 15 acyl sugars.

[0042] The polynucleotide of the invention comprises a long non-coding RNA (IncRNA) gene. Long non-coding RNA molecules are a type of RNA, generally defined as transcripts of at least 200 nucleotides that are not translated into protein. They are functional molecules that are usually involved in transcriptional control or in post-transcriptional regulation of gene expression (for example in splicing and / or translation of RNA molecules), and they have been shown to be regulators of various agronomically important traits in plants, such as phosphate starvation response, flowering time and interaction with symbiotic organisms. For most annotated IncRNAs no biological function or mode of action has so far been identified.

[0043] When compared to messenger RNA (mRNA) molecules that are translated into protein sequences, IncRNAs are generally expressed at a lower abundance, and they exhibit a higher tissue specificity and developmental stage specificity. This invention further relates to a marker for the identification of the polynucleotide of the invention, wherein the marker is selected from the group of SEQ ID no. 6 to SEQ ID no. 20. One marker selected from the group of SEQ ID no. 6 to SEQ ID no. 20 can be used to identify the polynucleotide of the invention. Preferably, more markers selected from this group are used to identify the polynucleotide of the invention.

[0044] In this context, the invention relates to the marker having SEQ ID no. 6 which is positioned on the publicly available genome sequence for Solcinum lycopersicum based on the tomato cultivar ‘Heinz’ (release SL4.0) and comprises the modification of T to C corresponding to the physical position 52.122.217 on chromosome 2, for the identification of the polynucleotide of the invention.

[0045] The invention also relates to the marker having SEQ ID no. 7 which is positioned on the publicly available genome sequence for Solanum lycopersicum based on the tomato cultivar ‘Heinz’ (release SL4.0) and comprises the modification of A to C corresponding to the physical position 52.122.746 on chromosome 2, for the identification of the polynucleotide of the invention.

[0046] The invention also relates to the marker having SEQ ID no. 8 which is positioned on the publicly available genome sequence for Solanum lycopersicum based on the tomato cultivar ‘Heinz’ (release SL4.0) and comprises the modification of T to G corresponding to the physical position 52.123.098 on chromosome 2, for the identification of the polynucleotide of the invention.

[0047] The invention also relates to the marker having SEQ ID no. 9 which is positioned on the publicly available genome sequence for Solanum lycopersicum based on the tomato cultivar ‘Heinz’ (release SL4.0) and comprises the modification of T to C corresponding to the physical position 52.123.740 on chromosome 2, for the identification of the polynucleotide of the invention.

[0048] The invention also relates to the marker having SEQ ID no. 10 which is positioned on the publicly available genome sequence for Solanum lycopersicum based on the tomato cultivar ‘Heinz’ (release SL4.0) and comprises the modification of A to G corresponding to the physical position 52.124.586 on chromosome 2, for the identification of the polynucleotide of the invention.

[0049] The invention also relates to the marker having SEQ ID no. 11 which is positioned on the publicly available genome sequence for Solanum lycopersicum based on the tomato cultivar ‘Heinz’ (release SL4.0) and comprises the modification of C to A corresponding to the physical position 52.125.839 on chromosome 2, forthe identification of the polynucleotide of the invention.

[0050] The invention also relates to the marker having SEQ ID no. 12 which is positioned on the publicly available genome sequence for Solanum lycopersicum based on the tomato cultivar ‘Heinz’ (release SL4.0) and comprises the modification of G to T corresponding to the physical position 52.129.083 on chromosome 2, forthe identification of the polynucleotide of the invention.

[0051] The invention also relates to the marker having SEQ ID no. 13 which is positioned on the publicly available genome sequence for Solanum lycopersicum based on the tomato cultivar ‘Heinz’ (release SL4.0) and comprises the modification of T to C corresponding to the physical position 52.129.886 on chromosome 2, for the identification of the polynucleotide of the invention.

[0052] The invention also relates to the marker having SEQ ID no. 14 which is positioned on the publicly available genome sequence for Solcinum lycopersicum based on the tomato cultivar ‘Heinz’ (release SL4.0) and comprises the modification of C to T corresponding to the physical position 52.130.942 on chromosome 2, for the identification of the polynucleotide of the invention.

[0053] The invention also relates to the marker having SEQ ID no. 15 which is positioned on the publicly available genome sequence for Solanum lycopersicum based on the tomato cultivar ‘Heinz’ (release SL4.0) and comprises the modification of A to C corresponding to the physical position 52.130.988 on chromosome 2.

[0054] The invention also relates to the marker having SEQ ID no. 16 which is positioned on the publicly available genome sequence for Solanum lycopersicum based on the tomato cultivar ‘Heinz’ (release SL4.0) and comprises the modification of T to C corresponding to the physical position 52.131.559 on chromosome 2, for the identification of the polynucleotide of the invention.

[0055] The invention also relates to the marker having SEQ ID no. 17 which is positioned on the publicly available genome sequence for Solanum lycopersicum based on the tomato cultivar ‘Heinz’ (release SL4.0) and comprises a deletion of an A nucleotide corresponding to the physical position 52.132.443 on chromosome 2, for the identification of the polynucleotide of the invention.

[0056] The invention also relates to the marker having SEQ ID no. 18 which is positioned on the publicly available genome sequence for Solanum lycopersicum based on the tomato cultivar ‘Heinz’ (release SL4.0) and comprises the modification of A to G corresponding to the physical position 52.133.315 on chromosome 2, forthe identification of the polynucleotide of the invention.

[0057] The invention also relates to the marker having SEQ ID no. 19 which is positioned on the publicly available genome sequence for Solanum lycopersicum based on the tomato cultivar ‘Heinz’ (release SL4.0) and comprises the modification of C to A corresponding to the physical position 52.134.631 on chromosome 2, forthe identification of the polynucleotide of the invention.

[0058] The invention also relates to the marker having SEQ ID no. 20 which is positioned on the publicly available genome sequence for Solanum lycopersicum based on the tomato cultivar ‘Heinz’ (release SL4.0) and comprises the modification of C to A corresponding to the physical position 52.137.590 on chromosome 2, forthe identification of the polynucleotide of the invention.

[0059] In a preferred embodiment, the invention relates to the marker having SEQ ID no.

[0060] 9, 10 and or 11 forthe identification of the IncRNA gene according to SEQ ID No: 2.

[0061] In a preferred embodiment, the invention relates to the marker having SEQ ID no.

[0062] 19 for the identification of the A P2c gene according to SEQ ID No : 3. This invention further relates to a marker for the identification of the SIASAT4 gene of the invention, wherein the marker is indicated with SEQ ID no. 27. This marker can be used to identify the SIASAT4 gene of the invention.

[0063] In this context, the invention relates to the marker having SEQ ID no. 27 which is positioned on the publicly available genome sequence for Solcinum lycopersicum based on the tomato cultivar ‘Heinz’ (release SL4.0) and comprises the modification of C to G corresponding to the physical position 86.023.940 on chromosome 1, for the identification of the SIASAT4 gene of the invention. The presence of the G nucleotide is related to the SIASAT4 gene of the invention. If the C nucleotide is present in the SIASAT4 gene, this is regarded as a SIASAT4 gene having an unfavorable allele, meaning that such a SIASAT4 gene does not confer glandular trichomes to produce an increased ratio of S4: 17 / S3: 15 acyl sugars.

[0064] A marker can be defined as a reference sequence that comprises the modification(s) that can be detected using any suitable method known. The term “marker”, "genetic marker" or “DNA marker” refers to a feature of an individual’s genome (e.g. a nucleotide or a polynucleotide sequence that is present in an individual’s genome) that is associated with one or more loci of interest. In some embodiments, a genetic marker is polymorphic in a population of interest. Genetic markers include, for example, single nucleotide polymorphisms (SNPs), indels (i.e. insertions / deletions), simple sequence repeats (SSRs), restriction fragment length polymorphisms (RFLPs), random amplified polymorphic DNAs (RAPDs), cleaved amplified polymorphic sequence (CAPS) markers, Diversity Arrays Technology (DArT) markers, and amplified fragment length polymorphisms (AFLPs), among many other examples. Genetic markers can, for example, be used to locate genetic loci containing alleles on a chromosome that contribute to variability of phenotypic traits. The term “marker” or “genetic marker” can also refer to a polynucleotide sequence complementary to a genomic sequence, such as a sequence of a nucleic acid used as a probe. The term “marker” then refers to a physical entity that can be used in molecular biological techniques for detecting the mutation.

[0065] In the context of the present invention, a marker allows detection of the polynucleotide of the invention and / or the SIASAT4 gene of the invention, and the selection of plants that show an increased density of type IV trichomes trait and / or plants that comprise glandular trichomes that produce an increased ratio of S4: 17 / S3: 15 acyl sugars at any stage of their life cycle, even when the plants are not yet bearing fruits, e.g. during the seedling stage. Marker- assisted breeding and selection greatly increases the speed with which a trait can be introduced into different genetic backgrounds, and with which it can be commercialized.

[0066] Methods for detecting markers and specific alleles are abundantly known in the field. In general, these methods allow to distinguish between two different alleles of a marker, on a specific chromosome. Detection of a polymorphism can be achieved by electrophoretic techniques, but the widespread availability of DNA sequencing often makes it easier to simply sequence amplified products directly. Once the polymorphic sequence difference is known, rapid assays for the detection of a polymorphism can be designed for progeny testing, generally involving some version of PCR amplification of specific alleles.

[0067] In particular examples, PCR detection and quantification is carried out using two labeled Anorogenic oligonucleotide forward primers and an unlabeled common reverse primer, for example, KASP™ (KB iosciences).

[0068] The invention further relates to the use of a marker for the identification of the polynucleotide of the invention, wherein the marker is selected from the group of SEQ ID no. 6 to SEQ ID no. 20.

[0069] The invention further relates to the use of a marker for the identification of the SIASAT4 gene of the invention, wherein the marker is the marker having SEQ ID no. 17.

[0070] This invention also relates to the use of a marker for the identification of a Solcinum lycopersicum plant showing an increased density of type IV trichomes, wherein the marker is selected from the group of SEQ ID no. 6 to SEQ ID no. 20. These markers can also be used for the identification of a Solanum lycopersicum plant that is resistant to Bemisia tabaci in case the SIASAT4 gene of the invention is also present in said plant.

[0071] This invention also relates to the use of a marker for the identification of a Solanum lycopersicum plant comprising glandular trichomes that produce an increased ratio of S4: 17 / S3: 15 acyl sugars, wherein the marker is the marker having SEQ ID no. 17. This marker can also be used for the identification of a Solanum lycopersicum plant that is resistant to Bemisia tabaci in case the polynucleotide of the invention is also present in said plant.

[0072] The invention further relates to the use of a marker in the development of a Solanum lycopersicum plant showing an increased density of type IV trichomes, wherein the marker is selected from the group of SEQ ID no. 6 to SEQ ID no. 20. These markers can also be used for the development of Solanum lycopersicum plants that are resistant to Bemisia tabaci. During the steps of introgressing and backcrossing of the polynucleotide of the invention, markers can be used to check if the polynucleotide of the invention is still (homozygously) present in selected plant.

[0073] The invention further relates to the use of a marker in the development of a Solanum lycopersicum plant comprising glandular trichomes that produce an increased ratio of S4: 17 / S3: 15 acyl sugars, wherein the marker is the marker having SEQ ID no. 17. This marker can also be used for the development of Solanum lycopersicum plants that are resistant to Bemisia tabaci. During the steps of introgressing and backcrossing of the SIASAT4 gene of the invention, the marker can be used to check if the SIASAT4 gene of the invention is still (homozygously) present in selected plant. This invention also relates to a seed comprising a polynucleotide comprising a IncRNA gene according to SEQ ID No: 2, or according to a sequence having at least 50% sequence identity to SEQ ID No: 2 and an AP2c gene according to SEQ ID No: 3, or according to a sequence having at least 50% sequence identity to SEQ ID No: 3, wherein the plant grown from said seed shows an increased density of type IV trichomes when compared to a control plant lacking the polynucleotide, as a result of the presence of the polynucleotide. Such a seed may further comprise a SIASAT4 gene according to SEQ ID No. 21, or a homologous sequence having at least 99.6% sequence identity to SEQ ID No. 21, which SIASAT4 gene confers an increased ratio of S4: 17 / S3 : 15 acyl sugars produced in glandular trichomes present on a plant grown from said seed.

[0074] The current invention also relates to propagation material capable of developing into and / or being derived from a plant comprising a polynucleotide comprising a IncRNA gene according to SEQ ID No: 2, or according to a sequence having at least 50% sequence identity to SEQ ID No: 2 and an AP2c gene according to SEQ ID No: 3, or according to a sequence having at least 50% sequence identity to SEQ ID No: 3, which polynucleotide confers an increased density of type IV trichomes, when compared to a control plant lacking the polynucleotide. Such propagation material may further be capable of developing into and / or being derived from a plant that also comprises a SIASAT4 gene according to SEQ ID No. 21, or comprising a homologous sequence having at least 99.6% sequence identity to SEQ ID No. 21, which SIASAT4 gene confers an increased ratio of S4: 17 / S3: 15 acyl sugars produced in glandular trichomes present on a plant, when compared to a control plant lacking the SIASAT4 gene.

[0075] Preferably, said propagation material is suitable for sexual reproduction, and is in particular selected from a microspore, pollen, an ovary, an ovule, an embryo sac and an egg cell, or is suitable for vegetative reproduction, and is in particular selected from a cutting, a root, a stem cell, and a protoplast, or is suitable for tissue culture of regenerable cells or protoplasts, which regenerable cells or protoplasts are in particular selected from a leaf, pollen, an embryo, a cotyledon, a hypocotyl, a meristematic cell, a root, a root tip, an anther, a flower and a stem, and wherein the propagation material comprises a polynucleotide comprising a IncRNA gene according to SEQ ID No: 2, or according to a sequence having at least 50% sequence identity to SEQ ID No: 2 and an AP2c gene according to SEQ ID No: 3, or according to a sequence having at least 50% sequence identity to SEQ ID No: 3, which polynucleotide confers an increased density of type IV trichomes, when compared to a control plant lacking the polynucleotide. In an embodiment, such propagation material also comprises a SIASAT4 gene according to SEQ ID No. 21, or comprising a homologous sequence having at least 99.6% sequence identity to SEQ ID No. 21, which SIASAT4 gene confers an increased ratio of S4: 17 / S3: 15 acyl sugars produced in glandular trichomes present on a plant, when compared to a control plant lacking the SIASAT4 gene. In one embodiment, said propagation material comprises a IncRNA gene comprising SEQ ID No: 2 and an AP2c gene according to SEQ ID No: 3.

[0076] In one embodiment, said propagation material is derived from a Solanum lycopersicum plant, representative seed of which was deposited with the NCIMB under deposit number NCIMB 44403, or from a progeny plant thereof.

[0077] The invention also relates to progeny of a plant, a cell, a tissue, or a seed of the invention, which progeny comprises the polynucleotide of the invention as defined herein, the presence of which polynucleotide confers an increased density of type IV trichomes. In another embodiment, such a progeny also comprises the SIASAT4 gene of the invention as defined herein, the presence of which SIASAT4 gene confers glandular trichomes to produce an increased ratio of S4: 17 / S3: 15 acyl sugars. Such progeny can in itself be a plant, a cutting, a seed, a cell, or atissue.

[0078] As used herein, ‘progeny’ is intended to mean the first and all further descendants, such as an Fl, F2, or further generation, from a cross with a plant of the invention, wherein a cross comprises a cross with itself or a cross with another plant, and wherein a descendant that is determined to be progeny comprises the polynucleotide of the invention as defined herein that confers an increased density of type IV trichomes. The plant of the invention that is used in this cross is optionally a plant grown from seed of deposit NCIMB 44403, or from progeny seed thereof which is a direct or further descendant through crossing a plant grown from the deposited seed with itself or with another plant for one or more subsequent generations, wherein the progeny seed has retained the polynucleotide of the invention and the SIASAT4 gene of the invention.

[0079] Progeny also encompasses a Solanum lycopersicum plant that carries the polynucleotide of the invention and the SIASAT4 gene of the invention and shows the increased density of type IV trichomes and the increased ratio of S4: 17 / S3: 15 acyl sugars produced by glandular trichomes and is resistant to Bemisia tabaci. and is obtained from the plant, or progeny of a plant, of the invention by vegetative propagation or another form of multiplication.

[0080] The present invention further relates to a method for identifying a plant from the Solanum genus showing an increased density of type IV trichomes and comprising glandular trichomes producing an increased ratio of S4: 17 / S3: 15 acyl sugars, wherein the method comprises screening a population of plants from the Solanum genus for the presence of a polynucleotide comprising a IncRNA gene according to SEQ ID No: 2, or according to a sequence having at least 50 % sequence identity to SEQ ID No: 2 and an AP2c gene according to SEQ ID No: 3, or according to a sequence having at least 50% sequence identity to SEQ ID No: 3 and for the presence of a SIASAT4 gene according to SEQ ID No. 21, or comprising a homologous sequence having at least 99.6% sequence identity to SEQ ID No. 21 and identifying the plant of the Solanum genus that comprises said polynucleotide and said SIASAT4 gene as a plant of the Solanum genus that shows an increased density of type IV trichomes and comprising glandular trichomes that produce an increased ratio of S4: 17 / S3: 15 acyl sugars Optionally, said method may also comprise the step of phenotypically screening for the presence of the increased density of type IV trichomes and the presence of glandular trichomes that produce an increased ratio of S4: 17 / S3: 15 acyl sugars. Furthermore, said method may also comprise the step of selecting the plant of the Solanum genus that shows the increased density of type IV trichomes and that comprises glandular trichomes that produce an increased ratio of S4: 17 / S3: 15 acyl sugars and the resistance to Bemisia tabaci.

[0081] The present invention also relates to a method for producing a plant of the Solanum genus showing an increased density of type IV trichomes and comprising glandular trichomes producing an increased ratio of S4: 17 / S3: 15 acyl sugars, comprising the step of introducing a polynucleotide comprising a IncRNA gene according to SEQ ID No: 2, or according to a sequence having at least 50 % sequence identity to SEQ ID No: 2 and an AP2c gene according to SEQ ID No: 3, or according to a sequence having at least 50% sequence identity to SEQ ID No: 3 and introducing a SIASAT4 gene according to SEQ ID No. 21, or comprising a homologous sequence having at least 99.6% sequence identity to SEQ ID No. 21, wherein the polynucleotide confers an increased density of type IV trichomes and wherein the SIASAT4 gene confers an increased ratio of S4: 17 / S3 : 15 acyl sugars produced in glandular trichomes when compared to a control plant of the Solanum genus lacking the polynucleotide and the SIASAT4 gene. In one embodiment, in the method for producing a plant of the Solanum genus, the polynucleotide and the SIASAT4 gene are introduced by cisgenesis. In one embodiment, the plant of the Solanum genus is a Solanum lycopersicum plant. In one embodiment, a plant showing said increased density of type IV trichomes and comprising glandular trichomes producing an increased ratio of S4: 17 / S3: 15 acyl sugars is further characterized by resistance to Bemisia tabaci. This resistance is observed on leaves of the plant, as described herein.

[0082] This invention also relates to a method for producing a plant of the Solanum genus showing an increased density of type IV trichomes and comprising glandular trichomes producing an increased ratio of S4: 17 / S3: 15 acyl sugars, said method comprising: a) crossing a plant of the Solanum genus comprising a polynucleotide comprising a IncRNA gene according to SEQ ID No: 2, or according to a sequence having at least 50 % sequence identity to SEQ ID No: 2 and an AP2c gene according to SEQ ID No: 3, or according to a sequence having at least 50% sequence identity to SEQ ID No: 3 and comprising a SIASAT4 gene according to SEQ ID No. 21, or comprising a homologous sequence having at least 99.6% sequence identity to SEQ ID No. 21, with another plant of the Solanum genus to obtain an Fl population; b) optionally performing one or more rounds of selfing and / or crossing a plant from of the Solanum genus from the Fl to obtain a further generation population; and c) selecting from the population a plant of the Solanum genus that comprises the polynucleotide and the SIASAT4 gene and that shows an increased density of type IV trichomes and comprises glandular trichomes producing an increased ratio of S4: 17 / S3: 15 acyl sugars.

[0083] In one embodiment, the crossing of step a) is between two plants of the Solanum genus of the same species, for example Solanum lycopersicum. Solanum galapagense, Solanum pimpinellifolium, Solanum chilense, Solanum peruvianum and Solanum hahrochaites . In another embodiment, the crossing of step a) is between two plants of the Solanum genus of different species, for example Solanum lycopersicum, Solanum galapagense, Solanum pimpinellifolium, Solanum chilense, Solanum peruvianum and Solanum hahrochaites, provided that the species are crossable.

[0084] The invention further relates to hybrid seed and to a method for producing said hybrid seed, comprising crossing a first parent plant with a second parent plant and harvesting the resultant hybrid seed, wherein the first parent plant and / or the second parent plant comprises the polynucleotide of the invention and the SIASAT4 gene of the invention. The resulting hybrid seed, and the hybrid plant showing the increased density of type IV trichomes and comprising glandular trichomes producing an increased ratio of S4: 17 / S3: 15 acyl sugars that can be grown from the hybrid seed, is also a part of the invention. In a preferred embodiment, the parent plants comprise the polynucleotide of the invention and the SIASAT4 gene homozygously.

[0085] This invention also relates to a method of growing a plant comprising a polynucleotide comprising a IncRNA gene according to SEQ ID No: 2, or according to a sequence having at least 50 % sequence identity to SEQ ID No: 2 and an AP2c gene according to SEQ ID No: 3, or according to a sequence having at least 50% sequence identity to SEQ ID No: 3 and comprising a SIASAT4 gene according to SEQ ID No. 21, or comprising a homologous sequence having at least 99.6% sequence identity to SEQ ID No. 21 which polynucleotide confers an increased density of type IV trichomes and which SIASAT4 gene confers an increased ratio of S4: 17 / S3 : 15 acyl sugars produced in glandular trichomes when compared to a plant lacking the polynucleotide and the SIASAT4 gene.

[0086] This invention further relates to a method of growing a Solanum lycopersicum plant comprising a polynucleotide that confers an increased density of type IV trichomes and comprising a SIASAT4 gene according to SEQ ID No. 21, or comprising a homologous sequence having at least 99.6% sequence identity to SEQ ID No. 21, wherein the polynucleotide and the SIASAT4 gene are as present in the genome of a Solanum lycopersicum plant, representative seed of which was deposited with the NCIMB under deposit number NCIMB 44403. The invention also relates to a method of growing a Solanum lycopersicum plant comprising the polynucleotide of the invention and the SIASAT4 gene of the invention, wherein said polynucleotide and said SIASAT4 gene are introgressed from NCIMB 44403 or from a progeny plant thereof. The invention further relates to a method for the production of a plant comprising the polynucleotide of the invention and the SIASAT4 gene of the invention, by using tissue culture or by using vegetative propagation.

[0087] The invention further provides a method for the production of a plant comprising the polynucleotide of the invention and the SIASAT4 gene of the invention by using a doubled haploid generation technique to generate a doubled haploid line that is completely homozygous, and therefore homozygously comprises the polynucleotide of the invention and the SIASAT4 gene of the invention, and that has an increased density of type IV trichomes and comprises glandular trichomes producing an increased ratio of S4: 17 / S3: 15 acyl sugars.

[0088] The invention further relates to a method for the production of a plant comprising the polynucleotide of the invention and the SIASAT4 gene of the invention, wherein the presence of said polynucleotide leads to increased density of type IV trichomes and wherein the presence of aid SIASAT4 gene leads to glandular trichomes producing an increased ratio of S4: 17 / S3 : 15 acyl sugars which method comprises growing a seed comprising said polynucleotide and said SIASAT4 gene into the said plant.

[0089] The invention further relates to a cell comprising the polynucleotide of the invention and the SIASAT4 gene of the invention as defined herein. A cell of the invention can be obtained from, or be present in, a plant of the invention. Such a cell may either be in isolated form, or a part of the complete plant, or from a part thereof, and still constitutes a cell of the invention because such a cell comprises the genetic information that determines the polynucleotide and the SIASAT4 gene as described herein. Each cell of a plant of the invention carries the polynucleotide the invention and the SIASAT4 gene of the invention, and thereby the genetic information that leads to increased density of type IV trichomes and glandular trichomes producing an increased ratio of S4: 17 / S3: 15 acyl sugars. A cell of the invention may also be a regenerable cell that can regenerate into a new plant of the invention. The presence of the genetic information in this context is the presence of polynucleotide of the invention and the SIASAT4 gene of the invention, wherein the polynucleotide and the SIASAT4 gene are as defined herein.

[0090] DEPOSIT

[0091] Seeds of Solcinum lycopersicum population S 23R.4240001, comprising the increased density of type IV trichomes trait of the invention homozygously on chromosome 2 and the glandular trichomes producing an increased ratio of S4: 17 / S3: 15 acyl sugars trait of the invention homozygously on chromosome 1, were deposited with NCIMB Ltd, Wellheads Place, Dyce, Aberdeen AB21 7GB United Kingdom on 11 July 2024 under deposit accession number NCIMB 44403. SEQUENCE INFORMATION

[0092] SEQ ID No: 1 Sequence comprising the transcript sequence of the IncRNA gene as comprised in the polynucleotide of the invention that is present in a Solcinum lycopersicum plant grown from a seed that is deposited under accession number NCIMB 44403.

[0093] AGTCAATTTTAAAAAAGAATATATTTTTTAATATTAAGTAATAATTTAATTATAAAATA TTAATTTTATTTTTAATGAAATAATTTACAACCATATAAGTTTTTTTCATTTATTTTAAT AATTTACAACCACATAAGTTTCTTTCATTTATTTTGAGTCACAAAACTTAAATTTTAAT TTTTTTAACATTAATATCGAATTAAACTACCTCACTAAAAATAAAACTGAGAAACTAT TTTCGTATCTTCGAAATTAAGAAGAGCCTCGTCGCATTCCATTATTAGACCGACGCGCC CATTTCTAGTTGGGTTTTGTTCGAAAGCCGAATATTGGGCAGTCTGATTGGGTCCACAT TTTACAAAATACTTCATACATCCAATTTATTTGCAAATTGCATTATACTCGAGCTCGCT GCGAAGATGGATATCACGAAAAATAACTTCTATGATCTACTGATTTGAGCCACGAATT TGGAACCACGACAAAAGAACTCTAACTCTGAACTTATTCTCGAAAGAATGACTCACTT TTCTTCTCTATATCTGTGTATGTGTGTATTT

[0094] SEQ ID No: 2 Genomic fragment of chromosome 2, comprising the genomic sequence of the IncRNA gene as comprised in the polynucleotide of the invention that is present in a Solanum lycopersicum plant grown from a seed that is deposited under accession number NCIMB 44403.

[0095] AGTCAATTTTAAAAAAGAATATATTTTTTAATATTAAGTAATAATTTAATTATAAAATA TTAATTTTATTTTTAATGAAATAATTTACAACCATATAAGTTTTTTTCATTTATTTTAAT AATTTACAACCACATAAGTTTCTTTCATTTATTTTGAGTCACAAAACTTAAATTTTAAT TTTTTTAACATTAATATCGAATTAAACTACCTCACTAAAAATAAAACTGAGAAACTAT TTTCGTATCTTCGAAATTAAGAAGAGCCTCGTCGCATTCCATTATTAGACCGACGCGCC CATTTCTAGTTGGGTTTTGTTCGAAAGCCGAATATTGGGCAGTCTGATTGGGTCCACAT TTTACAAAATACTTCATACATCCAATTTATTTGCAAATTGCATTATACTCGAGCTGGAA AAACAAATAGGGAAAATTATATTAAATACTAAATTATTAATTTAAATTAAATATTATA ATTACAATGTTATTTGATTTTAATCGGTAGCAAATATATTAATATTATTTCTCTCTAAA AAATTCATTTGTCATTTTCTCTCATCTTGTTTTATACAAATATAAATGTATAAATTATGT GTGTTTTTGTATAATACAAGAAAAAATTGGATATATAAATATATATCTCTTCTCCAACA TATTTATAATCATAGAATACAAAACTTCCTCTATCCAGTTTTCTTTTGCGTTTTCTGTTT TTTCGCTTTATAAAAACACAAATTATACAAAATATAATGTATAATTTGTATTTATATTA AGTGATACAACTGTTATAACTCTATTGTATGCAAATTCAAATTTTATGAAGTTATACAA AGGCAAAGACTATATACATACAAACACAATTTACATAGATACAAATGTCCGATAATCC GATAACTAAGTTTTATACAAATCAAAGCTACATTCGATTTTTAACAAATCTAATATTTG TGATAGAGCACAAATAGCAAAACTTTAGCTATAAAGCGCTATAATAATTTTTATGTTT

[0096] GATAAACCTAAAATTTACTCAAAATAAATTACTTTTTATATTTTTATATGGCTAAATCT

[0097] TATGCAAGAACACTATTATGGAAATATAAAGATCAATATCTTTTAGATTATTTCTTTAT

[0098] CGTTTTGTATTTGTTT TAT GAGAAAATTTTGCAGATTAAAAATGAAATAATAACGAA

[0099] TTACACATTTACAATGTTAAAATGTGAGGTGTAGAAGTTAATAATCACCCCCAAGAGG

[0100] CAAGATTGCGAATATAAAAAATTTGATCTACTTAAATTAATTAATTATTTAAAAAAGA

[0101] AAAAAAACGCAAAAATAAAGCATGATGATATGTCAAAAAAAAAGTCATAAATTTTGC

[0102] AGTATATGCAATTTACATGCTTAAACTTAGAGTAATTAATTATGTGCATCAATTTTGAT

[0103] ACCATTTATTATTATTATTATTATTTACAAATATATAATTATAAAAAAATAAAAAATTT

[0104] ATAGCAAACTTCAGATCAAACTTAAACTGTTTGACTTTCGAATAAAACTTCGTAACTCT

[0105] TTTATTTTTGTTCCAAACGAAAAAATTTTATGATGGTTTAAGTATGCCATTTAATAGTT

[0106] GTATGAGAAAGTTTTTAGTAGAAAAATATATTTTGCAGGTTTTGTTCCTTTTTAATCTC

[0107] TCTCCTTTTTTTTTCACTCCTCTACGCTATTTTTTATTTATCCCTCTCTTTATACAAAGTA

[0108] TTTGATATTATTTTTTTAATTTGAATAGATTTCATTACATAAATTTAAAATAAATCAAA

[0109] AATAAATCAAGTAAAAAAGAATATTATATTAAAATTAGCATTTCATTATTTCTTAAAA

[0110] TTAATATTAATTTTCCTTGACCTTTTTATAGTTCATAATACTCATTTATCAAAAATCAAA

[0111] TAGAGATAAATATATTGATACAAATTTAAATTTGAAGTCAAATAATTACATATTAAAA

[0112] AAGTTAAACTAAGATGTGTGCACTTAACCGCATAAAAATTAATTGTTTTAGTATTAGG

[0113] ACAAACATAAATAGCTATTTAAATTACATTAAACAATTTTTTTTTTCATTTTACTTTTTT

[0114] TTTTTCAAATTTCAAGCTATAAATATATTTTTTTCATATGCTAATATGATAACAATTGC

[0115] AACTTCTAAAATTTTTGGTTGAATTGCAACTTAGTGAAAAGGATTATTTATGAGCAAA

[0116] CAGTACAAGTTGTAGTTCTTCTGTTAATGTAAAAAATAAAAAATAAATATTTTAAAAT

[0117] ATTAATTAAATTTTATATTGTGTAATTTTAAGAATAACAAGATATCATATAAATTGACG

[0118] AAAGAGTATGTATATATTTATGTGTTTGTAAAAAAAAGTCCGATTTTGAATTACAATA

[0119] AGATAGTACAAAAACTTATGTAATTATTTGTCAAAATTATTAATGCTATTTCTATTATA

[0120] TTTTCTTATGATATTTGATGAATTTAAGTATATATGATGTCACACCCCTTTTTTTTCTCA

[0121] AATATTAGTTTATTTTTTGATTTTTTTATTTGTTTCGAAAGAAAAAAAAGTTTTTCCATT

[0122] AAAGTGACATTTTGAAAAGAGATAATTATTATACAGAGTCGCCACTTGGAATTAAGTT

[0123] TTGGTGTTCCAAGTCACCTTATTTGAATCCCTAATCAAAAGGAAATTTGACTCTATTTT

[0124] ATTGGTCTGCGAACTAGAAATTCGGATAAGGAATTCTGTTGACCGAGGGGAAGGTGTT

[0125] GGGCACCCTCGAGTCCCGTGGTTCTAGCACGGTCGCTTTATTGACTTTATTATGACTTA

[0126] ACTTAATTTTTGGACATTATATTATTTAACTTAATAATAAAAATATTTCTTTTTTTTTTA

[0127] CTTTATTCTCTCATTTATTTTAAACCTTTTAAAAATCGTCTTATTTATTTTAAATCTATT

[0128] GAAAATTATTTTATTATTTTTTTGTGTGTCCATTAATTCAAAATTTGAAACATTCATATT

[0129] TATTATTTATTATTTAATTATTTATTTTGCTTTTACATTTTTTACATATATTTTTTATATT

[0130] ATTATGGGGATGAATTTATATAGCTATTTTATAGTTAGAAAAAATTTAGAGTTCGAGTT AATTTAGGATTTTCTATTTATTTATTTTTTAATTTAAAAATAGAAAAATGAAATTTTAT GAGATTTGAAATTAGTTTTAAAATCACAAAAATTCTAAATAGATTAGGACTATGTTTA TTTTTAAGAAAGGTTTTAACTTTTTTTTTTTAATGCACGTAACAATTTTAACATTCTGAC

[0131] TCTTTTTTTTTTACGTTTTTTTTTGTTTTTTTTAATTATTTTTTTTTTACGTTTTTTTTTCTA TTTTTTTTCTCTAC ACGTTTTGTTTGATTAGAAATTTTTATATTTTTTTGTTTTCTGTAAA ATTTGTTTCTCTTATTTTTTTTTTATTTTTTTTATCATTATTTTTATTCTTATTATTATTTT

[0132] AAAGATTTTGTCTTCTTCTCCTTTTTTTTAAAATAATTTTTGTTACTCTTTTTTTAACGTT

[0133] TTTTTTTTCATTCATCTTTTTTTTTTTAATTTCTTGTTTTATATTTCTTTTTCATTCTGCTTCTTTCTTCTTCTTTTTTTTTTTTTAATTTTAATATTATTTTTTTTAGATTTTATATATTTGC CTTTTGTTTTGTTTTGTTTTGTTTTCTGTAATTCTATTTCTCTCTCTCTCTTTTTTTTGGGT

[0134] TATTTTCTGTTTCCTTTTTTTTAAAAAAGAAATATCTATTATTTTTATTTTCTTTTTTTTT CGTTTTTTTTTTATGTTTTAGGCTTTCTTGTTTTTATTTTTTTATTTTTTATTCTTTTGTTC TCTTTATACAACTTCTCAAAAAAATAATTATTATAATTATTAATAGTAATTATAATTAT

[0135] TTTATCATTAATCTCTAGATGAAATTACAATTCTGCCTAGATAAAAAAAAATTTACTAA

[0136] TTATTTATATATCATAATCTAACAAGTTTATAAATTTTATTTTATTTATTTTTTAGTGAA AAATTAGGCGCCAGAAAAACATTAGCTATTAAATATCACAAAATAAATTCTAATATAA ATTGCATCTAAAAAAGAATAACAGACACTGATAATAATGCAAAAAAAAAAATATGAT

[0137] ACATCTTTTATTTTGTATTATTTTTGTTAGCAAGAAACATTTTTTTTTCTTTTTTCTTTTT TTTTCAGATCGAGAGAAAAAAATAAAATAAATCTTAACATGCATCACATATTGATATA TAATAATAACAATACGATTCAAAGAACTTCATATTATTAGATTAAAATTTTCATATAA

[0138] CATTACATTTAAATAATGTGATAGTTTAGAATTACCTCGCTGCGAAGATGGATATCAC GAAAAATAACTTCTATGATCTACTGATTTGAGCCACGAATTTGGAACCACGACAAAAG AACTCTAACTCTGAACTTATTCTCGAAAGAATGACTCACTTTTCTTCTCTATATCTGTG

[0139] TATGTGTGTATTT

[0140] SEQ ID No: 3 Genomic fragment of chromosome 2, comprising the genomic sequence of the AP2c gene as comprised in the polynucleotide of the invention that is present in a Solanum lycopersicum plant grown from a seed that is deposited under accession number NCIMB 44403

[0141] ATGTGGGATCTAAATGAATCAGAAGAAGGTTGTTCTTCACCGATAGAGTTTGAAGGCG ATGATGAGAAAGGTAAACGGGTCGGATCCGTTTCGAATTCAAGTTCATCAGCTGTAGC TGTTGATGATATATCTGAAGAGGAATTAGATGGAGAAAGAGGGAAGAAAAAGAGAG

[0142] GTAAAATTTTCGGCTTCTCTATGGTGGGTCTGGGTAACGGCGATGAAGAACAGCCGGT GACCCGACAGTTTTTTCCGGTTGATGAGTCGGAAATGGGTGGTGTTGCTGCTGAAAAT GGATCCCCGAATTTTCCCAGAGCTCACTGGGTTGGAGTTAAATTTTACCAAACGGAGA

[0143] CACTTGGCAACACGGGATTGGCCAGGCCTGTAGATATGGTTCAGCAGCAACAGCAGC CTATTAAGAAGAGCCGCCGTGGACCAAGGTCTAGAAGCTCACAGTACCGTGGGGTTA

[0144] CCTTCTACCGGAGAACTGGCCGGTGGGAATCTCATATATGGTGAGCCATTTTACCCAA

[0145] AATTATTATTTCTCTTCACCTTCGAATTTTGGACTTTATTTTTTTTTACTAATTTGCTTTA

[0146] ATAATCTTACAGGGATTGTGGAAAACAAGTTTATCTAGGTATTTAATTTTCAATTAAGC

[0147] TATTTATTATTATTATTATTATTATTGTACATTAGAAAATTAATTTGGTCTTTATTTAAC

[0148] GGATTTCTCCTTTATAGGTGGATTTGATACTGCTCATGCAGCAGCTCGGTGAGCTTCTG

[0149] CTTCTTAGTCACCTTTTAATTTTTTCAACTCACAATTTCAAAATTGAAATTTATTTTTTA

[0150] AAATAAAATATACAGGGCATACGATAGGGCAGCCATAAAATTCCGGGGAGTAGAAGC

[0151] CGACATAAACTTTAGCTTAGAAGATTATGAGTCCGATTTAAAACAGGTAATACCATGA

[0152] TCAGATCAGTGCGACTGATTAGGAAAAAGTTTGTGTTTTATGATGTTCCTTTTTTAGAT

[0153] TTGATTTTATTTCTGTGTACCTGTCTAGTTGGAGTACTTTTATTCTAACACTCGTCTTAT

[0154] CGTGAACAGTATTTTTCACCATCAAATTGAAAAATTAAATTCTAATACATTATTTTTTA

[0155] ATGTATGATACAGATGACAAGTTTAACAAAGGAAGAATTTGTGCATGTGCTTCGAAGA

[0156] CAAAGTACTGGTTTTCCGAGGGGAAGCTCCAAGTATAGAGGGGTGACTTTGCACAAAT

[0157] GTGGTAGATGGGAAGCTAGAATGGGACAGTTCTTAGGCAAAAAGTAAAAAGAATATA

[0158] TTTATATTTATATATAAAGAAATTCATATCCTCATATTATTATTCTTCTCAATGGTAAA

[0159] ATCCATTTACTAAGGAATAATACTTTAGATGTCAATTAGTAACTTTTGTTCTTTCTTTG

[0160] AATAGGTACGTTTATTTGGGACTCTTTGATACTGAGATTGAAGCTGCCAGGTGTGTGG

[0161] TTGGCCGAATTTATTCTTCCTACTACTTAACCTCAATTACTTTTGAATTACTGAATATTT

[0162] AAGGTAGTTTATTTTCTTTTGTTAATACAGAGCTTATGATAAAGCTGCCATCAAGTGTA

[0163] ACGGGAAGGATGCAGTTACAAATTTTGATCCAAGCATTTATGAGAATGAACTAAACTC

[0164] AATTGGTAAATTGTACTATTCTTCTTTTATTTGCTTCTCGTTTTGTGTTTTGATGTGATT

[0165] ATTCAAACAGAAGATTCATTAATGTGATCTCCCAGAATCTACCGATAACGTGGCTGAT

[0166] CATAATCTGGATTTAAGTTTGGGTGGTTCAAGCTCAAAGCAAGGAAGTCAAGAATTGG

[0167] GGGATAATAGAGGTCAAAATTCTTCATCAATAATGCAATTGGATATTGATTGGCAGCG

[0168] CCATGGTTTAAGGCCTGAGGTATATTTACACGATCAATAGCTATATCTTGTTTTCGTTG

[0169] TGAAATTCAATTTTTACCTCTGATGCAAACATGTTTTAATTTCCAGAAGCAATCTGCCT

[0170] TAATTGATGCTCGAAGAAGAGAAAACAGGTACAATGAAACAGAAACGTTGCAACTTT

[0171] TAAGCCAGACGCACTTGCATTCTCCAGCCTCCTTGAAGCATAACAATAATAATAATAG

[0172] TCAAGTGCAACGGTTTGGCCAATTTATGAGACCTGGTGATCAATCCCATATGATTCAA

[0173] ATGTTTCCACAGCAATTTGGCTCATCAAATTATCAAGTGAATATCCTTTTTAATAATGT

[0174] ACTCTTTTTTTTTTCTTTTGAAATCTTGGTAGTTTTTTTTTTAAAAAAATGTTGTGGCAT

[0175] GTTACAGATTCAATTTCCAAGTGGCAGCAATGGTGGAAGAATTGGAGCTACAAATGTA

[0176] AGAGATCTATCGCTTGCGGCGACGAGTAATGGTTCTTCACAATGGCAATCCAATTTTC

[0177] CTCCTCAGATATTTGTAGCTGCTGCAGCATCATCAGGATTCCCTCAGCAGATAGTAAG

[0178] ACCTCAGAATTGGTCCTCAGAAAATGGTTTCCATCATTCACTCATGAGACCTTCTTGA SEQ ID No: 4 Sequence comprising the coding sequence (cds) of the AP2c gene as comprised in the polynucleotide of the invention that is present in a Solcinum lycopersicum plant grown from a seed that is deposited under accession number NCIMB 44403

[0179] ATGTGGGATCTAAATGAATCAGAAGAAGGTTGTTCTTCACCGATAGAGTTTGAAGGCG

[0180] ATGATGAGAAAGGTAAACGGGTCGGATCCGTTTCGAATTCAAGTTCATCAGCTGTAGC

[0181] TGTTGATGATATATCTGAAGAGGAATTAGATGGAGAAAGAGGGAAGAAAAAGAGAG

[0182] GTAAAATTTTCGGCTTCTCTATGGTGGGTCTGGGTAACGGCGATGAAGAACAGCCGGT

[0183] GACCCGACAGTTTTTTCCGGTTGATGAGTCGGAAATGGGTGGTGTTGCTGCTGAAAAT

[0184] GGATCCCCGAATTTTCCCAGAGCTCACTGGGTTGGAGTTAAATTTTACCAAACGGAGA

[0185] CACTTGGCAACACGGGATTGGCCAGGCCTGTAGATATGGTTCAGCAGCAACAGCAGC

[0186] CTATTAAGAAGAGCCGCCGTGGACCAAGGTCTAGAAGCTCACAGTACCGTGGGGTTA

[0187] CCTTCTACCGGAGAACTGGCCGGTGGGAATCTCATATATGGGATTGTGGAAAACAAGT

[0188] TTATCTAGGTGGATTTGATACTGCTCATGCAGCAGCTCGGGCATACGATAGGGCAGCC

[0189] ATAAAATTCCGGGGAGTAGAAGCCGACATAAACTTTAGCTTAGAAGATTATGAGTCCG

[0190] ATTTAAAACAGATGACAAGTTTAACAAAGGAAGAATTTGTGCATGTGCTTCGAAGACA

[0191] AAGTACTGGTTTTCCGAGGGGAAGCTCCAAGTATAGAGGGGTGACTTTGCACAAATGT

[0192] GGTAGATGGGAAGCTAGAATGGGACAGTTCTTAGGCAAAAAGTACGTTTATTTGGGA

[0193] CTCTTTGATACTGAGATTGAAGCTGCCAGAGCTTATGATAAAGCTGCCATCAAGTGTA

[0194] ACGGGAAGGATGCAGTTACAAATTTTGATCCAAGCATTTATGAGAATGAACTAAACTC

[0195] AATTGAATCTACCGATAACGTGGCTGATCATAATCTGGATTTAAGTTTGGGTGGTTCA

[0196] AGCTCAAAGCAAGGAAGTCAAGAATTGGGGGATAATAGAGGTCAAAATTCTTCATCA

[0197] ATAATGCAATTGGATATTGATTGGCAGCGCCATGGTTTAAGGCCTGAGAAGCAATCTG

[0198] CCTTAATTGATGCTCGAAGAAGAGAAAACAGGTACAATGAAACAGAAACGTTGCAAC

[0199] TTTTAAGCCAGACGCACTTGCATTCTCCAGCCTCCTTGAAGCATAACAATAATAATAA

[0200] TAGTCAAGTGCAACGGTTTGGCCAATTTATGAGACCTGGTGATCAATCCCATATGATT

[0201] CAAATGTTTCCACAGCAATTTGGCTCATCAAATTATCAAATTCAATTTCCAAGTGGCA

[0202] GCAATGGTGGAAGAATTGGAGCTACAAATGTAAGAGATCTATCGCTTGCGGCGACGA

[0203] GTAATGGTTCTTCACAATGGCAATCCAATTTTCCTCCTCAGATATTTGTAGCTGCTGCA

[0204] GCATCATCAGGATTCCCTCAGCAGATAGTAAGACCTCAGAATTGGTCCTCAGAAAATG

[0205] GTTTCCATCATTCACTCATGAGACCTTCTTGA SEQ ID No: 5 Sequence comprising the amino acid sequence of the AP2c gene as comprised in the polynucleotide of the invention that is present in a Solcinum lycopersicum plant grown from a seed that is deposited under accession number NCIMB 44403.

[0206] MWDLNESEEGCSSPIEFEGDDEKGKRVGSVSNSSSSAVAVDDISEEELDGERGKKKRGKI

[0207] FGFSMVGLGNGDEEQPVTRQFFPVDESEMGGVAAENGSPNFPRAHWVGVKFYQTETLGN TGLARPVDMVQQQQQPIKKSRRGPRSRSSQYRGVTFYRRTGRWESHIWDCGKQVYLGGF DTAHAAARAYDRAAIKFRGVEADINFSLEDYESDLKQMTSLTKEEFVHVLRRQSTGFPRG SSKYRGVTLHKCGRWEARMGQFLGKKYVYLGLFDTEIEAARAYDKAAIKCNGKDAVTN FDPSIYENELNSIESTDNVADHNLDLSLGGSSSKQGSQELGDNRGQNSSSIMQLDIDWQRH GLRPEKQSALIDARRRENRYNETETLQLLSQTHLHSPASLKHNNNNNSQVQRFGQFMRPG DQSHMIQMFPQQFGSSNYQIQFPSGSNGGRIGATNVRDLSLAATSNGSSQWQSNFPPQIFV

[0208] AAAASSGFPQQIVRPQNWSSENGFHHSLMRPS

[0209] SEQ ID No: 6 Genetic SNP marker that is able to identify the polynucleotide that confers the trait of the invention in a Solanum lycopersicum plant grown from a seed that is deposited under accession number NCIMB 44403. A genomic fragment of chromosome 2 is presented, wherein the position of a SNP comprising a change from T to C is indicated as [T / C] . This SNP mutation (i.e. the C nucleotide) is present in plants grown from seeds of the said deposit.

[0210] CAAAGTATTTTGAAAATCACCAAGAAAATAAATACTACTTGTACTGCCTATCACTGCA TAGGGAATTTCGAGTAATTCTTAACAAACGGAACGTGTCATA[T / C]TTTTAAGGTCCAA TTTTATTATTATATCAAAAAATTGATAGTAATGATTAATGACTATGTAAACGCCAAGG TTAGAGAGCCAATTGCAACTTTATCTACA

[0211] SEQ ID No: 7 Genetic SNP marker that is able to identify the polynucleotide that confers the trait of the invention in a Solanum lycopersicum plant grown from a seed that is deposited under accession number NCIMB 44403. A genomic fragment of chromosome 2 is presented, wherein the position of a SNP comprising a change from A to C is indicated as [A / C]. This SNP mutation (i.e. the C nucleotide) is present in plants grown from seeds of the said deposit.

[0212] AAACTAGTTGCTCCATGTGATTAATCTTGAAATAAGTTATACTGGGTATAAGTTATTTC ACCAAATATATGAGAAAATTTATCTTATCACTATAACAAAAATTGTAAGATAACTGAT TCAAGGACAGC[A / C]TAATACCTCCAACCAAATTTTATCCTAAATTTATTAGACTTTGT ACCTCACGGCTGAAACGTTTTGATTGATTGTTCTTTTCATTTAATTTTAGCAGAATTAG GTTTGTTTAAAGAAAAAGGGATGC SEQ ID No: 8 Genetic SNP marker that is able to identify the polynucleotide that confers the trait of the invention in a Solcinum lycopersicum plant grown from a seed that is deposited under accession number NCIMB 44403. A genomic fragment of chromosome 2 is presented, wherein the position of a SNP comprising a change from T to G is indicated as [T / G]. This SNP mutation (i.e. the G nucleotide) is present in plants grown from seeds of the said deposit. In this sequence, Y refers to any pyrimidine (C or T).

[0213] TATTTAGTTAGTTAAAGATCATTAGGAGCTATGGAACAAGGAAAGAGGATAAATCACT CCCATATATCCATAACCAGTTCTCAATAATTATAATTAAATTTGATGTGTTGCACGAAT TGAGAGATGGA[T / G]AGTATAGATGTATTTTCAATAATTYATAAATTTGTAAACTATAA TTTAATCAAGCACTTAGTTGAAATATAAGTTTTATATATCACTAAAAAAGCACTAA

[0214] SEQ ID No: 9 Genetic SNP marker that is able to identify the polynucleotide that confers the trait of the invention in a Solanum lycopersicum plant grown from a seed that is deposited under accession number NCIMB 44403. A genomic fragment of chromosome 2 is presented, wherein the position of a SNP comprising a change from T to C is indicated as [T / C] . This SNP mutation (i.e. the C nucleotide) is present in plants grown from seeds of the said deposit.

[0215] GAAATAATTTACAACCATATAAGTTTTTTTCATTTATTTTAATAATTTACAACCACATA AGTTTCTTTCATTTATTTTGAGTCACAAAACTTAAATTTTAATTTTTTTAACATTAATAT CGAATTAAA[T / C]TACCTCACTAAAAATAAAACTGAGAAACTATTTTCGTATCTTCGAA ATTAAGAAGAGCCTCGTCGCATTCCATTATTAGACCGACGCGCCCATTTCTAGTTGGG TTTTGTTCGAAAGCCGAATATTG

[0216] SEQ ID No: 10 Genetic SNP marker that is able to identify the polynucleotide that confers the trait of the invention in a Solanum lycopersicum plant grown from a seed that is deposited under accession number NCIMB 44403. A genomic fragment of chromosome 2 is presented, wherein the position of a SNP comprising a change from A to G is indicated as [A / G]. This SNP mutation (i.e. the G nucleotide) is present in plants grown from seeds of the said deposit. In this sequence Y refers to any pyrimidine (C or T), R refers to A or G, and W refers to A or T.

[0217] TTTAACAAATCTAATATTTGTGATAGAGCACAAATAGCAAAACTTTAGCTATAAAGCG CTATAATAATTTTTATGTTTGATAAACCTAAAATTTACTCAAAATAAATTACTTTTTATATTTTTATATG[A / G]CTAAATCTTATGCAAGAACACTATTATGGAAATATAAAGATCAATATCTTTTRGATTATTTCTTTATCGTTYTGTATTTGTTTTTATTGAGAAAATTTTGCAGA TTAAAAATGAAATAWTAACGAAT SEQ ID No: 11 Genetic SNP marker that is able to identify the polynucleotide that confers the trait of the invention in a Solcinum lycopersicum plant grown from a seed that is deposited under accession number NCIMB 44403. A genomic fragment of chromosome 2 is presented, wherein the position of a SNP comprising a change from C to A is indicated as [C / A], This SNP mutation (i.e. the A nucleotide) is present in plants grown from seeds of the said deposit.

[0218] AATGTAAAAAATAAAAAATAAATATTTTAAAATATTAATTAAATTTTATATTGTGTAA TTTTAAGAATAACAAGATATCATATAAATTGACGAAAGAGTATGTATATATTTATGTG TTTGTAAAAAAA[C / A]GTCCGATTTTGAATTACAATAAGATAGTACAAAAACTTATGTA ATTATTTGTCAAAATTATTAATGCTATTTCTATTATATTTTCTTATGATATTTGATGAAT TTAAGTATATATGATGTCACACCC

[0219] SEQ ID No: 12 Genetic SNP marker that is able to identify the polynucleotide that confers the trait of the invention in a Solanum lycopersicum plant grown from a seed that is deposited under accession number NCIMB 44403. A genomic fragment of chromosome 2 is presented, wherein the position of a SNP comprising a change from G to T is indicated as [G / T], This SNP mutation (i.e. the T nucleotide) is present in plants grown from seeds of the said deposit.

[0220] TAACTTGTGTCTGCCATATCCATTATAAAATTTTCCTCTACATCTTAAAGATAAAAAAA ATATACTTAAATTTTTATGAAACTTTATTCTGGGTGGACATATCTTCTTTCATCATTTCT CTACCTAAA[G / T]AAAAGTATCACATAATTTTAGTAATACAATTAATTAAATATTTCAA TTTTTTTATTTTCTTGGGGGATTTAATTTTTCACCTAATTTTTCATCAGTAATTAAAAAA TAATACTACTGAAAATTTAAA

[0221] SEQ ID No: 13 Genetic SNP marker that is able to identify the polynucleotide that confers the trait of the invention in a Solanum lycopersicum plant grown from a seed that is deposited under accession number NCIMB 44403. A genomic fragment of chromosome 2 is presented, wherein the position of a SNP comprising a change from T to C is indicated as [T / C] . This SNP mutation (i.e. the C nucleotide) is present in plants grown from seeds of the said deposit.

[0222] TAAAAATGTCTATGAAATTTAATGACCATAAGATATAAATGTTTAATCTGCTGTCATTG TACTAGTCATTAAAAGTAACATTTAAAAGGAACCAAACAATTGGGATTCCTAATTAAT ATTTGTGA[T / C]GCATTTTGCTCCTTTATTATAATATTTTAAGATATTTAATAATATTTGT TCAGTTTATAATATTAATGAATAATTTATTATATTTATTTTGAATATTTAGATAACCTAT ATTTTATAAGAGGGA SEQ ID No: 14 Genetic SNP marker that is able to identify the polynucleotide that confers the trait of the invention in a Solcinum lycopersicum plant grown from a seed that is deposited under accession number NCIMB 44403. A genomic fragment of chromosome 2 is presented, wherein the position of a SNP comprising a change from C to T is indicated as [C / T] . This SNP mutation (i.e. the T nucleotide) is present in plants grown from seeds of the said deposit. In this sequence Y refers to any pyrimidine (C or T), and M refers to A or C.

[0223] ATAAATCGCTTATTTTAAAAAAAGAAATTACTACGTACTACTTATTTTAGAAGTTGAAT GGAGTATTTGATAAAGTTTTCTTTTATGAATGGATGCAAGCGTTTTCAGATACG[C / T]A GTTAATAGTTYCAAACTTTGAAATAAAAGCTGAAAAAAGATATCMAAAAAATAAGTG GTCTTGATTTTGCATGTTCATATTTACCAAACTAATTACTCC

[0224] SEQ ID No: 15 Genetic SNP marker that is able to identify the polynucleotide that confers the trait of the invention in a Solanum lycopersicum plant grown from a seed that is deposited under accession number NCIMB 44403. A genomic fragment of chromosome 2 is presented, wherein the position of a SNP comprising a change from A to C is indicated as [A / C]. This SNP mutation (i.e. the C nucleotide) is present in plants grown from seeds of the said deposit. In this sequence Y refers to any pyrimidine (C or T).

[0225] ATAAATCGCTTATTTTAAAAAAAGAAATTACTACGTACTACTTATTTTAGAAGTTGAAT GGAGTATTTGATAAAGTTTTCTTTTATGAATGGATGCAAGCGTTTTCAGATACGYAGTT AATAGTTYCAAACTTTGAAATAAAAGCTGAAAAAAGATATC[A / C]AAAAAATAAGTGG TCTTGATTTTGCATGTTCATATTTACCAAACTAATTACTCC

[0226] SEQ ID No: 16 Genetic SNP marker that is able to identify the polynucleotide that confers the trait of the invention in a Solanum lycopersicum plant grown from a seed that is deposited under accession number NCIMB 44403. A genomic fragment of chromosome 2 is presented, wherein the position of a SNP comprising a change from T to C is indicated as [T / C] . This SNP mutation (i.e. the C nucleotide) is present in plants grown from seeds of the said deposit. In this sequence Y refers to any pyrimidine (C or T), R refers to A or G, and K refers to G or T.

[0227] AATTATTTACATTTGAATTGTTCTTTTAGTTAAAAGCTATATATTTTTAAAGCTTTTCCA GGCTAGCACTAAATGATGAATTACTACATACAAACTATAA[T / C]ATCCAGTTTCATCAC TTCTGAATCGGGGAATTATCGATGAATAATAAAATTAAAATGGATTTARATTTAATAT TCTRYTTAAATTTAAATKTTAAATTTA SEQ ID No: 17 Genetic SNP marker that is able to identify the polynucleotide that confers the trait of the invention in a Solcinum lycopersicum plant grown from a seed that is deposited under accession number NCIMB 44403. A genomic fragment of chromosome 2 is presented, wherein the position of a SNP comprising a deletion of an A nucleotide is indicated as [A / ] . This SNP mutation (i.e. the deletion of the A nucleotide) is present in plants grown from seeds of the said deposit. TTTACTCATTAATATATATCATAAATATTAATTAATAACAACTAATATTAAACGGATAT GATAATATTTAAAAAGAAATTTACATGTGACAATTGTGTAAATAAAAGGAAATGTAA GAGAGAGAATAAAGAGGA [A / ] GTTGTCTAACGTGGGATAGGATGAAGGAAAAGGAAA TGACAATTAGCGTAACCTAATAAAGGTAATGGAGTAGTAACTCATCCATTCACTCTAT TTCCATTACTAGATATTTTGTAATAAAATAA

[0228] SEQ ID No: 18 Genetic SNP marker that is able to identify the polynucleotide that confers the trait of the invention in a Solanum lycopersicum plant grown from a seed that is deposited under accession number NCIMB 44403. A genomic fragment of chromosome 2 is presented, wherein the position of a SNP comprising a change from A to G is indicated as [A / G]. This SNP mutation (i.e. the G nucleotide) is present in plants grown from seeds of the said deposit. In this sequence Y refers to any pyrimidine (C or T).

[0229] TTTCTATGTAAATATATAACTTAAGGTAATATGGGTGATTTTAGGTAATATTCTAAAAG TGGGCTAGTCCAAAGGTACAAGATCAAGTGAAATACTACAAGTAGTTAATAGTGGGA AACATATATCTATCATATCTAACCCT[A / G]TCTCTCTCCACTAACCTTTTTTATTTTTCTC CATCTTCCAACAACTTTATTAACCTCTCTATAAGTCGTAGGTTTCATGTTCTTTCAGCAT ACTYAAACACACACAACTTTTTTATTTTATCATTATATATATAT

[0230] SEQ ID No: 19 Genetic SNP marker that is able to identify the polynucleotide that confers the trait of the invention in a Solanum lycopersicum plant grown from a seed that is deposited under accession number NCIMB 44403. A genomic fragment of chromosome 2 is presented, wherein the position of a SNP comprising a change from C to A is indicated as [C / A], This SNP mutation (i.e. the A nucleotide) is present in plants grown from seeds of the said deposit.

[0231] AGATTATGAGTCCGATTTAAAACAGGTAATACCATGATCAGATCAGTGCGACTGATTA GGAAAAAGTTTGTGTTTTATGATGTTCCTTTTTT[C / A]GATTTGATTTTATTTCTGTGTAC CTGTCTAGTTGGAGTACTTTTATTCTAACACTCGTCTTATCGTGAACAGTATTTTTCACC ATCAAATTGAAAAAT SEQ ID No: 20 Genetic SNP marker that is able to identify the polynucleotide that confers the trait of the invention in a Solcinum lycopersicum plant grown from a seed that is deposited under accession number NCIMB 44403. A genomic fragment of chromosome 2 is presented, wherein the position of a SNP comprising a change from C to A is indicated as [C / A], This SNP mutation (i.e. the A nucleotide) is present in plants grown from seeds of the said deposit. In this sequence, R refers to A or G, M refers to A or C and N refers to A, C, G or T.

[0232] TTTTATTAGAANNTTTTTTTARAATAGGTATATCACTTGAAATTTTCTACAAAATTATTT TAAAAATTTAAAATCATCTTCAAGATCATATCAACGGCCA[C / A]ACCTAAGGGTCAAA GTGTATTTTCTAGATTTAGTCGTTTCTACMGCTAGTGTCTATAATTGTGTACTTTGTCA GAGGATGGAGTACAAAAATAGAAATTA

[0233] SEQ ID No: 21 Sequence comprising the coding sequence (cds) of the S1ASAT4 gene of the invention that is present in a Solanum lycopersicum plant grown from a seed that is deposited under accession number NCIMB 44403 ATGGTGAAACCCTCAGCTCCTACCCCGGATAATCTTCGGAGATTGAAGCTTTCCTTGTT CGATCAGATGGATATTGGTGCATATGTACCAATTGTCTTCAACTACTTGCCGAACAGC ACTTCATCATATGATCATGATGATAAGCTTGAAAAATCATTGTCGGAGACGCTAACCA AGTTTTACCCTTTTGCTGGAAGATTTAGAAAAGGCATTGATCCATTTTCCATCGACTGC AATGATGAAGGTATTGAATATGTTCGAACCAAAGTCAATGCAGACGATCTTGCCCAAT ATCTCCGTGGTCAAGCCCATAATGATATTGAGTCGTCTTTGATTGATCTTCTTCCTGTA ATGCATCGTCTACCATCAAGTCCATTATTTGGTGTTCAAGTGAATGTATTCAATAACGG AGGTGTAACCATAGGGATACAAATTTTACATATGGTATCTGATGCTTTCACTTTAGTAA AATTTGTAAATGAATGGGCGCACACCACCCTTACAGGGACGATGCCACTAGATAATCC CGGTTTTGGTCAATTGCCATGGCTATTTCCAGCAAGAGCGCTACCGTTTCCATTACCTG ATTTCAACACTACTACTGCCCCTAATTATAAGAATGTTACAAAGAGGTTTCTCTTTGAT GCTTTGGCAATAGAAAACCTCAGAAATACAATCAAAGCCAATGACATGATGATGAAG CAACCTTCTAGAGTGGTGGTCGTGATGTCCCTAATATGGAAGGTTCTTACACACATTTC TTCCGCCAAAAATAATGGAAATTCAAGGGACTCATCTTTAGTGTTTGTTGTTAATTTGA GGGGAAAACTGTCATGTACTGCACCGTCTTTAGAACACGTTGTAGGGAATTGTGTAAT ACCAGCAACTGCTAACAAGGAGGGCGATGAGGCAAGAAGAAAGGATGATGAGTTGA ATGATTTCGTTAAGTTGGTAAGAAATACAATACGGGACACATGTGAAGCCATTGGTAA GGCGGAAAGCGTTGATGATATTTCCTCTTTAGCATTTAACAATCTGACGAAATGTATA GAAAAAATTCTGCATGGAGACGAGATGGACTTCTATTCGTGCTCTAGTTGGTGCGGAT TCCCTTGGTATGAAGCTGACTTTGGTTGGGGAAAGCCATTCTGGGTGAGCTCAGTTAG TTTTGGTCATCATGGAGTAACTAATCTCATGGACACAAAAGATGGTGATGGAATACAA GTAACAATTTGTTTGAAGGAGAATGACATGATTGAGTTTGAGAGAGACCCTCACATTT

[0234] TGTCCTCCACTTCAAAACTAGCATTCCATTCCTTAGGATAA

[0235] SEQ ID No: 22 Sequence comprising the amino acid sequence of the S1ASAT4 gene of the invention that is present in a Solanum lycopersicum plant grown from a seed that is deposited under accession number NCIMB 44403.

[0236] MVKPSAPTPDNLRRLKLSLFDQMDIGAYVPIVFNYLPNSTSSYDHDDKLEKSLSETLTKFY

[0237] PFAGRFRKGIDPFSIDCNDEGIEYVRTKVNADDLAQYLRGQAHNDIESSLIDLLPVMHRLPS

[0238] SPLFGVQVNVFNNGGVTIGIQILHMVSDAFTLVKFVNEWAHTTLTGTMPLDNPGFGQLPW

[0239] LFPARALPFPLPDFNTTTAPNYKNVTKRFLFDALAIENLRNTIKANDMMMKQPSRVVVVM

[0240] SLIWKVLTHISSAKNNGNSRDSSLVFVVNLRGKLSCTAPSLEHVVGNCVIPATANKEGDEA

[0241] RRKDDELNDFVKLVRNTIRDTCEAIGKAESVDDISSLAFNNLTKCIEKILHGDEMDFYSCSS

[0242] WCGFPWYEADFGWGKPFWVSSVSFGHHGVTNLMDTKDGDGIQVTICLKENDMIEFERDP HILSSTSKLAFHSLG

[0243] SEQ ID No: 23 Sequence comprising the coding sequence (cds) of an S1ASAT4 allele

[0244] ATGGTGAAACCCTCAGCTCCTACCCCGGATAATCTTCGGAGATTGAAGCTTTCCTTGTT

[0245] CGATCAGCTGGATATTGGTGCATATGTACCAATTCTCTTCAACTACTTGCCGAACAGC

[0246] ACTTCATCATATGATCATGATGATAAGCTTGAAAAATCATTGTCGGAGACGCTAACCA

[0247] AGTTTTACCCTTTTGCTGGAAGATTTAGAAAAGGCATTGATCCATTTTCCATCGACTGC

[0248] AATGATGAAGGTATTGAATATGTTCGAACCAAAGTCAATGCAGACGATCTTGCCCAAT

[0249] ATCTCCGTGGTCAAGCCCATAATGATATTGAGTCGTCTTTGATTGATCTTCTTCCTGTA

[0250] ATGCATCGTCTACCATCAAGTCCATTATTTGGTGTTCAAGTGAATGTATTCAATAACGG

[0251] AGGTGTAACCATAGGGATACAAATTTTACATATGGTCTCTGATGCTTTCACTTTAGTAA

[0252] AATTTGTAAATGAATGGGCGCACACCACCCTTACAGGGACGATGCCACTAGATAATCC

[0253] CGGTTTTGGTCAATTGCCATGGCTATTTCCAGCAAGAGCGCTACCGTTTCCATTACCTG

[0254] ATTTCAACACTACTACTGCCCCTAATTATAAGAATGTTACAAAGAGGTTTCTCTTTGAT

[0255] GCTTTGGCAATAGAAAACCTCAGAAATACAATCAAAGCCAATGACATGATGATGAAG

[0256] CAACCTTCTAGAGTGGTGGTCGTGATGTCCCTAATATGGAAGGTTCTTACACACATTTC

[0257] TTCCGCCAAAAATAATGGAAATTCAAGGGACTCATCTTTAGTGTTTGTTGTTAATTTGA

[0258] GGGGAAAACTGTCATGTACTGCACCGTCTTTAGAACACGTTGTAGGGAATTGTGTAAT

[0259] ACCAGCAATTGCTAACAAGGAGGGCGATGAGGCAAGAAGAAAGGATGATGAGTTGA

[0260] ATGATTTCGCTAAGTTGGTAAGAAATACAATACGGGACACATGTGAAGCCATTGGTAA

[0261] GGCGGAAAGCGTTGATGATATTTCCTCTTTAGCATTTAACAATCTGACGAAATGTATA

[0262] GAAAAAATTCTGCATGGAGACGAGATGGACTTCTATTCGTGCTCTAGTTGGTGCGGAT

[0263] TCCCTTGGTATGAAGCTGACTTTGGTTGGGGAAAGCCATTCTGGGTGAGCTCAGTTAG TTTTGGTCATCATGGAGTAACTAATCTCATGGACACAAAAGATGGTGATGGAATACAA

[0264] GTAACAATTTGTTTGAAGGAGAATGACATGATTGAGTTTGAGAGAGACCCTCACATTT

[0265] TGTCCTCCACTTCAAAACTAGCATTCCATTCCTTTGGATAA

[0266] SEQ ID No: 24 Sequence comprising the amino acid sequence of an S1ASAT4 allele

[0267] MVKPSAPTPDNLRRLKLSLFDQLDIGAYVPILFNYLPNSTSSYDHDDKLEKSLSETLTKFYP

[0268] FAGRFRKGIDPFSIDCNDEGIEYVRTKVNADDLAQYLRGQAHNDIESSLIDLLPVMHRLPSS

[0269] PLFGVQVNVFNNGGVTIGIQILHMVSDAFTLVKFVNEWAHTTLTGTMPLDNPGFGQLPWL

[0270] FPARALPFPLPDFNTTTAPNYKNVTKRFLFDALAIENLRNTIKANDMMMKQPSRVVVVMS

[0271] LIWKVLTHISSAKNNGNSRDSSLVFVVNLRGKLSCTAPSLEHVVGNCVIPAIANKEGDEAR

[0272] RKDDELNDFAKLVRNTIRDTCEAIGKAESVDDISSLAFNNLTKCIEKILHGDEMDFYSCSS

[0273] WCGFPWYEADFGWGKPFWVSSVSFGHHGVTNLMDTKDGDGIQVTICLKENDMIEFERDP HILSSTSKLAFHSFG

[0274] SEQ ID No: 25 Sequence comprising the coding sequence (cds) of the SgASAT4 allele

[0275] ATGGTGAAACCCTCAGCTCCTACCCCGGATAATCTTCGGAGATTGAAGCTTTCCTTGTT

[0276] CGATCAGATGGATATTGGTGCATATGTACCAATTCTCTTCAACTACTTGCCGAACAGC

[0277] ACTTCATCATATGATCATGATGATAAGCTTGAAAAATCATTGTCGGAGACGCTAACCA

[0278] AGTTTTACCCTTTTGCTGGAAGATTTAGAAAAGGCATTGATCCATTTTCCATCGACTGC

[0279] AATGATGAAGGTATTGAATATGTTCGAACCAAAGTCAATGCAGACGATCTTGCCCAAT

[0280] ATCTCCGTGGTCAAGCCCATAATGATATTGAGTCGTCTTTGATTGATCTTCTTCCTGTA

[0281] ATGCATCGTCTACCATCAAGTCCATTATTTGGTGTTCAAGTGAATGTATTCAATAACGG

[0282] AGGTGTAACCATAGGGATACAAATTTTACATATGGTATCTGATGCTTTCACTTTAGTAA

[0283] AATTTGTAAATGAATGGGCGCACACCACCCTTACAGGGACGATGCCACTAGATAATCC

[0284] CGGTTTTGGTCAATTGCCATGGCTATTTCCAGCAAGAGCGCTACCGTTTCCATTACCTG

[0285] ATTTCAACACTACTACTGCCCCTAATTATAAGAATGTTACAAAGAGGTTGCTCTTTGAT

[0286] GCTTTGGCAATAGAAAACCTCAGAAATACAATCAAAGCCAATGACACGATGATGAAG

[0287] CAACCTTCTAGAGTGGTGGTCGTGATGTCCCTAATATGGAAGGTTCTTACACACATTTC

[0288] TTCCGCCAAAAATAATGGAAATTCAAGGGACTCATCTTTAGTGTTTGTTGTTAATTTGA

[0289] GGGGAAAACTGTCATGTACTGCACCGTCTTTAGAACACGTTGTAGGGAATTGTGTAAT

[0290] ACCAGCAACTGCTAACAAGGAGGGCGATGAGGCAAGAAGAAAGGATGATGAGTTGA

[0291] ATGATTTCGTTAAGTTGGTAAGAAATACAATACGGGACACATGTGAAGCCATTGGTAA

[0292] GGCGGAAAGCGTTGATGATATTTCCTCTTTAGCATTTAACAATCTGACGAAATGTATA

[0293] GAAAAAATTCTGCATGGAGACGAGATGGACTTCTATTCGTGCTCTAGTTGGTGCGGAT

[0294] TCCCTTGGTATGAAGCTGACTTTGGTTGGGGAAAGCCATTCTGGGTGAGCTCAGTTAG

[0295] TTTTGGTCATCATGGAGTAACTAATCTCATGGACACAAAAGATGGTGATGGAATACAA GTAACAATTTGTTTGAAGGAGAATGACATGATTGAGTTTGAGAGAGACCCTCACATTT

[0296] TGTCCTCCACTTCAAAACTAGCATTCCATTCCTTAGGATAA

[0297] SEQ ID No: 26 Sequence comprising the amino acid sequence of the SgASAT4 allele MVKPSAPTPDNLRRLKLSLFDQMDIGAYVPILFNYLPNSTSSYDHDDKLEKSLSETLTKFY PFAGRFRKGIDPFSIDCNDEGIEYVRTKVNADDLAQYLRGQAHNDIESSLIDLLPVMHRLPS SPLFGVQVNVFNNGGVTIGIQILHMVSDAFTLVKFVNEWAHTTLTGTMPLDNPGFGQLPW LFPARALPFPLPDFNTTTAPNYKNVTKRLLFDALAIENLRNTIKANDTMMKQPSRVVVVM SLIWKVLTHISSAKNNGNSRDSSLVFVVNLRGKLSCTAPSLEHVVGNCVIPATANKEGDEA RRKDDELNDFVKLVRNTIRDTCEAIGKAESVDDISSLAFNNLTKCIEKILHGDEMDFYSCSS WCGFPWYEADFGWGKPFWVSSVSFGHHGVTNLMDTKDGDGIQVTICLKENDMIEFERDP HILSSTSKLAFHSLG

[0298] SEQ ID No: 27 Genetic SNP marker that is able to identify the S1ASAT4 gene of the invention in a Solanum lycopersicum plant grown from a seed that is deposited under accession number NCIMB 44403. A genomic fragment of chromosome 1 is presented, wherein the position of a SNP comprising a change from C to G is indicated as [C / G]. The G nucleotide is present in plants grown from seeds of the said deposit.

[0299] ATGGTGAAACCCTCAGCTCCTACCCCGGATAATCTTCGGAGATTGAAGCTTTCCTTGTT CGATCAGATGGATATTGGTGCATATGTACCAATT[C / G]TCTTCAACTACTTGCCGAACA GCACTTCATCATATGATCATGATGATAAGCTTGAAAAATCATTGTCGGAGACGCTAAC CAAGTTTTACCCTTTTGCTGG

[0300] FIGURES

[0301] The invention will be further illustrated in the Examples that follow. In the description reference is made to the following figures:

[0302] Figure 1

[0303] Figure 1-1 shows a picture of the abaxial side of a tomato leaf showing a resistant phenotype to infection with Bemisia tabaci. The plant is clean or with only very few whitefly pupae.

[0304] Figure l-2a and figure l-2b show a picture of the abaxial side of a tomato leaf showing a resistant to slightly infested phenotype to infection with Bemisia tabaci. This is considered a very mild infection, only a few pupae can be found, scattered over a few leaves.

[0305] Figure 1-3 shows a picture of the abaxial side of a tomato leaf showing an intermediate resistance phenotype to infestation with Bemisia tabaci. A few leaves are quite densely covered or several leaves moderately covered with pupae. Figure 1-4 shows a picture of the abaxial side of a tomato leaf showing a susceptible phenotype to infestation with Bemisia tabaci. This is considered a quite severe infection with several leaves heavily covered with pupae and the rest of the plant moderately covered with pupae.

[0306] Figure 1-5 shows a picture of the abaxial side of a tomato leaf showing a very susceptible phenotype to infestation with Bemisia tabaci. Many leaves are found that are heavily covered with pupae. Plants are often starting to get covered by sooty mold.

[0307] EXAMPLES

[0308] EXAMPLE 1

[0309] Identifying increased type IV trichome density and resistance to whitefly in wild relatives of tomato A phenotypic screen was performed on multiple accessions of wild tomato relatives. The aim of this screen was to identify tomato plants and / or plants of wild relatives of tomato which showed increased type IV trichome density and resistance or improved resistance to Bemisia tabaci. The screen was successful, and an accession of tomato wild relative Solanum galapagense was selected. This plant showed an increased density of type IV trichomes on leaves, petioles and the stem together with a clear resistance to Bemisia tabaci,

[0310] A population of recombinant inbred lines (RILs) was generated and a mapping project was then initiated to elucidate which genetic variation caused the resistance to Bemisia tabaci as observed in the .S', galapagense accession. This plant was crossed with a Rijk Zwaan proprietary .S', lycopersicum line not showing an increased density of type IV trichome nor showing resistance to Bemisia tabaci. A QTL on chromosome 2 was identified. Fine-mapping of this QTL resulted in a 162.8kb interval on said chromosome on the Solanum lycopersicum reference genome. It was found that 18 genetic annotations were located in this region.

[0311] Available sequence data were compared for the resistant wild relative and the susceptible .S', lycopersicum proprietary line. First, the region was found to be highly syntenic between the two plants. However, a duplicated region upstream of the AP2c gene was found. In this intergenic region with a size of approximately 30 kb in the wild relative source, it appeared that a IncRNA was expressed. This transcript was not found in expression data of susceptible sources.

[0312] As other candidate genes were not expressed in the resistant source or were not deemed logical and relevant candidates, the focus was put on the duplicated region including the AP2c gene. Different recombinants were obtained in order to identify the causal gene. It appeared that the candidate genes upstream of the polynucleotide of the invention were not necessary for the phenotype of the invention. Also, recombinants within the polynucleotide of the invention were found. It appeared that only recombinants with both the IncRNA as well as the AP2c gene of the wild relative source showed the increased density of type IV trichomes and hence are resistant to Bemisia tabaci. Plants lacking the IncRNA but having the AP2c gene from the wild relative present, did not show the increased density of type IV trichomes and were not found resistant to Bemisia tabaci. Vice versa, if plants did comprise the IncRNA derived from the wild relative but comprised the AP2c gene from Solarium lycopersicum, they did neither show the increased type IV trichome phenotype nor the resistance to Bemisia tabaci.

[0313] Table 3 describes the marker scores and the increased type IV trichome density and Bemisia tabaci resistance phenotype for several plants described above. Markers 1 to 8 are able to distinguish SNPs observed between Solarium lycopersicum and the wild relative from which the polynucleotide of the invention is derived. These markers are located upstream of the polynucleotide (marker 1 approximately 43 kb and marker 8 approximately 1.3 kb) but are not associated with the observed phenotype. SEQ ID no. 6 to 20 are markers as disclosed herein and can be used for the identification of the polynucleotide. They span a region of less than 16 kb.

[0314] RZ proprietary line 2 is a line that does not show the increased type IV trichome density phenotype and is not resistant to Bemisia tabaci. All shown markers score A for this line, which means that the introgression comprising the polynucleotide of the invention is absent. When the introgression is present, including the polynucleotide, this results in B scores for all shown markers. The type IV trichome density is significantly increased and the plant is resistant to Bemisia tabaci.

[0315] Then a total of 7 recombinant lines are shown. These are derived from a cross between the RZ proprietary line 2 and the wild relative which is the source of the polynucleotide of the invention. Recombinant line 1 has a haplotype comparable to RZ proprietary line 2 and a corresponding phenotype. Recombinant line 2 comprises several markers relating to the introgression (marker 1 to SEQ ID no. 13), but not the complete polynucleotide is present. This line also does not show the phenotype of the invention. Recombinant line 3 comprises several markers relating to the introgression (SEQ ID no. 17 to SEQ ID no. 20) but also does not show the increased type IV trichome density phenotype and is not resistant to Bemisia tabaci. Recombinant line 4 does comprise the introgression and shows the phenotype of the invention. The haplotype of recombinant line 5 is almost comparable to recombinant line 3 and has a corresponding phenotype. Recombinant line 6 shows that a recombination between marker 8 and SEQ ID no. 6 is sufficient to show the increased type IV trichome density and the Bemisia tabaci resistance (provided that SEQ ID no. 6 to 20 score the B haplotype). Recombinant line 7 was an additional recombinant which shows a haplotype and a phenotype that is in line with the recombinant lines 4 and 6 mentioned above.

[0316]

[0317] Table 3: SNP marker scores for several plant lines and the observed phenotypes for the increased type IV trichome density and Bemisia tabaci resistance. An ‘A’ score corresponds to the nucleotide as present in Solanum lycopersicum, whereas the a ‘ B’ score corresponds to the nucleotide as present in the wild relative. The B scores are indicated with a grey color for easy reference.

[0318] EXAMPLE 2

[0319] Introduction of polynucleotide of the invention in Solatium lycopersicum plants results in plants that show an increased number of type IV trichomes

[0320] In order to demonstrate that introduction of the nucleotide of the invention results in Solanum lycopersicum plants that show an increased number of type IV trichomes, the polynucleotide was introduced in three proprietary Solanum lycopersicum lines that were used before for the creation of Fl hybrid Solanum lycopersicum plants. Such proprietary lines are genetically very stable, as the Fl hybrid variety obtained by crossing a father and mother parental line should be reliable as such varieties are used by growers.

[0321] Young plant tests

[0322] The young plant tests were performed in a greenhouse in De Lier, The Netherlands. Plants were sown on day 0 and the germination and first growth took place at 25 °C both day and night. On day 12 plants were planted out in single pots and on day 19 the plants were again planted out in larger pots. On day 20 the night temperature was lowered to 23°C. After inoculation with 50 whiteflies per plant on day 25, the temperature was maintained to 25°C during the day and 23°C during the night. The type IV trichome phenotype was observed on day 39, so the plants were approximately 6 weeks old. For the young plants, both the leaf and the petiole were observed separately.

[0323] A comparison was made of three distinctive Rijk Zwaan proprietary Solanum lycopersicum lines, designated line 1, line 2 and line 3, respectively, with and without the polynucleotide of the invention. The polynucleotide of the invention was introduced in each of the proprietary lines by introgression and backcrossed several times. The presence of the polynucleotide was confirmed by using the markers as disclosed herein. It was confirmed that the polynucleotide was present homozygously in the proprietary lines. In Table 4 below, the results of observing the type IV trichome density in Rijk Zwaan proprietary lines 1, 2 and 3 with and without the polynucleotide of the invention are shown. For each proprietary line with and without the polynucleotide of the invention 3 replicates (of 1 plant) were observed. Table 4: observation of the type IV trichome density on leaf and petiole of Rijk Zwaan proprietary Solcinum lycopersicum lines with and without the polynucleotide of the invention observed in young plant stage. Indicated are the average scores according to Table 1.

[0324] A further comparison was made between three distinctive Rijk Zwaan hybrids designated Alamina RZ, Ramyle RZ and Raymos RZ respectively, with and without the polynucleotide of the invention. These Solanum lycopersicum hybrids do not comprise the polynucleotide of the invention. By obtaining parental lines that do comprise the polynucleotide of the invention, comparable hybrids were obtained that do comprise the polynucleotide of the invention. This was confirmed by using the markers as disclosed herein. It was further confirmed that the polynucleotide was present homozygously in the Rijk Zwaan hybrids. In Table 5 below, the results of observing the type IV trichome density in Rijk Zwaan hybrids Alamina, Ramyle and Raymos with and without the polynucleotide of the invention are shown. For each hybrid without the polynucleotide of the invention 3 replicates (of 1 plant) were observed and for each hybrid with the polynucleotide 4 replicates (of 1 plant) were observed.

[0325] Table 5: observation of the type IV trichome density on leaf and petiole of Rijk Zwaan Fl Solanum lycopersicum hybrids with and without the polynucleotide of the invention, observed in young plant stage. Indicated are the average scores according to Table 1. Adult plant tests

[0326] The adult plant tests were performed in a plastic greenhouse (tunnel) in Almeria, Spain. Plants were sown and grown out to adult plants stages. The type IV trichome density was scored on either the leave or the petiole and no specific differences were observed.

[0327] A comparison was made of three distinctive Rijk Zwaan proprietary Solcinum lycopersicum lines, designated line 1, line 2 and line 3, respectively, with and without the polynucleotide of the invention. The polynucleotide of the invention was introduced in each of the proprietary lines by introgression and backcrossed several times. The presence of the polynucleotide was confirmed by using the markers as disclosed herein. It was confirmed that the polynucleotide was present homozygously in the proprietary lines. In Table 6 below, the results of observing the type IV trichome density in Rijk Zwaan proprietary lines 1, 2 and 3 with and without the polynucleotide of the invention are shown. For each proprietary line with and without the polynucleotide of the invention 12 replicates were observed: twelve plants per genotype were grown in a separate area.

[0328] Table 6: observation of the type IV trichome density on leaf and petiole of Rijk Zwaan proprietary Solanum lycopersicum lines with and without the polynucleotide of the invention observed in adult plant stage. Indicated are the average scores according to Table 1.

[0329] A further comparison was made between three distinctive Rijk Zwaan hybrids designated Alamina RZ, Ramyle RZ and Raymos RZ, respectively, with and without the polynucleotide of the invention. These Solanum lycopersicum hybrids do not comprise the polynucleotide of the invention. By obtaining parental lines that do comprise the polynucleotide of the invention, comparable hybrids were obtained that do comprise the polynucleotide nucleotide of the invention. This was confirmed by using the markers as disclosed herein. It was further confirmed that the polynucleotide was present homozygously in the Rijk Zwaan hybrids. In Table 7 below, the results of observing the type IV trichome density in Rijk Zwaan hybrids Alamina, Ramyle and Raymos with and without the polynucleotide of the invention are shown. For each hybrid with and without the polynucleotide of the invention 2 replicates were observed. One replicate means that twelve plants per genotype were grown in a separate area.

[0330] Table 7: observation of the type IV trichome density on leaf and petiole of Rijk Zwaan Fl Solcinum lycopersicum hybrids with and without the polynucleotide of the invention, observed in adult plant stage. Indicated are the average scores according to Table 1.

[0331] For both the young plant stage as well as the adult plant stage a clear increase in trichome IV density is observed between both proprietary lines and hybrids when the polynucleotide of the invention is present. The proprietary lines and hybrids that do not comprise the polynucleotide of the invention hardly show any type IV trichomes, but when the polynucleotide is introduced the number of trichomes significantly increases.

[0332] EXAMPLE 3

[0333] Introduction of polynucleotide of the invention in Solanum lycopersicum plants results in plants that have resistance to Bemisia tabaci.

[0334] After demonstrating that the presence of the polynucleotide of the invention resulted in an increased density of type IV trichomes in example 2, it was also studied whether the same plants were also resistant to infestation with Bemisia tahaci. Also here, Solanum lycopersicum plants in a young plant stage were tested separately from the corresponding plants in adult plant stages. All tests and data shown below correspond to non-preference (also no-choice) tests. This means that the Bemisia tabaci whiteflies were only allowed to roam on plants of the same genotype and could not select other plants.

[0335] Young plant tests

[0336] The young plant tests were performed in a greenhouse in De Lier, The Netherlands. Plants were sown on day 0 and the germination and first growth took place at 25 °C both day and night. On day 12, plants were planted out in single pots and on day 19 again the plants were planted out in larger pots. On day 20, the night temperature was lowered to 23 °C. The cages with plants were inoculated with approximately 50 whiteflies per plant on day 25, after which the temperature was maintained at 25 °C during the day and 23 °C during the night. The plants were pruned on day 40, meaning that the central leader and side shoots were removed. The first assessment of infestation with Bemisia tabaci was done on day 67 and the second assessment on day 74.

[0337] A comparison was made of three distinctive Rijk Zwaan proprietary Solcinum lycopersicum lines, designated line 1, line 2 and line 3, respectively, with and without the polynucleotide of the invention. The polynucleotide of the invention was introduced in each of the proprietary lines by introgression and backcrossed several times. The presence of the polynucleotide was confirmed by using the markers as disclosed herein. It was confirmed that the polynucleotide was present homozygously in the proprietary lines. In Table 8 below, the results of observing the Bemisia tabaci resistance phenotype in Rijk Zwaan proprietary lines 1, 2 and 3 with and without the polynucleotide of the invention are shown. For each proprietary line with and without the polynucleotide of the invention 3 replicates (of 1 plant) were observed.

[0338] Table 8: observation of the Bemisa tabaci resistance phenotype of Rijk Zwaan proprietary Solanum lycopersicum lines with and without the polynucleotide of the invention observed in young plant stage. Indicated are the average scores according to Table 2.

[0339] A further comparison was made between three distinctive Rijk Zwaan hybrids designated Alamina RZ, Ramyle RZ and Raymos RZ, respectively, with and without the polynucleotide of the invention. These Solanum lycopersicum hybrids do not comprise the polynucleotide of the invention. By obtaining parental lines that do comprise the polynucleotide of the invention, comparable hybrids were obtained that do comprise the polynucleotide nucleotide of the invention. This was confirmed by using the markers as disclosed herein. It was further confirmed that the polynucleotide was present homozygously in the Rijk Zwaan hybrids. In Table 9 below, the results of observing the Bemisia tabaci resistance phenotype in Rijk Zwaan hybrids Alamina, Ramyle and Raymos with and without the polynucleotide of the invention are shown. For each hybrid without the polynucleotide of the invention 3 replicates (of 1 plant) were observed and for each hybrid with the polynucleotide 4 replicates (of 1 plant) were observed.

[0340] Table 9: observation of the Bemisa tabaci resistance phenotype of Rijk Zwaan Fl Solcinum lycopersicum hybrids with and without the polynucleotide of the invention, observed in young plant stage. Indicated are the average scores according to Table 2.

[0341] Adult plant tests

[0342] The adult plant tests were performed in a plastic greenhouse (tunnel) in Almeria, Spain. Plants were sown and grown out to adult plants stages. The resistance to Bemisia tabaci was scored in two separate assessments.

[0343] A first comparison was made of three distinctive Rijk Zwaan proprietary Solanum lycopersicum lines, designated line 1, line 2 and line 3 respectively, with and without the polynucleotide of the invention. The polynucleotide of the invention was introduced in each of the proprietary lines by introgression and backcrossed several times. The presence of the polynucleotide was confirmed by using the markers as disclosed herein. It was confirmed that the polynucleotide was present homozygously in the proprietary lines. In Table 10 below, the results of observing the Bemisia tabaci resistance phenotype in Rijk Zwaan proprietary lines 1, 2 and 3 with and without the polynucleotide of the invention are shown. For each proprietary line with and without the polynucleotide of the invention one or more replicates were observed. One replicate means that twelve plants per genotype were grown in a separate area. This is indicated in the table. Table 10: observation of the Bemisa tabaci resistance phenotype of Rijk Zwaan proprietary Solcinum lycopersicum lines with and without the polynucleotide of the invention observed in adult plant stage. Indicated are the average scores according to Table 2.

[0344] A further comparison was made between three distinctive Rijk Zwaan hybrids designated Alamina RZ, Ramyle RZ and Raymos RZ, respectively, with and without the polynucleotide of the invention. These Solanum lycopersicum hybrids do not comprise the polynucleotide of the invention. By obtaining parental lines that do comprise the polynucleotide of the invention, comparable hybrids were obtained that do comprise the polynucleotide nucleotide of the invention. This was confirmed by using the markers as disclosed herein. It was further confirmed that the polynucleotide was present homozygously in the Rijk Zwaan hybrids. In Table 11 below, the results of observing the Bemisia tabaci resistance phenotype in Rijk Zwaan hybrids Alamina, Ramyle and Raymos with and without the polynucleotide of the invention are shown. For each hybrid with and without the polynucleotide of the invention 2 replicates were observed. One replicate means that twelve plants per genotype were grown in a separate area.

[0345] Table 11: observation of the Bemisa tabaci resistance phenotype of Rijk Zwaan Fl Solanum lycopersicum hybrids with and without the polynucleotide of the invention observed in adult plant stage. Indicated are the average scores according to Table 2. For both the young plant stage as well as the adult plant stage a clear increase in resistance to Bemisia tabaci is observed between both proprietary lines and hybrids when the polynucleotide of the invention is present. The proprietary lines and hybrids that do not comprise the polynucleotide of the invention show at best intermediate resistance phenotypes, but when the polynucleotide is introduced, the resistance to Bemisia tabaci significantly increases. This shows that the presence of the polynucleotide does not only result in an increase in type IV trichome density, but also in resistance to Bemisia tabaci.

[0346] EXAMPLE 4

[0347] Introduction of polynucleotide of the invention in combination with the SIASAT4 gene of the invention in Solanum lycopersicum plants results in plants showing type IV trichomes producing an increased ratio ofS4:17 / S3:15 acyl sugars that have resistance to Bemisia tabaci.

[0348] After the finding that the S1ASAT4 gene of the invention was responsible for the increased ratio of S4: 17 / S3: 15 acyl sugars plants comprising the polynucleotide of the invention and the favorable SIASAT4 allele or the unfavorable SIASAT4 allele. Resistance to Bemisia tabaci was tested in a young plant test as described above in Example 2 and 3 (only the second assessment is shown). Also, acyl sugars were measured according to the details provided below.

[0349] Acyl sugar measurements

[0350] Leaf disks were taken from leaves from the 3rd youngest internode from a new shoot. Acyl sugars were extracted by dipping 6 leaf disks (3.4 cm2) in an acetonitril / isopropanol / mQ (3:3: 1, v / v) solution with propyl 4-hydroxybenzoate (10 pg / ml) as internal standard. After 60-90 minutes, the supernatant was pipetted in a 1.5mL safe-lock tube (Eppendorf® Safe-Lock). The supernatant was then filtered on Acrodisc® 0.2 pm WWPTFE filter. 300 pl of filtrate was pipetted into a 1.5 mL glass vial with 300 pl insert.

[0351] Metabolites were analysed by an UHPLC system (W aters® Acquity H-class) coupled with a photodiode array detector. Using an auto-sampler set at 5°C, 2 pL of extract was injected into UHPLC column Waters Acquity® HSS Cl 8 100 mm x 2.1 mm, 1.8 pm maintained at 35 °C. The mobile phase solution was composed of 100% methanol (solution A) and 100% water with 0.15% formic acid (v / v) (solution B). The elution flow rate was 0.3 ml / min with the following gradient: B: 0-2.90 min, 95-50%; 2.90-16.17 min, 50-5%; 16.17-30.00 min, 5%; 30.00-35.00 min, 5-95%; and equilibration from 35.00-41.00 min, 95% B.

[0352] The UHPLC system was coupled with a time of flight-mass spectrometer (Waters® Xevo G-2S) equipped with an electrospray ionization source in negative mode. The source temperature was set at 100°C. The ion transfer capillary temperature was set at 300°C and needle voltage at 2.0 kV. Nitrogen was used as nebulizing gas with the desolvation gas flow rate at 800 1 / h and cone gas flow rate at 25 1 / h. The spectra were acquired within mass ranging from 50-1200 Da, with no collision in low energy and ramp collision from 15 to 45 eV in high energy. Data acquisition and processing were performed using MassLynxTM and TargetLynxTM software. Results were expressed in propyl 4-hydroxybenzoate equivalent in nanograms per cm2leaf area.

[0353] A comparison was made of three distinctive Rijk Zwaan proprietary Solcinum lycopersicum lines, designated line 1, line 2 and line 3, respectively, with and without the polynucleotide of the invention. Also, three distinctive Rijk Zwaan proprietary Solanum lycopersicum lines, designated line 4, line 5 and line 6, with and without the polynucleotide of the invention were included. Lines 1, 2 and 3 further comprised the SIASAT4 gene of the invention, being the favorable SIASAT4 allele. Lines 4, 5 and 6 comprised the unfavorable SIASAT4 allele

[0354] The polynucleotide of the invention was introduced in each of the proprietary lines by introgression and backcrossed several times. The presence of the polynucleotide was confirmed by using the markers as disclosed herein. It was confirmed that the polynucleotide was present homozygously in the proprietary lines.

[0355] The presence of the favorable or the unfavorable SIASAT4 allele was confirmed using the marker as disclosed herein.

[0356] In Table 12 below, the results of observing the Bemisia tabaci resistance phenotype in Rijk Zwaan proprietary lines 1 to 6 with and without the polynucleotide of the invention are shown. Also the total amounts of the S3: 15 and S4: 17 acyl sugars expressed in propyl 4- hydroxybenzoate equivalent in nanograms per cm2leaf area are indicated. Further, the marker score for the marker with SEQ ID No. 27 is indicated for the lines. For each proprietary line with and without the polynucleotide of the invention 4 replicates (of 1 plant) were observed.

[0357] Table 12: observation of the Bemisa tabaci resistance phenotype of Rijk Zwaan proprietary Solcinum lycopersicum lines with and without the polynucleotide of the invention observed in young plant stage. Indicated are the average scores according to Table 2. Further, results of acyl sugar measurements are provided for S3: 15 and S4: 17. Also the score for the marker with SEQ ID no. 27 is provided.

[0358] The proprietary lines 1, 2 and 3 that do comprise the SIASAT4 gene of the invention (the favorable allele) show, when the polynucleotide is introduced, in all genotypes a higher amount of acyl sugars present, but also a high conversion of S3: 15 to S4: 17 type acyl sugars. This also directly correlates with an increased resistance to Bemisia tabaci, because of the presence of the SIASAT4 gene of the invention as indicated with the marker score.

[0359] Further, the proprietary lines 4, 5 and 6 that do not comprise the SIASAT4 gene of the invention but the unfavorable allele, show a low or no conversion of S3: 15 to S4: 17 acyl sugars in comparison to the lines that do comprise the favorable allele. Although increased resistance to Bemisia tabaci is observed when the polynucleotide of the invention is introduced, for these plants a combination of the polynucleotide of the invention and the SIASAT4 gene of the invention is essential to obtain resistance to Bemisia tabaci.

Claims

CLAIMS1. Polynucleotide comprising a IncRNA gene according to SEQ ID No: 1, or according to a sequence having at least 50% sequence identity to SEQ ID No: 1 and an AP2c gene according to SEQ ID No: 2, or according to a sequence having at least 50% sequence identity to SEQ ID No: 2, which polynucleotide confers an increased density of type IV trichomes.

2. A plant comprising the polynucleotide as claimed in claim 1, which plant shows an increased density of type IV trichomes, when compared to a control plant not comprising the polynucleotide.

3. The plant as claimed in claim 2, wherein the increased density of type IV trichomes is observed on leaves and / or petioles and / or the stem of the plant.

4. The plant as claimed in claim 2 and 3, further comprising a SIASAT4 gene according to SEQ ID No. 21, or comprising a homologous sequence having at least 99.6% sequence identity to SEQ ID No. 21, which SIASAT4 gene confers an increased ratio of S4: 17 / S3: 15 acyl sugars produced in glandular trichomes present on said plant.

5. A plant as claimed in claim 4, which plant is resistant to Bemisia iabaci. when compared to a control plant not comprising the polynucleotide according to claim 1 and the SIASAT4 gene according to claim 4.

6. The plant as claimed in any of claims 2 to 5, which plant is a plant from the Solanaceae family, preferably a plant of the Solcinum genus, more preferably a plant of the Solanum lycopersicum species.

7. A Solanum lycopersicum plant as claimed in claim 6, wherein the polynucleotide as claimed in claim 1 and the SIASAT4 gene as defined in claim 4 are as present in the genome of a Solanum lycopersicum plant, representative seed of which was deposited with the NCIMB under deposit number NCIMB 44403.

8. A Solanum lycopersicum plant as claimed in claims 6 or 7, wherein the polynucleotide and the SIASAT4 gene, are introgressed from a plant grown from a seed deposited with the NCIMB under deposit number NCIMB 44403 or from a progeny plant thereof.

9. A marker for the identification of the polynucleotide as claimed in claim 1 and / or a marker for the identification of the SIASAT4 gene according to claim 4, wherein the marker for the identification of said polynucleotide is selected from the group of SEQ ID No. 6 to SEQ ID No. 20 and wherein the marker for the identification of said SIASAT4 gene is SEQ ID No. 27.

10. Use of a marker according to claim 9 for the identification of the polynucleotide according to claim 1 and / or for the identification of the SIASAT4 gene according to claim 4.

11. Use of a marker for the identification of a Solanum lycopersicum plant showing an increased density of type IV trichomes and comprising glandular trichomes producing an increased ratio of S4: 17 / S3: 15 acyl sugars, in particular a marker as claimed in claim 9.

12. Use of a marker in the development of a Solanum lycopersicum plant showing an increased density of type IV trichomes and comprising glandular trichomes producing an increased ratio of S4: 17 / S3: 15 acyl sugars, in particular a marker as claimed in claim 9.

13. Use of a marker for the identification or in the development of a Solanum lycopersicum plant as claimed in claim 11 and 12, which plant is resistant to Bemisia tabaci.

14. Seed comprising the polynucleotide according to claim 1 and the SIASAT4 gene as defined in claim 4.

15. Seed that can grow into a plant according to any of the claims 2 to 8.

16. Propagation material capable of developing into and / or being derived from a plant as claimed in any one of the claims 2 to 8, wherein the propagation material is suitable for sexual reproduction, and is in particular selected from a microspore, pollen, an ovary, an ovule, an embryo sac and an egg cell, or is suitable for vegetative reproduction, and is in particular selected from a cutting, a root, a stem cell, and a protoplast, or is suitable for tissue culture of regenerable cells or protoplasts, which regenerable cells or protoplasts are in particular selected from a leaf, pollen, an embryo, a cotyledon, a hypocotyl, a meristematic cell, a root, a root tip, an anther, a flower and a stem, and wherein the propagation material comprises the polynucleotide as claimed in claim 1 and the SIASAT4 gene as defined in claim 4.

17. Method for identifying a plant of the Solanum genus showing an increased density of type IV trichomes and comprising glandular trichomes producing an increased ratio of S4: 17 / S3: 15 acyl sugars, wherein the method comprises screening a population of plants of the Solanum genus for the presence of the polynucleotide according to claim 1 and the presence of the SIASAT4 gene as defined in claim 4, and identifying the plant of the Solanum genus that comprises said polynucleotide and said SIASAT4 gene as plant of the Solanum genus showing an increased density of type IV trichomes and comprising glandular trichomes producing an increased ratio ofS4: 17 / S3: 15 acyl sugars.

18. Method according to claim 17, further comprising the step of phenotypically screening for an increased density of type IV trichomes and for the presence of glandular trichomes producing an increased ratio of S4: 17 / S3: 15 acyl sugars.

19. Method for selecting a plant of the Solanum genus showing an increased density of type IV trichomes and comprising glandular trichomes producing an increased ratio of S4: 17 / S3: 15 acyl sugars, comprising performing the method of claim 17 or 18, further comprising the step of selecting a Solanum lycopersicum plant having resistance to Bemisia tabaci.

20. Method for producing a Solanum lycopersicum plant showing an increased density of type IV trichomes and comprising glandular trichomes producing an increased ratio of S4: 17 / S3: 15 acyl sugars, comprising the step of introducing the polynucleotide according to claim 1 and the SIASAT4 gene as defined in claim 4 in the plant, wherein the polynucleotide confers an increased density of type IV trichomes when compared to a control Solanum lycopersicum plant lacking the polynucleotide and wherein the SIASAT4 gene confers glandular trichomes producing an increased ratio of S4: 17 / S3: 15 acyl sugars when compared to a control Solanum lycopersicum plant without the S1ASAT4 gene.

21. Method according to claim 20, wherein the polynucleotide and the SIASAT4 gene are introduced by cisgenesis.

22. Method as claimed in claim 20 and 21 wherein the increased density of type IV trichomes and the glandular trichomes producing an increased ratio of S4: 17 / S3: 15 acyl sugars, together do result in resistance to Bemisia tabaci.

23. Method for producing a Solanum lycopersicum plant showing an increased density of type IV trichomes and comprising glandular trichomes producing an increased ratio of S4: 17 / S3: 15 acyl sugars, said method comprising: a) crossing a Solanum lycopersicum plant comprising the polynucleotide as claimed in claim 1 and the SIASAT4 gene as defined in claim 4, with another Solanum lycopersicum plant to obtain an Fl population; b) optionally performing one or more rounds of selfing and / or crossing a Solanum lycopersicum plant from the F 1 to obtain a further generation population; and c) selecting from the population a Solanum lycopersicum plant that comprises the polynucleotide that confers an increased density of type IV trichomes and the SIASAT4 gene that confers glandular trichomes producing an increased ratio of S4: 17 / S3: 15 acyl sugars.

24. Method for the production of hybrid seed of a plant belonging to the Solanaceae family comprising crossing a first parent plant and / or the second parent plant wherein the first and / or the second parent plant is a plant comprising a polynucleotide as claimed in claim 1 and the SIASAT4 gene defined in claim 4, and wherein the presence of said polynucleotide leads to an increased density of type IV trichomes in a plant that is grown from the hybrid seed, and wherein the presence of said SIASAT4 gene leads to glandular trichomes producing an increased ratio ofS4: 17 / S3: 15 acyl sugars25. Method as claimed in claim 24, wherein the hybrid seed is a seed of the species Solanum lycopersicum.

26. Method for growing a plant, in particular a plant of the Solanaceae family, that shows an increased density of type IV trichomes and that comprises glandular trichomes producing an increased ratio of S4: 17 / S3: 15 acyl sugars, the method comprising sowing a seed, comprising thepolynucleotide according to claim 1 and the SIASAT4 gene according to claim 4, and allowing the plant to grow.

27. Method according to claim 26, wherein the plant of the Solanaceae family is a Solcinum lycopersicum plant

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