Spontaneous mutational burst or mass occurrence of mutations in plants is extremely unusual phenomena in the nature. For culture of a tomato its description and studying to us are not known, that became the сause for research of casually arisen phenomenon in experimental selection work.

Technique and conditions of experiments. The not segregated line of a tomato of mutant origin, Novichock-mini-j-2 of kind Solanum lycopersicum L., characterized by genes sp (determinate habit), o (ovate fruit), j-2 (joint in pedicle is absent), u (uniform ripening fruit). It grew up in arid conditions of the Astrakhan area by individual selections of plants and studying of their posterities on standard technology of cultivation in conditions of an irrigation [4]. Feature arid conditions consists in years day time temperatures up to 44^о, heating of ground up to 60-65^оС, relative humidity of air at midday o'clock up to 13-15 %.

The structure of a flower of a studied tomato is characterized by a short, pistil, hided in jointing column of stamens that excludes cross-pollination. Criteria of selection of mutations was: 1) founding out of individual plants - new genetic form, in which one or several distinguishable traits from the initial form have been changed. Paid attention to fact that the mutant plant keeps the basic complex of attributes of the initial form, except for mutant. 2). In posterity of a mutant there are no other attributes not peculiar to the initial form. It excludes probability of a hybrid origin of a mutant.

For arising mutations at a tomato we use the corresponding symbols of genes accepted in genetics of a tomato for a designation of similar mutations in a world collection of mutants [22]. Verification of conformity of the mutant attributes which have arisen in experiences with their description in a genetic collection is led under full last published catalogue of the Center of genetic resources of a tomato [20]. All the found out mutant genes, except for 2, are used in breeding of a tomato, are present at collection samples and their identification does not represent difficulties. The binding, to chromosomes of the spontaneous mutant genes which have arisen in experiments is carried out according to a classical map of last full variant of its edition [22] with the correction which has concerned localization of a gene j-2 in 12, instead of in 11 chromosome, after works V.A. Тuinen et al [21] and editions TGC in 2006 y"s [14]. Localization of 4 genes on a chromosome 2 is resulted on the genetic map reconsidered and specified G. Bishop et al. [13]. Also conclusions Ku et al. [17] and Liu et al. [18] that the genes causing attributes oval, round, elongated and pear-shaped form of a fruit everyone concern to a locus ovate and mutations of the oval form conduct to the elongated and pear-shaped form are took into consideration.

Results of researches and discussion. In one of experiments of 2009y. in the mutant not segregated line of tomato Novichok-mini-j-2 of kind Solanum lycopersicum L. has been found out and within 4th years spontaneous mutational burst which has arisen is studied. It simultaneously with disease of plants by a tomato cluster deformations of a top of stem virus VKDVST (TCDTSV) is accompanied [3]. Mutational burst was inherited in 3 studied generations in which it is fixed 9 dominant - s+ (simple cluster), y+ (yellow skin), gs+ ( restored normal red color), j+ (pedicle with joint), u+ (green spot on the fruit base), o+ (round fruit), sp+ (indeterminate habit). cdv+ (resistance to TCDTSV) and dfs+ (diminished fruit size) and 12 recessive genes - c (potato leaves) , s (complicated cluster), y (colorless epidermis), bk (beak on the fruit), el (elongated fruit), j-2, sp, gs (green stripe), d-2 (shortened internodes and leaves) , ms (male sterility), efs (enlarged fruit size), psh (pear shape of fruit). The gene symbols of the traits - pear shaped fruit, diminished fruit, enlarged fruit , and resistance to TSDTSV (dominance is confirmed in F1) were bring into use by us, because of they there are not in the nomenclature list of genes. Genes with known localization are placed in sites of 6 different chromosomes: 1 (y, y+), 2 (s. s+, d, o+, el, bk), 6 (c, sp+), 7 (gs, gs+), 10 (u+), 12 (j+, j-2). Localization of the gene d was tied to chromosome 2 because of it characteristic traits. Localization of 4 genes, ms, cdv+, dfs+, efs , while is unknown. In the literature there are information that mutations of pollen sterility (ms) in many variants [15], mutations of enlarged and diminished fruit size [1] and also pear shape fruit [17,18] arise spontaneously. It was annually fixed 12 - 14 gene mutations: dominant into recessive, and recessive into dominant in 4 - 6 chromosomes.. Frequency of mutations in 2010-2012 y"s. changed from 16,75 up to 28,50 % and has on the average made 22,82 % or 1 on 4,4 plants. It is in 23-227 thousand times more, than is registered at eukaryote, including plants, at usual spontaneous mutagenesis [5,6].

On 4-th pairs genes s- s+, y- y+, gs- gs+, j-2- j+ states of alleles: dominant-recessive -dominant are noted. In the latter case the dominant mutation was sometimes accompanied by a new mutation. In relation to plants with opposite allele dominant mutations (reversions) on studied alleles arose much more often, than recessive. On the average distinctions have made 3,36 times.

As it is known the spontaneous mutagenesis , as rule, is caused by mobile genetic elements (MGE). The genetic analysis many spontaneous dominant and recessive mutations, types of their arising, frequent reversions, changes of alleles , their several consecutive alternation in posterity of a tomato with manifestation of mutant phenotype on the level of whole organism, give possibility conclude about participation in the mutagenesis the forms MGE, named by us transposons sexual cells (ТSC). Feature of transposons consists that, unlike retrotransposones they are moving directly as DNA elements of chromosomes [8]. The carried out experiments show, that TSC regulate a state of dominant and recessive genes or their allele changes at a level of the whole organism. Process of transpositions such TSC in plant genomes is already completed in the formed sexual cells and the zygote bears in itself mutations of the whole organism, as a rule, in a homozygous condition. On such genetic display TSC differ from earlier known controlling elements (КЭ), adjusting by type of himeric somatic spottiness tissue of plants - seeds, leaves, flowers [11,19]. The cytologic researches B. McClintock led on corn, it has been shown, that moving КЭ into site of dominant gene blocks it in recessive, and the subsequent moving the insertion КЭ restores them in dominant state.. Disappearance insertion in one locus of a chromosome is connected with the appearance of it in other locus [19.] Controlling elements in essence are somatic transposons [9]. In our experiments on the big number of dominant genes their blocking in recessive state has noted been, and in the subsequent generation arose reversion genes in a dominant state with frequent occurrence of new mutations. Somatic transposons (ST or KE) and TSC are inherited in generations by means of sexual process, but transposition SТ occurs in somatic cells at formation of tissue and organs at different stages of organogenesis.. As a result there are the spots reflecting mutations of genes of painting of a tissue. The transposition or moving TSC takes place in cells of a sexual way, changing a state of gene alleles in gametes, a zygote and all developing organism, as has caused given by us it the name.

At spontaneous mutational burst all 4 precisely presented recessive genes sp, u, o, j-2 an initial experimental line the Novichok -mini-j-2 muted in dominant alleles, that together with results of other carry out our experiments [2] gives the basis to draw a conclusion that TSC , basically, are localized in sites of recessive genes and, apparently, the more spontaneous recessive genes there are in the tomato, genome there are more and ТПК. Dominant alleles arise in recessive lines at spontaneous mutagenesis, as was mentioned, with greater frequency, than recessive mutations at dominant lines. It is possible to explain this phenomenon by that transposition TSC from site of recessive gene always is easily fixed by appearing of dominant gene. But a new insertion TSC can appear in a introning , neutral zone of chromosome or in a locus causing not noticed display new recessive gene that reduces frequency fixed recessive mutations. The conformity to natural low observable by us has opposite character in comparison with earlier noted for the general mutational process [6]. In our experiments mutations arose not only as recessive and dominant genes, but also as subordinated to recessive alleles,for example, by the form of a fruit - (o +) - o - el /psh.

Spontaneous mutagenesis it is characterized by frequent restoration recessive genes in dominant, repeatability of change alleles dominant- recessive with the appearing of new recessive mutations.

All early known and noted in our experiments 9 recessive mutations are presented at List of world mutant gene fund of a tomato, - in genetic collection of mutants and are fixed in it as spontaneously arisen [14,15,20]. For these genes natural repeatability of spontaneous occurrence is characteristic. About half in 1031 mutants known in a collection, are received only by a method inducing radioactive and chemical substances. Any of fixed by us in experiments spontaneous recessive mutants does not concern to this type of occurrence. The given fact proves distinction in mechanisms artificial and natural mutagenesis. In the latter case the mechanism of repeated changes of gene alleles can be useful in evolution of plants at cyclic change of environment conditions. The dominant genes created by evolution in the latent state are kept in posterity genome. We analyzed conditions of occurrence of the mutational burst. The role of exogenous virus TCDTSV accompanying occurrence of a stream of mutations, obviously, was the most significant and could consist in its integration into the tomato genome in the form of DNA. It could become the real reason of stressful infringement of genome balance and mass transposition ТSC. The integration of a virus or its fragments in the form of DNA into tomato genome indirectly is proved by acquisition of mutant plants the attributes peculiar to virus illness, or the genes of virus caused by influence, - a high bush and a complex flower cluster..Besides of the occurrence of inherited resistance mutant plants and lines to TCDTSV is fixed, that also could occur owing to integration of the virus into tomato genome in the form of DNA and expression genes of the latent virus. It is possible to assume, that the integrated provirus has appeared in the chromosome 2 in which the greatest quantity of mutations is noted, including s- gene.

In spite of the fact that the virus is itself the mutagen factor, and also is known as carrier and transmission factor of МГЭ [7,11], the main role in observable by us spontaneous mutagenesis belonged to TSC transposons . Such conclusion follows that spontaneous mutagenesis continued close frequency and a spectrum of mutations in the selected lines, resistant to TCDTSV..Under absence of display of virus activity in the form of disease of plants and lines. the violation of genome balance at healthy tomato plants was kept, as well as connected with it mutagenesis activity TSC. Participation retrotransposons in mutational burst also, possibly, took place, but they are not capable to horizontal moving on chromosomes. They are stabilized in genome and their move occurs at low frequency [12]. In occurrence mutational burst and mass moving transposons the decisive significance had not the fact of disease of a plant by virus infection, but genome interactions between virus and plant, which violated the genome balance of tomato plants.

The share of stabilized mutant posterities at mutational burst makes 10-20 %. Mutational burst can be used for acceleration and reduction in price of selection process in 2-2,5 times at creation of varieties and lines with new economic-valuable attributes, including resistance to virus illnesses.

The conclusion. Virus disease of plants of tomato TCDTSV accompanied mutational burst or the unusual phenomenon of splash in the spontaneous mutations, inherited during 3 generations with frequency of 17-28 %, and captured the genes localized in six chromosomes. As spontaneous mutagenesis, basically it is caused by moving of mobile genetic elements (МГЭ), noted types of occurrence of mutations, frequent reversions, change alleles, their alternation in posterities with display of a mutant phenotype at a level of the whole organism, have allowed to draw a conclusion on participation in mutagenesis in tomato of forms MGE named by us transposons of sexual cells (ТПК). ТПК regulate the dominant and recessive state of genes at transposition it in chromosomes of cells of a sexual way, unlike described at plants somatic transposons, named by controlling elements. Integration virus genome in the form of DNA into genome of the tomato, apparently, was a major factor of infringement genome balance of a plant and mutational burst. Presence of TCDTSV provirus in mutant genotypes indirectly is proved by mutant plants of the tomato, bearing the attributes peculiar to virus illness.

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