Annals of the Academy of Romanian Scientists  
Series on Agriculture, Silviculture and Veterinary Medicine  
Volume 15, Number 1/2026  
ISSN 2344-2085  
30  
THE MODERN CHALLENGES AND OPPORTUNITIES FOR  
IMPROVING THE GENETIC DIVERSITY OF TOMATOES  
Milania MAKOVEI1  
Abstract. This article designated modern climatic and genetic factors as causes of the  
intensification of genetic erosion in tomatoes (Solanum lycopersicum L). The possibility  
of expanding the genetic diversity of tomatoes through the use of mutant genes in  
breeding that control a wide range of economically valuable traits is being considered.  
The potential of mutant forms as a source of resistance to abiotic stress factors (high and  
low temperatures, drought) for use in breeding as donors is demonstrated. A description  
is provided of the phenotypic expression of mutant genes controlling plant habit and  
architecture, which determine the expression of various fruit traits, including resistance to  
the most common diseases. The possibility of using certain mutant genes to expand the  
range of recombinational variability in F2 populations and to obtain transgressive tomato  
forms with a completely new combination of mutant marker and economically valuable  
traits is demonstrated.  
Keywords: Tomato (Solanum lycopersicum L.), climatic and genetic factors, mutant  
genes, selection, genetic diversity  
DOI  
1. Introduction  
The tomato (Solanum lycopersicum L.) is one of the most widely cultivated  
vegetable crops in the world (approximately 5 million hectares) [1]. Its high share  
in the structure of gross vegetable production in various countries is due to its  
ecological adaptability and the nutritional value of its fruits, which contain sugars,  
organic acids, mineral salts, vitamin C, carotenoids, lycopene, fats, carbohydrates,  
fiber, and others [22]. These qualities facilitate its use in both fresh and processed  
forms.  
However, global and local climate change on the one hand, and one-sided  
breeding aimed at creating high-yielding, uniform, transportable tomato varieties  
and hybrids that better meet general standards on the other, have led to the  
intensification of its genetic erosion. Farmers and larger producers have also  
contributed to this in their pursuit of high yields, they have ceased to grow local  
varieties with superior flavor, leading to their loss as carriers of valuable traits  
1
Doctor Habilitatus in Biological Sciences, Associate Professor, Leading Researcher Milania  
MAKOVEI, Moldova State University, Institute of Genetics, Physiology and Plant Protection, 60,  
Alexei Mateevici str., MD 2009, Chisinau. Republic of Moldova. ORCID: 0009-0009-5039-6270  
E-mail: makoveimilania@gmail.com  
 
The Modern Challenges and Opportunities for  
Improving the Genetic Diversity of Tomatoes  
31  
characteristic of the ecological niche of the Republic of Moldova. The rapid  
evolution of pathogens and pests has also contributed to this. All these factors,  
identified as modern challenges (Figure 1), led to a deficit in the genetic diversity  
f the tomatoes cultivated gene pool, a decrease in the adaptability of modern  
varieties and hybrids, and their vulnerability to biotic, abiotic, and other stress  
factors, which ultimately affected crop yields and, in particular, their quality and,  
especially, the quality of the produce.  
Figure 1. Climatic and genetic factors increasing tomato erosion and opportunities to improve its  
genetic diversity  
32  
Milania Makovei  
The increasing deficit of tomato genetic diversity indicates that the search for  
opportunities to improve and expand it is becoming a strategic direction for  
modern research. This requires a reorientation of fundamental research and  
applied breeding toward the search for, identification of, and incorporation into  
the breeding process of sources of new germplasm with higher genetic diversity.  
These may include wild and semi-cultivated tomato varieties, landraces, as well as  
hybrid populations artificially created with their participation.  
Of particular interest in this regard are spontaneously occurring and artificially  
induced mutations [22, 17, 19]. The availability of a large number of easily  
identifiable tomato mutant marker genes it possible to address a wide and diverse  
range of research questions. These include, first and foremost, genome mapping  
[18]; localization of quantitative traits [3]; identification of the characteristics of  
the heterosis effect [8]; study of recombinational variability in F2-F3 hybrid  
populations [9, 11, 16]; study of biochemical and physiological processes of plant  
development [6, 18]; and others. There are examples of their effective use in  
practical breeding to obtain new varieties with a beneficial combination of  
economically valuable traits [9, 10, 16].  
A comprehensive approach to evaluating and identifying sources of new  
germplasm, combined with the use of classical breeding methods alongside  
gametic technologies, can also contribute to improving the genetic diversity of the  
tomato. According to Pfahler (1982), the male gametophyte of the tomato is more  
sensitive to environmental conditions and changes than the female gametophyte,  
which is covered by thick layers of somatic tissues [15]. This has been confirmed  
in the works of other authors who use male gametophyte traits as criteria for  
evaluating and selecting genotypes resistant to adverse environmental factors, not  
only in tomatoes but also in other agricultural crops [4, 5, 12]. High sensitivity  
and the presence of a large amount of pollen allow for the rapid testing of  
extensive collections and the identification of genotypes resistant to one or several  
stress factors at once [10].  
Hybridization plays a key role in expanding genetic diversity as one of the most  
important methods of classical breeding. It allows us to harness heredity as the  
greatest force that drives morphological development. Interspecific and  
intergeneric crosses are particularly effective in this regard, facilitating the  
production of plant forms with entirely new combinations of valuable traits [22].  
The combined use of classical breeding and pollen analysis methods allows for  
working with large hybrid populations using a streamlined approach and  
identifying genotypes that combine economically valuable traits with resistance to  
various abiotic stress factors [10].  
Farmers can also contribute to expanding the genetic diversity of tomatoes by  
selecting, under production conditions, the most valuable varieties that are less  
susceptible to current local challenges, differ in fruit characteristics, and possess  
The Modern Challenges and Opportunities for  
Improving the Genetic Diversity of Tomatoes  
33  
high flavor qualities. Selection, conservation, and active cultivation of these  
varieties represent one potential approach to enriching the gene pool of local  
varieties with ecotypes capable of realizing their genetically determined  
productivity potential under the conditions of the Republic of Moldova.  
As we can see, there are quite a few opportunities to overcome the negative  
impact of climatic and genetic factors on the intensification of tomato erosion.  
Proper organization and their combined use will allow for the intensification of  
the search, identification, and selection of the most valuable genotypes that  
combine economically valuable traits with resistance to abiotic and biotic stress  
factors, thereby contributing to the expansion of tomato genetic diversity.  
The aim of the research was to comprehensively study the genetic potential of  
mutant marker genes of tomato for their effective use in practical breeding,  
improving varietal diversity and replenishing collections with genes controlling a  
wide range of economically valuable traits  
2. Materials and Methods  
The experimental material consisted of 125 mutant tomato lines carrying 187 mutant  
marker genes, more than 50% of which are of practical value for addressing current  
challenges in practical breeding.  
2.1. Phenotypic expression of mutant genes.  
The nature of expression and the degree of phenotypic expression of the mutant genes  
were determined in accordance with the tomato gene nomenclature [22, 20].  
Studied the nature and characteristics of the expression of marker genes controlling  
plant habit, architecture, and fruit traits (shape, color, taste).  
2.2. Screening of mutant tomato lines for resistance to abiotic stress factors.  
Testing for resistance to abiotic stress factors (high and low temperatures, drought) was  
conducted based on male gametophyte (pollen) traits using artificially simulated stress  
conditions in vitro [10]. Fresh pollen collected from the flowers of each mutant sample  
served as a control, it was germinated on an artificial nutrient medium (15% sucrose  
and 0.006% boric acid) under in vitro conditions (3 hours) at a temperature of 25°C.  
Pollen viability-germination (p, %) and pollen tube length (l, µm) were determined.  
Resistance to high (45°C) and low (6°C) temperatures, as well as drought (simulated by  
high sucrose concentration), was assessed based on the ability of pollen from  
experimental samples to germinate (pollen germination resistance), and the ability of  
germinated grains to form pollen tubes (resistance based on tube length) of a length  
sufficient for fertilization (three pollen grain diameters) [7] under the influence of these  
stressors. For each sample and in each experimental treatment, 500 pollen grains were  
evaluated in 3 replicates. Pollen preparations were examined under a microscope (Zeiss  
model, 7×20). The resistance of each genotype was determined by the ratio of the  
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Milania Makovei  
indices of the studied pollen traits in the experimental variants (E) to the control (C),  
expressed as a percentage (%).  
2.3. The expression of morpho biological and economically valuable traits in  
homozygous and heterozygous offspring.  
The morphological, biological, and economically valuable traits, including yield and  
fruit quality, were assessed in accordance with UPOV recommendations (Solanum  
lycopersicum L.) [21] and standard methods [14]. Hybridization was carried out  
according to schemes specifically developed by us. More than 50 F1 hybrid  
combinations were obtained. The nature of expression and the degree of phenotypic  
expression of traits controlled by mutant genes in the F1 and F2 progeny populations  
were determined following the gene nomenclature [22]. Statistical analysis of the raw  
data was performed using the computer software programs Excel and Statistica 7.  
3. Results and Discussions  
The success of breeding new tomato lines, varieties, and hybrids depends directly on  
the availability of thoroughly studied donor plants with the desired traits in collections.  
However, concepts for organizing such collections remain undeveloped, a situation  
observed not only in local collections but also in global ones [23]. According to the  
author, of the one million samples collected in global collections, less than 1% have  
phenotypic characteristics. This creates enormous difficulties for breeders, as they must  
repeatedly review large amounts of material in search of sources of the desired traits.  
Consequently, a comprehensive study of available genetic resources not only for  
immediate use but also for supplementing existing and creating new trait collections –  
is a top priority. In this regard, we conducted an assessment of the genetic potential of  
mutant tomato forms, aimed at identifying genes that control a wide range of traits at  
different stages of ontogenesis, including resistance to abiotic and biotic stress factors,  
for the subsequent expansion of existing and creation of new traits collections.  
3.1. The potential of mutant tomato varieties for resistance to abiotic stress factors,  
as determined by male gametophyte (pollen) characteristics.  
Testing of an extensive collection of materials revealed significant heterogeneity among  
mutant forms in terms of the level and type of pollen resistance to high and low  
temperatures and drought.  
First and foremost, marked differences were observed between genotypes within the  
collection regarding indicators of traits characterizing the quality of freshly collected  
pollen (control). Depending on their genotypic characteristics, the “viability-  
germination” trait varied widely (from 1.5% to 78.2%). Regarding the ability of  
germinated pollen grains to form pollen tubes of a certain length under in vitro  
conditions, the variation among them was even greater (from 11 µm to 150 µm). This  
The Modern Challenges and Opportunities for  
Improving the Genetic Diversity of Tomatoes  
35  
indicates that the same plant cultivation conditions had different effects on pollen  
quality, highlighting the genotype-specific characteristics of each.  
The differences between mutant forms were more pronounced in terms of their pollen’s  
response to high and low temperatures and drought. Pollen response was inconsistent,  
even within a single genotype, as measured by two different traits:  
“pollen germination resistance” and “resistance to pollen tube length”. The genotype-  
specific response of pollen to the effects of various factors, both individually and in  
combination, allowed for their classification into groups based on the type and level of  
resistance of each genotype to a specific stress factor (Figure 2).  
Figure 2. Characteristics of mutant tomato forms by type and level of resistance to high and low  
temperatures and drought  
Twenty-two mutant forms were identified whose pollen germinated rapidly and with  
high efficiency (54.6%97.8%), while also forming long pollen tubes (58 µm150 µm)  
under the influence of all three negative factors. This indicates the high resilience of  
their male gametes, which are capable of germinating and forming long pollen tubes  
under conditions that significantly from optimal ones.  
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Milania Makovei  
In contrast, another group of mutant forms (26 genotypes) exhibited pollen that was  
highly sensitive to all three stress factors. Both pollen germination rates and pollen tube  
lengths were very low in this group, ranging from 0.4% to 9.7% and 1.0 µm to 20.4  
µm, respectively. They were classified into the group of unstable genotypes (Figure 2).  
The third, fourth, and fifth groups comprised mutant forms (28 genotypes) whose  
pollen exhibited high resistance to only one of the stress factors studied, but was highly  
sensitive to the other two. Conversely, mutant forms (Group 6) were identified whose  
pollen germinated well, simultaneously forming very long pollen tubes under the  
influence of high temperature and drought, but exhibited high sensitivity to the low-  
temperature factor. A similar pollen response is also characteristic of the mutant forms  
distributed in groups 7 and 8 (Figure 2).  
This is undoubtedly a unique gene pool of particular value for use as a source of  
resistance to abiotic stress factors in the male gametophyte during the breeding of new  
tomato varieties and hybrids, as well as for expanding existing genetic collections and  
establishing new trait-based genetic collections.  
3.2. Genetic diversity of mutant genes that determine the growth habit and  
architecture of tomato plants.  
Plant habit is a key indicator for any variety or hybrid, determining its intended use and  
agronomic suitability.  
An evaluation and analysis of the potential of mutant forms based on the phenotypic  
expression of genes affecting plant habit revealed a high degree of diversity, but only a  
few of them have practical value.  
This diversity is mainly related to plant size, which is controlled by the genes bls, d, dd,  
dmp, dmd, sd, ssp, sp, sp+ and sp±. Morphological differences between them are  
determined both by the intensity of lateral shoot formation (ls) and the degree of their  
branching (atn, bu, br, cg, bip).  
A special group within the collection consists of forms with mutant genes ssp, br, com,  
sd, d, dd and bls, which increase the plant’s compactness. They exhibit a distinctive  
phenotype caused by mutational changes in the genome (Figure 3). These are compact  
varieties with short internodes the plants are stunted in growth. Twenty-four forms have  
been identified, which are grouped by a series of mutant genes exhibiting similar  
phenotypic manifestations.  
The clear phenotypic expression of mutant marker genes under various growing  
conditions (open field, greenhouse) made it possible to include some of them in  
hybridizations (Mo 113, Mo 443, Mo 500, Mo 504, Mo 632, Mo 755, and Mo 791).  
Crossbreeding them with lines from our own breeding program and known  
The Modern Challenges and Opportunities for  
Improving the Genetic Diversity of Tomatoes  
37  
Figure 3. Some varieties of mutant tomato plants with a bush-type growth habit  
varieties (L111, L1185, Fakel, Zagadka, Barnaulsky Konservny) and subsequent  
analysis of heterozygous F1 progeny and segregating F2 populations demonstrated  
the effectiveness of their use. Eleven F2 populations from different hybrid  
combinations were analyzed. High recombination and transgressive variability  
was established for marker traits. The pronounced phenotypic expression of traits  
such as compactness, dwarfism, limited growth, and leaf surface characteristics  
allowed for the evaluation of extensive hybrid populations at early stages of  
ontogenesis (in the seedling phase). This made it possible to accelerate the  
research by conducting it on small plots and to identify a wide variety of forms  
with genetically determined traits.  
The targeted use of some of these has made it possible to accelerate the breeding  
process and, in a short period of time, create more than 20 lines and a number of  
ornamental varieties Prichindel, Cireaska, MiracolMak, Dimetra and Ilika.  
These are intended for cultivation on balconies, loggias, and terraces, and are  
homologated in the Republic of Moldova [2].  
At the same time, the collection was found to contain genes controlling  
indeterminate (sp+), determinate (sp), and superdeterminate (ssp) plant growth  
types. Of particular interest are plant varieties with a semi-determinate (sp±)  
growth type (Mo 147, Mo 341, Mo 432, Mo 446, Mo 544, Mo 620, Mo 723, La  
1159, La 2529, La 2921, La 3179), which are widely sought after in breeding for  
heterosis to create early-maturing F1 tomato hybrids.  
Such a high degree of diversity among mutant genes that control plant growth  
patterns opens up broad opportunities for their versatile use in breeding to develop  
tomato varieties and hybrids intended for cultivation both in the field and in  
greenhouses.  
However, a significant factor reducing the efficiency of tomato cultivation in  
protected ground conditions is the excessive formation of lateral shoots, the  
removal of which requires considerable labor and financial costs. The creation of  
varieties and hybrids with low suckering potential, short internodes, and a longer  
fruiting period may help solve this problem.  
This is possible by incorporating the ls (lateral suppressor) and br (brachytic)  
genes into the breeding program. However, their direct use is complicated by their  
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Milania Makovei  
linkage to traits of the reproductive system (small fruit size, partial sterility). We  
were able to partially overcome these difficulties using a specially designed  
crossing scheme that included mutant lines (Mo 341, Mo 835, Mo 443, La 2529,  
La 3179), semi-mutant forms (111, 11069, 1751), and cultivated-type lines (L28,  
L186, L187, L828, L556). Analysis of F1 hybrids showed 100% dominance of the  
tillering ability of the cultivated parental forms in all combinations. In contrast, in  
the segregating F2 populations, high recombination variability was observed for  
the ls and br genes, confirming the conclusions of other authors regarding the role  
of mutant genes in activating genetic recombination and broadening the spectrum  
of intrapopulation variability, which facilitates the selection of new forms with  
beneficial combinations of economically valuable traits in the F2F3 generations  
[22, 9, 16].  
The complex nature of the expression of the traits “internode length” and “number  
of lateral shoots per stem” in F2 inbred populations necessitated their  
classification into a cluster complex based on breeding value in combination with  
other traits (flower structure, fruit weight, shape, and color). This allowed us to  
identify more than 50 recombinant forms with a completely new combination of  
mutant marker and economically valuable traits. Some of them (23 lines) already  
demonstrate stable expression of traits in the F3F4 generations. Eight of these  
have a semi-determinate growth habit and are characterized by early fruit ripening  
(94…109 days). Most of these lines are distinguished by weak tillering ability,  
short internodes, and uniform inflorescences, including in fruit color, shape, and  
weight. Some genotype variants with different levels of tillering ability are clearly  
shown in Figure 4.  
Figure 4. Varieties of tomato genotypes based on the pattern of lateral shoot (stepson) formation  
and internode length: 1 plants with short internodes but intense lateral shoot formation; 2 short  
internodes, single reduced lateral shoots; 3 short internodes and absence of lateral shoots; 4 –  
long internodes and absence of lateral shoots  
The Modern Challenges and Opportunities for  
Improving the Genetic Diversity of Tomatoes  
39  
The rich genetic potential of the collection of tomato mutant forms, particularly in  
terms of genes influencing plant habit and architecture, and the potential for their  
effective use in breeding to develop varieties and hybrids with novel and  
beneficial combinations of economically valuable traits demonstrates the potential  
for their use in creating trait collections and expanding genetic diversity.  
3.3. Mutant genes controlling various fruit traits and their significance for  
expanding the genetic diversity of tomatoes.  
Genes controlling fruit traits (color, taste, shape, fruit weight, fruit surface  
structure, pericarp thickness, number of seed chambers, etc.) play a special role in  
enriching the tomato’s cultivated gene pool.  
An assessment of the potential of mutant forms based on the nature of their  
expression and the degree of phenotypic expression of the genes they carry  
revealed a high degree of diversity, including unique combinations of genes that  
influence different fruit traits.  
A total of 32 gene carriers controlling unusual fruit coloration were identified:  
Abg, dg, gs, gf, r, t, y, sh, o, B, Del. Marked differences between mutant forms  
carrying the same genes were established based on the nature of expression and  
the intensity of coloration in the ripening fruit, depending on their combination  
with other genes within the same genome (Figure 5).  
Figure 5. Characteristics of fruit coloration in the mutant tomato lines Mo 922, Mo 120, and Mo  
632, which carry the t (tangerine) gene, under different combinations of this gene with other genes  
in their genomes  
Fruit coloration is determined by several genes, with the R-red flesh being  
dominant over the r-yellow, the Y-yellow skin over the y-colorless, and the P-  
smooth, shiny, hairless skin over the p-dull, hairy surface. The recessive t gene is  
responsible for orange fruit color. Most mutants in the collection (93 forms) carry  
the dominant RTY genes and have red flesh and yellow skin. Mutants with the  
double recessive genotype rrtt have orange-colored fruits. The homozygous  
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Milania Makovei  
genotypes RRTT, rrTT, and RRtt have red, yellow, and orange fruits, respectively.  
Fruit color is also influenced by the B gene. Some authors believe that the B, Bc,  
and Bog genes, which control lycopene and β-carotene content, simultaneously  
influence the color, taste, and marketability of the fruit [22, 9]. The B gene causes  
an increased β-carotene content in the fruit only in the presence of the dominant R  
gene. Therefore, the RRTTBB genotype produces β-orange or pink fruits, whereas  
the RRTTB+B+ genotype produces red ones. Yellow-fruited tomatoes have the  
genotype rrTTB+B+. In some mutant forms, the yellow flesh of the fruit may be  
determined by recessive atat genes.  
Mutant forms carrying the hp, hp-1 and Ip genes have also been described and  
isolated, these are of interest for the development of varieties characterized by  
high levels of carotenoids, ascorbic acid (vitamin C), and dry matter. According to  
available information [13], genes of the high pigment series increase the content  
of lycopene in fruits by 3545%, β-carotene by 8090%, and vitamin C by 20–  
50%, making their use essential in tomato breeding for taste qualities.  
The gs gene also imparts a distinctive coloration to the fruit. Tomatoes with this  
gene have a striped surface on both green and ripe fruit. The gs gene is easily  
transmitted, especially to large-fruited plants from its carriers. The coloration of  
the radial stripes on ripe fruits varies: golden on pink fruits; pink-red on orange  
fruits; and yellow or orange on red fruits (Figure 6).  
Figure 6. Expression patterns of traits controlled by the gs gene.  
The presence of the Del and sh genes in the tomato genome gives the fruit flesh a  
reddish-orange color, which is associated with a decrease in lycopene content and  
an increase in β-carotene.  
Varieties with the Abg and gf genes, which influence the chlorophyll content in  
the fruit, have higher levels of lycopene and β-carotene. This gives them a dark  
hue, the intensity of which depends on their interaction with other genes (Fig. 7).  
A number of other genes may also influence fruit color, but those whose  
combination can ensure high levels of vitamin C, β-carotene (provitamin A), and,  
overall, high processing and taste qualities of the fruit are of particular value for  
breeding. Of the genotypes shown in Figures 6 and 7, five are the result of our  
own breeding program using the gs, Del, and gf genes in crosses.  
The Modern Challenges and Opportunities for  
Improving the Genetic Diversity of Tomatoes  
41  
Figure 7. Characteristics of trait expression in genotypes with the Del, sh, Abg and gf genes  
The appearance of the fruit is most appealing when the fruit ripens simultaneously  
and uniformly across its entire surface. Mutant forms with the u gene, which is  
recessive to the U gene that controls the presence of a green spot at the base of  
the fruit, exhibit uniform coloring. Sometimes the green spot remains even on  
ripe fruits, which leads to a decrease in product quality. The proportion of  
mutant tomato forms with a green spot at the base of the fruit in the collection is  
quite high, at 53.6%.  
One of the most important characteristics that largely determines the direction  
and intended use of the product is the shape of the fruit. In contrast to the  
commonly accepted round fruit shape, the mutant collection includes genotypes  
with flat, ellipsoidal, pear-shaped, heart-shaped, oval, cylindrical, and other  
shapes. The high diversity of the collection is due to the presence of forms with  
different fruit surfaces (ranging from smooth to highly ribbed) (Figure 8). In  
some forms, the fruit surface is smooth, but the apical part is elongated into a  
more or less pointed tip. The nature of the manifestation and the degree of  
phenotypic expression of these and a number of other traits are controlled by the  
presence in the collection of a large number of the genes Ol, o, obl, el, f, bk, n  
and n-2.  
Figure 8. Varieties of mutations in the shape and surface of the fruit with the n, n-2, bk, el and f  
genes  
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Milania Makovei  
Mutant genes rin, nor and alc are of particular value for breeding, as their  
presence in the tomato genome prevents fruit overripening (Figure 9). These  
genes contribute to increased total yield and affect the viscosity of fruit juice,  
which is twice as high in varieties and hybrids containing these genes compared  
to conventional ones, ensuring high marketability [9].  
By using them in crosses with inbred lines with high combinatorial potential, we  
were able to obtain more than 24 combinations of F1 hybrids.  
Figure 9. Phenotypic expression of traits controlled by the nor, rin and alc genes  
Most of these exhibited high heterosis in terms of yield and fruit quality. Some  
(F1 Ingstar and F1 Rozamak) successfully passed state testing and were  
homologated in the Republic of Moldova [2]. A number of others are currently  
undergoing competitive testing. However, to improve the efficiency of breeding  
F1 heterotic hybrids involving these genes, it is recommended to use earlier-  
maturing varieties and lines with high taste qualities and large fruits as one of  
the parental forms (preferably the maternal parent).The intermediate inheritance  
of these traits by F1 hybrids balances the negative influence of the parent with  
mutant genes. In such combinations, the first-generation offspring typically  
exhibit heterosis in terms of overall productivity and fruit quality.  
The genes that control the degree of pubescence in various parts of the tomato  
plant including the fruits namely Wom, Ln, p, Vi confer exceptional ornamental  
value and special significance on the plants.  
The phenotypic expression of these genes is observed from the cotyledon leaf to  
the mature fruit, which is covered with a light fluff down (Figure 10). Mature  
fruits have a beautiful, presentable appearance and excellent taste, due to their  
high content of dry matter, sugars, and vitamin C. The fruits are intensely red  
with dense flesh, multi-chambered, and have seeds that form only in the apical  
portion of the fruit. As the fruit grows and develops, the pubescence on it  
decreases and becomes slightly visible in mature fruit (Figure 10).  
Carriers of the Wom and Ln genes (Mo 835 and Mo 341) have been incorporated  
into the breeding program. Direct and reciprocal hybrids (16 combinations) were  
obtained from crosses with homozygous lines (L8/234, L33/241, L1033) and  
The Modern Challenges and Opportunities for  
Improving the Genetic Diversity of Tomatoes  
43  
cultivated-type lines (L187, L556, L828) with high combinatorial ability, which  
are currently undergoing genetic and biochemical studies for their effective use  
in practical breeding and replenishment of collections with useful genes.  
Figure 10. The nature of fetal phenotypic expression at different stages of development,  
controlled by the Ln and Wom genes  
A high degree of heterogeneity was observed among the mutant forms, as  
evidenced by fruit weight, pericarp thickness, and the number of seed chambers.  
Fruit weight is a complex polygenic trait that varies depending on environmental  
conditions [22]. Significant differences in fruit weight were observed among the  
mutant forms: 24.6% of the forms had large and very large fruits (100…300 g);  
very small fruits (0.4…20 g) were observed in 17.8%; the remaining 57.6% of  
mutant tomato forms had fruit weights ranging from 30 to 100 grams.  
Pericarp thickness and the number of seed chambers are traits that directly  
determine fruit quality and marketability. The collection describes a large  
number of mutant genes controlling pericarp thickness, shape and seed chamber  
structure Gr, bs, bs-2, el, ck, fie, Ip, lo, loc, Ol, ptb, ss, and others. The  
phenotypic expression of some of these is shown in Figure 11.  
Figure 11. Some varieties classified by characteristics such as number, shape, color, and structure  
of the seed chambers, which are controlled by the Lc, Ip, lo, loc, Ol, bs-2 genes  
3.4. Genetic sources of resistance to certain diseases.  
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Milania Makovei  
The presence of genes controlling resistance to pathogens may help address  
problems associated with the rapid evolution of pathogens. From the total number  
of tomato mutant forms studied, 14 genotypes were identified that carry genes  
controlling resistance to Mi Meloidogyne incognita; Tm, Tm2a, Tm-2 to  
Tomato mosaic virus; F to Fusarium; Ve to Verticillium; Cf, Cf-2, Cf-3, Cf-4  
to Cladosporium. The genomes of these mutant forms contain, in various  
combinations, genes that simultaneously control resistance to a particular  
pathogen, different fruit traits, and plant growth type (primarily indeterminate and  
semi-determinate). They can be used to enrich working and long-term collections  
with valuable genes.  
Сonclusions  
(1). It is shown that the genetic potential of mutant forms, which carry a large  
number of marker genes controlling important economically valuable traits in  
tomatoes, represents a unique source of new germplasm for enhancing genetic  
diversity through the creation of new breeding materials, lines, varieties, and  
hybrids of tomato, as well as for supplementing existing and creating new trait  
collections.  
(2). A high degree of heterogeneity was observed among mutant tomato lines in  
terms of resistance to abiotic stress factors (high and low temperatures, drought).  
Genotypes with different levels and types of resistance, as determined by  
characteristics of the male gametophyte of the tomato, have been identified; under  
conditions of climate change, these are of particular value for use in breeding  
programs as sources of resistance.  
3). It was found that mutant genes, when combined with cultivated tomato lines,  
stimulate genetic recombination processes, leading to an expansion of the  
spectrum of intrapopulation transgressive variation and the emergence of new  
forms with unique combinations of mutant marker and economically traits,  
thereby contributing to the enrichment of the tomato crop gene pool and the  
expansion of its genetic diversity.  
Acknowledgments  
The research was carried of the Proiect in the field of science and innovation  
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