CHAPTER II

REVIEW OF RELATED LITERATURE

Tomato Plant

Tomatoes are summer’s fruit, yet too much light can be lethal to cultivated tomato

(Solanum lycopersicum) plants. A single genomic region, however, can make plants tolerant to

continuous light, and increase crop yields significantly, according to a study in Nature

Communications. In the 1920s, researchers discovered that the leaves of modern tomato plants

yellowed and decayed when exposed to constant light. But other plants, such as

peppers, Arabidopsis, and even species related to tomatoes showed no signs of such injuries. So

Aarón Vélez Ramírez of Wageningen University in the Netherlands and his colleagues analyzed

genomic and gene expression data from eight wild varieties of tomato, and found that the

presence of one gene on chromosome 7 was critical to the light-hardiness of these species

(Madhusoodanan, 2014).

The genus Lycopersicon includes both self-incompatible and self-compatible species,

with the latter varying in their degree of outcrossing. Tomato is self-pollinating, but can easily

hybridize within the species or cross with wild relatives under appropriate conditions, thus

permitting gene introgression from wild relatives. Moreover, tomato is considered a model

research organism because it has a relatively short life-cycle, with a new cultivar “Microtom”

being even shorter-lived (70-90 days from sowing to fruit ripening) (Dan et al. 2006).

Tomato is the center piece system for genetic and molecular research in the family

Solanaceae. Tomato has emerged as a model for fleshy fruit ripening, in part due to its ease of use

as a model system resulting from facilitating attributes including simple genetics, numerous

characterized mutants, crossfertile wild germplasm to promote genetic studies and routine
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transformation technology. Recently it has been taken for genome sequencing by an international

consortium currently funded and supported by ten contributing countries (Giovannoni et al.,

2006).

Originating from the Andes, tomatoes (Solanum lycopersicum L.) were imported to

Europe in the 16th century. At present, this plant is common around the world, and has become an

economically important crop. Furthermore, this plant is a model species for introducing

agronomically important genes into dicotyledonous crop plants (Paduchuri et al. 2010). The

tomato is considered a protective food because of its particular nutritive value, as it provides

important nutrients such as lycopene, beta-carotene, flavonoids, Vitamin C and hydroxycinnamic

acid derivatives. Furthermore, this crop has achieved tremendous popularity especially in recent

years with the discovery of lycopene’s anti-oxidative activities and anti-cancer functions. Thus,

tomato production and consumption are constantly increasing. It is noteworthy that tomatoes are

not only sold fresh, but also processed as soups, sauces, juices or powder concentrates. The

tomato ranks 7th in worldwide production after maize, rice, wheat, potatoes, soybeans and

cassava, reaching a worldwide production of around 160 million tons on a cultivated area of

almost 4.8 million hectares in 2011(Wu et al. 2011; Raiola et al. 2014)).

Health benefits of eating tomatoes emerge

Of all the non-starchy 'vegetables,'* Americans eat more tomatoes and tomato products

than any others. Researchers Britt Burton-Freeman, PhD, MS, and Kristin Reimers, PhD, RD of

the National Center for Food Safety & Technology, Illinois Institute of Technology and ConAgra

Foods, Inc., looked at the current research to discover the role tomato products play in health and

disease risk reduction. The researchers found that tomatoes are the biggest source of dietary

lycopene; a powerful antioxidant that, unlike nutrients in most fresh fruits and vegetables, has
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even greater bioavailability after cooking and processing. Tomatoes also contain other protective

mechanisms, such as antithrombotic and anti-inflammatory functions. Research has additionally

found a relationship between eating tomatoes and a lower risk of certain cancers as well as other

conditions, including cardiovascular disease, osteoporosis, ultraviolet light-induced skin damage,

and cognitive dysfunction (Freeman, 2010).

Tomatoes are widely available, people of all ages and cultures like them, they are cost-

effective, and are available in many forms. "Leveraging emerging science about tomatoes and

tomato products may be one simple and effective strategy to help individuals increase vegetable

intake, leading to improved overall eating patterns, and ultimately, better health." Research

underscores the relationship between consuming tomatoes and reduced risk of cancer, heart

disease, and other conditions," the authors conclude. "The evidence also suggests that

consumption of tomatoes should be recommended because of the nutritional benefits and because

it may be a simple and effective strategy for increasing overall vegetable intake." The article is

particularly timely since the recently released Dietary Guidelines for Americans 2010 moved

tomatoes to a newly established category of "orange/red" fruits and vegetables to encourage

higher consumption of these healthy foods (Reimers, 2010).

Botanical Point of View of Tomato

From botanical point of view, the tomato is a fruit. Nevertheless, it contains a much lower

sugar content compared to other fruits. It is a diploid plant with 2n = 24 chromosomes. The

tomato belongs to the Solanaceae family, which contains more than 3,000 species, including

plants of economic importance such as potatoes, eggplants, tobacco, petunias and peppers (Bai

and Lindhout 2007). In 1753, Linnaeus placed the tomato in the Solanum genus (alongside with

potato) under the specific name S. lycopersicum. In 1754, Philip Miller moved it to its own
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genus, naming it Lycopersicum esculentum (Foolad 2007; Perlata and Spooner 2007).

Nevertheless, the designation of the tomato was for a long time a subject of consideration and

discussion by many scientists.

The use of molecular data (genome mapping) and morphological information allowed for

the verification of of the Solanaceae classification when the genus Lycopersicon was re-

introduced in the Solanum genus in the Lycopersiconsection. Thus, after almost two centuries, the

description of Linnaeus was confirmed. Due to the use numerous citations from recent references

and in order to be consistent with much of the literature, the Linnaeus classification is followed in

this review (Foolad 2007).

Several reports indicate Peru as the centre of diversity for wild relatives of tomato. It

seems justified to considerS. lycopersicum cerasiforme as an ancestor of the cultivated tomato

because of its abundant existence in Central America (Bai and Lindhout 2007). Nevertheless,

recent extensive genetic studies have revealed that the closest relative of the tomato is Solanum

pimpinellifolium (The Tomato Genome Consortium 2012). It turned out that the genome

sequences of both of the above-mentioned tomatoes as well as domesticated cultivars and S.

pimpinellifolium showed only a 0.6 % nucleotide divergence. Domestication has triggered a

range of traits (morphological and physiological) that distinguish domesticated crops from their

wild ancestor. Studies on the domestication process, not only of tomatoes but also in the cases of

maize and rice revealed that rapid phenotypic divergence is often controlled genetically by a

relatively small number of loci (Koenig et al.2013).

Adaptation and climatic requirements of tomato

Tomato is native to tropical Central and South America, where it was cultivated in pre –

Columbian times. Its wild progenitor is thought to have been the cherry tomato, L. esculenton var.

cerasiforme, which grows wild in the Peru – Ecuador area though tomatoes were probably

domesticated from weedy forms which had spread as far as north Mexico. Tomatoes shown a

wide climatic tolerance and can be grown in the open wherever there is more than three months

of frost free weather. Tomato is more successful where there are long sunny 6 periods. The

optimum growing temperatures are 21°C to 24°C. At these temperatures good quality seeds will

take about seven days to emerge. Temperature affects flowering and pollination. The hot and dry

weather leads to drying of the flowers and stops pollination. If temperatures are below 15°C or

above 29°C, pollen release is restricted resulting in incomplete fertilization of ovules. This causes

collapsed fruit walls and formation of deep indentation in the fruit, a phenomenon called catface

(Bok et.al.,2006).

Economic Importance of Tomato

Among the members of family Solanaceae, many species of economic importance such

as tomatoes, potatoes, tobacco, peppers and eggplants can be distinguished. In recent years,

interest of scientists in the tomato as a model plant has significantly increased, also due to the fact

that its genome has been sequenced (The Tomato Genome Consortium 2012). The tomato is an

excellent model for both basic and applied research programs. This is due to it possessing a

number of useful features, such as the possibility of growing under different cultivation

conditions, its relatively short life cycle, seed production ability, relatively small genome (950

Mb), lack of gene duplication, high self-fertility and homozygosity, easy way of controlling

pollination and hybridization, ability of asexual propagation by grafting and possibility to

regenerate whole plants from different explants (Bai and Lindhout 2007).

Among the existing tomato genotypes, cv. Micro Tom is considered to be a model system

due to the aforementioned unique characteristics (Kobayashi et al. 2013). This dwarf tomato

cultivar was created for ornamental purposes and originated by crossing two cultivars (Florida

Basket and Ohio 4013-3), and shows small and ripened fruits as well as dark-green and rugose

leaves. Additional, in contrast to other model organisms such as Arabidopsis or rice, the tomato

has many interesting features. For example, tomato plants produce fleshy fruits that are important

for the human diet. The tomato has sympodial shoots, and it is the only model plant with

compound leaves. Furthermore, there exists a large pool of tomato mutants, which were either

spontaneous or induced by chemicals or irradiation that are available at the Tomato Genetic

Resource Center (Lozano et al. 2009).

Incandescent light bulb

Incandescent light bulb is a controlled fire on display. When electrical current makes

contact with the base of the bulb, electricity enters and heats the tungsten filament housed inside.

And when the filament heats up, “incandescence” is created, which is light produced by heat. The

light in an incandescent light bulb is really just that effect occurring in a contained, controlled

environment. As the filament continues to burn, particles fly off the filament. And when there are

no more particles to burn, the light bulb burns out, which typically takes place 800-1,200 hours

into the life of the incandescent light bulb. Incandescent light bulbs, in fact, produce 90 percent

heat and 10 percent light. If you have ever touched a lit incandescent bulb, you have experienced

the 90/10 heat-to-light ratio. (Cyr, 2016) 

Artificial light sources for photosynthesis

Artificial lighting should provide plants with energy and information required for

development. For this purpose, fluorescent lamps, particularly those having enhanced blue and

red spectra (i.e. cool fluorescent white lamps), are widely used in growth chambers, together with

additional light sources to achieve the sustained photosynthetic photon fluence necessary for high

productivity. However, the spectrum and intensity of fluorescent lights are not stable over a long

time (Massa GD, Emmerich JC, Morrow RC, Bourget CM, Mitchell CA. 2006).

High intensity discharge (HID) lamps, such as metal halide and high-pressure sodium

lamps, have relatively high fluence (max. 200 lumens per watt) and high photosynthetically

active radiations (PARs) efficiency (max. 40%), and are typically used in greenhouses and plant

growth rooms. Among artificial lighting systems, LEDs present the maximum PAR efficiency.

LEDs emitting blue, green, yellow, orange, red and far red are available and can be combined to

provide either high fluence, or special light wavelength characteristics, thanks to their narrow-

bandwidth light spectrum. Their long life expectancy and ease of control make them ideal for

greenhouses in use all year round. The LED technology is predicted to replace fluorescent and

HID lamps in horticultural systems and to revolutionize controlled growth environments. (Yeh N,

Chung JP. 2009)

Incandescent, compact fluorescent and LED lamps

Current LED lamps are five times more efficient than incandescent lamps and have a

much longer lifespan than compact fluorescent light bulbs. To evaluate a product’s true

efficiency, however, its entire life must be taken into consideration, from manufacture through

disposal. The study “Life Cycle Assessment of Illuminants: A Comparison of Light Bulbs,

Compact Fluorescent Lamps and LED Lamps”, examined the energy required for each product

over its entire lifespan, including raw material production, manufacture and assembly,
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transportation, use, and disposal. Also examined were environmental impacts such as resource

depletion, human toxicity, and greenhouse-gas generation. The report’s conclusions include:

Current-generation LED lamps need less than 2% of their energy consumption for their

manufacture; the remaining 98% is turned into light. LED lamps use 80% less energy than

incandescent lamps over their lifetime. LED lamps’ impact on the environment is consistently

less than that of incandescent lamps. Worldwide, lighting accounts for approximately 19% of

electricity consumption, corresponding to 2.4% of energy consumption. Replacing incandescent

with LEDs would theoretically reduce global electricity consumption for lighting by 50%. (Kille,

2013)

The Benefits of Sunlight vs. Artificial Light for Plant Growth

All plants need light to grow. Plants use light to create the energy needed to make the

food they need to grow and a plant’s growth rate and longevity is dependent upon how much light

it receives, according to the Texas A&M Agricultural Extension website. Living in a house or

apartment with limited access to sunlight does not mean you cannot grow plants. Artificial lights

can be used to supplement natural sunlight or, in some cases, replace it completely. (Dean, 2017)

Sunlight

Sunlight is the best light for plants. The rainbow effect created by holding a prism up to

sunlight is a visual description of the colors in the light spectrum. Plants need the red and blue

parts of the light spectrum, with red being more important than blue. Indoor plants placed within

10 feet of a sunny window receive the natural balance of red and blue rays they need. ( Dean,

2017)
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How do plants grow toward the light?

Plants have developed a number of strategies to capture the maximum amount of sunlight

through their leaves. As we know from looking at plants on a windowsill, they grow toward the

sunlight to be able to generate energy by photosynthesis. Now an international team of scientists

has provided definitive insights into the driving force behind this movement – the plant hormone

auxin. The growth of plants toward light is particularly important at the beginning of their

lifecycle. Many seeds germinate in the soil and get their nutrition in the dark from their limited

reserves of starch and lipids. Reaching for the surface, the seedlings rapidly grow upwards

against the gravitational pull, which provides an initial clue for orientation. With the help of

highly sensitive light-sensing proteins, they find the shortest route to the sunlight – and are even

able to bend in the direction of the light source. (Research News, 2013)