Annals of the Academy of Romanian Scientists
Series on Agriculture, Silviculture and Veterinary Medicine Sciences
ISSN 2344-2085
Volume
15
, Number
2
/
2026
79
SPECIFICS OF PRESENTATION FOR PHYSIOLOGICAL
DISORDERS IN TOMATOES
Vasile BOTNARI
19
Abstract
.
Disorders and difficulties in plant growth and development can also arise from
factors
other
than
those
caused
by
infectious
contamination.
Tomatoes,
like
other
agricultural crops, are often exposed to inappropriate environmental conditions. During
the growth period, extreme temperatures, heat stress, insufficient or excessive light, frost,
soil moisture deficit, nutrient deficiencies or excesses, as well as some effects resulting
from inappropriate application of plant protection products frequently affect plants.
Although tomatoes tolerate a relatively wide range of climatic conditions - allowing
cultivation in diverse production environments - obtaining high yields requires specific
growth and development conditions considered optimal. Whether grown on an industrial
scale,
on
small
plots
or
in
home
gardens,
tomato
cultivation
requires
systematic
monitoring of plant health. Understanding the conditions that trigger physiological
disorders and ways to strengthen plant health are of particular importance for increasing
resistance
to
potential
attacks
by
infectious
diseases
and
pests.
Knowledge
of
the
symptoms of physiological disorders under different production conditions contributes to
obtaining high yields and quality products.
Keywords
:
Physiological disorders, tomato symptoms, environmental conditions.
DOI
10.56082/annalsarsciagri.2026.2.79
1.
Introduction
Tomatoes are widely cultivated in open fields, polytunnels, greenhouses, and other
types of protected environments, using natural soil, sand, or artificial growing
substrates. Increasing consumer demands, combined with environmental concerns
and a shift toward organic production, have led to significant changes in tomato
cultivation technologies. The focus has shifted toward sustainable production,
which aims to achieve a competitive, high-quality harvest while reducing the use of
polluting inputs. In this context, it should be noted that tomato crops are susceptible
to
various
diseases
and
pests,
which,
under
conditions
favorable
to
their
development, can cause irreparable damage. The ability to prevent the emergence
and spread of infections, or to mitigate the impact of such outbreaks, becomes the
primary factor determining success in this type of endeavor. Furthermore, the
19
Dr. Hab. Eng., Principal Scientific Investigator: Resistance Genetics Laboratory Plants within
Institute
of
Genetics,
Physiology
and
Plant
Protection
of
the
State
University
of
Moldova,
Chisinau. Republic of Moldova. Honorary Member of the Academy of Romanian Scientists,
Bucharest, Romania. E-mail: vasilebotnari52@gmail.com, vasilebotnari@yahoo.com
80
Vasile
Botnari
potential effects of climate change, such as an extended growing season, lower
temperatures in the spring, temperatures exceeding the optimum in the autumn and
winter, or irregular precipitation patterns—cannot be ignored, as these factors can
influence the onset and progression of certain diseases [3, 27].
Physiological disorders and plant growth difficulties can also be caused by factors
other
than
infectious
pathogens.
Throughout
the
growing
season,
plants
are
frequently affected by extreme temperatures, insufficient or excessive light, frost,
and soil moisture deficits [9]. Industrial development, the growing number of means
of transport, and other social and economic factors characterized by high energy
consumption have led to an increase in the volume of gases emitted into the
atmosphere, as well as in the amount of harmful compounds released into water and
soil [37]. Instances where plants suffer damage, whether due to acid precipitation
[16], the use of contaminated water for irrigation, or the improper application of
treatments against diseases and pests, are becoming increasingly frequent.
Conducting a comprehensive analysis of the effects of climate change on the
emergence of diseases and the control of their progression is becoming increasingly
difficult. This is because, in addition to the phenotypic diversification resulting
from the use of new genetic technologies in plant breeding, the instability of abiotic
factors triggers various physiological disorders, manifesting as symptoms that
resemble those caused by certain pathogens [8]. There is already clear evidence that
global warming and climate change
[9]
will influence the occurrence and severity
of
harmful
agents
affecting
plants.
This
will
impact
disease
management,
particularly regarding the timing of agrotechnical operations, the application and
efficacy of chemical, physical, and biological control measures, and how these are
integrated into plant protection strategies. The complexity of the risks involved in
managing phytosanitary status can be significantly reduced if one also considers the
symptoms
characteristic
of
physiological
disorders,
disorders
that
arise
when
growing conditions do not align with the requirements of the plants' ontogeny.
Whether tomatoes are grown on an industrial scale, on smaller plots, or in home
gardens, a systematic assessment of the plants' condition is essential. Although
symptoms
caused
by
bacterial,
fungal,
and
viral
diseases
are
extensively
documented in the specialized literature [11, 13], they are frequently associated
with,
and
can
easily
be
confused
with,
morphological
changes
and
various
disturbances in growth and development resulting from growing conditions that
deviate from the optimum. There are numerous situations in which plants exhibit
etiolation, yellowing, the wilting of certain parts or organs, or various types of spots
or lesions, symptoms that, at first glance, resemble those caused by infections [15,
22, 27]. The inability to manage and maintain technical parameters in accordance
with optimal requirements, based on a diagnosis of the plants' physiological state,
can
lead
to
the
excessive
use
of
plant
protection
products,
resulting
in
Specifics of Presentation for Physiological Disorders in Tomatoes
81
environmental pollution and the contamination of agricultural produce with harmful
substances
Climate
change
alters
the
epidemiology
of
diseases
and
necessitates
the
development of adaptive strategies for integrated disease management [21]. Under
current circumstances, preventive measures are of particular importance, being
considered the most appropriate and relevant approach in plant protection. In this
context,
recognizing
symptoms,
understanding
the
conditions
that
trigger
physiological disorders, and knowing how to strengthen plant health are crucial for
increasing resistance to potential attacks by various pathogens and pests; this, in
turn, contributes to achieving high yields and quality produce.
2.
Materials and methods
Phenological data were collected through multifactorial experiments conducted
under various pedoclimatic conditions at the following institutions: the Institute of
Genetics,
Physiology
and
Plant
Protection;
the
Scientific-Practical
Institute
of
Horticulture and Food Technologies (Chisinau); the "Porumbeni" Institute of Crop
Science
(Pașcani);
and
the
Institute
of
Vegetable
Growing
and
Irrigation
Technologies (Tiraspol).
Over
the
years,
specific
experimental
phases
were
evaluated
to
analyze
and
determine the responses of various tomato genotypes exposed, throughout the entire
growing season, to thermal and water stress, as well as to a wide range of nutritional
levels.
This
evaluation
included
the
characterization
of
morphological
traits,
documented through photographs taken under diverse plant cultivation conditions.
Throughout
the
growing
season,
phenological
observations
were
conducted
regarding plant growth and development, as well as morphological changes and
variations in the coloration of vegetative organs and fruits during the ripening stage.
The underlying causes of physiological disorders and reduced plant resistance to
abiotic
stressors
were
analyzed
visually
and
examined
based
on
symptoms
described in the specialized literature.
The assessment of physiological disorders in plants involves the use of a set of
visual and biochemical techniques designed to identify deviations from normal
metabolism. In this context, visual diagnosis is the most accessible method [22], as
it relies on the correlation between external symptoms and internal biochemical
disturbances within the plant. Changes in leaf color, such as yellowing (chlorosis)
along the margins or in the areas between the veins, often indicate a deficiency in
nitrogen,
magnesium,
or
iron
[15].
Reddish
or
purplish
hues
may
signal
a
phosphorus
deficiency
or
thermal
stress
caused
by
unsuitable
temperatures.
Malformations and necrosis, manifesting as leaf curling or crinkling, thinning of
leaf tips, or the appearance of dry spots (in the absence of spores or parasites), point
to hydrothermal shock or chemical burns [27].
82
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3. Results and Discussions
3.1. Physiological disorders caused by imbalances in environmental factors
Environmental factors are of particular importance for the growth and development
of tomato plants; among these, temperature, light, water availability, and nutrient
supply play a primary role [3, 10]. Like other agricultural crops, tomatoes are
constantly exposed to fluctuations in environmental factors. Throughout the growing
season, plants are frequently affected by extreme temperatures, frost, soil moisture
deficits, as well as nutrient deficiencies or excesses.
To
achieve
higher
yields,
plants
require
specific
conditions
for
growth
and
development, conditions considered relatively optimal. Although tomato plants can
tolerate significant fluctuations in climatic factors with relative ease [4, 12], a trait
that allows them to be cultivated across a variety of geographical regions and
production environments, it should be noted that extreme temperatures, water deficits
(in both the soil and the atmosphere), excessive or insufficient solar radiation, intense
heat,
drought,
and
frost
all
represent
risk
factors.
These
elements
limit
the
productivity, stability, and quality of tomato harvests.
In general, minor, short-term fluctuations in climatic factors do not cause significant
disruptions to the physiological functions of the plants. The term “moderately
unfavorable conditions” refers to factors whose intensity and duration, while not
immediately lethal, trigger a wide range of long-term consequences for physiological
processes, ultimately having a negative impact on final productivity
[35,
36].
The temperature
of the soil
or growing medium directly influences germination,
root growth, and the functionality of the root system. Seed germination occurs at
temperatures above 10–12 °C, but below 35 °C. The development of the root system
is
largely
determined
by
the
temperature
of
the
growing
medium
[28,
29].
Temperatures below 5–6 °C cause root function to cease, while at values below 0 °C,
the roots die. Root growth and development are also inhibited at temperatures
exceeding 30–35 °C. Extreme temperatures in the root zone affect other plant organs,
as well as the plant as a whole.
At temperatures below 5 °C, water in the soil becomes unavailable; however, plants
regain their absorption capacity as the temperature rises above 12–15 °C, although
this capacity decreases again at temperatures above 25 °C [4
].
Nutrient uptake occurs within temperature ranges similar to those required for water
absorption. The ammoniacal form of nitrogen (NH₄
) is absorbed more readily at
⁺
relatively low temperatures (8–14 °C), whereas the nitrate form (NO₃
) is assimilated
⁻
more easily as the temperature increases (24–34 °C) [1]. Phosphorus (in the form of
P₂O₅) is assimilated more readily at temperatures above 20 °C. Air temperatures
below 10 °C can negatively affect plant growth, leading to a negative bioconversion
balance [12]. At higher temperatures, synthesized substances are primarily directed
Specifics of Presentation for Physiological Disorders in Tomatoes
83
from the leaves toward the fruits, at the expense of root growth, whereas at lower
temperatures, they are directed more toward the roots than the fruits. Consequently, at
temperatures below 18 °C, tomato plants yield smaller harvests compared to optimal
levels [10].
During the ripening period, fruit color is largely determined by temperature. Low
temperatures inhibit the synthesis of lycopene, which dictates the intensity of the red
color—while temperatures above 30 °C significantly reduce the formation of this
pigment. Under such conditions, the synthesis of beta-carotene predominates, giving
the fruits an orange hue. The inhibition of lycopene synthesis at temperatures above
30 °C (and of carotene at temperatures exceeding 40 °C) results in uneven ripening,
characterized by the appearance of yellow or yellow-green patches against the red
background of the fruit [18]. High temperatures also inhibit the synthesis of enzymes
responsible for tissue softening; at the stage of biological maturity, this often leads to
fruits with a firmer, harder consistency, which significantly diminishes their eating
quality. In field conditions, tomato plants can be negatively affected by large
temperature fluctuations, as well as by spring and autumn frosts. For vigorous
growth, tomato plants require daytime temperatures between 19 and 24 ºC and
nighttime temperatures of 16–18 ºC. Air temperatures below 10 ºC inhibit vegetative
growth
and
development,
reduce
fruit
set,
and
impair
ripening;
conversely,
temperatures exceeding 35 ºC decrease the number of fruits and diminish their color
intensity.
Well-hardened seedlings, once planted in the soil, can withstand short-term exposure
to temperatures as low as –1.5 to –2 ºC; however, such conditions severely affect the
plants, reducing their productivity by 30–50% and delaying the onset of fruiting by
8–10 days. It is a well-known fact that young plants are more sensitive to low
temperatures than mature ones. Even light frosts or prolonged periods of low
temperatures can cause flower drop and lead to the appearance of translucent or
waxy-looking spots on leaves and young fruits, which subsequently rot.
The destruction of tomato seedlings can also occur in protected environments.
Damage to seedlings due to frost—especially after planting in unheated greenhouses,
is a critical issue, often resulting in irreversible damage. In unheated greenhouses
covered with polyethylene film, when the ambient air temperature drops to –4 ºC, the
temperature at the plant surface can fall to –1.5 ºC, while the temperature in the
spaces between the plants reaches 0.5–1 ºC.[6] At such temperatures, 30–40% of the
seedlings perish (Fig. 1).
If the growing point and the first inflorescences are damaged, tomato plants may
sometimes develop lateral shoots that continue vegetative growth; however, in such
cases, the efficiency of an early crop is significantly reduced. At temperatures of –3
ºC, tomato plants suffer severe damage—though they may occasionally recover, but
such
crops
require
additional
effort
and
expense
for
restoration,
rendering
the
recovery and maintenance process unprofitable. Initial symptoms of frost damage in
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tomato plants appear within a few hours, but a definitive assessment of the extent of
the damage and the resulting losses can only be made after 2–3 days.
a.
before planting
b
. after planting
Fig. 1
.
Damage to plants due to frost
Tomato plants grown from early direct sowing in open fields are relatively more
resistant to frost compared to crops established via transplants. In field conditions,
seedlings at the stage of having one true leaf can withstand short-duration frosts
down to –4 ºC [6]. Ripe fruits are particularly sensitive to frost; they lose their
flavor and processing quality at temperatures of –0.4 to –0.5 ºC.
Light
plays a crucial role in the growth and development of tomato plants, largely
determining their flowering capacity, fruit set, fruit formation, and fruit quality.
While its influence is not as profound as that of temperature, insufficient light
during the seedling stage can lead to the total failure of the crop, as the excessive
elongation of seedlings significantly increases risk. A lack of light during this
phase also affects the development of reproductive organs
[18,
36].
Low
light
levels
can
significantly
impair
plant
growth
and
development,
especially in protected cultivation environments during winter and spring. Under
conditions of low solar radiation, fruits from the first inflorescences may develop
internal cavities, exhibit a lower dry matter content, and possess a thinner skin
that is prone to cracking upon maturity (Fig. 2).
Light
intensity
and
quality
are
of
particular
importance
regarding
the
accumulation
and
redistribution
of
substances
assimilated
during
the
photosynthesis process. When plants are grown under low-light conditions, the
quantity of photosynthetic products in the leaves is lower; consequently, these
products are redirected toward increasing the assimilation surface area—which
has a low photosynthetic yield—at the expense of flower initiation, fruit set, and
fruit development [24]. This explains why flowers may abort when exposed to
insufficient light intensity over an extended period.
Specifics of Presentation for Physiological Disorders in Tomatoes
85
Fig. 2.
Symptoms characteristic of light deficiency in the context of low temperatures
In
greenhouse
conditions,
insufficient
light
leads
to
irregularities
in
fruit
development;
the
fruit
skin
becomes
thinner
and
prone
to
cracking
when
environmental parameters fluctuate. Fruit cracking occurs most frequently at the
beginning of the ripening stage. In some cases, fruit cracking is caused by high
nitrogen
levels
in
the
soil.
Under
conditions
of
chronic
light
deficiency
in
enclosed spaces, the negative impact on photosynthetic processes can be partially
mitigated
by
enriching
the
atmosphere
with
carbon
dioxide
[18].
Unlike
temperature, light has a less significant influence on fruit ripening, yet it directly
affects the content of dry matter, carotene, and other biochemical compounds
(Fig. 3).
Fig. 3.
Cracked fruits caused by insufficient light.
A
prolonged
lack
of
light
during
the
fruit
development
phase
leads
to
the
formation of a much thinner skin, which tends to crack easily on very sunny days,
a factor that negatively affects fruit quality. In such cases, fruits located at the
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base of the cluster, as well as those that developed during periods characterized by
frequent cloud cover and low light levels, are the most severely affected.
Sunscald occurs primarily in determinate (bush-type) tomato plants with small
leaves. On green fruits, whitish spots appear; these areas dehydrate and eventually
become sunken. On ripe fruits, yellow spots appear. The affected areas lose
turgor, wilt, and can lead to tissue rot on the fruit's surface (Fig. 4). To mitigate
sunscald damage in tomatoes, it is essential to ensure conditions that promote
vigorous vegetative growth in open-field cultivation and to use shade netting in
greenhouses during the summer. Thanks to advances in plant breeding, varieties
and hybrids have been developed that are adapted to the climatic conditions of
various geographic regions, as well as to different cultivation systems, such as
greenhouses, polytunnels, or open-field production. Currently, there are genotypes
specifically developed for greenhouse cultivation during the winter, which require
light intensities of 8,000–12,000 lux, as well as genotypes intended for open-field
cultivation, which require 14,000–25,000 lux to achieve higher yields.
Fig. 4
.
Sunscald on green and ripe fruits
Water
, as a factor in plant growth and development, plays a primary role in
achieving
higher
tomato
yields.
Requirements
for
atmospheric
humidity
are
relatively
moderate,
while
those
for
soil
moisture
are
significantly
higher.
Although tomato plants can, to some extent, tolerate short-term atmospheric or
soil drought, such conditions negatively impact productivity—particularly if the
moisture deficit occurs during the fruit-setting and development stages. This
effect is less significant when the plant already bears a sufficient number of fruits
that are in the ripening phase. These observations indicate that tomato plants have
varying water requirements depending on their developmental stage [2].
Research conducted over several years at the Institute for Plant Breeding and
Vegetable Growing in Tiraspol suggests maintaining a minimum soil moisture
Specifics of Presentation for Physiological Disorders in Tomatoes
87
level of 70% of field capacity up until the flowering stage, increasing to 80%
during the fruit formation and ripening periods [34].
The optimal soil moisture level depends largely on plant density, soil porosity,
and the degree of root system development. Plants grown from direct field sowing
are more resilient regarding water availability because they develop taproots that
penetrate deeper and extract water from a larger volume of soil; in contrast, plants
raised from transplants develop fibrous root systems concentrated primarily in the
upper soil layers. Fluctuations in root-zone water content, ranging from maximum
to minimum levels and vice versa, are particularly hazardous, as they induce water
stress, leading to various physiological disorders and negatively impacting both
yield and produce quality. If seed germination is triggered but followed by low
soil moisture, the germinating seeds or seedlings may die during the early stages
of development.During the seedling growth stage, excess moisture, combined
with high temperatures and/or low light intensity, leads to plant elongation. If soil
moisture
is
low,
the
plants
wilt.
In
some
cases,
this
physiological
state
is
reversible, allowing the plants to gradually recover; in more severe instances, the
plants die.
Fig. 5
.
Non-infectious leaf curling
.
Due to their well-developed root systems, mature plants are less susceptible to
water stress. However, significant fluctuations between low and high moisture
levels,
particularly
in
protected
cultivation
environments,
often
result
in
the
formation
of
cracks
at
the
base
of
the
stem,
which
facilitate
the
entry
of
pathogens.
Initially, water stress does not produce clearly visible symptoms on the leaves.
Nevertheless,
under
conditions
of
acute
and
prolonged
drought,
the
plant's
metabolism, the redistribution of assimilates, and the structure of its tissues and
organs are all affected. In the absence of sufficient moisture, leaves wilt; in certain
88
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genotypes, they also curl around the veins. In some varieties, leaf curling is a
genetically determined trait that enhances the plant's adaptation to atmospheric
and/or soil drought (Fig. 5).
Inadequate water supply over an extended period leads to pollen grain infertility,
as well as the shedding or abortion of flowers. At the same time, excessive air
humidity creates unfavorable conditions for the opening of anthers and the release
of pollen. In some cases, fruit cracking is caused by sudden changes in the
moisture level of the soil or growing medium [19]. During periods of high
temperatures and sunny weather, the soil can overheat and dry out. If heavy
irrigation
is
applied
under
these
conditions,
large
quantities
of
water
are
transported
to
the
fruits;
their
walls
cannot
withstand
the
resulting
osmotic
pressure and consequently crack (Fig. 6). To prevent fruit cracking, it is necessary
to maintain optimal soil moisture and an appropriate plant nutrition regimen
Fig. 6.
Fruit cracking caused by a sudden change in soil moisture
Fig. 7
.
Fruits with reduced quality due to physiological disorders caused by environmental factors
Specifics of Presentation for Physiological Disorders in Tomatoes
89
Excess moisture during fruit ripening, following a relatively dry period, leads to
fruit cracking and quality degradation, even under open-field conditions (Fig. 7).
Some
recommendations
suggest
reducing
soil
moisture
levels
during
the
biological ripening stage and harvesting the fruit at the "waxy" phase or at the
onset of the transition to red. It is well established that fruit ripened under
conditions of limited water supply possess superior flavor, owing to a higher dry-
matter content and a more balanced ratio of sugars to acids.
Air.
Intense air circulation combined with high temperatures and low humidity
causes excessive flower drop (Fig. 8). Relative air humidity is considered optimal
when it ranges between 55–60% during the first vegetative stage and 65–70% during
the second. Excess air humidity promotes the development and spread of diseases
affecting foliage and fruit. To prevent sudden fluctuations in air humidity, airflow
within the greenhouse should be maintained at a rate of 1–1.5 m/s.
Fig. 8
.
Absence of flowers and young fruits caused by low air humidity
Carbon dioxide is a component of the air (0.03%) of particular importance, as it
enables the process of photosynthesis. CO
2
levels are actively managed when
cultivating tomatoes in protected environments [18]. It has been observed that
increasing
the
CO
2
concentration
to
0.09–0.14%
enhances
photosynthesis,
resulting in more vigorous plant growth and development. During the day in the
winter season, it is recommended to maintain the CO
2
concentration in the
greenhouse air at 800–1,000 ppm. Excessively high carbon dioxide levels can lead
to negative effects, impacting both the plants and pathogenic organisms.
90
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The relationship between plants and the air becomes increasingly significant when
the
air's
composition
undergoes
substantial
changes
due
to
uncontrolled
gas
emissions and excessive pollution. Following a prolonged period of atmospheric
drought, light precipitation can trigger the appearance of various spots or rust-like
lesions on leaves, stems, and fruits, not only in tomatoes but also in other plant
species, which are frequently mistaken for symptoms of infectious diseases.
The role of oxygen in biological processes is well-recognized and documented
from various perspectives. Tomato plants are sensitive to root asphyxiation caused
by low oxygen wqlevels in the soil. A reduction in oxygen content within the root
zone, often occurring in compacted or waterlogged soils, impairs the function of
active roots, leading to premature aging and even root death. When oxygen is
insufficient, the recovery of the root system is a lengthy process.
3.2. Soil Requirements
Tomatoes yield higher harvests when grown in chernozem, sandy-clay, or clay-
sandy soils that are rich in humus, well-structured, and well-managed, provided
the groundwater table is at a depth of more than 4 meters. Satisfactory results can
also be achieved in well-drained heavy soils. Light soils have a lower water-
retention capacity, which exacerbates moisture deficits, leading to water stress
during periods of drought or insufficient irrigation. Heavy soils warm up more
slowly and, if compacted or excessively wet, can cause root system asphyxiation.
Tomatoes are sensitive to root asphyxiation resulting from soil compaction and an
excess of water that reduces soil porosity. Prolonged asphyxiation can trigger
symptoms
of
various
nutrient
deficiencies,
such
as
those
of
phosphorus,
magnesium, or nitrogen, which increase the plants' vulnerability to infectious
diseases.
For tomato cultivation, the soil must be well-supplied with organic matter and
essential nutrients, and it should have a pH between 6 and 7. The pH level
specifically influences nitrogen uptake [1], depending on whether the nitrogen is
in nitrate (NO
3
) or ammonium (NH
4
) form. If the pH drops below 5 or rises above
7, there is a significant reduction in stem growth and overall plant habit (Fig. 9).
Higher pH values in the soil or substrate inhibit the uptake of phosphorus, boron,
copper,
iron,
magnesium,
and
zinc,
whereas
lower
pH
values
reduce
the
availability of molybdenum [31].
Specifics of Presentation for Physiological Disorders in Tomatoes
91
Fig. 9
.
Disorders
caused
by high acidity of the nutrient substrate
Tomatoes exhibit moderate tolerance to soil salinity (2.5 dS m
¹); however, it is
⁻
preferable for soils to have salinity levels within a normal range, specifically, they
should be free of chlorides and sodium [25, 28]. The adverse effects of excessive
salinity are most likely to occur when crops are established on land with a high
water table, or when irrigation water containing salt concentrations exceeding 1
g/l is used extensively over a prolonged period.
3.3. Physiological disorders caused by nutritional status
Plant
nutrition
is
a
complex
physiological
process
consisting
of
the
uptake,
primarily by the roots, and to a lesser extent by the leaves, of accessible mineral
substances
from
the
root
zone.
These
substances
are
then
transported
and
metabolized into organic compounds essential for growth. Subsequently, they are
incorporated into various structural components, most notably carbohydrates and
proteins, which, through degradation, provide the energy required for vital life
processes.
Along with water and other environmental factors, nutrients are frequently in
short
supply
relative
to
the
plant's
requirements.
Consequently,
ensuring
an
optimal supply of nutrients, balanced according to the specific needs of the plant,
is a prerequisite for achieving high yields and superior quality. A deficiency or an
excess of nutrients triggers various physiological disorders [32], which negatively
impact plant vigor, resistance, productivity, and the quality of the produce.
Under conditions of water scarcity, root growth is impaired, resulting in a weaker
root
system;
in
such
cases,
the
uptake
of
nutrients
becomes
more
difficult.
Extreme
temperatures
also
disrupt
plant
metabolism,
which
can
lead
to
the
emergence of various nutrient deficiencies and physiological disorders.
Assessing the nutritional status of plants based on visible symptoms is a primary,
yet subjective, method of evaluation. This assessment is based on identifying
states of nutrient deficiency or toxicity, manifested through symptoms appearing
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on various plant organs. These symptoms are specific to each element and its
particular mode of expression, while also being influenced by the biological
characteristics
of
the
varieties
and
the
conditions
under
which
the
crop
is
maintained. Through this approach, one can evaluate several aspects, including:
the general condition of the plants at different stages of growth and development;
the ratio between the root system and the aerial parts; the growth rate; stem length
and leaf orientation; the positioning of inflorescences along the stem; the number
of
flowers
and
fruits
per
cluster;
fruit
size
and
internode
length;
and
leaf
coloration, using a standard green hue as a reference for a healthy nutritional
status, among other factors.
In some instances, an excess or deficiency of multiple nutrients relative to the
plant's
requirements
may
result
in
complex,
interrelated
symptoms
[14,15],
sometimes exacerbated by unfavorable environmental or production conditions. In
such cases, determining the plant's nutritional status is best achieved through
chemical analysis of plant tissues and the soil, an effective method for assessing
the level of nutrient supply.
Managed
plant
nutrition
entails
ensuring
optimal
conditions
for
growth
and
development. Occasionally, symptoms of physiological disorders stemming from
nutritional imbalances can be mistaken for signs of pathogen-induced infections,
potentially
leading
to
the
application
of
treatments
based
on
an
incorrect
diagnosis. Accurately diagnosing nutritional imbalances and specific nutritional
requirements is of great importance for optimizing plant growth and development.
When applying supplementary fertilizers, this can help mitigate the impact of
disturbances caused by environmental factors (light, temperature, humidity, air
composition), while simultaneously strengthening the plants' resistance to various
diseases.
Nitrogen
is the most important nutrient for plant life, playing a decisive role in
increasing tomato yields. A plant's nitrogen requirement varies throughout its
growth cycle. In the early stages of development, plants have relatively low
nitrogen demands within the root zone; the greatest amounts are assimilated
during the period of intensive vegetative growth and fruit formation.
Due to its high mobility and capacity for translocation within the plant, symptoms
of nitrogen deficiency first appear on mature leaves at the base of the stem or
shoots, subsequently spreading toward the upper leaves. Insufficient nitrogen
supply in the soil, or poor nitrogen uptake, visibly reduces plant vigor. In such
cases, the leaves take on a lighter color (Fig. 10), and the fruits become smaller
and of inferior quality. Easily observable symptoms, which generally indicate a
deficiency of this nutrient, include a loss of green color intensity, etiolation, short
stems with thin tips, small leaves, and sparse or delayed flowering. Nitrogen
deficiency leads to reduced vegetative growth rates and a shortened growth cycle,
Specifics of Presentation for Physiological Disorders in Tomatoes
93
both of which negatively impact productivity. The fruits exhibit inferior flavor,
reduced flesh firmness, and poor resistance to transport and storage [1].
Fig. 10.
Symptoms characteristic of nitrogen deficiency
In cases of severe deficiency, the main stems are shorter and have few branches;
the leaves turn yellow and then brown, starting at the tips and margins—before
wilting and drying out. This yellowing may be followed by premature defoliation
[15]. In some varieties, the main veins and petioles of the lower leaves take on a
red or anthocyanin-like coloration. An excess of nitrogen during the seedling
growth stage causes the plants to elongate, a response similar to that triggered by
a lack of light, or an excess of water or heat, which makes it difficult to diagnose.
Under such conditions, it becomes more challenging to properly "harden off" the
seedlings before transplanting.
Excessive nitrogen fertilization during the growing season has negative effects: it
promotes vigorous stem and leaf growth at the expense of fruit production and
early maturity, while also reducing the plants' resistance to extreme environmental
conditions and pathogens. Symptoms appear on mature leaves at the base of the
stem; as temperatures rise, these leaves may resemble those damaged by frost or
"scalded," showing burns at the tips and margins.
In mature plants, a moderate excess of nitrogen can be identified by thick, rigid
stems, lush foliage with an intense green color, and leaves that appear curled or
puckered, often accompanied by a lack of inflorescences. Tomato plants are
particularly sensitive to an excess of ammoniacal nitrogen, which inhibits calcium
uptake. Ammonium toxicity leads to the yellowing of the leaves.
Phosphorus.
This
element
plays
a
primary
role
in
plant
metabolism,
being
involved in the most important physiological processes. Phosphorus participates
in a range of processes related to the formation of organic compounds, such as
nucleic acids and phospholipids, contributes to the development and quality of
fruit, stimulates root system growth, and enhances plant resistance to adverse
conditions, including extreme temperatures, moisture deficits, diseases, and pests.
94
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Phosphorus has low mobility in the soil, combined with a very high retention
capacity. The total content of this element remains relatively stable [31]. Most
soils have the capacity to retain phosphorus, with the exception of sandy soils. To
prevent phosphorus deficiency in plants, it is essential to ensure the availability of
this element, in forms that are easily accessible to the plant, starting from the
earliest stages of growth.
A deficiency of phosphorus in the soil or nutrient substrate leads to various
physiological disturbances (Fig. 11) and a slowdown in plant growth rates. When
plants are under-supplied with phosphorus, the mobility of this nutrient ion within
the plant triggers a redistribution of phosphorus from mature organs to younger
ones. Consequently, signs of deficiency first appear on older leaves, which remain
small and rigid, and develop a purplish hue on their undersides. In cases of more
severe phosphorus deficiency, fruit cracking may occur.
Fig. 11.
Symptoms characteristic of Pdeficiency Phosphorus deficiency
associated with an excess of Cu, Mg, or Zn
An excess of phosphorus leads to the accumulation of its mineral form within
plant organs. Plants exhibit slowed growth and premature aging, which results in
lower yields. Furthermore, an excess of phosphorus can affect the availability of
other elements, such as zinc and copper, and can disrupt the uptake of these
nutrients by the plant.
Potassium
plays a particularly important role in plant nutrition and is absorbed by
plants in large quantities compared to other nutrients, with the exception of
nitrogen. In some cases, the total potassium content in chernozem soils exceeds
the amount required to sustain plants throughout their entire growing season;
however, it should be noted that only a small fraction of the soil's potassium is
actually available for plant nutrition. Nevertheless, to achieve higher yields and
better quality, tomatoes require additional potassium fertilization, even in soils
already rich in this element.
Specifics of Presentation for Physiological Disorders in Tomatoes
95
Available potassium (K₂O) is the form most accessible to plants, as it is absorbed
by the roots from the soil solution. The rate of potassium uptake depends on
several factors: the type of crop (direct-seeded, transplanted, or grown in a
greenhouse on various substrates, etc.), the development of the root system, the
stage of the growing season, the level of potassium present in the active root zone,
and the presence of other cations that may inhibit the uptake of this element.
As an essential element for plant metabolism, potassium, when assimilated in
sufficient
quantities,
enhances
drought
resistance,
stimulates
chlorophyll
synthesis, and boosts the rate of photosynthesis [26]. Potassium deficiency in
tomatoes
significantly
reduces
both
plant
productivity
and
fruit
quality.
Inadequate
supply
of
accessible
potassium
impairs
the
enzymatic
processes
responsible for converting sugars into starch, which leads to the accumulation of
sugars in the fruit and a decline in overall quality. Potassium is involved in
regulating the water balance within a plant; a deficiency causes stomata to remain
locked in an open position. Consequently, potassium helps reduce water loss
through transpiration and enhances plant resistance to water stress and extreme
temperatures.
When this element is deficient, tomato plants exhibit symptoms resulting from
metabolic disruption, which leads to premature aging and reduced resistance to
various infections, as well as to adverse temperature and humidity conditions.
Potassium deficiency manifests primarily on mature leaves, as the element is
translocated,
much
like
nitrogen,
from
mature
tissues
to
younger
ones.
The
deficiency
appears
as
yellow-brown
spots,
starting
at
the
leaf
margins
and
spreading inward across the leaf blade; these eventually take on the appearance of
burns (Fig. 12).
Fig. 12
. Symptoms characteristic of potassium deficiency
96
Vasile
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Under conditions of potassium deficiency, fruits ripen unevenly, develop irregular
shapes, are prone to the formation of internal cavities, and exhibit inferior taste quality.
Such symptoms may occur when tomatoes are grown in clay or sandy soils, or when
using
growing
media
that
are
inadequately
supplied
with
potassium.
In
certain
situations, sodium can substitute for potassium and perform some of its secondary
functions; however, this process negatively affects the taste quality of the fruits.
Excessive potassium fertilization can lead to toxicity, manifesting as leaf necrosis and
stunted plant growth. This occurs because the uptake of nutrients, such as nitrogen,
phosphorus,
magnesium,
and
zinc,
is
blocked,
leading
to
various
physiological
disorders.
Calcium
is essential for plants and plays a direct role in physiological processes,
serving
primarily
as
a
structural
component
in
the
formation
of
pectocellulosic
membranes [5]. Calcium supports the development of the root system, strengthens the
supporting tissues within the stem, promotes vigorous plant growth, and enhances
resistance to breakage as well as to stressors caused by unfavorable growing conditions.
Plants assimilate calcium through their root system by absorbing the relevant ions from
the soil solution. A calcium deficiency in the soil, or limited availability due to an
excess of another element, such as potassium, can lead to various physiological
disorders that reduce both the yield and the quality of the fruit.
Calcium, sulfur, and magnesium, classified as secondary macronutrients, play a
significant role, as plants require them in relatively large quantities. Since these
elements are generally present in the soil in substantial amounts, it is often assumed that
the plants' requirements are largely met; consequently, they are not typically applied as
standalone fertilizers, unlike nitrogen, phosphorus, or potassium. However, in tomato
cultivation, particularly when artificial substrates are used, situations frequently arise
where it becomes necessary to supplement the levels of these elements through the
application of specialized fertilizers.
In intensively irrigated fields and protected cultivation environments (greenhouses and
polytunnels), calcium is easily lost through leaching. A deficiency manifests as stunted
growth and curled, light-green leaves, while the growing tip (apical meristem) may dry
out. Foliar application of calcium helps plants recover from stressful conditions,
fostering greater stress tolerance while also enhancing the nutritional value and quality
of
the
produce
[30].
Furthermore,
foliar
fertilization
improves
fruit
quality
by
increasing sugar content and enhancing transportability and shelf life [20].
Compared to other nutrients, calcium deficiency is less pronounced; nevertheless, it can
lead
to
nutritional
imbalances,
manifesting
in
various
ways
depending
on
environmental
factors
and
production
conditions.
Deficiency
symptoms
typically
appear in young plants, affecting the roots, stems, and leaves. Roots become coated in a
mucilaginous substance, while young stems and leaves exhibit deformities, often
accompanied by a brownish-yellow discoloration along the margins. Under drought
conditions combined with extreme temperatures, calcium deficiency can lead to pollen
sterility, widespread flower abortion, and a reduced fruit-setting capacity. Fruits,
including those that are still green, may develop blackening at the tip (Fig. 13). An
Specifics of Presentation for Physiological Disorders in Tomatoes
97
excess of calcium can inhibit the uptake of certain micronutrients, specifically iron,
manganese, magnesium, and potassium, with which it maintains an antagonistic
relationship.
Fig. 13
. Symptoms characteristic of calcium deficiency
Sulfur
is an important element for plants; it is widely distributed in nature and is
absorbed by plants through their root systems, primarily in the form of SO₄²
ions,
⁻
which are subsequently transported and utilized in the synthesis of sulfur-containing
compounds [31]. In small quantities, this element can also be taken up through the
leaves from the atmosphere in the form of SO₂ oxides; however, high concentrations of
this form of sulfur in the air can cause toxicity.
Sulfur is required by plants from the very early stages of growth for the synthesis of
sulfur-containing amino acids and proteins. Certain compounds synthesized through
sulfur metabolism help counteract the negative effects of abiotic stress caused by the
presence of free radicals within the plant [17]. Sulfur deficiency is less common than
deficiencies in nitrogen, phosphorus, or potassium. Nevertheless, in cases of severe
sulfur insufficiency, plant growth is stunted, and leaves remain small, exhibiting a
greenish-yellow to whitish coloration due to a reduction in chlorophyll content. Sulfur
deficiency resembles nitrogen deficiency; however, because sulfur has limited mobility
within the plant, it is not translocated from mature leaves to younger ones. Deficiency
symptoms first appear on young leaves and gradually spread toward the base of the
plant (the reverse of the pattern seen with nitrogen). The leaf veins appear lighter in
color
than the rest of the leaf (Fig. 14)
High concentrations of sulfur oxides in the atmosphere accelerate plant aging, cause
necrosis of leaf tissues, and reduce the ascorbic acid content in fruits. During prolonged
droughts, the concentration of sulfur oxides in the atmosphere increases significantly;
98
Vasile
Botnari
when these are absorbed by light precipitation, they can lead to acid rain and various
forms of toxicity in plants
.
a.
sulfur
b.
magnesium
Fig. 14
. Symptoms characteristic of sulfur and
magnesium
deficiency
Magnesium
is recognized as an essential element for plant growth and development,
as it is a component of chlorophyll and plays a role in balancing plant nutrition,
particularly in relation to nitrogen and potassium. Indirectly, magnesium influences
fruit quality, as well as the suitability of the fruit for storage and transport.
Due to its high mobility, magnesium facilitates the uptake and translocation of
phosphorus within plants; there is a positive correlation between phosphorus and
magnesium
content
in
various
plant
tissues.
Furthermore,
in
conjunction
with
potassium, magnesium helps regulate cell turgor. A magnesium deficiency leads to a
reduction in chlorophyll content, which in turn diminishes the rate of photosynthesis
and overall productivity.
Symptoms of magnesium deficiency occur primarily in indeterminate varieties grown
in greenhouses on acidic substrates characterized by low potassium and excess calcium
levels. Because magnesium is highly mobile within the plant, much like nitrogen, it
migrates from mature leaves to younger ones; consequently, when the element is in
short supply, deficiency symptoms first appear on the mature leaves. Such deficiencies
Specifics of Presentation for Physiological Disorders in Tomatoes
99
are most frequently observed when tomatoes are grown in greenhouses or polytunnels,
resulting in stunted growth and the appearance of chlorotic spots between the leaf veins.
Symptoms of magnesium deficiency manifest as discoloration in the areas between the
leaf veins, which turn light green to yellowish due to a reduction in chlorophyll content,
while the veins themselves remain green. In some cases, leaf deformations occur, along
with localized swellings between the veins and the development of orange or purple
coloration. Subsequently, necrosis may appear in the affected areas of the leaves,
which, in more severe cases, leads to the breakdown of the tissue.
Microelements
are nutritional components that plants require in very small quantities
for normal growth and development. When tomatoes are grown in open fields, the
plants' requirements for manganese, iron, copper, zinc, boron, and molybdenum are
generally met by the levels of these elements already present in the soil [7]. Initially,
symptoms resulting from a deficiency or an excess of a specific element are very
difficult to detect. Disruptions in plant growth and development caused by a lack of
microelements
occur
more
frequently
when
tomatoes
are
cultivated
in
artificial
substrates.
Manganese
and
boron
play
a
particularly
important
role
in
the
synthesis
of
carbohydrates and in the initiation of reproductive organ formation and fruit set; zinc,
meanwhile, is essential for regulating respiratory processes and synthesizing auxins.
The deficiency of any given microelement in tomato plants manifests differently
throughout the growing season, though the severity of the impact varies depending on
the specific physiological stage.
Boron is involved in regulating the plant's water balance and in mitigating the toxicity
of other elements present in the soil solution or artificial substrate. A boron deficiency
leads to the necrosis of meristematic tissues (Fig. 15), most notably in the apical buds—
as well as the shedding of flower buds, blossoms, and young fruit.
Fig. 15.
Shedding of flower buds caused by a boron deficiency
100
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In
plant
nutrition,
the
limited
availability
of
an
element
dictates
the
course
of
physiological processes and crop yield.
Iron
is an essential element that can become
unavailable to plants, particularly in calcareous and alkaline soils, leading to various
physiological disturbances. The primary visual symptoms of iron deficiency in plants
include the yellowing of leaves, or chlorosis, resulting from a decline in chlorophyll
content; this occurs because iron serves as an essential cofactor for chlorophyll
synthesis. Since iron is not translocated to other plant organs, the deficiency manifests
primarily in young leaves [23]. It presents as a discoloration of the leaf blade, which
turns a whitish-yellow due to the reduction in chlorophyll, a condition similar to
magnesium deficiency (Fig. 16).
Fig. 17
. Characteristic symptoms of iron deficiency
Compared to other microelements,
manganese
is present in plants in larger
quantities, particularly in the leaves, and to a lesser extent in the fruits. The
manganese ion enhances plant resistance to drought by reducing transpiration.
Manganese deficiency resembles that of iron or sulfur, though the discoloration of
the leaves creates a more mottled, mosaic-like appearance. Leaf growth is stunted,
and the foliage gradually turns a grayish-yellow, eventually developing necrotic
spots [32 33,]. An excess of manganese can also cause physiological disorders,
which may manifest as dark streaks along the leaf veins and the deformation of
fruits.
Copper
plays an important role in plant metabolism. Within the leaves, it is
located in the chloroplasts. Together with other microelements, it participates in
metabolic
reactions
and
contributes
to
maintaining
the
balance
between
carbohydrates and proteins; it also promotes an increase in the content of sugars
and
vitamins.
Copper
prevents
the
physiological
aging
of
leaves,
thereby
prolonging their photosynthetic activity.
Copper
deficiency
leads
to
disruptions
in
the
synthesis
of
amino
acids
and
proteins. Tomatoes exhibit moderate sensitivity to an excess of copper. In more
pronounced cases, this causes the atrophy of the root system and color changes in
the
leaf
tissue
between
the
veins,
at
supra-optimal
levels,
this
phenomenon
manifests as a chlorotic, pale-green appearance.
Specifics of Presentation for Physiological Disorders in Tomatoes
101
Fig. 17.
Characteristic symptoms of copper deficiency
Certain hues, caused by anthocyanins, may appear on some leaves, especially when an
excess of copper is accompanied by an excess of other microelements, such as
manganese or zinc. In cases of more severe toxicity, copper can lead to the premature
drying of the leaves (Fig. 17).
Although zinc is present in plants only in small quantities, it plays an important role in
the synthesis of several enzymes and the regulation of respiratory processes
[38].
it also
exerts a positive influence on chlorophyll formation and the fruit-setting process. Visual
symptoms of zinc deficiency may appear when tomatoes are grown in calcareous
(carbonate-rich) soils, a condition exacerbated by the application of high doses of
phosphorus, and manifest as interveinal chlorosis (discoloration of the leaves between
the veins), the formation of short internodes, small leaves arranged in a rosette pattern,
and the development of deformed fruit (Fig. 18).
Along with other microelements, molybdenum is of major importance for the balanced
nutrition of tomatoes, particularly when they are cultivated in acidic soils or substrates.
A deficiency in molybdenum leads to the accumulation of nitrates within the plant's
tissues. Under conditions of insufficient molybdenum, characteristic symptoms emerge,
including leaf discoloration, stunted growth, and reduced fruit production.
102
Vasile
Botnari
Fig. 18
. Characteristic symptoms of zinc deficiency
Physiological disorders in plants can also arise from the improper use of pesticides.
These disorders depend on factors such as the type of herbicide, the active ingredient,
the dosage, the application conditions, and the plant's developmental stage. In some
instances, it is difficult to determine whether the observed symptoms are caused by
herbicide exposure, seed quality issues, or a viral infection. It is also common for
pesticides applied to a preceding crop to have a residual effect; residues remaining in
the soil can significantly impair plant growth and productivity
4. Conclusions
(1). Difficulties in plant growth and development can arise from factors other than
infectious con-tamination. During the growing season, tomatoes are often exposed to
unfavorable environmental conditions, such as extreme temperatures, insufficient or
excessive light, and moisture deficits, which, either individually or through complex
interactions, can lead to physiological disorders with serious consequences for yield
quantity and quality.
(2). Tomatoes are sensitive to soil acidity and compaction levels. Reduced porosity
combined
with
excess
soil
moisture
leads
to
root
asphyxiation.
Prolonged
asphyxiation can trigger symptoms re-sembling various nutrient deficiencies—such
as those of phosphorus, magnesium, or nitrogen—thereby increasing the plants'
vulnerability to infectious diseases.
(3) Nutrient deficiencies or excesses cause various physiological disorders that impact
plant vigor and resistance, as well as productivity levels and fruit quality. Assessing
plant nutritional status based on visible symptoms is a primary, subjective method of
evaluation. This assessment relies on identifying states of deficiency or toxicity
Specifics of Presentation for Physiological Disorders in Tomatoes
103
manifested through specific signals on plant or-gans—signs that vary according to the
specific nutrient, the nature of the manifestation, the bio-logical characteristics of the
variety, and crop management conditions.
(4). Accurately diagnosing nutritional imbalances and specific nutritional needs is
crucial for opti-mizing nutritional regimens, regulating tomato plant growth and
development, and applying sup-plementary fertilizers; such measures help mitigate
the impact of environmental stressors (light, temperature, humidity, air composition)
while simultaneously strengthening the plants' resistance to various pathogens.
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