The Journal of Horticultural Science and Biotechnology

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Physiological disorders in tomato and some

methods to avoid them

Margit Olle & Ingrid H. Williams

To cite this article: Margit Olle & Ingrid H. Williams (2016): Physiological disorders in tomato
and some methods to avoid them, The Journal of Horticultural Science and Biotechnology, DOI:
10.1080/14620316.2016.1255569

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Published online: 15 Dec 2016.

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THE JOURNAL OF HORTICULTURAL SCIENCE AND BIOTECHNOLOGY, 2016
http://dx.doi.org/10.1080/14620316.2016.1255569

REVIEW

Physiological disorders in tomato and some methods to avoid them

Margit Ollea and Ingrid H. Williamsb
a
Estonian Crop Research Institute, Department of Plant Breeding, Jogeva alevik, Estonia; bEstonian University of Life Sciences, Institute
of Agricultural and Environmental Sciences, Tartu, Estonia

ABSTRACT ARTICLE HISTORY

Tomato can be injured by two very important physiological disorders, namely blossom end Accepted 26 October 2016
rot (BER) and fruit cracking (FC). The purpose of this review is to describe these physiological
KEYWORDS
disorders and to assess whether cultivar and cropping system can minimise their symptoms. Blossom end rot; fruit
Physiological Ca deficiency is usually related to the inability of the plant to translocate cracking; growing systems;
adequate Ca to the affected plant part, rather than to insufficient Ca levels in the growing tomato; cultivar; weather
medium. The first visible symptom of BER is a small darkened or water-soaked area around
the blossom end of the fruit, appearing about the time the fruit begins to ripen. The spot
darkens, enlarges, and becomes sunken as the fruit matures. Large lesions may show in
concentric rings. FC is the splitting of the epidermis around the calyx or stem scar. FC is a
physiological disorder, which mainly occurs when there is a rapid net influx of solutes and
especially water into the fruit, while at the same time ripening or other factors reduce the
strength and elasticity of the tomato skin. This review article gives an overview of the causes
of BER and FC and summarizes growing methods aimed at reducing these physiological
disorders of tomatoes.

Introduction uptake of nutrients by the roots, the transport of Ca2+

to and within the fruit or a varying demand for Ca2+
Tomato (Lycopersicon esculentum Mill.) is a very
depending on the growth rate of fruits (Saure, 2001).
important greenhouse crop. It is a major world food
FC is of particular importance as cracked fruit
vegetable, and its cultivated area and production are
causes serious loss of marketable yield, particularly
constantly increasing (FAO Statistics, 2006). Today,
in unheated greenhouses in some years. It often
tomato is grown commercially in 159 countries. The
appears when conditions drastically change the rate
major producers of tomatoes in 2009 were China, the
of growth, such as wide fluctuations in temperature
United States, India, Turkey, Egypt, Italy, and Iran
and moisture (Guichard, Bertin, Leonardi, & Gary,
(Ibrahim, Moharum, & Abd El-Ghany, 2015).
2001) although the environmental causes are not well
Tomato can be injured by two very important
understood (Peet, 2007).
physiological disorders, namely blossom end rot
Tomato cultivars differ in their susceptibility to
(BER) and fruit cracking (FC). Occurrence of these
BER (Ho et al., 1995; Ho, Belda, Brown, Andrews,
disorders results in unmarketable fruit and increased
& Adams, 1993; Kirkby, 1979; Nonami, Fukuyama,
wastage in the supply chain.
Yamamoto, Yang, & Hashimoto, 1995; Nukaya, Goto,
BER limits tomato production and quality (Adams
Jang, Kano, & Ohkawa, 1995) as well as to FC
& El-Gizway, 1988; Adams & Ho, 1989, 1995; Adams
(Abbot, Peet, Willits, Sanders, & Gough, 1986;
& Holder, 1992). This disorder was first identified
Fernandez-Munoz, Cuartero, & Gomez-Guillamon,
more than 100 years ago (Saure, 2001) and there is
1995; Maroto, Lopez, Bardisi, Pascual, & Alagarda,
still no totally effective method of controlling it. The
1995; Mullins & Straw, 1992; Pascual, Bardisi, Lopez-
unpredictability of its occurrence (Battey, 1990) and
Galarza, Alagarda, & Maroto, 1998; Sperry, Davis, &
the lack of control procedures make the problem very
Sanders, 1996).
serious. Estonian tomato grower Voldemar Kamenik
The purpose of this review is to describe the phy-
has found that up to 15% of production can show
siological disorders of BER and FC and to summarise
BER symptoms. Many researchers have attributed the
the influence of cultivar and cropping system on the
occurrence of BER in tomato to a deficiency of Ca2+
occurrence of both disorders.
in the fruit or parts of the fruit in connection with the

CONTACT Margit Olle margit.olle@gmail.com

© 2016 The Journal of Horticultural Science & Biotechnology Trust
2 M. OLLE AND I. H. WILLIAMS

Blossom end rot and some methods to avoid
it
Physiological Ca deficiency is usually related to the
inability of the plant to translocate adequate Ca to
the affected plant part, rather than to insufficient
Ca levels in the growing medium. It is a common
problem for vegetable growers. Many vegetables
develop specific symptoms: for example, black-
heart of celery (Geraldson, 1954; Pardossi,
Bagnoli, Malorgio, Campiotti, & Tognoni, 1999),
tipburn of chervil (Kleemann, 1999; Kleemann,
2002; Olle & Bender, 2009), tipburn of lettuce
(Collier & Tibbits, 1982; Cox, McKee, &
Dearman, 1976; Kleemann, 2004), tipburn of
Chinese cabbage (Aloni, 1986; Aloni, Pashkar, &
Libel, 1986), BER of tomato (Adams & Ho, 1995;
Adams & Holder, 1992; Saure, 2001), and Ca defi-
ciency of glasshouse cucumber (Bakker &
Sonneveld, 1988). The first visible symptom of
BER is a small darkened or water-soaked area
around the blossom end of the fruit, appearing
about the time the fruit begins to ripen (Saure,
2001). The spot darkens, enlarges, and becomes
sunken as the fruit matures. Large lesions may
show concentric rings. The affected tissue is leath-
ery and firm unless invaded by secondary decay
organisms. BER usually causes the fruit to ripen
prematurely and to be inedible. Quite commonly,
the affected fruit areas become infected with sec-
ondary pathogens, which appear as a black, felt-like
growth on the fruit. The affected area may be a
mere speck or it may involve more than half of the
fruit.
In conventional production in the USA the incidence
of BER (Figure 1) was higher in August than in
September (Elmer & Ferrandino, 1991). The reason
could be that weather plays a big role for tomatoes
grown in unheated greenhouses. The temperature in
September is much lower than in August. However,
some researchers have found that with lower
Figure 1. Picture of blossom end rot on tomato.
temperatures the incidence of Ca deficiency is lower
(Misaghi & Grogan, 1978; Nelson & Niedziela, 1998;
Saure, 2001).
BER seems to occur when stress exceeds stress
tolerance, most frequently in young fruit at the begin-
ning of cell enlargement (Saure, 2001). Ca uptake was
highly correlated with solar radiation and root tem-
perature. The cause of BER is usually an interaction
between the effects of irradiance and ambient tem-
perature on fruit growth, and the effects of environ-
mental stress on Ca uptake and distribution within
the whole plant (Adams & Ho, 1993). BER in fruits
increased at the highest light intensity and with light-
ing at night and during weekends (Stadler, 2012).
When less water is available to the plant, Ca uptake is
reduced because Ca flows through the plant with the
water flow of transpiration. Transpiration is also
depressed in conditions of high humidity when Ca
uptake by plants may be restricted. Increased light
period and/or light intensity increases growth rate and
this higher growth rate will increase the demand for Ca.
The tissue level of Ca may fall below the critical level
and induce Ca deficiency symptoms in plants
(Kleemann, 1999).
Tomato cultivars differ in the incidence of BER
in their fruits with some being more susceptible to
BER than others (Ho et al., 1993, 1995; Kirkby,
1979; Nonami et al., 1995; Nukaya et al., 1995).
Early cultivars are more susceptible to BER
(Magan, Gallardo, Thompson, & Lorenzo, 2008).
Susceptible cultivars tend to have a higher fruit
load and larger fruit (Ho et al., 1995). These differ-
ences between cultivars result from cultivar differ-
ences in the daily rates of Ca uptake, and the
difference is more consistent in older plants (Ho
et al., 1995). Differences in Ca translocation may
correspond to differences in fixation through vary-
ing oxalate content (Wiersum, 1979). In contrast,
other researchers indicate that difference in BER
incidence between cultivars cannot simply be
explained by the absorption of Ca ions in plants
but that a genetic control metabolism is involved in
BER (Nonami et al., 1995). Furthermore, Ho and
White (2005) found that the susceptibility of dif-
ferent cultivars to BER appears to be related to the
development of the xylem network and the rate of
cell expansion during early fruit development.
Determinate cultivars stop producing new fruit
trusses early in the season, and Ca then tends to
move through the plant easily with the water flow
of transpiration. Saure (2001) postulated that when
growth rate is lower the incidence of BER is also
lower, and Wiersum (1979) found that high growth
rate is associated with reduction of Ca content in
fruits. Similarly, Hanger (1979) reported that with
rapid growth, the import of photosynthates and
nutrients through the phloem increases rapidly,
 THE JOURNAL OF HORTICULTURAL SCIENCE AND BIOTECHNOLOGY
whereas Ca uptake falls to very low levels.
Commercial tomato production tends to be based
on indeterminate or semi-indeterminate tomato cul-
tivars rather than determinant ‘bush’ cultivars.
Determinant cultivars produce large flushes of fruit
late in the season. This heavy fruit set requires large
quantities of Ca that are very difficult for the plant to
supply on a steady basis. Indeterminate and semi-
indeterminate cultivars set fruit over a longer period
of time, and it is easier for the plant to supply enough
Ca to fewer fruit at any given time. If those last two
mechanisms are in place at the same time, the result
can be that BER incidence is not affected by growth
types.
Many researchers (Grassbaugh, Regnier, &
Bennett, 2004; Hudu, Futuless, & Gworgwor, 2002;
Schonbeck & Evanylo, 1998) have found that mulches
increase the yield of soil-grown tomatoes. Organic
mulches have been found to decrease the incidence
of Ca deficiency in plants (Elmer & Ferrandino, 1991;
John et al., 2005; Magnusson, 2002). Mulching results
in adequate soil moisture throughout the season and
thereby helps to reduce BER. Mulching increases soil
temperature, and Adams and Ho (1993) reported that
an increase in root temperature stimulated Ca uptake
in proportion to water uptake. Increased Ca uptake
will decrease the incidence of BER. However, mulch
may overheat the soil and therefore lead to an
increase in the incidence of BER. If these last two
mechanisms are in the place at the same time, the
result can be that BER incidence is not affected by
mulch treatment.
Olle (2015) found that some of the best methods
for preventing Ca deficiency in chervil and lettuce
involve growing plants in negative DIF conditions
and growing plants under a far-red filter. Both meth-
ods may also help prevent BER in tomatoes.
Negative DIF is when night temperature is higher
than day temperature. In such conditions plants
remain more compact. Fraszczak, Kaluzevicz,
Krzesinski, Lisijeka, and Spizewski (2011) showed
that basil plant height was enhanced by positive DIF
and was inhibited by negative DIF. Under negative
DIF conditions if plants stay more compact they con-
tain more Ca and there are fewer Ca deficiency symp-
toms in plants (Olle, 2015).
The use of negative DIF in young plants mainly
affects their vegetative growth, while a reproductive
plant hardly shows any reaction to the temperature
regime as long as the 24-hour temperature remains
constant (Heuvelink, 1989). Lowering the day tem-
perature and raising the night temperature in the
plant production phase or early in the season results
in more compact plants with firmer main and truss
stems (De Koning, 1994). Accordingly, stem elonga-
tion was remarkably restricted by negative DIF treat-
ments (Ito et al., 1995). Similarly, transplants have
3
greater economic value when their stem and/or
petiole elongation are prohibited in negative DIF
conditions. This type of environmental control is
easily achievable in a closed system with artificial
lighting (Kozai, 1998).
The yield was equal or higher when a negative DIF
was used; the development rate was not affected but
the fruits became larger (De Koning, 1994). Negative
DIF for 8 days decreased leaf area and chlorophyll
content. Leaf number did not differ in negative DIF
and control treatments. Relative growth rate, net
assimilation rate, and leaf area ratio decreased slightly
by negative DIF treatment.
Transpiration rate decreased considerably in the
light period, and increased markedly during the
dark period in the negative DIF plants (Ito et al.,
1995). Low transpiration flow to tomato fruits can
lead to low Ca levels and the development of BER
(Ho et al., 1993). Therefore, it can be concluded that
a marked increase of transpiration rate in the dark
period can lead to higher content of Ca in fruits and
prevent BER.
Lettuce plants have been grown under far-red fil-
ters; these filter out 45% of red light and 90% of far
red light (increasing the proportion of red light avail-
able to plants). Far-red light is responsible for plant
elongation. If a far-red filter is used to remove much
of the far-red light then plants are more compact.
Fletcher, Tatsiopoulu, Mpezamihigo, and Hadley
(2005) found that the far-red filtering films giving
the highest red:far red (R:FR) ratios reduced plant
height in Impatiens. Under a far-red filter, plants are
more compact in growth, contain more Ca, and show
fewer Ca-deficiency symptoms (Olle, 2015).
Light quality manipulations have been used to
decrease the need for chemical growth regulators.
One of these methods has involved filtering studies
to remove far-red light (Rajapakse & Shahak, 2007).
Different covers have been used to increase the R:FR
ratio, which decreases stem elongation (Rajapakse,
Young, McMahon, & Oi, 1999). In the 1980s, a
CuSO4 liquid filter was nominated as the best growth
regulating cover for tomato (Mortensen & Strømme,
1987), because there was a effective reduction of stem
elongation and internode length. However, CuSO4
covers did not become popular, because they were
difficult to handle and expensive. They also had phy-
totoxic properties which made their use hazardous
(Rajapakse et al., 1999).
The next step was the development of cheaper,
easier to handle materials, which replaced CuSO4
covers. Photoselective films made from plastic had
promising results (Lia, Rajapaksea, Youngb, & Oic,
2000). These dyed photoselective films had red and
far-red absorbing qualities, which enabled their use to
manipulate the R:FR ratio inside greenhouses (Lia
et al., 2000). They were of reasonable cost and
4 M. OLLE AND I. H. WILLIAMS

therefore were recommended especially for organic

growers (Rajapakse et al., 1999), who were not
allowed to use chemical growth regulators. For con-
ventional growers, the inconvenient delay in flower-
ing caused by far-red absorbing films, and the short
life span of the films themselves, hinders their com-
mercial use (Rajapakse & Shahak, 2007).
The content of Ca was higher in plants grown
under a far-red absorbing film. Olle (2015) compared
the effect of light quality (red and far-red light) and
DIF (positive and negative DIF). She found that stem
and shoot elongation are decreased and flowering
increased in negative DIF conditions and also in
conditions of much red light and less far-red light.
On the one hand, negative DIF conditions lead to
a markedly increased transpiration rate at night and
therefore increase the content of Ca in fruits and
prevent BER. On the other hand, stem and shoot
elongation are decreased and flowering increased in
negative DIF conditions and also in conditions of
much red light and less far-red light. Therefore, it
can be concluded that also under far-red absorbing
film the content of Ca increases and BER disorder is
prevented.
In summary, the options for reducing BER in Figure 2. Picture of concentric cracking of tomato.
greenhouse tomato production are:

(1) Avoid too high temperatures;

(2) Avoid too high light intensity;
(3) Apply organic mulches to decrease the inci-
dence of Ca deficiency in soil-grown plants;
(4) Grow plants in negative DIF conditions;
(5) Grow plants under far-red filters.

Fruit cracking and some methods to avoid it

FC (Figures 2 and 3) is a physiological disorder,
which mainly occurs when there is a rapid net influx
of solutes and especially water into the fruit, while at
the same time ripening or other factors reduce the
strength and elasticity of the tomato skin (Leonardi,
Guichard, & Bertin, 2000; Maroto et al., 1995; Peet,
1992). FC is the splitting of the epidermis around the
calyx or stem scar (Guichard et al., 2001).
There are three types of FC in tomatoes: (1) con-
centric cracking (Figure 2), which is a splitting of the
epidermis in circular patterns around the stem scar;
(2) radial cracking (Figure 2), which is a splitting of
the epidermis from the stem scar towards the blos-
som end; and (3) fruit cuticle cracking (Dorais,
Demers, Papadopoulus, & Ieperen, 2004), which has Figure 3. Picture of radial cracking of tomato.
several other names including russeting, hair crack-
ing, swell cracking, shrink cracking, rain check, craz-
ing, and cuticle blotch. Cracking downgrades the and reduces shelf life, while concentric and radial
quality of tomato, because cuticle cracking causes fruit cracking immediately renders the fruit unmar-
poor appearance (roughened skin and corky tissue) ketable for fresh consumption (Dorais et al., 2004).
 THE JOURNAL OF HORTICULTURAL SCIENCE AND BIOTECHNOLOGY
Cuticle cracking is a common problem in many crops
including sweet pepper, apple, pear, grape, and sweet
cherry (Dorais et al., 2004). The appearance of cuticle
cracking occurs in the last phase of fruit growth. FC
can occur at all stages of fruit growth but as fruits
mature they become more susceptible, especially as
colour develops (Olson, 2004). The earlier the fruit
cracks, the deeper the cracks become. Generally, fruit
cracking is most common on the large, beefsteak-type
tomatoes. The cracked areas can allow entry of bac-
terial and fungal pathogens that cause fruit rots. FC
of cherry tomatoes and was high after morning har-
vest, declined at noon and was low after evening
harvest (Lichter et al., 2002). This suggests that cherry
tomatoes should be harvested in the evening.
Tomato cultivars vary in the incidence of cracking
(Abbot et al., 1986; Fernandez-Munoz et al., 1995;
Maroto et al., 1995; Mullins & Straw, 1992; Pascual
et al., 1998; Sperry et al., 1996). Genotypic differences
in composition, skin anatomy, and cell morphology
are related to tomato fruit cracking. More susceptible
cultivars crack in the mature green stage and more
tolerant cultivars at later stages. Peet (1992) reported
that anatomical characteristics of crack-susceptible
cultivars are: (1) large fruit size Snapp, Huang, &
Warncke, 2002), (2) low skin tensile strength and/or
low skin extensibility at the turning to pink stage of
ripeness, (3) thin skin, (4) thin pericarp, (5) shallow
cutin penetration, (6) few fruits per plant, and (7)
fruit not shaded by foliage. The more resistant a
cultivar is, the later in maturity the fruit cracking
may occur. Sadhankumar, Rajan, and Peter (2001)
reported that the crack-susceptible cultivars had
higher juice content, acidity, total sugar content,
and reducing sugar content, and lower insoluble
solid content, pectin, skin and pericarp thickness,
and penetrance of fruits compared to the crack-resis-
tant cultivars. Crack-resistant cultivars are generally
associated with a thick cuticle and a compact and
resistant skin type as evidenced by high penetrance
values (Sadhankumar et al., 2001). Avdeyev and
Ivanova (2000) showed that one pair of dominant
genes controls resistance. The dominant genes cause
total resistance of fruit to radial and concentric crack-
ing. Kasai, Hayama, Kashimura, Kudo, and Osanai
(2008) demonstrated that induced accumulation of
an apple expansion gene in pericarp reduced the
susceptibility of fruit cracking in apples. Some culti-
vars of tomatoes have an epidermis that stretches well
and will have very little or no circular cracking.
Others do not stretch well and have a lot of cracking.
Some cultivars tend to have larger fruits than
others. Ehret, Hill, Raworth, and Estergaard (2008)
and Emmons and Scott (1998) observed that larger
tomato fruit tend to show more than do smaller ones.
Plants with a low fruit load produce larger fruit with
an increased incidence of cuticle cracking.
5
Consequently, the skin of large fruit should be sub-
jected to a greater stress than the skin of small fruit,
and therefore should be more likely to crack.
Most cultivars with a determinate growth habit
have been found to have low concentric cracking,
while most cultivars with an indeterminate growth
habit were found to have diverse concentric cracking
(from none to severe) (Hu, Wang, Chen, & Yang,
2012). Indeterminate and semi-indeterminate culti-
vars set fruit over a longer period of time, and it is
easier for the plant to supply enough nutrients to
fewer fruit at any given time.
Dorais et al. (2004) found that cultivars less sus-
ceptible to cracking are of Dutch origin and were
developed for growing conditions characterised by
low light intensities such as those prevailing in north-
ern Europe. Dutch cultivars tend to have thicker skin,
are more transportable, and have a lower incidence
of FC.
Ehret et al. (2008) found that, in tomato, FC
started 2 weeks after the time of maximum fruit
growth rate, increasing steadily thereafter until har-
vest. This suggests that rapid growth of plants may be
the reason for FC (why?).
Ehret et al. (2008) found that older tomato plants
grown under the same conditions, as younger plants
are more prone to FC in July and August. The reason
why the incidence of FC is lower in September than
in August could be that the weather plays a big role
for tomatoes grown in unheated greenhouses.
In the Northern Hemisphere, higher temperatures
and higher light intensities occur more in summer,
and during early autumn the temperatures and light
intensities are lower. Therefore, in summer the
increase in fruit temperature increases gas and hydro-
static pressures of the pulp on the skin, resulting in
immediate cracking in ripe fruit or delayed cracking
in green fruit. In addition, in summer high light
intensity may have a role in increasing cracking
apart from its association with high temperatures
(Peet, 1992). Under high light conditions, fruit solu-
ble solids and fruit growth rates are higher. Both of
these factors are sometimes associated with increased
cracking (Peet, 1992).
The temperature in September is much lower than
in August. However, Peet (1992) found that with
lower temperatures the incidence of FC is lower.
It could be assumed that conditions, which are
suitable for cuticle cracking, are suitable also for FC.
Dorais et al. (2004) reported that cuticle cracking of
greenhouse tomato fruit progressively increases as the
natural light level increases in spring and summer,
and decreases with the decline in light level during
the autumn. Increase in air temperature increases
fruit temperature. Increasing fruit temperature
increases gas pressure inside the fruit, which could
cause the fruit to expand in volume, thus stretching
6 M. OLLE AND I. H. WILLIAMS
the skin of the fruit, but this is unlikely to be the
primary cause of cracking. Alternatively, cuticle
cracking can occur due to higher temperature
because of an increase of assimilate supply to the
fruit and an increase in fruit growth (Dorais, et al.,
(2004). Kiyofumi, Hironobu, Shyuichi, and Hisamitsu
(2006) demonstrated that the sharp fluctuations of
temperature and humidity of the greenhouse were
closely associated with FC. The reason could be that
cracking could be related to high variations in fruit
growth and water influx under changing conditions
of temperature (Guichard et al., 2001).
In summary, the options for reducing FC in green-
house tomato production are:
(1) Avoid too high temperatures;
(2) Avoid too high light intensity;
(3) Avoid fluctuations in humidity.
Summary
Tomato can be injured by two very important
physiological disorders, namely blossom end rot
(BER) and fruit cracking (FC). Physiological Ca
deficiency is usually related to the inability of the
plant to translocate adequate Ca to the affected
plant part, rather than insufficient Ca levels in
the growing medium, and has been linked to
BER symptoms. The first visible symptom of
BER is a small darkened or water-soaked area
around the blossom end of the fruit, appearing
about the time the fruit begins to ripen. The spot
darkens, enlarges, and becomes sunken as the
fruits mature. Large lesions may show concentric
rings. The affected tissue is leathery and firm
unless invaded by secondary decay organisms.
BER usually causes the fruit to ripen prematurely
and to be inedible. FC is a physiological disorder,
which mainly occurs when there is a rapid net
influx of solutes and especially water into the
fruit at the same time that ripening or other fac-
tors reduce the strength and elasticity of the
tomato skin. FC is the splitting of the epidermis
around the calyx or stem scar. Tomato cultivars
differ in their susceptibility to BER and FC.
Organic mulches decrease the incidence of BER
on soil-grown plants. In conventional production
in the Northern Hemisphere, the incidence of FC
is lowest in July and higher in August than in
September. FC starts 2 weeks after the time of
maximum fruit growth rate, increasing steadily
thereafter until harvest. FC can occur due to
higher temperature because of an increase of
assimilate supply to the fruit and an increase in
fruit growth. The options for reducing BER in
greenhouse tomato production are: avoid too
high temperatures, avoid too high light intensity,
apply organic mulches to decrease the incidence of
Ca deficiency of soil grown plants, grow plants in
negative DIF conditions, or grow plants under far
red filters. The options for reducing BER in green-
house tomato production are: the first two options
for avoiding BER, and also to avoid fluctuations in
humidity.
Acknowledgements
The former institute (Jogeva Plant Breeding Institute) of
Estonian Crop Research Institute funded the literature
study and composition of the review article. Preparation
of the manuscript was further supported by institutional
research funding IUT36-2 of the Estonian Ministry of
Education and Research.
Disclosure statement
No potential conflict of interest was reported by the
authors.
Funding
The work was supported by the Institutional research
funding of the Estonian Ministry of Education and
Research. [Grant Number IUT36-2].
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