Article

Why calcium deficiency is not the cause of blossom-end rot in tomato and pepper fruit – a reappraisal

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Abstract

From a review of the relevant literature it is concluded that Ca2+ deficiency is not the cause but a result of blossom-end rot (BER) in tomato and pepper fruit. Actually, a depletion of the apoplastic pool of water-soluble Ca2+ in fruit has been observed only after the symptoms of BER were already visible, whereas in fruit at the early stages of BER development, the distribution and concentration of Ca2+ was still similar to that in healthy fruit. The actual causes of BER are obviously the effects of abiotic stress, e.g. by salinity, drought, high light intensity, heat, and ammonia nutrition, resulting in an increase of reactive oxygen species (ROS), high oxidative stress and finally cell death. Cell death results in a disintegration of the plasma membrane and tonoplast and a breakdown of the endoplasmic reticulum, thus not following but preceding ion leakage, including Ca2+ leakage, and loss of turgor. Bioactive gibberellins (GAs) reduce the accumulation of Ca2+ but increase the susceptibility to stress and the risk of BER, while abscisic acid (ABA) has the opposite effect. Ca2+ stabilizes cell structures and may thus limit cell expansion. It is usually sufficiently available for plant development and therefore Ca2+ deficiency is rare in nature. Application of GA biosynthesis inhibitors, such as prohexadione-Ca, and of GA antagonists, such as ABA, may completely inhibit the development of BER even at very low availability of Ca2+. With this approach, a better understanding and a more efficient control of BER in tomato and pepper fruit is envisaged.

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... Decreasing the sink/source ratio increased the delivery of phloem-borne assimilates in comparison to xylem-born Ca into fruits, so the low density of Ca in the blossom-end of fruits with a rapid growth rate induced BER (Ho & White, 2005), as well as promoting the strong and positive correlation between BER incidences and the fruit growth rate (Fig. 7). Yoshida et al. (2014), Saure (2005Saure ( , 2014, and Turhan et al. (2006) also reported that the incidence of BER was increased during the condition consisting of the high growth rate of fruit. High growth rates could increase the risk of the tissue content of Ca falling below the critical level required for cell wall stabilization and membrane integrity (Ho and White, 2005). ...
... Total tissue Ca concentration was not an accurate cause of BER development, because fruit with BER symptoms had similar or even higher Ca concentrations in comparison to healthy fruits (Ho & White, 2005). As a low total fruit Ca concentration did not lead to BER symptoms, Saure (2014) suggested that Ca partitioning and distribution at the cellular level could cause BER. ...
... As 1F and 2F fruits showed the highest relative growth rate (Fig. 1A), they had a limited time to uptake Ca for rapidly expanding cells and Ca concentration might fall below the critical level for cell wall stabilization and membrane integrity. Saure (2014) also reported that at low Ca supply, BER did not occur when the growth rate was low. The results obtained for the control treatment in our study confirmed this conclusion. ...
Article
Blossom‐end rot is generally considered a calcium‐related physiological disorder. The results of the previous studies show that several factors such as plant conditions can be effective on the blossom‐end rot incidence. Therefore, the present study was undertaken to investigate the effect of the sink/source ratio on the incidence of the blossom‐end rot of two greenhouse tomato (Solanum lycopersicum L.) cultivars: ‘Grandella' and ‘Isabella'. To this end, four treatments were applied: saving one fruit per truss (1F), two fruits per truss (2F), three fruits per truss (3F), and no fruit pruning (control). The results showed that the tomato cultivar ‘Isabella’ was more susceptible to the blossom‐end rot than ‘Grandella’. Decreasing the sink/source ratio increased the incidence of the blossom‐end rot and the relative fruit growth rate. The correlation between the blossom‐end rot incidence and the relative fruit growth rate showed that the fruit growth rate could be regarded as an important factor in the incidence of this disorder. Endogenous auxin and cytokinin concentrations acted as the regulators of the fruit growth rate and influenced it. Slowing down the relative growth rate by keeping proper sink/source ratio based on tomato cultivar is, therefore, an effective, cheap and healthy way to control the incidence of the blossom‐end rot, especially in organic farming. This article is protected by copyright. All rights reserved.
... Physiological disorders are one of the main factors reducing tomato productivity, and blossom-end rot (BER), together with fruit cracking, is the most important one (Olle and Williams, 2017), as it may occur in all tomato producing-areas worldwide, resulting in unmarketable fruits and wastage in the supply chain that generate important economic losses. Visible symptoms of this physiopathyresulting in a large water-soaked dark spot at the end of the fruit (Fig. 1)have been associated to a local plant nutritional disorder characterized by increased permeability and deterioration of fruit cell membranes, with subsequent loss of turgor and leakage of cell liquids into the cellular space (Ho and White, 2005;Saure, 2001Saure, , 2014. Typical symptoms occurs 2-4 weeks after anthesis, when the young fruit is in rapid expansion (Gutteridge and Bradfield, 1983;Saure, 2001;Sonneveld et al., 1991). ...
... Chiu and Bould (1976) firstly stated that no decrease in BER occurrence was detected at reduced Ca +2 supply, while Nonami et al. (1995) observed that fruits with BER symptoms had similar distribution and concentration of Ca 2+ ions compared to the healthy ones. These findings suggested that Ca 2+ deficiency in fruits may not be the only direct determinant of BER in tomato plants, and subsequently this cause-effect relationship was questioned by Saure (2014Saure ( , 2001. According to this author, BER incidence is not directly linked to Ca 2+ deficiency. ...
... Rather, BER would be induced by oxidative stress (e.g. triggered by unfavorable environmental conditions): the leakiness of plasma membranes would be an effect of an increased reactive-oxygen species (ROS) concentration that could lead to loss of apoplastic Ca 2+ ions (Breusegem and Dat, 2006;Saure, 2014). According to this explanation, low concentration of calcium in fruit apoplast should not be considered as the cause, but an effect of BER, whose occurrence is actually related to multiple factors, including environmental, genetic and physiological aspects (Fig. 2). ...
Article
Blossom-end rot (BER) is a physiological disorder causing severe losses in tomato crops. Despite its economic importance and the large collection of studies concerning its onset, BER occurrence is still poorly understood, making its prediction and prevention very difficult. Currently, two theories are accredited to explain the BER onset: the first one identifies a reduced root uptake and an aberrant regulation of cellular partitioning of calcium as the major physiopathy agent, while the second one hypothesizes a primary role to abiotic stresses, as they induce reactive oxygen species (ROS) production in the plant, leading to membranes disintegration and loss of cell turgor. To date, there are no unequivocal proofs that allow us to definitively go beyond one of the two hypotheses. Rather, a multitude of genetic, physiological and environmental factors form a complex network of interactions and synergies contributing to BER occurrence. This is why the "multi-disciplinary approach" is maybe the most appropriate one to understand this physiopathy and to develop new and effective BER-contrasting tools with genetic and agronomic methods. This review adopts this kind of approach to investigate the causes of BER and to describe the practices preventing its occurrence, possibly providing the most complete compendium about this disorder to date.
... Following the new soilless cropping techniques, where the producer provides the necessary nutrient levels that the plant requires at any time, the possibility that mild Ca 2+ deficiency alone is not the main cause of these physiological disorders is beginning to be assessed [18,26]. Several authors speculate on the cause-effect relationship of Ca 2+ deficiency in both disorders, observing that on many occasions, fruits with these symptoms contained equal or higher concentrations of Ca 2+ in their tissues [26]. ...
... Following the new soilless cropping techniques, where the producer provides the necessary nutrient levels that the plant requires at any time, the possibility that mild Ca 2+ deficiency alone is not the main cause of these physiological disorders is beginning to be assessed [18,26]. Several authors speculate on the cause-effect relationship of Ca 2+ deficiency in both disorders, observing that on many occasions, fruits with these symptoms contained equal or higher concentrations of Ca 2+ in their tissues [26]. Other studies indicated that either low levels or high levels of Ca 2+ in the nutrient solution led to the appearance of BER in fruits of various species [27]. ...
... Many authors, in their eagerness to predict and act on time against these problems, have centered their attention on the study of the main triggers of these physiological disorders. These studies distinguish different agronomic, physiological, and genetic factors that favor the induction of BER and TB [15,18,20,26,28]. In the following sections, we will succinctly describe all these factors and their relationships. ...
Article
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In the current scenario of human-driven climate change, extreme weather events will likely affect agricultural production worldwide. Soilless production systems have recently arisen as a solution to optimize the use of natural resources, such as water and soil, and hence will contribute to reducing the environmental impact of agriculture. However, nutritional imbalance due to adverse environmental factors, such as drought, high temperatures, and salinity, might produce calcium-related physiological disorders during plant growth, such as blossom-end rot (BER) in fruits and tipburn (TB) in leaves, which are a serious problem in crop production. Here, we discuss the different agronomic, physiological, and genetic factors that favor the induction of BER in tomato and TB in lettuce and anticipate the use of an integration of breeding and technological approaches to alleviate nutritional disorders in soilless production systems.
... Thus, the nutritional content and dry weight under each treatment were re-calculated as a percentage relative to each control condition (of the corresponding experiment, in vitro or in pots), and the nutritional content was expressed as a percentage of plant dry weight, as indicated in Table S6. As observed in Figure 6, the control conditions for in vitro and in-pot treatments correlate with each other and also with nitrate excess; this was expected as nitrate should not have a negative impact on plant nutrition when compared to Ca deficit or ammonium treatment [33]. In the case of ammonium treatment, it correlated with N content and was negatively correlated with potassium content, which is an indication of the competition with the excess of NH4 + [34]. ...
... This is in accordance with the experiments in vitro. It has been previously described that plants with insufficient Ca in the soil show higher levels of p in roots, a nutrient that promotes root development, allowing a greater contact surface with the soil [33,43] and thus extracting as much Ca as possible. The balance between nutrients is key for proper plant development. ...
... This is in accordance with the experiments in vitro. It has been previously described that plants with insufficient Ca in the soil show higher levels of p in roots, a nutrient that promotes root development, allowing a greater contact surface with the soil [33,43] and thus extracting as much Ca as possible. ...
Article
Full-text available
Both nitrogen and calcium fertilization management are vital for crops, where an imbalance of these elements can cause both physiological and yield problems. It has been proposed that nitrogen absorption, particularly ammonium, is in part dependent on calcium supply. Moreover, the balance between these two nutrients could be a key indicator of plant growth in some species. Tomato, one of the most cultivated crops worldwide, can also be widely affected by nutritional imbalance. Using large amounts of N fertilizers could lead to an imbalance with other nutrients and, thus, detrimental effects in terms of plant development and yield. Here we show that ammonium excess has a negative impact on plant development and results in calcium deficiency. Moreover, a deficit in calcium nutrition not only affects calcium concentration but also leads to a restriction in N uptake and reduced N concentration in the plant. These effects were evident at the seedling stage and also during flowering/fruit set. Using PCA analysis, we integrated both phenotypic and nutritional imbalances in seedlings and grown plants. Interestingly, the Ca/N ratio appears to be a key indicator to monitor appropriate N and calcium nutrition and more importantly the balance between both. Maintaining this balance could be an essential element for tomato crop production.
... These results indicate that Apogee and ABA triggered the expression of a range of transcription factors up-regulating gene networks involved in stress responses that resulted in fruit tissue becoming more resistant to BER compared to GA 4 + 7 -treated fruit. Other studies suggested that fruit susceptibility to BER is mainly dependent on its ability to deal with reactive radicals naturally produced in response to stresses such as salinity, drought, high light intensity, heat, and ammonia nutrition (Ho and White 2005;Saure 2014). Bioactive GAs may increase fruit susceptibility to stress and BER development, while Apogee and ABA could have the opposite effect (Saure 2014). ...
... Other studies suggested that fruit susceptibility to BER is mainly dependent on its ability to deal with reactive radicals naturally produced in response to stresses such as salinity, drought, high light intensity, heat, and ammonia nutrition (Ho and White 2005;Saure 2014). Bioactive GAs may increase fruit susceptibility to stress and BER development, while Apogee and ABA could have the opposite effect (Saure 2014). ...
... Therefore, comparing fruit with higher and lower susceptibility to BER, it was possible to conclude that mechanisms inhibiting BER development were possibly involved in increasing fruit tissue resistance to oxidative stress and toxic compounds. Indeed, studies have shown that fruit susceptibility to BER is enhanced by abiotic stress conditions, such as salinity, drought, high light intensity, heat, and ammonia nutrition (Ho and White 2005), which increase ROS and other toxic compounds that eventually can lead to cell death and BER symptoms (Saure 2014). ...
Article
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The objectives of this study were to analyze changes in gene expression and identify candidate genes and gene networks potentially inhibiting or triggering blossom-end rot (BER) in tomatoes treated with plant growth regulators. ‘Ace 55 (Vf)’ tomato plants were grown in a greenhouse and sprayed with Apogee (300 mg L⁻¹), abscisic acid (ABA) (500 mg L⁻¹), water (control), or gibberellins 4 + 7 (GA4 + 7) (300 mg L⁻¹) weekly after pollination. The BER incidence rate was zero in Apogee- and ABA-, medium in water-, and high in GA4 + 7-treated plants from 26 to 40 days after pollination (DAP). At 26 DAP, healthy blossom-end fruit tissue still not showing visible BER symptoms was used for transcriptome analysis. Candidate genes potentially inhibiting or triggering BER were identified through a correlation analysis between gene expression levels at 26 DAP and BER incidence rate from 26 to 40 DAP. Genes inhibiting BER should be up-regulated in Apogee- and/or ABA-treated fruit and down-regulated in GA4 + 7-treated fruit. Genes triggering BER should be down-regulated in Apogee- and/or ABA-treated fruit and up-regulated in GA4 + 7-treated fruit. Most of the candidate genes inhibiting BER have functions leading to higher resistance to oxidative stress and toxic compounds, whereas most of the candidate genes triggering BER have functions leading to higher levels of oxidative stress and cell death. The results suggest that Apogee and ABA inhibited BER possibly by increasing fruit tissue resistance to reactive oxygen species (ROS) and other toxic compounds, whereas GA4 + 7 triggered BER possibly by increasing the levels of fruit oxidative stress.
... Many studies suggested that BER is a symptom of Ca deficiency physiological disorder in the distal fruit tissue. However, Saure (2014) implied that Ca deficiency is not the cause but a result of BER in tomato fruit because depletion of the apoplastic water-soluble Ca contents in fruit was observed only after BER symptoms were visible. In this study, the total and watersoluble Ca content in the distal fruit tissue were similar in IL5-4 and 'M82' (Fig. 6). ...
... In this study, the total and watersoluble Ca content in the distal fruit tissue were similar in IL5-4 and 'M82' (Fig. 6). These results were in agreement with the suggestion of Saure (2014), and our results indicated that a higher incidence of BER in IL5-4 fruit might not be related to both total and watersoluble Ca contents in distal fruit tissue. However, to elucidate this, it would be necessary to investigate the localization of water-soluble Ca because it is suggested that water-soluble apoplastic Ca content is related to BER incidence (Ho and White, 2005). ...
... However, to elucidate this, it would be necessary to investigate the localization of water-soluble Ca because it is suggested that water-soluble apoplastic Ca content is related to BER incidence (Ho and White, 2005). The development of BER requires several steps; abiotic stress increases the production of reactive oxygen species (ROS) and causes lipid peroxidation with an increase in membrane leakiness (Saure, 2014). In our study, the high incidence of BER in IL5-4 fruit may not have been caused by abiotic stress, as both IL5-4 and 'M82' plants were grown under the same conditions. ...
Article
Fruit Brix is an important indicator in determining the quality of tomatoes (Solanum lycopersicum), and increasing it is an important objective. The production of high Brix tomatoes requires breeding and genetic studies of fruit. During domestication S. lycopersicum lost genetic variation of some wild tomato relative that could be useful for breeding. In this study, we investigated introgression lines (ILs) from a cross between the wild relative Solanum pennellii and the cultivated tomato S. lycopersicum ‘M82’. While there are many genetic and physiological studies that demonstrate the usefulness of tomato S. pennellii ILs, few have investigated the high Brix values of IL fruit. Accordingly, we attempted to detect tomato ILs that resulted in high Brix ripening fruit, in order to obtain valuable genetic and genomic resources for the investigation of phenotypes originating in the S. pennellii genome. IL5-4 may be a line that carries an S. pennellii chromosome segment on chromosome 5 of ‘M82’. Previous research indicated that IL5-4 fruit have higher Brix levels than ‘M82’ fruit. Our results corroborated these findings and revealed Brix changes in fruit during development. We also found that IL5-4 plants showed a higher incidence of blossom-end rot (BER), a major physiological disorder in tomatoes. Therefore, we investigated the physiological mechanism responsible for the higher incidence of BER in IL5-4, by focusing on calcium content, which may be related to BER occurrence. The total and water-soluble Ca contents of fruit tissues were significantly lower in IL5-4 than in ‘M82’ in the proximal part, while no differences were observed in the distal part. Thus, our results suggested that a higher incidence of BER in IL5-4 fruit may not be related to both total and water-soluble Ca contents in the distal fruit tissue, and genetic factors originating in the S. pennellii chromosome may induce high BER incidence in IL5-4. The characterization of IL5-4 in this study showed that it is a valuable genetic and genomic resource for high-Brix breeding stock and for the investigation of novel BER mechanisms.
... Thus, the low concentration of Ca 2+ , especially in the apoplast, negatively affects the integrity and function of the cellular membranes (Ho and White, 2005;Hocking et al., 2016). In the most extreme cases, Ca 2+ deficiency can cause cell membrane disruption, followed by the extravasation of the solutes and cellular plasmolysis, resulting in the formation of BER symptoms in fruits (Saure, 2001(Saure, , 2014. In addition, the mobility of Ca 2+ in the plant is extremely restricted, as it is basically transported via the xylem through transpirational water flow, which explains why the leaves are the main site of Ca 2+ accumulation in the detriment of the fruits (Saure, 2005;Olle and Bender, 2009;Hocking et al., 2016). ...
... In addition, the mobility of Ca 2+ in the plant is extremely restricted, as it is basically transported via the xylem through transpirational water flow, which explains why the leaves are the main site of Ca 2+ accumulation in the detriment of the fruits (Saure, 2005;Olle and Bender, 2009;Hocking et al., 2016). Indeed, due to fruits' low Ca 2+ sink strength, they present the smallest accumulations of Ca 2+ and greater susceptibility to the appearance of Ca 2+ deficiency symptoms (Saure, 2014;Kumar et al., 2015;Hocking et al., 2016). ...
... Recently, several studies have shown that these Ca 2+ transporters are under a complex and intricate hormonal regulation (De Freitas et al., 2011bDe Freitas et al., 2014). For instance, evidence has been raised indicating gibberellin (GA) and abscisic acid (ABA) crosstalk in the coordination of CAX activity in the tonoplast and BER development symptoms (De Freitas et al., 2011a, 2014Saure, 2014). In tomato plants, cv. ...
Article
Blossom end rot is one of the main physiological disorders of the tomato crop. Usually, the blossom end rot (BER) occurrence is associated with fruit calcium (Ca2+) deficiency; however, increasing evidence of Ca2+ disorders and hormonal regulation has emerged. Therefore, the present work was undertaken to investigate the role of gibberellin (GA) signaling on the control of tomato plant responses to Ca2+ deficiency. In this study, we used the tomato mutant procera (pro), which has a constitutive response to GA, and its isogenic line cv. Micro-Tom (MT) grown in a hydroponic system with or without Ca2+. Subsequently, the analysis of the growth of the shoot and the root system was performed, and the contents of photosynthetic pigments and oxidative stress were also analyzed. Furthermore, the fruit production, BER incidence, accumulation and metabolism of Ca2+ were also analyzed. The deficiency of Ca2+ reduced both shoot and root growth in both genotypes. Likewise, photosynthetic pigment degradation and oxidative stress were induced by Ca2+ deficiency. However, only pro plants exhibited the inhibition of fruit dry mass, as well as a higher incidence of BER, when exposed to Ca2+ deficiency compared to MT plants. In accordance with the greater pro susceptibility to Ca2+ deficiency, the efficiency of the transport, use and accumulation of Ca2+ in fruits was lower in pro when compared to the MT. Thus, we conclude that the higher sensitivity to GA in pro plants impairs Ca2+ metabolism and favors the occurrence of BER in tomato fruits, opening new approaches for the GA signaling on BER response.
... In this research, applying 5 mL•L -1 of iodine resulted in the highest firmness value (3.26 N) with respect to the control, while the values obtained with 200 mg•L -1 of calcium prohexadione and 3 mL•L -1 of iodine were the lowest for this variable (Table 3). Saure (2014) indicates that obtaining adequate fruit growth requires increasing the synthesis and concentration of gibberellins (GB), of which the most important in the pericarp of the tomato fruit are those of the GB1 and GB20 types (Bohner, Hedden, Bora-Haber, & Bangerth, 1988). The GB1 content during fruit growth and development is high, and the Ca 2+ concentration is reduced; this is generated by the plant to allow cell expansion and increase membrane permeability, which is restored when the fruit has reached physiological de coadyuvar al sistema de defensa antioxidante en contra de especies reactivas de oxígeno, ocasionadas por factores abióticos (presencia de sales o metales pesados) (Caffagni et al., 2012). ...
... Entre los parámetros del calidad de fruto más apreciados por el consumidor se encuentra la firmeza de la pulpa (Figueroa-Cares et al., 2018), la cual se relaciona con sus maturity (Marschner, 1995). In this sense, applying a GB synthesis inhibitor, such as calcium prohexadione, is usually an alternative to reduce the risk of calcium deficiency in the early stages of fruit growth and development (Saure, 2014). ...
... For its part, iodine is a non-essential microelement for plants (except for some aquatic species); however, it does have some beneficial effects (Caffagni et al., 2012) as it has been reported that it can induce a decrease in the respiration process and ethylene synthesis in cherry tomato fruits (Islam et al., 2018), which maintains the integrity of the cell wall (Dhall & Singh, 2013;Saure, 2014). This is desirable if the fruit's shelf life is to be increased. ...
Article
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Palabras clave: Solanum lycopersicum L., etefón, prohexadiona de calcio, yodo, selenito de sodio. Abstract S olanum lycopersicum L. is one of the most consumed horticultural products in the world, due to its wide versatility in use (fresh and processed) and high nutraceutical value. The objective of this research was to evaluate the effect of preharvest spraying of ethephon, calcium prohexadione, iodine and sodium selenite on some physicochemical quality parameters in greenhouse-grown saladette tomato fruits. The experimental design was completely randomized and the evaluated variables were color, weight, equatorial and polar diameter, roundness index, firmness, total soluble solids (TSS), titratable acidity (TA) and lycopene concentration. Individual application of ethephon (1.6 mL•L-1) and iodine (5 mL•L-1) significantly increased fruit weight (133.71 g) and firmness (3.26 N), with respect to the control (80.36 g and 0.95 N). Fruits that showed the highest TA (0.34 % citric acid) were those sprayed with 125 mg•L-1 of sodium selenite. On the other hand, the equatorial and polar diameter, TSS and lycopene concentrations, brightness, hue and roundness index did not differ statistically among treatments. Preharvest foliar application of ethephon, iodine and sodium selenite could be considered as an agronomic management alternative in greenhouse tomato production systems. Resumen S olanum lycopersicum L. es una de las hortalizas de mayor consumo a nivel mundial, debido a la amplia versatilidad en su uso (fresco y procesado) y por su alto valor nutracéutico. El objetivo de esta investigación fue evaluar el efecto de la aspersión precosecha de etefón, prohexadiona de calcio, yodo y selenito de sodio sobre algunos parámetros de calidad fisicoquímica en frutos de tomate saladette cultivados en invernadero. El diseño experimental fue completamente al azar y las variables evaluadas fueron color, peso, diámetro ecuatorial y polar, índice de redondez, firmeza, sólidos solubles totales (SST), acidez titulable (AT) y concentración de licopeno. La aplicación individual de etefón (1.6 mL•L-1) y yodo (5 mL•L-1) incrementó significativamente el peso (133.71 g) y firmeza (3.26 N) de los frutos, respectivamente, con respecto al testigo (80.36 g y 0.95 N). Los frutos que presentaron mayor AT (0.34 % de ácido cítrico) fueron los asperjados con 125 mg•L-1 de selenito de sodio. Por su parte, el diámetro ecuatorial y polar, la concentración de SST y de licopeno, la brillantez, la tonalidad de color y el índice de redondez no difirieron estadísticamente entre los tratamientos. La aplicación foliar en precosecha de etefón, yodo y selenito de sodio podría ser considerada como una alternativa de manejo agronómico en los sistemas de producción de tomate en invernadero.
... Since Ca 2+ is thought to play a key role, BER is termed a "calcium-related disorder" [93]. BER incidence is related to environmental factors, like high salinity, water scarcity, high temperature, and ammonia nutrition, which contribute to Ca 2+ deficiency [91,94,95]. However, a close relationship between calcium levels and BER cannot always be demonstrated [90]. ...
... This situation is also aggravated when plants are grown vigorously, when GAs levels are high and when ABA is low. All these are typical BER symptoms [94]. Thus final Ca 2+ deficiency can be considered a result, but not the cause, of only BER. ...
... The enriched CO 2 increased plant growth only when there was high nutrient availability [23,24]. Adding Ca fertilizer can promote plant growth and significantly increase plant yield in tomatoes [50] and cucumber and melon [51]. This may be connected with the findings that Ca promotes N's absorption by plants, increases NO 3 enzyme activity in the leaf, and enhances plant photosynthetic capacity [50,52]. ...
... Adding Ca fertilizer can promote plant growth and significantly increase plant yield in tomatoes [50] and cucumber and melon [51]. This may be connected with the findings that Ca promotes N's absorption by plants, increases NO 3 enzyme activity in the leaf, and enhances plant photosynthetic capacity [50,52]. But 15 N tracer of N results confirmed that rising of Ca levels forms antagonistic effects under high levels, affecting N absorption, distribution, or utilization in plants, and finally influencing the plant growth of apples [52]. ...
Article
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The vegetative growth, photosynthetic, and stomatal characteristics were investigated in Phalaenopsis Queen Beer ‘Mantefon’ to determine light’s influence with calcium ammonium nitrate (CAN) levels under 800 μmol·mol−1 CO2. Two lights (150 ± 20 and 300 ± 20 μmol·m−2·s−1) and CAN levels were employed for 40 weeks: calcium, ammonium, and nitrate levels by 0.90, 0.55, and 2.97 mmol·L−1 (CAN1), 8.63, 1.11, and 6.05 mmol·L−1 (CAN2), 12.80, 1.72, and 9.13 mmol·L−1 (CAN3), and 18.80, 2.27, and 12.20 mmol·L−1 (CAN4), respectively. The number of leaves increased in the plants grown at 300 ± 20 μmol·m−2·s−1 with CAN1 compared to control. Plants grown at 300 ± 20 μmol·m−2·s−1 with CAN4 had the lowest number of leaves among all plants. The time to the mature leaf span decreased in the plants grown at 300 ± 20 μmol·m−2·s−1 with CAN1. The net CO2 uptake was higher in the plants grown at 300 ± 20 μmol·m−2·s−1 than those grown at 150 ± 20 μmol·m−2·s−1 with CAN1–3 conditions. The water-use efficiency is higher in the plants grown with CAN1 than those with CAN2–4 at 300 ± 20 μmol·m−2·s−1. The maximum stomatal aperture was the largest in the plants grown at 300 ± 20 μmol·m−2·s−1 with CAN1–2 among all plants. Consequently, light levels of 300 ± 20 μmol·m−2·s−1 in Phalaenopsis Queen Beer ‘Mantefon’ must be accompanied by nutrient CAN1 to improve photosynthesis and stomatal activity and promote leaf growth under 800 μmol·mol−1 CO2 conditions.
... BER is a physiological disorder of tomato fruit that is now considered to be driven by abiotic stress conditions, including increased temperatures, relative humidity, and soil water fluctuations, resulting in an increase in reactive oxygen species that cause high oxidative stress, and finally cell death (Saure, 2014). Stress may interfere with calcium (Ca 2+ ) allocation or uptake by the fruit, making the observed low Ca 2+ in symptomatic BER fruit a secondary consequence of the stress that causes the disorder (Saure, 2014). ...
... BER is a physiological disorder of tomato fruit that is now considered to be driven by abiotic stress conditions, including increased temperatures, relative humidity, and soil water fluctuations, resulting in an increase in reactive oxygen species that cause high oxidative stress, and finally cell death (Saure, 2014). Stress may interfere with calcium (Ca 2+ ) allocation or uptake by the fruit, making the observed low Ca 2+ in symptomatic BER fruit a secondary consequence of the stress that causes the disorder (Saure, 2014). The production system · grafting · cultivar interaction found in 2016 showed that the percentage of total yield that was unmarketable due to BER was significantly higher for 'Tribute' high tunnel fruit (5.7%) than for 'Tribute' open field fruit (0.6%), while no significant difference was identified for 'Garden Gem'. ...
Article
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Although grower interest in high tunnel tomato ( Solanum lycopersicum ) production has increased in recent years, systematic high tunnel research conducted in humid, subtropical regions has been limited. The potential of tomato grafting to mitigate biotic and abiotic stresses makes it complementary to high-value production systems in high tunnels. In this 2-year study, grafted vs. nongrafted organic tomato production in high tunnels and open fields was investigated to determine possible synergistic effects of these two technologies. In 2016, high tunnels resulted in a significant increase of total and marketable yields, by 43% and 87%, respectively, over open field production. Grafting also significantly increased total and marketable yields over nongrafted plants by 34% and 42%, respectively. Cultivar effects demonstrated greater benefits with the implementation of high tunnel and grafting technologies for ‘Tribute’ (a beefsteak-type tomato) than for ‘Garden Gem’ (a plum-type tomato), as the increase in marketable yield was 33% greater for ‘Tribute’ in high tunnels and 45% greater for ‘Tribute’ with grafting. In 2017, a delayed effective transplanting date and the lack of high tunnel summer season extension produced results that were generally cultivar specific. While grafting increased the total yield of both cultivars (by 18%), marketable yield was increased by grafting only for ‘Tribute’ in high tunnels (by 42%). Additionally, high tunnels improved marketable yield of ‘Tribute’ by 129% but had no effect on ‘Garden Gem’. This demonstrated the consistent trend of the beefsteak-type tomato benefiting more from the combination of high tunnel and grafting technologies than the plum-type tomato. High tunnels reduced fruit decay and cracking by up to 71% compared with open field production. Stink bug (Pentatomidae) damage had the greatest impact on marketable yields each season, reaching 13% and 34% of total yields in 2016 and 2017, respectively, and was unaffected by high tunnel production or grafting. This study revealed the benefits of integrating high tunnel and grafting technologies for enhancing organic production of fresh-market tomato in the humid subtropics, and demonstrated more research is warranted to establish regional planting dates and further optimize this high-value cropping system.
... Thus, a number of studies have shown that Ca deficiency in tomato fruits is a major factor for BER. By contrast, however, it was also suggested that a relationship between Ca deficiency and the occurrence of BER is not always obvious (Saure, 2014). This may be due to the combined involvement of a variety of other factors, such as salinity, other nutrient status, temperature, and humidity, in BER's onset (Ikeda and Kanayama, 2015;Taylor and Locascio, 2004). ...
... This may be due to the combined involvement of a variety of other factors, such as salinity, other nutrient status, temperature, and humidity, in BER's onset (Ikeda and Kanayama, 2015;Taylor and Locascio, 2004). These abiotic stresses can induce oxidative stress, which is also considered a possible factor for BER (Saure, 2014). In bell pepper (Capsicum annuum L.), which belongs to the same family as tomato, Silber et al. (2005) reported a negative correlation between fruit manganese (Mn) concentration and BER incidence. ...
Article
Fruit blossom-end rot (BER) is a serious physiological disorder that can cause significant yield losses in tomato (Solanum lycopersicum). Although many studies have suggested that calcium (Ca) deficiency in tomato fruits is a major factor for BER, its onset mechanism has not been fully elucidated. Ionomics is a high-throughput elemental profiling of living organisms that can be applied to understand how differences in plant's physiological status involving inorganic elements. In this study, we examined ionomic differences between the tomato cultivar M82 and its introgression line IL8-3, which contains a short chromosome segment from its wild relative Solanum pennellii on chromosome 8 of M82, and has a low incidence of fruit BER. Among the essential elements, Ca showed marked different behavior between the two lines. IL8-3 showed preferential Ca partitioning to fruits compared with M82. The slow growth rate and high Ca concentration observed in IL8-3 fruit during the early growth stages may also be responsible for the low BER incidence in this line. Although Ca ions bind to cell wall pectin and membrane phospholipids, and contribute to cell structure stability, these components showed no significant differences between fruits of the two lines. The fruit ionome differed considerably between M82 and IL8-3, and was not affected by available Ca status in the field. The M82 fruit had higher concentrations of many elements such as magnesium, potassium, boron, and sulfur than did IL8-3, and this trend was also observed in rotten fruit. This suggests that the influence of the leaf (source), rather than the fruit (sink), could be involved in the onset mechanism of BER.
... If necrosis occurs at the end of the fruit, it is difficult to distinguish IB from BER. There have been many studies of BER (de Freitas et al., 2012;Ho and White, 2005;Saure, 2014) and it was reported that a local deficiency in Ca induces cell necrosis. In recent studies about BER, it was suggested that Ca 2+ transport was an essential factor in the lower incidence of BER using a tomato introgression line (Ikeda et al., 2017); the incidence of BER under salinity was reduced by organo-mineral fertilizer (Kataoka et al., 2017); the cultivar difference in the susceptibility to BER was likely explained by the difference in the growth rate of young fruit affecting watersoluble Ca in the distal part of tomato fruit (Vinh et al., 2018); the involvement of reactive oxygen species as a major protagonist in BER appearance; and BERresistant cultivars showed a larger increase in their ROS scavenging capacity (Rached et al., 2018). ...
... Our anatomical research also showed that IB was due to Ca deficiency, the same as BER. BER is caused by the effects of abiotic stress, resulting in an increase in reactive oxygen species, high oxidative stress and finally cell death (Saure, 2014). Further study is necessary to clarify what causes cell death around the vascular bundle in IB fruit. ...
Article
Internally brown (IB) tomato fruit is a physiological disorder in which the inside of the fruit turns brown or black. The mechanisms underlying the development of IB are not well understood. In this study, we examined the incidence of IB using hydroponics, and investigated the anatomical features and the ratio of dimeric rhamnogalacturonan II-borate (dRG-II-B) to total rhamnogalacturonan II (RG-II) (boron cross-linking ratio) in cell walls that decreased with boron deficiency. IB fruit developed when the growth of the stem and leaves were normal and the micro-element concentrations were low. The IB region was observed to be brown inside the fruit at an early stage, and this changed to black in the mature fruit. It was detected around the pericarp, locular gel, placenta and columella in the tomato fruit. An abnormality was observed around the vascular bundle in IB fruit. The interior of the cells near the vascular bundle was changed to a substance with a high electron density, and a disintegrated image was observed. Few Ca precipitates were observed on plasma membranes or cell walls in the collapsed cells by an antimonite precipitation method. It is suggested that Ca deficiency appears to be related to the cell collapse in IB. There was no significant difference in the boron cross-linking ratio between the IB and normal fruit. This suggested that IB was not related to B deficiency directly. We concluded that IB was caused by necrosis of cells around the inner vascular bundle due to Ca deficiency.
... The effect of the improvement in water relations on reducing BER incidence may be related to the reduction in generation of reactive oxygen species (ROS), which may increase fruit susceptibility to BER (Schmitz-Eiberger & Noga, 2003;Saure, 2014). Thus, the varieties that are more tolerant to stress conditions should present lower susceptibility to BER. ...
... However, speculation is that this physiological disorder is controlled by several factors, not only the nutrient content in the fruit but also the factors that affect its distribution between leaves and fruit, such as leaf stomatal conductance and leaf transpiration. In fact, our results identified the roles of several factors acting either to inhibit or trigger BER in different varieties, in addition to tolerance to high temperatures, as observed by other authors (Saure, 2014). ...
Article
Full-text available
Calcium (Ca2+) is a nutrient in tomato plants, of which deficiency usually causes several problems including a physiological disorder known as blossom-end rot (BER) in the fruit. The objective of this study was to evaluate and identify morphological and physiological characteristics related to the susceptibility of tomato varieties to BER. The varieties studied were 'Amalia', 'IPA-6', 'M-82', 'Mara', and 'Nagcarlan', presenting different fruit formats. Physiological parameters that negatively correlated with BER were plant water potential, leaf area, plant dry mass, relationship between proximal/distal Ca 2+ , K + content in the proximal and distal portions of the fruit, and proximal Ca 2+ content. Physiological parameters that positively correlated with BER were number of trichomes in the abaxial and adaxial leaf portions, leaf stomatal conductance, distal Ca 2+ content bound to the cell wall, leaf transpiration, and fruit length. Our results showed that 'Mara' and 'Nagcarlan', 'Amalia' and 'IPA-6', and 'M-82' presented low, medium, and high susceptibility to BER, respectively. We also found that total fruit Ca 2+ concentration, particularly in the distal fruit tissue, was not the only factor responsible for the development of BER; rather, the balance between factors that increase and decrease the susceptibility of each variety affected development of this disorder.
... The use of BRs has been reported to activate plant antioxidant defenses such as a generation of compounds such as ascorbic acid or even to activate enzymes such as superoxide dismutase (SOD), ascorbate peroxidase (APX), and catalase (CAT) that inactivate ROS (Yadav et al. 2012;Maia et al. 2018). Thus, stress tolerance conferred through the exogenous application of this hormone is different from all the mechanisms already analyzed in studies on Ca 2+ disorders (Saure 2001(Saure , 2014. ...
... For many years, researchers have considered BER as a disorder caused only by Ca 2+ depletion in the tissues in the distal portion of the fruit. However, more recently, it has been concluded that localized Ca 2+ deficiency may lead to membrane leakage, which results in BER symptom development (Saure 2014). ...
Article
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Blossom-end rot (BER) is a physiological disorder believed to be triggered by low Ca²⁺ content in the distal fruit tissue. However, many other factors can also determine fruit susceptibility to BER. It is possible that during fruit growth, Ca²⁺ imbalance can increase membrane leakiness, which may trigger the accumulation of reactive oxygen species, leading to cell death. Brassinosteroids are a class of plant hormones involved in stress defenses, specially increasing the activity of antioxidant enzymes and the accumulation of antioxidant compounds, such as ascorbic acid. The objective of this study was to understand the mechanisms by which 24-epibrassinolide (EBL) reduces fruit susceptibility to BER. Tomato plants ‘BRS Montese’ were cultivated in a greenhouse and were weekly sprayed with water (control) or EBL (0.01 µM) after full bloom. Plants and fruits were evaluated at 15 days after pollination (DAP). According to the results, EBL treatment inhibited BER development, increased fruit diameter, length, and fresh weight. EBL-treated fruit showed higher concentrations of soluble Ca²⁺ and lower concentrations of cell wall-bound Ca²⁺. EBL-treated fruit also had higher concentrations of ascorbic acid and lower concentrations of hydrogen peroxide, compared to water-treated fruit. EBL treatment increased the activity of the three main antioxidant enzymes known as ascorbate peroxidase, catalase, and superoxide dismutase. According to the results, EBL treatment maintained higher soluble Ca²⁺ and antioxidant capacity, reducing fruit susceptibility to BER.
... Fruit cracking in cherry tomatoes is linked with high water potential in developing fruits, which is enhanced when Ca-chelating agents are added, while it is decreased by the addition of CaCl 2 (Lichter et al. 2002). Blossom end rot has been linked to Ca deficiency (Saure 2014). An adequate provision of Ca in the fruit has been demonstrated to enhance fruit firmness and shelf life (Dorais et al. 2001). ...
Article
Tomato fruits of high quality and yield are of interest to greenhouse growers. As potassium (K) and calcium (Ca) have been associated with fruit quality, we studied the effect of the balance between such cations and the electrical conductivity (EC) on fruit yield and quality. Plants were fertigated with K (Kext) and Ca (Caext) balances in the nutrient solution varying from 0.54–1.22 and 0.69–1.44 at ECs of 2.3 ± 0.1 dS m⁻¹ and 2.7 ± 0.1 dS m⁻¹, respectively. Optimum balance for fruit yield was 0.82–0.85, and it was associated with high fruit K internal (Kint) concentration (~ 200 mmol kg⁻¹). Higher EC increased firmness when the balance was ≤ 1.00, with the optimum at 0.85; firmness was correlated with Caint when EC was 2.3 dS m⁻¹; however, at 2.7 dS m⁻¹, increasing Kext increased firmness and intensified the effect of Caext. Increasing a* values were associated with high lycopene and uniform maturation when that balance was 0.82 and 1.00 at low and high EC, respectively. Highest a* values were observed when Kint in the fruit was ~ 200 mmol kg⁻¹, while 91 to 130 mmol kg⁻¹ were associated with lower a* values. Fruits contained higher starch (+ 20%), total (+ 12%), and reducing sugars (+ 29%) at high EC. The optimum K and Ca balance, 0.82–0.85, is independent of their concentration and was associated with high yield and firmness, improved fruit color, and increased lycopene and sugar concentrations.
... Blossom end rot is a necrosis appearing at the 246 blossom end of the fruit (in tomato, pepper, apple…). Although usually attributed to calcium 247 deficiency, it may rather result from complex interactions between environmental factors and involve 248 secondary oxidative stress (Saure, 2014). The fact that solutions found so far to prevent the appearance 249 of such disease are largely empirical indicates that more mechanistic studies integrating metabolism 250 and growth conditions are needed. ...
Thesis
The interest of the tomato (Solanum lycopersicum) fruit has spread in plant science where it is used as the model for fleshy fruit. The valuable advantages of the tomato fruit are numerous: an ease of culture, a short generation time, a high knowledge with important resources, a sequenced genome, an ease for transforming…. The development of tomato fruit is a complex regulated process, divided in four main steps: cell division, cell expansion, turning and ripening. Each step is characterized by a phenotype resulting from changes at different cellular levels. Thus, gene expression, protein abundance, enzyme activities, metabolic fluxes and metabolite concentrations show significant changes during these steps. Thanks to recent technologies advances and in particular the development of ‘omics techniques’, such as genomic, transcriptomic, proteomic, metabolomic, the main cell components can now be analysed by high-throughput. In this context, the objective of my PhD was to perform a quantitative proteomic analysis of the tomato fruit development and then integrate omics data both by statistical analyses and by mathematical modelling. The first chapter focused on results obtained for the quantitative proteomic developed in collaboration with the PAPPSO platform (INRA, Gif-sur-Yvette). Samples were harvested at nine stages of tomato fruit development, total proteome was extracted and quantified by label-free LC-MS/MS. Then I searched for the most appropriate method, testing a set of filters on the data, to obtain an absolute label-free protein quantification from ion intensities (XIC). Finally, I obtained the absolute quantification of 2494 proteins using a method based on peptides intensity modelling. The quantification of proteins by LC-MS/MS was then validated by comparison with 32 enzymatic capacities used as proxy for protein abundance. The second chapter was dedicated to the results of integrative omics analyses throughout tomato fruit development. First, transcriptomic has been performed in collaboration with Genotoul GeT (Toulouse) and Usadel‘lab (RWTH Aachen University, Germany). Using spikes in the experimental design, more than 20000 transcripts have been quantitatively determined at the nine stages of development. Then, this absolute quantification of the tomato transcriptome has been cross-validated with 71 transcripts previously measured by qRT-PCR. Finally, we integrated the four omics datasets- transcriptome, proteome, metabolome and activome – in order to identify key variables of the tomato fruit development. For the four levels, analyses confirmed that the entrance in maturation phase was accompanied by major changes, and revealed a great similarity between the end and the beginning of development, especially in the energy metabolism. The third chapter focuses on modelling results of the protein translation based on the absolute quantification of transcriptomic and proteomic. To explain the decreasing correlation observed between proteins and transcripts concentration throughout development, we proposed a mathematical model of protein translation based on an ordinary differential equation and involving two rate constants (for synthesis and degradation of the protein). The resolution of this equation, validated by a quality criteria based on a closed confidence interval, led to the estimation of the rate constants for more than 1000 proteins. These results were then compared with previous published data reported for plants and more widely in eukaryotic cells. Finally, the last chapter describes all the materials and methods used to obtain the results presented in the manuscript.In the systems biology context, this work illustrates how integration of multiple omics datasets and mechanistic modelling based on absolute omics quantification can reveal new properties of cellular component.
... Finally, as some authors claimed the low Ca concentration in the fruit tip, accompanying BER, to be a consequence of a metabolic disorder, related with an increase of reactive oxygen species (De Freitas et al., 2011;Saure, 2014), it is possible that microorganisms and/or DOM have interacted with the tomato plant metabolism and promoted Table 2 Average nutrient concentrations in the hydroponic (HP) and aquaponic (AP) slabs. resistance against BER via other ways than by only enhancing Mn uptake (Canellas et al., 2015;Mangmang et al., 2015). ...
Article
Full-text available
Decoupled aquaponic systems (DAPS) use the wastewater of recirculated aquaculture systems (RAS) as water source for plant production in recirculated hydroponic systems. RAS wastewater is complemented with macro- and micronutrients to obtain equivalent concentrations and pH as in standard hydroponic nutrient solutions (NS). Unlike in single recirculating aquaponic systems, optimal growth conditions can be established in each production part of a DAPS (i.e. fish and plant parts) avoiding compromises. DAPS design seems more adapted for commercial farming operations but feasibility studies on large-scale systems are lacking. Therefore, the production of tomatoes (Solanum lycopersicum L., cv. Foundation) grown in a NS based on complemented pikeperch RAS wastewater (i.e. AP treatment) has been compared to that of tomatoes grown in conventional hydroponic NS (i.e. HP treatment), in semi-practice conditions. During 3 consecutive years, tomatoes were grown on rockwool slabs, in a large-scale Venlo-type climate-controlled greenhouse, using a recirculated drip irrigation system identical to the ones used by the professionals of the hydroponic tomato sector. While the electroconductivity was significantly higher in the AP treatment due to the presence of NaCl in the RAS wastewater, no significant differences for the total and marketable fruit yields, fruit number, and size were found between the AP and HP treatments. However, while the level of blossom-end rot (BER) varied substantially (0.9–18.6 %) in the HP treatment, it was remarkably constant and low (0.2-0.4%) over the years in the AP treatment, suggesting a beneficial effect of RAS wastewater. Our results clearly indicate the suitability of complemented pikeperch RAS wastewater as feeding water for professional HP tomato production using drip irrigation for DAPS. As RAS water contains a diversity of microorganisms and dissolved organic matter, it is assumed that some of these acted as plant biostimulants and mitigated the salinity stress and the BER symptoms.
... In the fruit metabolism, calcium is also considered an important element which could affect, among others, pectin deesterification during fruit development, ripening, and senescence (Hocking et al. 2016). Pectin-related disorders, such as bitter bit in apple or blossom-end rot in tomato and pepper, could be prevented by calcium preharvest application (Saure 2014) recognizing calcium as an important element for fruit biology. ...
Article
Full-text available
Heat and calcium treatments are used to improve postharvest fruit longevity; however, the exact mechanism remains poorly understood. To characterize the impact of these treatments on sweet cherries metabolism, ‘Lapins’ fruits were treated with heat or CaCl2 solutions and their combination and subsequently were exposed at room temperature, for up to 4 days, deined as senescence period. Single and combined heat and calcium treatments partially delayed fruit senescence, as evidenced by changes in fruit colour darkening, skin penetration force, and respiration activity. Calcium content was noticeably increased by heat in Ca-treated fruit. Several primary metabolites, including amino acids, organic acids, and alcohols, were decreased in response to both treatments, while many soluble sugars and secondary metabolites were increased within 1 day post-treatment. Changes of several metabolites in heat-treated fruits, especially esculetin, peonidin 3-O-glucoside and peonidin 3-O-galactoside, ribose, pyroglutamate, and isorhamnetin-3-O-rutinoside, were detected. The metabolome of fruit exposed to calcium also displayed substantial modulations, particularly in the levels of galactose, glycerate, aspartate, tryptophan, phospharate rutin, and peonidin 3-O-glucoside. The expression of several genes involved in TCA cycle (MDH1, IDH1, OGDH, SUCLA2, and SDH1-1), pectin degradation (ADPG1) as well as secondary (SK1, 4CL1, HCT, and BAN), amino acids (ALDH18A1, ALDH4A1, GS, GAD, GOT2, OPLAH, HSDH, and SDS), and sugar (PDHA1 and DLAT) metabolism were afected by both treatments. Pathway-speciic analysis further revealed the regulation of fruit metabolic programming by heat and calcium. This work provides a comprehensive understanding of metabolic regulation in response to heat and calcium during fruit senescence.
... These factors result in an abnormal acccumulation and partitioning of Ca 2+ in the cells leading to blossom end rot occurrence. Therefore, poor supply of Ca which has an important role in the stability of the plasma membrane as well as cell wall [12] is frequently associated with Blossom end rot in tomatoes [13]. ...
Article
Full-text available
Blossom end rot (BER) is one of the physiological disorders of economic importance in tomato farming since it significantly reduces yield and thus affects profit margins. Most tomato disorders are due to mineral deficiencies and unbalanced nutrition. Improving the supply of specific nutrients and uniform soil moisture can reduce their occurrences. This study was conducted to evaluate the influence of watering regimes, Calcium (Ca) and Potassium (K) on blossom end rot occurrence in two tomato varieties in Maslah and Guul sites. The trials were laid out in a randomized complete block design (RCBD) in split-split plot arrangement with watering regimes (daily, thrice and twice a week) as main plots, tomato varieties (Riograde and Rionex) as sub plots, and 3 levels of Ca and K (0 Kg/ha, 25 Kg/ha, 50 kg/ha) as the sub-sub plots and replicated three times. Calcium treatments had the lowest score of blossom end rot compared to control. In Guul, the highest BER score (2.83) was observed under the control treatment while the lowest score (1.06) was recorded on the 50 kg/ha, Ca rate. Similar results were observed in Maslah with the control having the highest score of BER (3.22) while Ca 50 kg/ha scored lowest (1.11). No statistical differences were observed in the K treatments in the two study sites, however it was notable that lower rates of K reduced the blossom end rot incidences. Water stress led to increase in severity of the BER in the two study sites. In Guul, the highest score was in minimal watering regime (twice a week) of 2.36 score and lowest was at optimal watering regime (daily) of 1.08 score whereas in Maslah the highest blossom end rot score was in minimal watering regime (twice a week) of 3.19 and the lowest score of 1.19 on medium watering regime (Thrice a week).Therefore, optimal application of Ca, K, at 50 kg/ha with adequate and uniform soil moisture can improve management of blossom end rot in tomatoes thus raising farmer’s returns.
... Albahou (1999) reported that high salinity contributed to the decrease in the blossom-end rot incidences. However, various researchers are of the opinion that blossom-end rot increases with increasing salinity and NH4 content (Pill and Lambeth, 1980;Nonami et al., 1995;Saure, 2014 ...
Article
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The study investigates the effect of water deficit conditions on blossom-end rot (BER) in tomato. Drip irrigation was used in irrigation. Therefore, experiment was conducted as a randomized complete block design (three replicates), in a 2 × 5 factorial arrangement, corresponding to two different water resources [treated wastewater (TWW) and freshwater (FW)] and five different irrigation practices. The irrigation strategies comprised full irrigation (100%), deficit irrigation, which involves the use of 75% and 50% of the water used in full irrigation, and partial root drying (PRD). According to the study results, in the full irrigation applications, blossom-end rot according to the fruit number values were 10.85% in fresh water irrigation while it was the lowest with 8.57% for treated wastewater. Also, in full irrigation applications, blossom-end rot values according to fruit weight were found to be lower compared to those in other applications. It was determined that there was a negative relation between blossom-end rot and Ca value. The lowest blossom-end rot incidence was observed in the full irrigation applications, while it was lower in the PRD applications than in the D applications. In the production of marketable tomato, the PRD applications can be more advantageous than the D applications under water shortage conditions. Keywords: BER, wastewater, tomato, PRD, deficit irrigation
... Traditionally it has been considered that BER was caused by a Ca 2+ deficiency, but this assumption is being disputed. In fact Ca 2+ deficiency seems to be the result, not the cause, of BER, which is rather the occurrence of a stressful condition (Saure, 2014). Actually, Ca 2+ content diminution has been detected only after BER symptoms have already developed in fruits but not in early steps of the physiopathy where fruits have similar distribution and concentration of Ca 2+ than healthy fruits (Ho & White, 2005;Nonami, Fukuyama, Yamamoto, Yang, & Hashimoto, 1995). ...
Article
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Pollination and therefore fruit set in tomato (Solanum lycopersicum) is very sensitive to temperature. Parthenocarpy can be a very useful trait in tomato breeding in sustainable agriculture faced with global warming. Within a collection of Moneymaker tomato mutants a monogenic and recessive mutant with high fruit number was identified in heat stress conditions and named high fruit set under stress (hfs). No morphological alterations in vegetative and reproductive organs were observed except the bigger size of ovary. hfs is parthenocarpic, pollination not being required for fruit set, although this trait was not absolute since it produced some under‐seeded fruit. When plants were grown under extreme temperatures (higher than 35°C), hfs exhibits higher fruit yield than wild‐type (WT) due to increased fruit number. Another very interesting characteristic of hfs is its improved fruit quality under heat stress, exhibiting a better sweetness/acidity balance than WT. Interestingly, hfs was also tolerant to the combination of heat and salt stress, and the positive effect on production was due to both components of yield, fruit number and fruit weight. The generation of mostly seedless fruit and the high productivity and fruit quality under extreme temperatures make hfs a very interesting mutant to obtain new breeding high‐yield lines in adverse environmental conditions.
... In addition, Nonami et al., (1995) suggested that Ca defficiency in the fruit may not be the direct cause of the occurrence of BER in tomato, because fruit that had just started having BER had a similar distribution and concentration of Ca ion as normal fruit. Saure (2014) in a reappraisal concluded that the actual causes of BER are obeviousely the effect of a biotic stress, as salinity, drought, heat, increase in reactive oxygen species (ROS) . Exogenous application of calcium can improve plant growth under evironmental stress (Tuna et al., 2007). ...
... Plants absorb calcium through the root, which is then distributed via the xylem sap flux by physiological and molecular mechanisms involved in the ion absorption and transport in the plant (Singh et al., 2015). Because of its dependence on the transpiration flux, the calcium concentration is much lower in the fruits than in the leaves (Saure, 2014). Calcium is important in melon because it participates in the regulation of fruit softening (Madani & Forney, 2015), consequently affecting the product quality and its potential shelf life. ...
... However, they can also result in oxidative stress and ultimately cell death. Blossom end rot is a type of necrosis appearing at the blossom end of the fruit (e.g. in tomato, pepper and apple) and is usually attributed to calcium deficiency, but it may result rather from complex interactions between environmental factors and involve secondary oxidative stress (Mestre, 2012;Saure, 2014). Solutions for preventing the appearance of such a Downloaded from https://academic.oup.com/aob/article-abstract/122/1/1/4987336 by INRA (Institut National de la Recherche Agronomique) user on 24 July 2018 disease have so far been largely empirical, indicating that more mechanistic studies integrating metabolism and growth conditions are needed. ...
Article
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Background: One of the key goals of fruit biology is to understand the factors that influence fruit growth and quality, ultimately with a view to manipulating them for improvement of fruit traits. Scope: Primary metabolism, which is not only essential for growth but is also a major component of fruit quality, is an obvious target for improvement. However, metabolism is a moving target that undergoes marked changes throughout fruit growth and ripening. Conclusions: Agricultural practice and breeding have successfully improved fruit metabolic traits, but both face the complexity of the interplay between development, metabolism and the environment. Thus, more fundamental knowledge is needed to identify further strategies for the manipulation of fruit metabolism. Nearly two decades of post-genomics approaches involving transcriptomics, proteomics and/or metabolomics have generated a lot of information about the behaviour of fruit metabolic networks. Today, the emergence of modelling tools is providing the opportunity to turn this information into a mechanistic understanding of fruits, and ultimately to design better fruits. Since high-quality data are a key requirement in modelling, a range of must-have parameters and variables is proposed.
... En este sentido Lazcano (2000), indica que cuando hay disponibilidad de calcio en el suelo, se incrementa la producción de sustancias en las plantas que dificultan la penetración de patógenos al fruto. Por otra parte posiblemente no hubo rotura enzimática de las protopectinas que componen el armazón de la pared celular, la cual no fue degrada por agentes oxidantes, y de esta forma dichas paredes no se reticulan, las células no pierden compartimentación y el fruto no pierde firmeza (Dhall & Singh, 2013;Saure, 2014;Kiferle et al., 2013;Landini, Gonzali, & Perata, 2011), dificultando así el ataque de patógenos al fruto. Los resultados de esta investigación difieren a los reportados por Zamban et al. (2018) Peso de fruto. ...
Article
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RESUMEN En aras de contribuir a la seguridad alimentaria y a la motivación de los pequeños productores de hortalizas del departamento de Sucre (Colombia), se implementó este ensayo bajo la premisa de que el calcio en el suelo contribuye a tener plantas con pared celular mucho más resistentes al ataque fúngico y con ello se disminuye la aparición de enfermedades. Con base en lo anterior, se estableció este ensayo en cultivo de tomate, en la que se establecieron cinco (5) dosis de calcio (0, 1, 2, 3 y 4 ton.ha1) y tres repeticiones en un diseño completamente al azar. Como fuente de calcio se utilizó un producto comercial que contiene 37% de CaO y 5,3% de MgO. El objetivo fue evaluar el efecto de diferentes dosis de calcio en los componentes del rendimiento del cultivo del tomate Chonto en condiciones semicontroladas en Sucre – Caribe colombiano. El suelo fue de reacción ácida y de baja oferta nutricional. Se utilizó un diseño completamente al azar con tres repeticiones. La información fue almacenada en tablas Excel y procesada con el programa estadístico SAS 9.0, se hizo análisis de varianza y de regresión. Se encontró que las dosis 2 y 3 ton.ha-1 favoreció el crecimiento de las plantas de tomate, aumentando dimensiones del fruto, peso del fruto y rendimiento y los modelos que explican esas tendencias son de orden lineal.
... Ca deficiency in tomato causes a decrease in Ca pectates, which leads to tissue deterioration and deformation (Islam et al., 2016). In addition, Ca deficiency leads to oxidative stress (Saure, 2014). Among preventive measures, spraying with Ca is recommended and is often applied as salts, such as calcium chloride (CaCl 2 ) (Schmitz-Eiberger et al., 2002), since tomato is not known as a Clsusceptible specie (Komosa and Gó rniak, 2012). ...
Article
Abstract Background: The occurrence of calcium (Ca)‐related disorders is common in tomato production and may be due to factors such as excess nitrogen (N) fertilization. Thus, Ca sprays are often applied as preventive measure to avoid the occurrence of blossom‐end rot. In addition, evidence indicates that Si can act beneficially on tomatoes and relieve Ca deficiency. Aims: We aimed to verify the effects of Ca‐EDTA and Si application on tomatoes experiencing moderate Ca deficiency. Methods: Tomato plants were grown in nutrient solutions with Ca sufficiency (+CaR) or Ca deficiency (–CaR) or under conditions of –CaR plus 25 mM Ca‐EDTA shoot sprays (–CaR +CaS), –CaR plus 7.1 mM Si shoot sprays (–CaR +SiS) or –CaR +CaSiS, with a total of nineteen sprays being supplied after anthesis, per foliar treatment. Results: Plants grown under –CaR had smaller leaves and fruits, and new shoots were deformed. Plants that were sprayed with Ca‐EDTA and Ca‐EDTA plus Si in the same solution developed more blossoms than those grown without Ca; however, they showed chlorosis and necrosis on the edges of older leaves, i.e., Ca‐EDTA was toxic. However, the plants sprayed with Si alone were the most similar to control plants, showing only deformations in the younger leaves. Conclusion: Ca‐EDTA at 25 mM sprayed twice a week after anthesis, totaling 19 sprays, caused toxicity in MT tomatoes. Sodium silicate (7.1 mM Si) sprayed twice a week after anthesis, totaling 19 sprays, made MT tomatoes less sensitive to Ca deficiency. Thus, we highlighted the potential of Si spraying to alleviate the slight Ca deficiency, which is relatively common in horticulture.
... The use of PP in plant production does not cause the formation of sludge in irrigation systems, which often results in irregular substrate moisture in hydroponic crops [23,24]. Reduced nutrient solution flow in the drip system, especially in the period of high temperature under covers, may be the cause of increased share of fruit with dry BER rot in such vegetables as tomato or pepper [25]. ...
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The aim of this study was to investigate the effect of phosphorus application in the form of polyphosphates on the yield and quality of sweet pepper fruits grown with LED (light-emitting diodes) assimilation lighting. Phosphorus is absorbed by the root system of plants mainly in the form of orthophosphates ions. The availability of phosphorus depends, among other things, on the pH of the substrate and the temperature. Two cultivars of sweet pepper with red fruits were tested in hydroponic cultivation on a mineral wool substrate. The plants were fertilized with one of three schedules, each of the same concentration of components, but differing only in the form of the applied phosphorus: polyphosphates (PP) and orthophosphates (OP). In the experiment, stem length extensions and number of leaves, chlorophyll concentration in leaves and fluorescence of the chlorophyll in a leaf were measured. The number and weight of fruits in total as well as marketable and non-commercial fruits with symptoms of dry rot (BER-blossom end rot) were studied. The concentration of dry matter and selected chemical components in fruits were examined and the sensory quality of fruits was evaluated using the QDA (Quantitative Description Analysis) method. The nutrient status of the pepper plants was also examined. Polyphosphates used in the medium increase the activity of photosynthetic apparatus of leaves and have a positive effect on the share of marketable yield of the total yield in the cultivar susceptible to BER. Fertigation in hydroponic cultivation with medium containing 30% phosphorus in the form of polyphosphates increased the uptake of calcium in pepper plants growing with LED lighting. The pepper cultivars tested differ in, among other things, the susceptibility to BER and the quality attributes of the fruit.
... improve the disease resistance of tomato (Saure 2014). At present, many reports are available on the effects of different nutrient couplings on tomato growth, yield, and physiological metabolism under soil cultivation conditions (Liu et al. 2012, Wang and Xing 2017, Chamurliev et al. 2019. ...
Article
The experiment aimed to obtain a quadratic regression mathematical model of the comprehensive evaluation score of yield, quality, and four macroelements (N, K, P and Ca). The suitable nutrient solution was chosen and verified via computer simulation of the model and the highest comprehensive score in all treatments. Results showed that P, K and Ca had a positive effect on the comprehensive evaluation value of tomato, whereas N showed a negative effect. The optimal formula calculated using the regression equation could promote high-yield and high-quality tomato. The single-plant yield, soluble protein, vitamin C, total sugar, lycopene, and elemental utilisation of K and Ca in the tomato were 13.93, 78.95, 3.29, 20.98, 51.91, 16.69 and 24.14% higher than those in the special formula treatment of Japanese Yamazaki tomato, respectively. In summary, the optimal nutrient solution formula of tomato cultivation was obtained, in which the N, P, K and Ca levels were 24.83, 4.50, 9.49 and 5.73 mmol/L, respectively.
... This can lead to membrane decomposition and cell wall failure; in fruits, this has been proposed to give rise to disorders such as Blossom end rot (De Freitas and Mitcham 2012). Whether calcium deficiency is the cause of such disorders or calcium deficiency is a result of these conditions has been recently debated (De Freitas et al. 2013;Saure 2014). ...
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Calcium (Ca) content of fruit is related to fruit quality. However, total content alone is insufficient to judge fruit quality; calcium partitioning and chemical form are also important. There are multiple validated methods for determining total calcium content. However, methodologies to measure calcium fractions are not widely known and tend to be complex, affecting their accuracy. Here, based on previous work, we aim to adapt and validate a method for the extraction of the various forms of calcium in fruits. The main modification was freezing samples to minimize the loss of sample material in the procedure, reducing the centrifugation time and speed, and the addition of nitric acid (HNO3) to improve digestion and eliminate interference from the matrix. An experiment with table grape berries cv. Thompson seedless in harvest and Postharvest (35 days) was carried out. The results of the total calcium were compared with the conventional methodology of calcination and inductively coupled plasma (ICP) measurement. We detected no and no significant differences between the two tested methods. Electron Microscope - Energy-dispersive X-ray spectroscopy analysis (SEM-EDS) and the presence of calcium oxalates were verified.
... Plants absorb calcium through the root, which is then distributed via the xylem sap flux by physiological and molecular mechanisms involved in the ion absorption and transport in the plant ( Singh et al., 2015). Because of its dependence on the transpiration flux, the calcium concentration is much lower in the fruits than in the leaves ( Saure, 2014). Calcium is important in melon because it participates in the regulation of fruit softening ( Madani & Forney, 2015), consequently affecting the product quality and its potential shelf life. ...
Article
The aim of this work was to study the effect of preharvest foliar spraying with calcium nitrate solutions on the fruit melon quality. Treatments included a combination of five solutions with Ca(NO3)2 (0.0; 1.3; 2.6; 5.2 and 10.5 g/L) and two postharvest storage environments [(24.0 ± 1.5 °C and 6.4 ± 0,8 mbar of vapor pressure deficit (VPD), and 10.0 ± 0.5 °C and 3.0 ± 0.5 mbar (VPD)]. Fruits stored at 24 ˚C for 11 days had the lowest water loss with treatments 1.3 and 2.6 g/L Ca (NO3)2. It was observed that at both 24 °C and 10 °C, fruits that had a greater firmness at the 2.6 and 5.2 g/L of Ca(NO3)2 treatments, respectively. No differences (P>0.05) were observed in total soluble solids (TSS) between treatments and storage environments. However, spotting of the epidermis of the fruits increased as the Ca(NO3)2 concentration also increased.
... However, no changes were observed in the distal part of tomato fruits that had been subjected to treatments ( Figure S6). 32 It is also known that in the same plant, healthy fruits and fruits with BER may occur. However, the distribution of Ca 2+ in healthy fruits and fruits with BER was different. ...
Article
In this study, we aimed to shed light on the effects of the Ni−Ca interaction on the activity of the urease enzyme and the incidence of blossom end rot (BER) in tomato (Solanum lycopersicum) fruits. Therefore, Micro-Tom tomato plants were grown under different levels of Ni (0.0 and 1.0 mg dm −3) and Ca 2+ (0.35 and 0.70 g dm −3), both alone and in combination. Increased yields and urease activity, as well as inhibition of the incidence of BER, were observed for tomato fruits grown with Ni, even under low Ca availability (0.35 g dm −3). In addition, a microprobe X-ray fluorescence spectroscopy characterization of the distal part of the fruits revealed a sharp decrease in Ca content in the fruits that present BER symptoms.
... In addition, other authors also consider that blossom problems occur either due to reduced transport of Ca to rapidly growing distal fruit tissue or to increased demand of the distal fruit tissue for Ca, caused by accelerating fruit expansion. 5 And other authors, such as Saure, 6 consider that the lower concentration of Ca is the result of this condition. ...
Article
Background: Sweet pepper fruit quality disorders have been related mainly to an unbalanced nutrient supply and non-optimal growth conditions. Increases in the atmospheric CO2 concentration ([CO2 ]) have been associated with a reduction of transpiration, which can affect calcium (Ca) uptake as it is linked closely to water uptake. We investigated whether foliar application of Ca can counterbalance the effects of saline water and elevated [CO2 ]. Results: High CO2 favored generative growth instead of vegetative growth. Foliar Ca supply did not affect the marketable yield, but reduced the total yield when combined with salinity and 400 µmol mol(-1) CO2 . Salinity affected negatively the total yield but this was overcome when CO2 was applied. The B and K concentrations were reduced by foliar Ca application, while Ca and Mn were increased at 400 µmol mol(-1) CO2 . Salinity increased the Mn, Cl, and Na concentrations, regardless of the [CO2 ], and decreased K at 800 µmol mol(-1) CO2 . The total protein was affected negatively only by elevated [CO2 ], and the total free amino acid concentration was reduced by all treatments. Conclusion: The effect of Ca application differed according to the other treatments applied. This procedure should be optimized to overcome future climate impacts on fruit quality.
... Traditionally, research studies have related bitter bit with calcium deficiency and the balance between calcium and other nutrients such as nitrogen, potassium, or magnesium (Ferguson & Watkins, 1989;Amarante et al., 2013;de Freitas et al., 2015). However, some works have shown a lack of relationship between calcium content level and bitter pit, indicating that the disorder could be affected by other causes such as climate and/or growing conditions (Saure, 1996(Saure, , 2005Lotze et al., 2008;Saure, 2014;Torres et al., 2017b). ...
Article
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Aim of study: A portable VIS/NIR spectrometer and chemometric techniques were combined to identify bitter pit (BP) in Golden apples. Area of study: Worldwide Material and methods: Three different classification algorithms – linear discriminant analysis (LDA), quadratic discriminant analysis (QDA) and support-vector machine (SVM) –were used in two experiments. In experiment #1, VIS/NIR measurements were carried out at postharvest on apples previously classified according to 3 classes (class 1: non-BP; class 2: slight symptoms; class 3: severe symptoms). In experiment #2, VIS/NIR measurements were carried out on healthy apples collected before harvest to determinate the capacity of the classification algorithms for detecting BP prior to the appearance of symptoms. Main results: In the experiement #1, VIS/NIR spectroscopy showed great potential in pitted apples detection with visibly symptoms (accuracies of 75–81%). The linear classifier LDA performed better than the multivariate non-linear QDA and SVM classifiers in discriminating between healthy and bitter pitted apples. In the experiment #2, the accuracy to predict bitter pit prior to the appearance of visible symptoms decreased to 44–57%. Research highlights: The identification of apples with bitter pit through VIS/NIR spectroscopy may be due to chlorophyll degradation and/or changes in intercellular water in fruit tissue.
... This indicates a difference in total fresh weight of paprika fruits at different temperatures ( Fig. 1). High temperature also increases the production of reactive oxygen species (ROS) in the fruit, leading to cell death and BER incidence (Saure, 2014). Since ROS production is also increased by salinity stress (Aktas et al., 2005), ROS may be one of the reasons for the differences in the incidence of BER between temperature and KNO 3 concentrations (Fig. 2). ...
Article
Blossom-end rot (BER) in paprika (Capsicum annuum L.), as in tomato fruits, is a physiological disorder caused by calcium (Ca) deficiency in fruits. While studies have revealed that excessive nitrogen and potassium fertilization and high temperatures affect Ca transport to fruits and induce BER, few studies have investigated the effects of other elemental concentrations in paprika fruit and their association with BER occurrence. This study aimed to investigate the relationship between the changes in the elemental composition and incidence of BER in paprika fruits grown at different potassium nitrate (KNO3) concentrations and different temperature conditions. Paprika plants were grown in rockwool blocks in a greenhouse at 25°C and 30°C for approximately four months and irrigated with liquid fertilizer and three different concentrations (0, 27.9, and 74.4 mM) of KNO3 were added. Subsequently, the total fresh weights of ripe paprika fruits and the incidence of BER were measured, and the elemental compositions in the pericarp of the first and last sampled fruits were analyzed. The total fresh weights significantly decreased and the incidence of BER increased with increasing KNO3 concentrations and temperature. The profiles of 11 mineral elements in the pericarps of paprika fruits revealed a significant positive or negative correlation between not only the concentration of Ca, but also that of several other elements including boron (which crosslinks pectin in the cell wall like Ca) and the incidence of BER. These results suggest that Ca deficiency may not be the only cause of BER occurrence, and that several elements may also be involved. The insights from this study will contribute to help predict the incidence of BER and stabilize crop production by improving fertilizer application and environmental control.
... Many disorders have been related to Ca nutrition, but the mechanisms that control their occurrence are not clearly understood yet. Any condition that affects Ca uptake by the plant and its distribution to the fruit can lead to Ca deficit disorder [7][8][9][10]. In ...
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In this study, the rate of absorption and transport of calcium (Ca) in orange fruits (Citrus sinensis cv. Fukumoto) after surface treatment with 45Ca was evaluated by supplying treatments at different dates between fruit set and mid-phase II of growth (i.e., 30, 44, 66 and 99 days after full bloom, DAFB). 45Ca absorption was always detected, but 45Ca application at the fruit set was associated with the highest rates of Ca transport into the pulp (39%) compared to the other treatments (17–19%). Scanning electron microscopy SEM) images of the fruit surface showed the occurrence of stomata along with the entire rind at all the developmental stages evaluated. However, from the beginning of stage I, stomata began to collapse and develop plugs, and this became frequent as the fruit age increased. From 44 DAFB, oil gland density increased, and Ca oxalates (CaOx) were found in the fruit flavedo and albedo. Fruit Ca (not labelled) concentration increased from fruit set (30 DAFB) to 99 DAFB, although oil gland formation and rind growth and differentiation likely hindered Ca transport to internal fruit tissues (pulp). The total Ca concentration in the pedicel was always higher than that in the fruit, with no differences between the first three treatments. The information obtained in this study may be essential for improving Ca fertilizer efficacy in citrus by spray applications.
... For example, extreme heat causes pre-harvest physiological disorders, resulting in fruit cracking and blossom drop in tomato plants. For such extreme heat, temporary equipment and/or manual control is required to lower the temperature in the greenhouse (Liebisch et al., 2009;Saure, 2014). Therefore, for greenhouse cultivation, there is a need to continuously and adequately manage the environmental conditions inside greenhouses. ...
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While the high year-round production of tomatoes has been facilitated by solar greenhouse cultivation, these yields readily fluctuate in response to changing environmental conditions. Mathematic modeling has been applied to forecast phenotypes of tomatoes using environmental measurements (e.g., temperature) as indirect parameters. In this study, metabolome data, as direct parameters reflecting plant internal status, were used to construct a predictive model of the anthesis rate of greenhouse tomatoes. Metabolome data were obtained from tomato leaves and used as variables for linear regression with the least absolute shrinkage and selection operator (LASSO) for prediction. The constructed model accurately predicted the anthesis rate, with an R ² value of 0.85. Twenty-nine of the 161 metabolites were selected as candidate markers. The selected metabolites were further validated for their association with anthesis rates using the different metabolome datasets. To assess the importance of the selected metabolites in cultivation, the relationships between the metabolites and cultivation conditions were analyzed via correspondence analysis. Trigonelline, whose content did not exhibit a diurnal rhythm, displayed major contributions to the cultivation, and is thus a potential metabolic marker for predicting the anthesis rate. This study demonstrates that machine learning can be applied to metabolome data to identify metabolites indicative of agricultural traits.
... Therefore, the BER disorder "can be triggered by mechanisms that reduce: 1) plant Ca 2+ uptake from the soil, 2) fruit Ca 2+ uptake from the plant, and 3) Ca 2+ translocation within the fruit. These factors will result in an abnormal accumulation and partitioning of Ca 2+ in the cells" (Bonomelli, Arias, and Villalobos 2018;Saure 2014). Bitter Pit in apples is closely associated with the Ca content of the fruit (Fallahi et al. 1997;Fallahi, Righetti, and Raese 1988;Ferguson, Reid, and Prasad 1979;Ferguson and Watkins 1989;Perring and Pearson 1986); a relatively high fruit Ca content at harvest can reduce the incidence of this disorder (Hewett and Watkins 1991). ...
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This study was designed to optimize calcium-45 (⁴⁵Ca) sample preparation method in a Liquid Scintillation Counter (LSC) for use with vegetables and soil samples. Different tests were conducted on fruit and soil samples to determine the best conditions related to drying, time and centrifuge speed (rpm) after calcination at 450°C and 550°C for vegetable and soil samples, respectively. The homogeneity of samples, linearity, detection and quantification limits were also determined. The quench curve was calculated to establish the parameters to program the LSC. Six ⁴⁵Ca sample replicates were measured with a coefficient of variation (CV) of 1.74%, which indicates good sample homogeneity. The method is linearity over the range of activity studied for 0–30,000 disintegrations per minute (dpm). Our results suggest that the optimum homogenized sample preparation for calcination purposes was lyophilization. Techniques using ⁴⁵Ca applied to soil and plant tissues are promising tracers of Ca in plants, which can be very useful for the study of Ca kinetics in plants and can serve to guide efficient fertilization programs.
Article
It is known that poorly performed fertigation directly impacts on tomato production and biometric components. In addition, consumers are also affected by interrelated characteristics that interfere with the acceptability of the fruit, such as the physicochemical parameters and nutrients in the fruit. Thus, eco-friendly technologies, such as irrigation with ultra-low frequency electromagnetic treated-water, which attenuates the inadequate management of fertigation, are essential to improve marketable fruit yields. Thus, the objective of the present work was to investigate the impact of treated water with very low-frequency electromagnetic resonance fields in physical, chemical and nutritional parameters at different nutrient solution strengths in tomato fruits. In this study, experiments were carried out in randomized blocks and five doses of fertigation were used (1.5; 2.5; 4.0; 5.5; and 7.0 dS m−1), employing two types of water: electromagnetically treated and untreated. It can be seen that the fertigation affected some parameters, mainly the number of fruits with blossom-end rot, fruit size, and weight. Variance analysis (ANOVA) was performed with the subsequent use of the Tukey test. In all statistical tests, a confidence level of 95% was considered. The soluble solids content increased by 28% as a function of the fertigation doses. The electromagnetically treated water reduced the number of fruits with blossom-end rot by 35% (p < 0.05). Overall, electromagnetic water improved the physicochemical quality parameters and the nutritional status of tomato fruits. Thus, this study demonstrated that green technology could leverage tomato fruit production and quality.
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Water stress in tomato plants may cause the incidence of blossom-end rot. This study aimed to analyze the effect of abscisic acid leaf application for increasing the calcium uptake in irrigated tomato (‘Santa Clara' cultivar) in the field, as a possible mechanism of blossom-end rot inhibition. The treatments consisted of four irrigation levels (25 %, 50 %, 75 % and 100 % of the crop water requirements to fulfil the crop evapotranspiration) and two abscisic acid doses (0 mg L-1 and 500 mg L-1). The fruits were harvested at 15 and 30 days after the anthesis and evaluated for calcium content and percentage of blossom-end rot. The application of abscisic acid increased the calcium partition to the distal region of the fruits at 30 days after the beginning of flowering, as well as reduced the incidence of blossom-end rot by 86 %, when compared with plants not treated with abscisic acid. It is possible to conclude that the foliar application of abscisic acid can significantly reduce the incidence of blossom-end rot.
Chapter
In a greenhouse, an artificial environment is created inside a covering and crops are produced therein so as to cope with changes in the weather. However, cooling is more difficult to achieve than heating, and it is thus necessary to develop technology that addresses high temperatures. Tomato is a major vegetable grown in greenhouses. In tomato cultivation under high-temperature conditions, pollen fertility decreases, fertilization fails, and poor fruit sets result in decreased fruit yields. The incidence of physiological disorders (e.g., fruit cracking, blossom-end rot, leaf marginal necrosis, and yellow shoulder fruit) also increases. The mechanism underlying the occurrence of each physiological disorder and management techniques for suppressing these disorders are introduced in this chapter. Heat countermeasure technologies used in greenhouses in Japan, such as night cooling using a heat pump, fog cooling, and the use of appropriate covering materials, are also explained.
Article
Calcium (Ca) deficiency disorders in apple fruit have been associated with high gibberellins (GAs) activity in the tree. This study was carried out to assess the effects of treatments of ‘Braeburn’ apple trees with prohexadione-calcium (ProCa, an inhibitor of GAs biosynthesis) or gibberellins (GA4+7) on vegetative growth of the trees and postharvest incidence of Ca-related physiological disorders and decay in the fruit. ProCa (300 mg L⁻¹) or GA4+7 (300 mg L⁻¹) treatments were applied post-bloom (PB) and preharvest (PH). PB treatments started 15 days after full bloom (DAFB), with one application every week and six applications in total. PH treatments started five weeks before anticipated harvest (125 DAFB), with one application every week and four applications in total. Control trees were left untreated. When applied PB, ProCa reduced and GA4+7 promoted vegetative growth of the trees. ProCa PB delayed the impairment of xylem functionality (at the proximal region of the fruit) during fruit growth on the tree. Treatments had no effect on fruit weight, pectinmethylesterase (PME) activity or the expression of CAX3, CAX6 and V-ATPase (transporters of Ca into the vacuole) genes assessed in the external cortical tissue at the distal end of the fruit at harvest. When sprayed PB, ProCa increased the total Ca content and reduced K/Ca, Mg/Ca, N/Ca, (Mg + K)/Ca and (K + Mg + N)/Ca ratios in the flesh at the distal portion of the fruit, compared to the treatment with GA4+7, but without differing from the control. In general, ProCa application PB or PH reduced the expression of Ca-ATPase1, Ca-ATPase2, H⁺-PPase and CAX2 (Ca transporters into the vacuole) genes, increased total water‐soluble Ca and reduced electrolyte leakage in the fruit at harvest. After two months of cold storage followed by five days of shelf-life, the incidence and severity of bitter pit (BP) was lower on fruit from trees treated with ProCa PH, and higher on fruit from trees treated with GA4+7 PB. GA4+7 PB also increased the incidence of skin cracking and decay in the fruit. The results showed that ProCa application represents a feasible tool to reduce the incidence of BP. However, ProCa is more effective to reduce BP if applied weekly for five week before harvest.
Article
Abstract This study aimed to investigate influences of calcium chloride (CaCl2) concentrations combined with nano-chitosan coating on the quality of strawberry during postharvest storage. The fruit were dipped in different concentrations of calcium chloride (1 %, 2 %, 3 %, 4 %) before being coated with 0.2 % nano-chitosan. Physico-chemical analysis including the overall quality index, weight loss, firmness, titratable acidity, total soluble solid, l-ascorbic acid content, antioxidant capacity, total phenolic content, total anthocyanin content, and malondialdehyde content were performed in 3 d intervals until fruit became unmarketable. Among six examined treatments, a combination of 3 % CaCl2 and nano-chitosan (NCTS) was the most effective one as maintaining the highest score of overall quality index of strawberry stored at 4 °C up to 15 d. The treatment also significantly reduced weight loss, preserved l-ascorbic acid, total anthocyanin contents, antioxidant capacity, and retarded malondialdehyde production. The scanning electron microscope image showed a smooth surface of strawberries coated with 3 % CaCl2 combined 0.2 % nano-chitosan. There has no bitterness detected in the treated strawberries after being stored 15 d at 4 °C. The major volatile compounds determined in the initial day were remained until the 15th d of storage.
Chapter
Sweet and pungent peppers (Capsicum spp.) are globally important vegetable and spice commodities as they are valued for their nutritional qualities, antioxidant compounds, flavors, pungency, brilliant colors, and textures. Capsicum. has extraordinary variability in its germplasm both in cultivated and wild species. This review presents an account of research done over several decades in the context of crop improvement. Key developments with reference to linkage analyses, DNA-based markers, the identification of quantitative trait loci for complex traits, transcriptomes of ripening fruit, and genome sequences are summarized. Prospects are excellent for using conventional, biotechnological, and genomic approaches to improve fruit yield, fruit quality, and biotic stress tolerance so that productivity in this specialty crop could be sustained, despite the changing climate. However, more research is needed to build resources to improve peppers for tolerance to abiotic stress factors.
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Blossom-end rot is a physiological disorder causing significant losses in the tomato industry each year. Accumulation of reactive oxygen species has been established as a key characteristic of blossom-end rot development. An increase in peroxidase activity and lignin precursor content are also associated with blossom-end rot symptoms, leading to the hypothesis that lignification may be occurring during blossom-end rot development. To investigate the potential involvement of lignification, hydrogen peroxide content, catalase activity, and peroxidase activity were measured in the top, bottom, and blossom-end rot affected tissue of blossom-end rot affected fruit, and the top and bottom of healthy fruit. Lignin was assayed using histochemical staining, autofluorescence, and thioglycolic acid degradation methods. Hydrogen peroxide content was increased in blossom-end rot affected and blossom-end rot adjacent tissues compared to healthy fruit and the top of blossom-end rot affected fruit. Catalase activity was significantly reduced and ferulic acid peroxidase activity was increased in the blossom-end rot affected and unaffected tissue at the bottom of blossom-end rot affected fruit compared to the bottom of healthy fruit. Lignin analyses showed increased lignin content in blossom-end rot affected tissue compared to tissue from the bottom of healthy fruit. These results show that lignification occurs during blossom-end rot development, likely through a peroxidase-mediated pathway.
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As a crucial element for plants, calcium (Ca) is involved in both nitrogen (N) absorption and assimilation. Plants tend to exhibit lower nitrogen use efficiency (NUE) under Ca-deficient conditions. Improving NUE in apple production can reduce the negative effects of the excessive use of N fertilizer. However, the role of Ca in the regulation of nitrate uptake and reduction remains unclear. Herein, we investigated the growth response and nitrate (NO3⁻) metabolism of apple dwarf rootstock seedlings (M9T337) for Ca²⁺ concentrations of 0, 2, 4, 6, 8, 10, and 12 mM Ca²⁺ using ¹⁵N isotope labeling tracer, non-invasive micro-testing technology, and quantitative real-time polymerase chain reaction (qrt-PCR). Results showed that Ca accumulation rate in the plant tissue increased with increasing Ca supply levels; however, the plant biomass, photosynthesis, root activity, and ¹⁵NUE peaked for 6 mM of Ca²⁺ supply. Further investigation revealed that for 6 mM of Ca²⁺ supply, nitrate reductase (NR) activities were relatively high, the transcription of nitrate transporter (MdNRT1.1; MdNRT2.1) up-regulated and NO3⁻ maximum influx rate in the roots occurred. Furthermore, seedlings treated with 6 mM Ca²⁺ had a significantly lower NO3- concentration in leaves and roots, a higher NH4⁺ concentration in leaves, and a higher amino acid compounds concentration. Moreover, both Ca deficiency and excessive Ca inhibit N absorption and utilization, and the adverse effects of Ca deficiency on seedling growth and N metabolism were greater than those associated with excessive Ca²⁺ supply. Conclusively, the results of this study indicate that appropriate Ca²⁺ supply (6 mM) was optimal as it increased NUE by enhanced photosynthesis, N metabolizing enzyme activities, NO3⁻ uptake and transport.
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The macronutrient potassium (K) has vital physiological functions in plants and its availability can strongly impact quality of crops like tomato. The impact of K nutrition on conventional tomato fruit quality parameters has been described several times, but detailed investigations on the effect of K supply on the fruit metabolite profile are still rare. To fill this gap, we investigated the influence of K fertilization on the metabolite profile of tomato fruits. For this purpose, an outdoor pot experiment with three different cocktail tomato cultivars was performed. A fertilization regimen with five K levels was applied, ranging from deficiency to sufficient supply. Fruit samples were analyzed by untargeted GC×GC-MS to cover the primary metabolite profile as well as some secondary metabolites. As verified using ICP-OES, fruit K content was highly proportional to the supplied amount of K. At the metabolite profile level, the most prominent and cultivar-independent effect of increased K fertilization was the rise of tricarboxylic acid (TCA) cycle intermediates. Further effects were more cultivar-specific, for example an increase of the mobile nitrogen pool (e.g. amines like putrescine and amides like asparagine), changes in the profile of minor sugars (especially disaccharides) as well as higher levels of some secondary metabolites. Pronounced response patterns were mainly observed in the cultivars Primavera and Yellow Submarine that were recently characterized as higher yielding, demanding a stronger consideration of cultivar differences in future studies.
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Calcium (Ca) is a crucial element in plant growth and development. Studies focused on the Ca deficiency in Prunus are still limited that triggered us to examine tolerance of peach plants against Ca deficiency using physiological and histological methods. Peach cultivar Rich May plants grafted onto GF 677 and Garnem rootstocks were planted in 10 L pots containing perlite. Plants were irrigated with Hoagland’s nutrient solution. After growth for two months in nutrient solution, plants were exposed to Ca deficient Hoagland solution (except control). Plants were exposed to mineral deficient conditions for 4 months. End of the study, Ca deficiency depressed plant growth, damaged cell membranes and impaired chlorophyll biosynthesis. Concentrations of chlorophyll and chlorophyll precursors were significantly decreased by Ca deficiency in both rootstocks. Cortical cell diameter decreased in both rootstocks under Ca deficiency conditions. However, cortex thickness increased in Garnem and decreased in GF 677. Increments in xylem thickness, xylem conduits length and number of xylem conduits were found in Ca deficient Garnem plants and the parameters decreased in GF 677 under Ca deficiency. Pith autolysis in midrib tissue was not observed in the current experiment. Garnem was found more tolerant rootstock than GF 677 to Ca deficiency. We consider that larger cortical cells and hyperplasy and hypertrophy of xylem vessel elements increased stomatal conductance to uptake more Ca that mitigated Ca starvation in Garnem.
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This research aimed to analyze the visual and nutritional symptoms in nopal (Opuntia ficus indica (L.), through induction of nutritional deficiencies by the missing element technique under hydroponic conditions. In order to meet this objetive, we planned to work in a greenhouse of the Graduate College in the State of Mexico, from June 2008 until the plants searched nutritional deficiencies were observed. Plant materials used were cladodes a year from the production area of Tlalnepantla, Morelos, which were dried and transplanted into pots with tezontle for promoting their rooting. 11 treatments of different nutrient solutions were tested with three replicates each, each pot was an experimental unit and treatments were distributed according to a completely randomized design. Data were analyzed through an ANOVA with means comparison test of Tukey (α= 0.05). In the results, it is quite important that the plants showed nutritional deficiencies at 13 months into the experiment; observing morphological differences in the presence or absence of subulate leaves and their coloring, highlighting the treatment that did not contain nitrogen. Also, significant difference in number of pads, dry weight and nutrient concentration were observed. It is concluded that this methodology works for inducing macro-nutrient deficiency.
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Root restriction has been reported to reduce fruit yield, the incidence of blossom end rot (BER) and K concentration in tomato (Lycopersicon esculentum L. 'F121') plant organs. The objectives of the present work were to study the effect of root restriction, and combination of K and Ca solution concentrations, on greenhouse tomato fruit yield, quality and cation uptake. Root restriction reduced total yield but improved fruit quality by increasing the dry matter concentration and reducing the incidence of BER. Increasing the K concentration from 5.0 to 10 mmol · L -1 reduced the marketable yield, due to increased incidence of BER. Root restriction decreased K concentration and K/Ca ratio in tomato plant organs, but had no effect on K uptake rate per unit root fresh weight. Increasing K concentration from 2.5 to 10 mmol. L -1 increased the K concentration in plant organs and K uptake rate, but reduced that of Ca. In contrast, increasing Ca concentration in the solution had no effect on K concentration in plant organs and K uptake rate. The incidence of BER correlated well with K/Ca concentration ratio in the leaves, whereas a poor correlation was obtained with K/Ca concentration ratio in ripe fruit.
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Plant hormones are small molecules that regulate plant growth and development, as well as responses to changing environmental conditions. By modifying the production, distribution or signal transduction of these hormones, plants are able to regulate and coordinate both growth and/or stress tolerance to promote survival or escape from environmental stress. A central role for the gibberellin (GA) class of growth hormones in the response to abiotic stress is becoming increasingly evident. Reduction of GA levels and signalling has been shown to contribute to plant growth restriction on exposure to several stresses, including cold, salt and osmotic stress. Conversely, increased GA biosynthesis and signalling promote growth in plant escape responses to shading and submergence. In several cases, GA signalling has also been linked to stress tolerance. The transcriptional regulation of GA metabolism appears to be a major point of regulation of the GA pathway, while emerging evidence for interaction of the GA-signalling molecule DELLA with components of the signalling pathway for the stress hormone jasmonic acid suggests additional mechanisms by which GA signalling may integrate multiple hormone signalling pathways in the response to stress. Here, we review the evidence for the role of GA in these processes, and the regulation of the GA signalling pathway on exposure to abiotic stress. The potential mechanisms by which GA signalling modulates stress tolerance are also discussed.
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The mechanisms regulating Ca2+ partitioning and allocation in plants and fruit remain poorly understood. The objectives of this study were to determine Ca2+ partitioning and allocation in tomato plants and fruit in response to whole-plant and fruit-specific abscisic acid (ABA) treatments, as well as to analyse the effect of changes in Ca2+ partitioning and allocation on fruit susceptibility to the Ca2+ deficiency disorder blossom-end rot (BER) under water stress conditions. Tomato plants of the cultivar Ace 55 (Vf) were grown in a greenhouse and exposed to low Ca2+ conditions during fruit growth and development. Starting 1 day after pollination (DAP), the following treatments were initiated: (i) whole plants were sprayed weekly with deionized water (control) or (ii) with 500mg l−1 ABA; or fruit on each plant were dipped weekly (iii) in deionized water (control) or (iv) in 500mg l−1 ABA. At 15 DAP, BER was completely prevented by whole-plant or fruit-specific ABA treatments, whereas plants or fruit treated with water had 16–19% BER incidence. At 30 DAP, BER was prevented by the whole-plant ABA treatment, whereas fruit dipped in ABA had a 16% and water-treated plants or fruit had a 36–40% incidence of BER. The results showed that spraying the whole plant with ABA increases xylem sap flow and Ca2+ movement into the fruit, resulting in higher fruit tissue and water-soluble apoplastic Ca2+ concentrations that prevent BER development. Although fruit-specific ABA treatment had no effect on xylem sap flow rates or Ca2+ movement into the fruit, it increased fruit tissue water-soluble apoplastic Ca2+ concentrations and reduced fruit susceptibility to BER to a lesser extent.
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Blossom-end rot (BER) is a fruit physiological disorder that occurs under salinity stress and is known to be associated with impairment of calcium translocation to the fruit tip. In the present study we investigated the possibility that oxidative stress is part of the causative mechanism of BER in pepper (Capsicum annuum L.). Pepper plants of two cultivars ('Silica', not sensitive to BER and '107', sensitive) were grown in a greenhouse and irrigated with nutrient solution made up with either desalinated water (control - E.C 1.9), or saline water (salinity - E.C. 3.7- 7.0). Irrigation was applied by a circulation system. BER symptoms were observed in '107' throughout the experiment while in 'Selica' the amount of BER was minimal. BER occurrence was significantly increased under salinity in the sensitive but not in the insensitive cultivar. The fruit calcium concentration in both sensitive and resistant cultivars was not changed by salinity. The oxidative stress-related enzymatic activities in the fruit were dependent on the fruit developmental stage. Under salinity, oxygen radical and hydrogen peroxide production, and NAD(P)H oxidase activity were all enhanced at the most sensitive developmental stage. In contrast, the anti-oxidative activities of superoxide dismutase (SOD) and catalase were not affected by salinity. The results suggest that generation and scavenging of oxygen free radicals may be part of the mechanism that leads to the appearance of BER symptoms in pepper fruits under saline conditions. We suggest that measures to alleviate oxidative stress in greenhouses would reduce the risks of blossom-end rot in peppers.
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We investigated the possibility that oxidative stress contri-butes to blossom-end rot (BER) initiation in bell pepper (Capsicum annuum L.) grown under high salinity. Pepper plants (cv. Mazurka, Rijk Zwaan, the Netherlands) were grown in a greenhouse and irrigated with nutrient solution made up with either desalinated water (control — rising from E.C. 1.9 to 2.4dSm À1) or saline water (salinity – rising from E.C. 3.2 to 7.0 dS m À1). Irrigation was by a circulation system. BER symptoms were observed throughout the experi-ment but were highly enhanced in the salinity–grown plants during the spring and summer. The fruit calcium concentration was not affected by salinity, but manganese concentrations in both leaves and fruits were significantly reduced under these conditions. Under salinity there was an enhancement of apoplast reactive oxygen species (ROS) production, which was partly a result of increase in NAD(P)H oxidase activity in the pericarp of pepper fruit at the stage that it was most sensitive to BER. Apoplast ROS production and extracted NAD(P)H oxidase activity were inhibited by manganese, zinc and to a lesser extent by calcium. These cations also negated the enhancement of ROS production caused by incubation of fruit pericarp discs in NaCl solutions. Manganese, zinc and calcium also inhibited NAD(P)H oxidase activity, extracted following their infiltra-tion into fruit pericarp discs. The results suggest that generation and scavenging of oxygen free radicals in the apoplast may contribute to the appearance of BER symptoms in pepper fruits under saline conditions. It is suggested that manganese may serve as antioxidant in pepper fruit and that manganese addition to peppers grown under salinity may alleviate BER symptoms in the fruits.
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The aim of the present research was to study changes in the major anti-oxidative enzyme activities known to be associated with the apoplast, during the induction of blossom-end rot (BER) in bell pepper (Capsicum annuum L.) fruits grown under saline conditions. Pepper plants of the BER-sensitive cv. 'Mazurka' and less-sensitive cv. 'Selica' were grown under control, low-and high-salinity irrigation regimes. Fruits were harvested continuously and, after approx. 7 months under these conditions, BER symptoms started to appear (in the Spring season) and fruits were sampled for biochemical measurements. The intercellular washing fluid (IWF; apoplast fraction) of the fruit pericarp was prepared and enzyme activities in that fraction were determined. The results showed that production of reactive oxygen species (ROS) in the fruit apoplast was higher in 'Mazurka' than in 'Selica'. Apoplastic peroxidase and superoxide dismutase (SOD) activities, and the concentration of ascorbic acid, increased during 'Selica' fruit development, whereas the concentration of H 2 O 2 decreased. During the stage of rapid growth, when fruits are most susceptible to BER, apoplastic peroxidase activity was greatly enhanced in the healthy pericarp of BER-affected fruits compared to healthy fruits. Under saline conditions, apoplastic peroxidase and SOD activities, as well as ascorbic acid and H 2 O 2 levels, increased in healthy fruits of the resistant cultivar 'Selica'; whereas, in healthy fruit of the BER-sensitive cultivar 'Mazurka', apoplastic peroxidase activity decreased while the level of ascorbic acid increased, but to a much lesser extent than in 'Selica'. SOD activity and H 2 O 2 levels increased similarly in both cultivars with increasing levels of salinity. It is proposed that apoplast-associated peroxidase activity and ascorbic acid concentration are increased coordinately in response to salt stress and may be part of a protective anti-oxidation mechanism that determines cultivar sensitivity to BER. T omato and pepper are often affected by blossom-end
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Although gibberellins (GAs) have been shown to induce development of the physiological disorder blossom-end rot (BER) in tomato fruit (Solanum lycopersicum), the mechanisms involved remain largely unexplored. BER is believed to result from calcium (Ca) deficiency, but the relationship between Ca content and BER incidence is not strong. Our objectives were to better understand how GAs and a GA biosynthesis inhibitor affect BER development in tomato fruit. Tomato plants of two BER-susceptible cultivars, ‘Ace 55 (Vf)’ and ‘AB2,’ were grown in a greenhouse environment and subjected to Ca-deficiency conditions. Plants were treated weekly during fruit growth and development with 300mgL−1GA4+7, 300mgL−1 prohexadione-calcium (Apogee®, a GA biosynthesis inhibitor), or water beginning 1day after flower pollination. GA4+7 treatment induced an increase in BER incidence in both cultivars up to 100%, whereas ‘Ace 55 (Vf)’ and ‘AB2’ plants treated with Apogee did not show BER incidence. The number of functional xylem vessels was higher in the placental and pericarp tissue of tomato fruit treated with Apogee at the early stages of fruit growth. Treatment with Apogee also increased fruit pericarp Ca concentration. GA4+7 treatment enhanced the expression of the putative CAX and Ca-ATPase genes, that code for proteins involved in Ca movement into storage organelles. The lowest water-soluble apoplastic Ca concentration and the highest membrane leakage values were observed in the pericarp of GA4+7-treated fruit. These results suggest that GAs consistently reduced fruit Ca uptake and water-soluble apoplastic Ca concentration, leading to leakier plasma membranes and an increase in BER development in fruit tissue of both tomato cultivars. Keywords Solanum lycopersicum L.–Ca–Blossom-end rot–BER–Apoplast–Membrane
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Calcium (Ca) uptake into fruit and leaves is dependent on xylemic water movement, and hence presumably driven by transpiration and growth. High leaf transpiration is thought to restrict Ca movement to low-transpiring tomato fruit, which may increase fruit susceptibility to the Ca-deficiency disorder, blossom end rot (BER). The objective of this study was to analyse the effect of reduced leaf transpiration in abscisic acid (ABA)-treated plants on fruit and leaf Ca uptake and BER development. Tomato cultivars Ace 55 (Vf) and AB2 were grown in a greenhouse environment under Ca-deficit conditions and plants were treated weekly after pollination with water (control) or 500 mg l(-1) ABA. BER incidence was completely prevented in the ABA-treated plants and reached values of 30-45% in the water-treated controls. ABA-treated plants had higher stem water potential, lower leaf stomatal conductance, and lower whole-plant water loss than water-treated plants. ABA treatment increased total tissue and apoplastic water-soluble Ca concentrations in the fruit, and decreased Ca concentrations in leaves. In ABA-treated plants, fruit had a higher number of Safranin-O-stained xylem vessels at early stages of growth and development. ABA treatment reduced the phloem/xylem ratio of fruit sap uptake. The results indicate that ABA prevents BER development by increasing fruit Ca uptake, possibly by a combination of whole-plant and fruit-specific mechanisms.
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Plant growth retardants (PGRs) reduce the shoot growth of plants by inhibiting gibberellin biosynthesis. In this study, we performed detailed analyses of the inhibitory effects of PGRs on Arabidopsis abscisic acid (ABA) 8'-hydroxylase, a major ABA catabolic enzyme, recently identified as CYP707As. In an in vitro assay with CYP707A3 microsomes expressed in insect cells, uniconazole-P inhibited CYP707A3 activity more effectively than paclobutrazol or tetcyclacis, whereas the other PGRs tested did not inhibit it significantly. Uniconazole-P was found to be a strong competitive inhibitor (K(i)=8.0 nM) of ABA 8'-hydroxylase. Uniconazole-P-treated Arabidopsis plants showed enhanced drought tolerance. In uniconazole-P-treated plants, endogenous ABA levels increased 2-fold as compared with the control, and co-application of GA(4) did not suppress the effects, indicating that the effects were not due to gibberellin deficiency. Thus uniconazole-P effectively inhibits ABA catabolism in Arabidopsis plants. We also discuss the structure-activity relationship of the azole-type compounds on ABA 8'-hydroxylase inhibitory activity.
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A high incidence of blossom-end rot (BER) fruits is a serious problem in hydroponics of tomato, especially in hot summer season. We examined the morphological aspects based on light and electron microscopic observations. The fluid-soaked area appeared on the surface of fruits at first, finally the color changed to black and corky dark brown. The cells that exhibited abnormal structure appeared at the part of epidermal and subepidermal layers at first. Electron microscopic observation showed that (1) disruption of plasma membrane and tonoplast occurred, (2) the cell wall was wavy-shaped, (3) the endoplasmic reticulum was broken down, (4) the plastids became swollen, etc. Plasma membrane and tonoplast are the sensitive parts to cell death affected by BER. The peripheral cells around necrotic tissues of BER had normal internal structure, but the plasma membrane was detached from the cell wall, suggesting occurrence of plasmolysis. The phenomenon of plasmolysis was rarely observed at other parts of fruits distant from the part of BER and in the cells of normal fruits. Those findings suggested that the distal part of fruits was under osmotic stress when BER appeared. In developed BER fruits, the sub-epidermal cells were broken down. Some cells around broken cells showed deposition of lignin, and peripheral cells showed meristematic structure like callus. It is thought that they are the response of wound healing of tomato fruits.
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Water and calcium uptake are important factors affecting the incidence of fruit blossom-end rot (BER) in tomato and pepper. In the present study an attempt was made to manipulate these factors by severe root pruning and to examine the effect on BER in greenhouse-grown bell pepper (Capsicum annuum L., cv. Mazurka). Pepper plants were transplanted, with the root system split into four separated compartments, each containing a single root quarter. Removal of half, or three quarters, of the root from fruit-bearing plants significantly reduced fruit BER incidence compared with plants with intact roots, especially in fruits which were at the rapid expansion stage at the beginning of the treatments. Removal of three quarters of the root reduced midday leaf water potential, stomatal conductance, and plant height. The number and weight of fruits were not affected by these treatments. Root pruning caused only a slight reduction in stem sap flow, as measured by the heat pulse technique. Calcium concentrations in the distal part of fruits from quarter-root-plants were higher than in fruits from non-pruned plants, whereas magnesium and potassium concentrations were not affected. On the other hand, in the leaves, calcium, magnesium and potassium concentrations were all reduced by root pruning. The K/Ca ratio decreased in the blossom-end of the fruits and increased in the leaves of root-pruned compared with control plants. The results suggest that root pruning did not affect the total uptake of calcium, apparently driven by transpiration, but did enhance calcium partitioning to the developing fruit. Root pruning also affected calcium distribution within the fruit and therefore attenuated BER incidence.
Chapter
Understanding the mechanisms of calcium (Ca2+) deficiency disorder development in plants has been a challenge for more than a 100 years. Previous studies support the hypothesis that Ca2+ deficiency disorders can be triggered by mechanisms that reduce plant Ca2+ uptake from the soil, fruit Ca2+ uptake from the plant, and Ca2+ translocation within the fruit, and also result in abnormal regulation of cellular Ca2+ partitioning. Plant Ca2+ uptake can be determined by Ca2+ content and availability in the soil, root growth, activity of apoplastic and symplastic pathways of root Ca2+ uptake, as well as uptake competition between Ca2+ and other nutrients. Fruit Ca2+ uptake is determined by Ca2+ content in the xylem sap, and xylem/phloem ratio of fruit sap uptake, which is affected by the rates of leaf and fruit transpiration and growth. Calcium translocation to distal fruit tissue, containing the lowest fruit Ca2+ content and the highest susceptibility to Ca2+ deficiency disorders, is potentially dependent on the cell wall Ca2+ -binding capacity and symplastic Ca2+ uptake by the tissue at the peduncle end of the fruit, abundance of functional xylem vessels connecting peduncle and distal fruit tissues, as well as the hydrostatic gradient required for Ca2+ translocation towards the distal tissue. Cellular Ca2+ partitioning is defined by the activity of Ca2+ channels, Ca2+ ATPases, and Ca2+ exchangers present in cellular membranes, as well as the capacity of the cell wall to bind Ca2+, and the formation of Ca2+ precipitates in different cellular compartments. Therefore, Ca2+ deficiency disorders in fruit may not be caused by a single factor, but most likely by a combination of mechanisms that lower Ca2+ concentration at a specific tissue and cell localization, leading to Ca2+ deficiency symptoms. © 2012 Wiley-Blackwell. Published 2012 by John Wiley & Sons, Inc.
Article
気温, 送風, 媛化剤散布が, 発育, 体内無機栄養状態, 尻ぐされ果 (BER) 発生率に及ぼす影響を調査した.気温を28℃から24℃に下げることにより個体重, 葉重, 体内成分含量は増加し, BER発生率は著しく減少した.また, 5ppmウニコナゾール散布によりBER発生率が減少したが, 各器官乾物重と果実収量も減少した.BERは高温, 上位葉への送風で高い発生率を示した.BER発生率は果房送風によりやや減少した.果径肥大は開花後13日~22日の間に肥大速度が高くなり, その時期にBERが多発した.果実初期生育時の果径増加速度の最高値は高温, 上位葉への送風により増加した.ウニコナゾール散布により成長速度は低下し, 果径増加速度の最高値が低下した.BER発生率の増加に伴って果実のK含有率が増加し, Ca含有率は減少する傾向がみられた.ウニコナゾール散布により果実のK含有率が減少し, Caの含有率は増加した.BER発生率の増加に伴い果実のK/Ca, N/Ca比は増加し, Ca/Mg比は低下した.以上より, 果実の肥大初期の高温や蒸散流の低下により, 果実中へのCaの流入が抑制されると, 果実の無機栄養状態が好ましくない状態になりBERが発生すると考えられる.また, 開花後0~25日の果房への冷風送風と開花前後のウニコナゾールの葉面散布がBERの発生を防止するうえで有効であった.
Article
Tomato (Lycopersicon esculentum cultivar ‘Summer Flavor’) seedlings received a single uniconazole soil drench at 0, 12.5, 25, 50, 100, 200 or 400 μg per plant at the five-leaf stage. After 2 weeks of growth, plant height, length and area of the fifth leaf, and plant fresh and dry weights progressively decreased with increasing rates of uniconazole. The rate of leaf production was unaffected at low rates of the retardant. Another group of seedlings received soil drenches at the above rates at the onset of flowering. Uniconazole applied at both stages resulted in short final plant height at all rates. Total fruit yield was reduced by uniconazole at the 12.5 μg per plant rate, with further reduction as rates increased. Yields of large fruits (≥ 110 g) increased at the two lower rates of uniconazole. The incidence of blossom-end rot (BER) was nearly eliminated by this retardant at all rates. Foliar application of uniconazole at the flowering stage with rates of 0, 25, 50, 100, 250, 500, 750 or 1000 μg per plant had less control on plant height compared with their respective rates as a soil drench. Total fruit yield was not affected until the rate exceeded 100 μg per plant. The incidence of BER decreased sharply with increasing rates of uniconazole. The number and weight of fruits ≥ 110 g were doubled at rates between 25 and 250 μg per plant. Plant fresh and dry weights were lower as a result of uniconazole foliar spray. Fruit number was reduced regardless of the rate and stage of uniconazole application.
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The transition of T' phase to orthorhombic phase of Sm1.85Ce0.15CuO4-y was discovered by high temperature X-ray diffraction in vacuum at 715°C. The orthorhombic phase obtained by annealing at high temperatures in vacuum is a superconducting phase (Tce = 10 K, a = 0.396(1), b = 0.395(0), c = 1.201(4) nm) which has a slight deviation from T' phase. XPS analysis indicated that there are mixing valences of Cu1+ and Cu2+ in the samples. Oxygen deficiencies can be produced by reducing treatment and annealing at high temperature in vacuum. The appropriate amount of oxygen deficiencies may play an important role in the superconductivity of both electron type and hole type materials. A phenomenological structural model of electron transfer is proposed to explain some experimental results.
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GA1, GA8, GA17, GA19, GA20 and GA29 were identified by combined gas chromatography-mass spectrometgry (GC-MS) in immature seeds and pericarp of Lycopersicon esculentum Mill. (tomato). Higher levels of these GAs were present in the seeds than in the pericarp; seeds in addition contained GA15, GA24, GA25, and GA44. Fruits of the Lycopersicon pimpinellifolium Mill. mutant I were smaller and contained lower GA1 concentrations, but higher GA20 concentrations, than those of mutants III and IV. In contrast, differences in fruit size in L. esculentum due to position on the truss did not correlate with GA1 concentration in either the pericarp or seeds.
Article
Calcium mobility in tomato plants grown in sand- and water-cultures with little calcium and other cations was studied using 45Ca as a tracer. Radioautographs and 45Ca activity measurements of plant parts showed that 45Ca was “fixed” near the base (root or stem) of plants grown in low-Ca nutrient solutions, compared with plants grown with adequate Ca in which 45Ca was readily translocated to the tops. When plants were grown with little Ca and other cations (i.e. low K, Mg and micronutrients), 45Ca was also readily translocated to the tops. Calcium stress during the fruiting stage caused serious blossom-end rot of fruits, indicating that Ca absorbed by plants before fruiting was not readily available subsequently for fruit development. Leaf total-Ca is not a reliable index for predicting fruit-Ca deficiency in tomatoes.
Article
The effects of paclobutrazol treatment on plasma membrane lipid composition and ATPase activity of bell pepper fruit (Capsicum annuum) subjected to chilling temperatures were assessed. Application of the growth regulator paclobutrazol affected plant growth and fruit morphology. The plants were more compact and the fruits were less elongated than control fruits. There was about 60% more plasma membrane on a fresh weight basis from treated fruits. At harvest there was no difference in sterol to phospholipid ratio, or in phospholipid fatty acid composition of control compared with paclobutrazol treated fruit. However, plasma membrane ATPase acitivity of treated fruit was two times higher than that of control fruit. After storage at chilling temperature (2°C), the control fruit developed more chilling iniury, and had greater weight loss and a higher rate of K+ leakage than paclobulrazol treated fruit. Plasma membrane phospholipid content decreased and saturation of phospholipid fatty acids was higher than in control fruit. These two changes were largely absent in plasma membrane from treated fruit. At harvest antioxidant levels in the plasma membrane of paclobutrazol treated peppers were higher than in those of controls and changed little during storage, whereas levels in control fruit plasma membrane decreased 66%. ATPase activity increased and then decreased in control fruit held at low temperature, whereas in treated fruit activity was constant. The protective effect of paclobutrazol against chilling injury of pepper fruit may result from a combination of its effect on fruit morphology, and protection of the lipids against oxidative stress.
Article
Many researchers have attributed the occurrence of blossom-end rot (BER) in tomato to a deficiency of Ca2+ in the fruit or parts of the fruit in connection with the uptake of nutrients by the roots, the transport of Ca2+ to and within the fruit or a varying demand for Ca2+ depending on the growth rate of fruits. However, a critical concentration of Ca2+ in the fruit has not yet been found, and the influence of favourable or unfavourable growing conditions on the development of BER is still poorly understood. The symptoms of BER are caused by a disintegration of the cell membranes and an increased ion permeability. It is proposed that not a single factor but the sequence of two equally important factors are involved: (1) a higher susceptibility to various stresses due to an increase in physiologically active gibberellins (GAs) and a resulting decrease in Ca2+, causing the enhanced permeability of cell membranes, (2) some kind of stress, e.g. by soil water deficit, high salinity, or high NH4+ activity, causing the deterioration of cell membranes with subsequent loss of turgor and leakage of cell liquids. BER seems to occur when stress exceeds stress tolerance, most frequently in young fruit at the beginning of cell enlargement. At this stage, usually the highest amount of physiologically active GAs and the lowest amount of Ca2+ are found, i.e. stress tolerance is lowest. Ca2+ per se is considered neither a primary nor an independent factor in the development of BER. This hypothesis is advanced with the objective of stimulating further research into the causes of a physiological disorder that, as yet, remains stubbornly obscure.
Article
Blossom-end rot (BER) of tomato ( Lycopersicon esculentum) fruits is considered to be a physiological disorder caused by calcium deficiency. We attempted to clarify the localization of calcium in the pericarp cells and the ultrastructural changes during the development of BER. Calcium precipitates were observed as electron-dense deposits by an antimonate precipitation method. Some calcium precipitates were localized in the cytosol, nucleus, plastids, and vacuoles at an early developmental stage of normal fruits. Calcium precipitates were increased markedly on the plasma membrane during the rapid-fruit-growth stage compared with their level at the early stage. Cell collapse occurred in the water-soaked region at the rapid-fruit-growth stage in BER fruits. There were no visible calcium precipitates on the traces of plasma membrane near the cell wall of the collapsed cells. The amount of calcium precipitates on plasma membranes near collapsed cells was smaller than that in the cells of normal fruits and normal parts of BER fruits, and the amount on cells near collapsed cells was small. The amount of calcium precipitates on the plasma membranes increased as the distance from collapsed cells increased. On the other hand, calcium precipitates were visible normally in the cytosol, organelles, and vacuoles and even traces of them in collapsed cells. The distribution pattern of the calcium precipitates on the plasma membrane was thus considerably different between normal and BER fruits. On the basis of these observations, we concluded that calcium deficiency in plasma membranes caused cell collapses in BER tomato fruits.
Article
Today no one questions the assertion that Ca2+ is a crucial regulator of growth and development in plants. The myriad processes in which this ion participates is large and growing and involves nearly all aspects of plant development (recent reviews in Harper et al., 2004; Hetherington and Brownlee, 2004; Hirschi, 2004; Reddy and Reddy, 2004; Bothwell and Ng, 2005). Despite this wealth of research, the concept of Ca2+ as an intracellular regulator is relatively recent and within the professional life span of many people who are still active and working on this topic today. The aim of this essay is to identify those lines of thought and research that led to the idea that Ca2+ is a second messenger in plant cell growth and development. This essay thus focuses primarily on work starting in the mid sixties and extending to the mid eighties. I do not provide an exhaustive review of the history of Ca2+ research, nor do I attempt to treat modern aspects of Ca2+ research. However, I do strive to identify the roots of modern Ca2+ research and to chart the origin of the current revolution.
Article
Reactive oxygen species (ROS) are versatile molecules mediating a variety of cellular responses in plant cells, including programmed cell death (PCD), development, gravitropism, and hormone signaling. A picture showing how ROS function in signal transduction networks has started to emerge as the result of recent studies providing genetic, cell biological, and physiological evidence describing roles for ROS in signaling (Apel and Hirt, 2004; Laloi et al., 2004; Mittler et al., 2004; Mori and Schroeder, 2004). However, further efforts are necessary to characterize the targets and molecular functions of ROS, as well as the complex interplay of ROS-generating and ROS-scavenging mechanisms. Moreover, the interactions of nitric oxide with other ROS species in hormone signaling is a subject of interest (Desikan et al., 2004; Wendehenne et al., 2004; Guo and Crawford, 2005; Bright et al., 2006). Due to limited space, in this Update article we focus on recent progress made in understanding the roles of ROS in hormone signaling.
Article
Plant growth is adaptively modulated in response to environmental change. The phytohormone gibberellin (GA) promotes growth by stimulating destruction of the nuclear growth-repressing DELLA proteins [1-7], thus providing a mechanism for environmentally responsive growth regulation [8, 9]. Furthermore, DELLAs promote survival of adverse environments [8]. However, the relationship between these survival and growth-regulatory mechanisms was previously unknown. Here, we show that both mechanisms are dependent upon control of the accumulation of reactive oxygen species (ROS). ROS are small molecules generated during development and in response to stress that play diverse roles as eukaryotic intracellular second messengers [10]. We show that Arabidopsis DELLAs cause ROS levels to remain low after either biotic or abiotic stress, thus delaying cell death and promoting tolerance. In essence, stress-induced DELLA accumulation elevates the expression of genes encoding ROS-detoxification enzymes, thus reducing ROS levels. In accord with recent demonstrations that ROS control root cell expansion [11, 12], we also show that DELLAs regulate root-hair growth via a ROS-dependent mechanism. We therefore propose that environmental variability regulates DELLA activity [8] and that DELLAs in turn couple the downstream regulation of plant growth and stress tolerance through modulation of ROS levels.
Biochemical changes in pepper (Capsicum annuum L.) fruit associated with blossom-end rot (abstract) HortScience 36, 485. Bangerth, F., 1973. Investigations upon Ca related physiological disorders
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Aloni, B., Karni, L., 2001. Biochemical changes in pepper (Capsicum annuum L.) fruit associated with blossom-end rot (abstract). HortScience 36, 485. Bangerth, F., 1973. Investigations upon Ca related physiological disorders. J. Phy-topathol. 77, 20–37.
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Goodwin, P.B., 1978. Phytohormones and fruit growth. In: Letham, D.S., Goodwin, P.B., Higgins, T.J.Y. (Eds.), Phytohormones and Related Compounds – A Compre-hensive Treatise. Elsevier, Amsterdam, pp. 175–214.
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