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Anatomy of the vascular bundles. LM images of cross sections taken through rachis (A and B) and pedicels (C and D) of H (A and C) and BS (B and D) samples. Sections were stained with Toluidine Blue (A and B) and Safranin-Astra Blue (C and D). Lignified secondary xylem cell walls stain greenish with Toluidine Blue, whereas cellulosic primary cell walls stain dark blue (A and B). Double staining with Safranin-Astra Blue stains lignified secondary xylem cell walls reddish, whereas cellulosic primary cell walls turn blue (C and D). Arrows indicate primary rays. Abbreviations: C-cambium; P-pith; PP-primary phloem, PPFprimary phloem fibers; PX-primary xylem; SPsecondary phloem; SX-secondary xylem. Scale bars: 100 μm. (For interpretation of the references to colour in this figure legend, the reader is referred to the web version of this article.)
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Berry Shrivel (BS) is a post-veraison physiological ripening disorder of grapevine berries. Its symptoms encompass low pH, reduced content of sugars and anthocyanins, and loss of turgor leading to berries shriveling. Evidence for the primary causes of BS is still speculative and anatomical studies are scarce. So far, anatomical studies have determi...
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... x V. berlandieri) rootstock) between 2013 and 2015. Samples (3-5 mm in length) for LM and TEM were collected from basal, middle and distal parts of rachis (having a diameter of ∼4 mm) and randomly selected pedicels of H and BS af- fected berry clusters in 2013 and 2014 ( Supplementary Fig. S1). In 2013, samples were collected from eight H and eight BS affected clusters taken from 16 individual plants and in 2014 from seven H and 11 BS affected clusters taken from 18 plants. ...
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... anatomical structure of the vascular system in rachis and ped- icel of H and BS affected grapes were analyzed on sections at LM level ( Fig. 1, Supplementary Fig. S1). Among the three staining methods applied (Toluidine Blue, Safranin-Astra Blue and FCA) the Toluidine Blue gave the best results. Conductive tissues were arranged into dis- crete vascular bundles separated by primary rays (Fig. 1). The vascular cylinder of rachis consisted of pith, primary and secondary xylem, cambium, secondary ...
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... in rachis and ped- icel of H and BS affected grapes were analyzed on sections at LM level ( Fig. 1, Supplementary Fig. S1). Among the three staining methods applied (Toluidine Blue, Safranin-Astra Blue and FCA) the Toluidine Blue gave the best results. Conductive tissues were arranged into dis- crete vascular bundles separated by primary rays (Fig. 1). The vascular cylinder of rachis consisted of pith, primary and secondary xylem, cambium, secondary phloem, and primary phloem with distinct pri- mary phloem fiber caps in some larger bundles ( Fig. 1A and B; Sup- plementary Fig. S1). The zone of primary phloem fibers was absent in pedicels ( Fig. 2C and D; Supplementary Fig. S1). ...
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... and FCA) the Toluidine Blue gave the best results. Conductive tissues were arranged into dis- crete vascular bundles separated by primary rays (Fig. 1). The vascular cylinder of rachis consisted of pith, primary and secondary xylem, cambium, secondary phloem, and primary phloem with distinct pri- mary phloem fiber caps in some larger bundles ( Fig. 1A and B; Sup- plementary Fig. S1). The zone of primary phloem fibers was absent in pedicels ( Fig. 2C and D; Supplementary Fig. S1). Comparing H (Fig. 1A and C) and BS (Fig. 1B and D) clusters, no anatomical differences in the vascular tissue organization could be determined with LM cross sec- ...
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... gave the best results. Conductive tissues were arranged into dis- crete vascular bundles separated by primary rays (Fig. 1). The vascular cylinder of rachis consisted of pith, primary and secondary xylem, cambium, secondary phloem, and primary phloem with distinct pri- mary phloem fiber caps in some larger bundles ( Fig. 1A and B; Sup- plementary Fig. S1). The zone of primary phloem fibers was absent in pedicels ( Fig. 2C and D; Supplementary Fig. S1). Comparing H (Fig. 1A and C) and BS (Fig. 1B and D) clusters, no anatomical differences in the vascular tissue organization could be determined with LM cross sec- ...
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... vascular bundles separated by primary rays (Fig. 1). The vascular cylinder of rachis consisted of pith, primary and secondary xylem, cambium, secondary phloem, and primary phloem with distinct pri- mary phloem fiber caps in some larger bundles ( Fig. 1A and B; Sup- plementary Fig. S1). The zone of primary phloem fibers was absent in pedicels ( Fig. 2C and D; Supplementary Fig. S1). Comparing H (Fig. 1A and C) and BS (Fig. 1B and D) clusters, no anatomical differences in the vascular tissue organization could be determined with LM cross sec- ...
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... zone of primary phloem fibers was absent in pedicels ( Fig. 2C and D; Supplementary Fig. S1). Comparing H (Fig. 1A and C) and BS (Fig. 1B and D) clusters, no anatomical differences in the vascular tissue organization could be determined with LM cross sec- tions. ...
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... of rachis consisted of pith, primary and secondary xylem, cambium, secondary phloem, and primary phloem with distinct pri- mary phloem fiber caps in some larger bundles ( Fig. 1A and B; Sup- plementary Fig. S1). The zone of primary phloem fibers was absent in pedicels ( Fig. 2C and D; Supplementary Fig. S1). Comparing H (Fig. 1A and C) and BS (Fig. 1B and D) clusters, no anatomical differences in the vascular tissue organization could be determined with LM cross sec- ...
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... develop and function only during one vegetation period. Primary xylem and phloem develop from procambial cells derived from apical meristem and later in the season, secondary xylem and phloem are developed from fascicular cambial cells. In our observations, primary phloem and xylem were similarly developed in H and BS affected rachis and pedicels (Fig. 1). Additionally cambial ac- tivity can be assumed as secondary phloem and xylem, which are de- veloped by cambium, are present in rachis and pedicels of H and BS samples. Previous studies have described alternating bands of sec- ondary hard and soft phloem in peduncles of BS affected berries orga- nized in tangentially continuous rows ( ...
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Wood, a type of xylem tissue, originates from cell proliferation of the vascular cambium. Xylem is produced inside, and phloem outside, of the cambium¹. Morphogenesis in plants is typically coordinated by organizer cells that direct the adjacent stem cells to undergo programmed cell division and differentiation. The location of the vascular cambium...
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... These helped to elucidate the primary [9][10][11][12] and the secondary metabolism [13], including different aroma compounds [14], towards the effects of abiotic stress, e.g., drought [15,16] or heat stress [17,18], as well as changed profiles due to pathogen infections [19]. Attention one of the hypotheses of berry shrivel induction but results also hint toward a disturbed transport of assimilated and nutrients towards berries in association with reduced cell viability in the rachis and berries [43,45,46,51,52]. Cell death in berries and rachis of BS symptomatic grape clusters have been observed [43,53], and recently it was shown that cell death in the berries precedes the ones in the rachis [46]. ...
... Cell death in berries and rachis of BS symptomatic grape clusters have been observed [43,53], and recently it was shown that cell death in the berries precedes the ones in the rachis [46]. Collapsed cells as well as a localized thickening of cell walls in the secondary phloem have been observed on the cellular level in the rachis and pedicels of BS grape clusters [45]. These dramatic changes point toward processes involving cell wall modification and degradation in either BS induction, BS symptoms development, or both. ...
... In a previous study, we could show that pedicels of BS-affected grape clusters were thinner, which may indicate a reduced growth with an impact on assimilate transport [47]. On the other hand, no obvious major anatomical rearrangements were observed in the vascular tissue organization in the rachis and pedicels of BS grape clusters with light microscopy [45], while a localized cell wall thickening of the secondary phloem in the rachis and pedicels of BS grapes, as well as degraded and collapsed cells near cambium cells, were observed [45]. All these observations point towards cell wall modification and degradation in BS symptomatic grape rachis, for example, due to the activity and altered expression of EXPA6 (VIT_06s0004g04860), XTH32 (VIT_06s0061g00550), BXL1 (VIT_05s0077g01280), and PME3 (VIT_09s0002g00320), confirmed by qPCR. ...
Berry shrivel (BS) is one of the prominent and still unresolved ripening physiological disorders in grapevine. The causes of BS are unclear, and previous studies focused on the berry metabolism or histological studies, including cell viability staining in the rachis and berries of BS clusters. Herein, we studied the transcriptional modulation induced by BS in the rachis of pre-symptomatic and symptomatic clusters with a custom-made microarray qPCR in relation to a previous RNASeq study of BS berries. Gene set analysis of transcript expression in symptomatic rachis tissue determined suppression of cell wall biosynthesis, which could also be confirmed already in pre-symptomatic BS rachis by CESA8 qPCR analyses, while in BS berries, a high number of SWITCH genes were suppressed at veraison. Additionally, genes associated with the cell wall were differently affected by BS in berries. A high percentage of hydrolytic enzymes were induced in BS grapes in rachis and berries, while other groups such as, e.g., xyloglucan endotransglucosylase/hydrolase, were suppressed in BS rachis. In conclusion, we propose that modulated cell wall biosynthesis and cell wall assembly in pre-symptomatic BS rachis have potential consequences for cell wall strength and lead to a forced degradation of cell walls in symptomatic grape clusters. The similarity to sugar starvation transcriptional profiles provides a link to BS berries, which are low in sugar accumulation. However, further studies remain necessary to investigate the temporal and spatial coordination in both tissues.
... This decreased rate may be due to the sugar accumulation disorder, which is currently being studied for its exact causes. In literature this discrepancy in harvest time has been attributed in a series of reasons among them is the pathogens effect [28], the anatomical problems caused by nutrient metabolism [29], and to weather conditions [30], reasons that can act separately of synergistic. In the present experiment, the different weather conditions recorded among the three cultivation years, exhibiting different specific characteristics for the different phenological periods should be the major reason. ...
Grape is considered as one of the world's most important crop. Climatic and soil parameters can significantly affect the yield and quality of grapes. The aim of this study was to analyse and assess this impact of soil and climatic conditions on table grape yield and quality. The experiments took place on a commercial table grape vineyard in Greece for three successive years (2015, 2016 and 2017). The climatic conditions were found to affect significantly the table grape yield and all the tested quality parameters apart from the berry density and titratable acidity. In addition, the climatic conditions affected the table grape yield and quality but in a different degree per phenological stage. The effect of soil conditions on table grape yield and quality parameters differed significantly per annum due to the climatic conditions variation in terms of temperature and precipitation. The conducted analysis highlighted that the information regarding the combined effect of weather and soil conditions on the spatiotemporal variability of the production of table grapes in terms of their quantity and quality is an essential component of the modern precision viticulture. Based on the contemporary principles of the precision viticulture, appropriate cultivation practices can be planned and implemented, reducing thus the impact of climate change on the vine production efficiency and quality.
... According to McCarthy (1999), for this type of shriveling, berries lose weight due to water loss, and sugars are concentrated. Although studies about berry shrivel have increased recently (Bondada, 2016;Crespo Martinez et al., 2019;Hall et al., 2011;Keller et al., 2016), the causes of berry shrivel are still unclear. ...
Late-season dehydration is known as a type of berry shrivel and it is often associated with wine grapes. This study is different from other berry shrivel studies as it focuses on seedless grape cv. Sultan 7 (Vitis vinifera L.) and aims to determine the effects of different pruning levels (90, 150, 180 buds per vine), leaf removal (LR) (25%, 50% per shoot) treatments on vine yield, and the number of harvested clusters. Additionally, we aim to find the effects of treatments on the number of unhealthy clusters with shriveled (S) berries. Although treatments were conducted under the same irrigation practices and determined close leaf water potential values, S berries were not obtained from the vines with 90 buds per vine treatment. Thus, S berries were determined in the lower side of unhealthy clusters together with non-shriveled (NS) berries in treatments of 150 and 180 buds per vine. It was found that the total soluble solids content and titratable acidity of S berries were higher than NS. While high pruning levels per vine enabled to obtain more clusters, the expected yield was negatively affected due to the quantity of the unhealthy cluster. Significant polynomial regression was observed between the number of harvested and unhealthy clusters in 2016 (R 2 = 0.69) and 2017 (R 2 = 0.76). In this research, it was hypothesized that the water losses in S berries of unhealthy clusters in overloaded vines (150, 180 buds per vine) were caused by obtaining more harvested clusters compared to the 90 buds per vine treatment.
... Dissimilar from bunch stem necrosis, rachis and pedicels stay viable and green with no obvious symptoms on the surface. Nevertheless morpho-anatomical study observed cell death in the rachis of symptomatic grape clusters of Cabernet Sauvignon (Bondada and Keller 2012b;Hall et al. 2011), TEM analysis showed collapsed cells and cell wall thickenings in the secondary phloem (Crespo-Martinez et al. 2019) and light microscopy and SEM analyses revealed higher rates of callose deposition on sieve plates (Bondada 2016;Crespo-Martinez et al. 2019). Reduced cell viability was also observed in grape berries leading to the assumption that the loss of cell membrane integrity is an important factor for water loss and berry shrinkage (Krasnow et al. 2008(Krasnow et al. , 2012. ...
... Dissimilar from bunch stem necrosis, rachis and pedicels stay viable and green with no obvious symptoms on the surface. Nevertheless morpho-anatomical study observed cell death in the rachis of symptomatic grape clusters of Cabernet Sauvignon (Bondada and Keller 2012b;Hall et al. 2011), TEM analysis showed collapsed cells and cell wall thickenings in the secondary phloem (Crespo-Martinez et al. 2019) and light microscopy and SEM analyses revealed higher rates of callose deposition on sieve plates (Bondada 2016;Crespo-Martinez et al. 2019). Reduced cell viability was also observed in grape berries leading to the assumption that the loss of cell membrane integrity is an important factor for water loss and berry shrinkage (Krasnow et al. 2008(Krasnow et al. , 2012. ...
The process of grape berry ripening follows three phases with distinct metabolic processes and complex regulations via phytohormones. The physiological ripening disorder berry shrivel (BS) is characterized by reduced sugar accumulation, low anthocyanin contents, and high acidity in affected berries. The processes leading to BS induction are unknown, but recent transcriptional data on reduced expression of switch genes hint towards a disturbed ripening onset. Herein we investigated the phytohormone composition throughout grape berry ripening in healthy and BS berries in Vitis vinifera L. cultivar Blauer Zweigelt. Thereby we hypothesize that phytohormones are key players for BS induction and suppress the expression of switch genes at veraison. The presented metabolomics and RNAseq data describe two distinct phytohormone profiles in BS berries, differing between pre- and post-veraison with a clear ethylene precursor (aminocyclopropane-1-carboxylic acid, ACC) peak before veraison. Exogenous application of ACC led to BS symptoms, while ethephone application led to berry abscission. During post-veraison, we observed high ABA-glucose ester (ABA-GE) and low indole-3-acetate aspartate (IAA-Asp) and isopentenyladenine (iP) contents in BS berries and the transcriptional induction of several phytohormone pathways. The presented descriptive data provide valuable knowledge to further decipher the role of phytohormones in BS induction and BS symptom development.
... Past studies focused on morpho-anatomical issues (Hall et al. 2011;Bondada and Keller 2012a;Bondada 2014;Crespo-Martínez et al. 2019), nutritional aspects (Bachteler et al. 2015;Bondada 2016;Griesser et al. 2017), viticultural practices (Raifer et al. 2014) or on a restricted list of genes . ...
... Previous studies showed reduced cell viability in the rachis of BS grapes possibly leading to a reduced transport function, although obvious necrosis on the outside of the rachis is not observed in our case (Hall et al. 2011;Bondada 2016). Recently, in-depth microscopic study of symptomatic BS Zweigelt grapes showed cell deformations and signs of total or more likely partial occlusion of the phloem sieve elements (Crespo-Martínez et al. 2019). A drawback of all anatomical studies so far is that presymptomatic analyses were not possible, as up to date no method exists to predict BS clusters in vineyards as well as a reliable method to induce BS. ...
Key message
The lower expression at veraison of several ripening master regulators “switch genes” can play a central role in the induction of the berry shrivel ripening physiological disorder in grapevine.
Abstract
Berry shrivel (BS) is a ripening physiological disorder affecting grape berry with visible symptoms appearing after veraison. Berry shrivel leads to shrinking berries with a reduced weight and a lower content of sugars and anthocyanins. In this study, for the first time a transcriptomic analysis coupled with selected metabolites quantification was undertaken to understand the metabolic modifications induced by the disorder. Different stages of berry development were considered including pre- and symptomatic berries. No metabolic alterations in the berry transcriptome and in the metabolite content was observed in pre-symptomatic and pre-veraison samples. Interestingly, at veraison, with still not visible symptoms appearing on the berry, a subset of genes, called switch genes previously suggested as master regulators of the ripening onset in grape berries, were strongly lower expressed in BS. Later during the ripening phase and with visible symptoms of the disorder, more than 3000 genes were differentially expressed. The genes up-regulated were related to hormone biosynthesis, response to stress and the phenylpropanoid pathway, while the genes down-regulated during ripening belonged mainly to the flavonoid pathway, and the sugar metabolism. In agreement, BS berries showed lower content of sugars and anthocyanins from the onset of veraison onward, while the amount of acids was not significantly affected. In conclusion, these results highlight a pivotal role of the switch genes in grapevine ripening, as well as their possible contribution to induce the ripening disorder berry shrivel, although it remains unclear whether this is part of the cause or consequences of the BS disorder.
Grapevine berry shrivel, a ripening disorder, causes significant economic losses in the worldwide wine and table grape industries. An early interruption in ripening leads to this disorder, resulting in shriveling and reduced sugar accumulation affecting yield and fruit quality. Loss of sink strength associated with berry mesocarp cell death is an early symptom of this disorder; however, potential internal or external triggers are yet to be explored. No pathogens have been identified that might cause the ripening syndrome. Understanding the underlying causes and mechanisms contributing to berry shrivel is crucial for developing effective mitigation strategies and finding solutions for other ripening disorders associated with climacteric and non-climacteric fruits. This review discusses alterations in the fruit ripening mechanism induced by berry shrivel disorder, focusing primarily on sugar transport and metabolism, cell wall modification and cell death, and changes in the phytohormone profile. The essential open questions are extracted and analyzed, thus identifying the critical knowledge gaps and key challenges for future research.
Banana (Musa spp.) is one of the main fruits consumed worldwide. However, finger drop, is a physiological disorder that causes many postharvest problems, which eventually reduces market value and consumer acceptance. Therefore, the objective of the study was to evaluate the anatomical changes that occur in the pedicel rupture area (drop zone) of bananas diploids (BB França) and tetraploid (BRS Pioneira) in three ripening stages. The levels of gene expression involved in the natural ripening process and in the development of finger drop, was also investigated. The accumulation of their mRNAs and those of expansin (EXP1), pectate lyase (PEL1) and xyloglucan endotransglucosylase/hydrolase protein (XTH4) genes already isolated from bananas were measured by quantitative polymerase chain reaction in three ripening stages. BB França presented a higher resistance to finger drop due to the presence of some specific morphoanatomical characteristics, such as larger parenchymal cells and greater deposition of lignin. In contrast, there was degeneration of the pedicel parenchymal tissue of the BRS Pioneira genotype, forming large empty spaces during the ripening of the fruits, mainly in stage 6, which contributed to the finger drop. The diploid BB França is a strong candidate for use in banana breeding programs aimed at fruit drop resistance. This will certainly improve the quality of banana varieties. Moreover, PEL1 proved to be an excellent candidate gene for functional studies of finger drop in bananas.
Grapevine yield is defined as the quantity of harvest, expressed as either grape mass or wine volumeunits, which has been collected per surface unit are and per crop cycle. The information about current and future yield, termed a yield assessment in this paper, is an essential decision-making element for the grape and wine industry. Crop management, wine-making, commercial, accounting and strategic operations are all adapted to and all impact on the expected yield of the current vintage. Numerous yield assessment approaches have been proposed in the scientific literature. However, only a few of them have considered their adaptation to the operational context under the constraints, needs and strategies of commercial vineyards and wineries. The few studies that have worked on the operational implementation of yield assessment methods have only partially addressed this issue, concentrating their improvement efforts on a single step in the yield assessment process. This paper first proposes to review the characteristics of yield development in an operational context that must be taken into account by yield assessment methodsThese characteristics are consolidated into three main challenges for yield assessment methods: (i) addressing the complex temporality of yield development, (ii) ensuring a local monitoring of yield development and (iii) fitting to the operational needs and constraints to allow for relevant decision support systems in the field. The approaches of yield estimation, prediction and forecast are discussed in the context of these challenges. In a second step, the paper proposes a generic framework for the yield assessment process, including a review of the variables that are used to explain grapevine yield. Issues and proposals from the literature associated respectively with measurement, sampling and yield modelling are reviewed and the need for improved modelling of relationships between explanatory variables and the desired, reported yield variable is discussed. The yield assessment methods found in the literature are categorised and compared according to measurement, estimation and modelling approaches, and then according to the three challenges identified for yield assessment in operational conditions, such that the yield assessment method is adapted to commercial needs and not to research objectives. In conclusion, concrete proposals for new grape yield assessment methods are discussed in order to investigate the as yet unexplored opportunities for the improvement in yield assessment in operational contexts that have been identified in the paper. These considerations could easily be transposed to other perennial crops.
Many dyes produced during the 19th century were named after locations. Manufacturers proliferated the number of synonyms used and in time, the original names were forgotten. Therefore, in the headings below, the original names are followed by the current name(s) in parentheses. The stories of some of these dyes that survived into the 21st century are recounted here. Numerical identity data are provided. Chemical structures also are provided and for simplicity, ionic structures, which can be multiple and pH variable, are presented as their parents.
