Mandalari , D. Carughi Published 22 December Medicine Plants Epidemiological and clinical studies have indicated positive outcomes related to tree nut consumption. Here, we review the production, nutrient, phytochemical composition and emerging research trends on the health benefits of pistachio nuts Pistacia vera L. Pistachios are a good source of protein, fiber, monounsaturated fatty acids, minerals and vitamins, as well as carotenoids, phenolic acids, flavonoids and anthocyanins.
Polyphenols in pistachios are important contributors to the… Expand. View PDF. Save to Library Save. Create Alert Alert. Share This Paper. Figures and Tables from this paper. International journal for vitamin and nutrition research. These findings have important implications for the horticultural management of this fruit species and can help in the breeding strategies for selecting the parents to cross the pistachio tree.
Materials and Methods 2. N , in 27th of June and in 22nd of July Bearing branches showed from 40 to 50 fruits; non-bearing had no fruits. The 12 samples were loaded into one lane of an Illumina flow cell, and clusters were created by Illumina Bot. De Novo Assembly, Evaluation and Annotation The quality of the raw sequences generated from transcriptome sequencing was assessed with FastQC version 1.
The total pre-processed reads from all 12 samples were then de novo assembled using Trinity [21] version 2. The complete assembly statistics and evaluation statistics is given in Table S1. The complete workflow of the Pistachio de novo transcriptome assembly and annotation were summarized in Figure S1. The comparison selected for the study is given in Table 1. Genes represented with an adjusted p-value FDR lower than 0.
In addition, the functional-enrichment analysis was performed to identify which gene ontology GO terms and metabolic pathways that were significantly enriched in differentially expressed genes DEGs.
Table 1. Results 3. De Novo Transcriptome Assembly and Annotation High-throughput sequencing technology has provided an excellent opportunity for transcriptome survey in non-model plant species, including P. To examine the inflorescence bud abscission phenomenon of P.
The picking dates of the material coincide with a period of the initial competition between fruit and inflorescence buds, not causing inflorescence bud drop, and a period of strong completion, causing the drop of inflorescence buds. The sequencing of the data of June produced million raw reads 60 Gb of data , whereas July produced million reads 59 Gb of data as a single-end. The high-quality single-end reads with an average quality score of 38 were selected for the transcriptome assembly after trimming off the low-quality bases and adapters from the June and July data sets Table S2.
The assembly was evaluated with BUSCO to assess the transcriptome assembly by measuring the completeness of the transcriptome based on evolutionary present universal single-copy orthologs. The number of up- and down-regulated genes, along with the total number of genes obtained in each sample comparison, are listed in Table 1.
We used RSEM for the quantification of the genes. The count matrix generated we then taken as the input by edgeR. The downstream analysis resulted in the identification of a total of 14, genes in which were up-regulated and were down-regulated.
For each of the analysis, the total number of genes ranged from to The number of genes up-regulated was in a range of to and down-regulated genes were spans from to It is likely that the cv.
Bianca faces a limitation of resources around the third week of June, when the first sampling of the plant material was made and, that in a month, it reaches its maximum peak, corresponding to the second sampling period when the drop of inflorescence buds started. The figure shows 37, genes were common among all the seasons which might have a role in the developmental process rather than the alternate bearing.
This comparison can also avoid factors like physiological and developmental changes that might occur in the bud during the two different time points June and July. Table S4. These enzymes are involved in the formation of carbohydrate reserves [29].
Figure 2. Figure shows the Mapman pathways in sucrose-starch metabolism. Individual genes were represented by small squares. The color scale indicates the log2 FC value. On the contrary, the expression of S-adenosyl-L-methionine-dependent methyltransferases was repressed. Genes , 11, 8 of 20 3. In contrast, we identified some GO-terms that were up-regulated in response to alternate bearing, such as nutrients ion transport, ABA catabolic process, gibberellin catabolic process, amino acid transmembrane transport and carbohydrate metabolic process Figure S4.
The genes involved in hormone-related categories are summarized in Figure 3. Figure 3. Relating to ABA there was an up-regulation in abscisic acid insensitive 3, lipid transfer protein 3, shaker potassium ion channel, SNF1, potassium transport 3, phosphotransmitter 4 and Carotenoid cleavage dioxygenase 1.
Genes , 11, 9 of 20 3. Figure 4. Similarly, we also found that the inflorescence buds of bearing and non-bearing pistachio shoots differed in their carbohydrate storage and mobilization patterns, suggesting that the in-season carbon mobilization might influence the flower bud abscission directly or indirectly linked to the alternate bearing.
Figure 5. The red circle represents the value of log2 fold change. The line indicates the effects on carbohydrate levels driven by differential expression of different CHO metabolism genes. Discussion The growth of the endocarp of the cultivar Bianca is from the first week of May to the end of June, while the growth of the embryo is from the first week of July to the end of August [1].
The relationships between the flower bud drops linked to alternate bearing and the carbohydrate storage have been mentioned in several studies [2,5,13,30]. The role of individual sugars in the process of inflorescence bud abscission has not yet been investigated [2]. The raffinose family of oligosaccharides has a wide range of predicted functions and are currently emerging as crucial molecules during stress response in plants [34], because of their membrane-stabilizing, antioxidant and, perhaps, predictable signaling functions [35].
They participate in several cellular functions, such as transport and storage of sugars [36], signaling molecule following pathogen attack and wounding [37], signal transduction [38], membrane trafficking [39] and mRNA export [40].
MIOX2 plays a prominent role in the oxidation of inositol for the needs of the plant in different tissues and it is involved in the biosynthesis of nucleotide sugar precursors for cell-wall matrix polysaccharides [41]. In source leaves, T6P fine-tunes sucrose levels by adjusting sucrose synthesis, while it regulates Sucrose consumption in sink organs, probably acting via multiple mechanisms, including inhibition of the SnRK1 gene [43]. T6P regulates growth in relation to sucrose supply by adjusting biosynthetic reactions and through regulating hormone signaling like auxin either directly or indirectly [44].
In the pistachio tree, it has been demonstrated that the accumulation of nitrogen, phosphorus and potassium is greater in inflorescence buds of non-bearing branches, compared to the ones of the bearing branches [10], and that the nutrient contents of the trees and annual nutrient consumption are influenced by the alternate bearing [10].
Competition between flower buds and developing nuts for N might play an important role. However, the information on the effects of fruiting on nutrient concentrations of different organs of pistachio trees relative to bud abscission is limited. Sugar phosphates transformed into sucrose and transport to fruit. This can reduce the sugar phosphates in the source tissues of trees with strong sink tissue such as fruit. A similar decline in photosynthesis due to leaf aging has been reported for apple trees [49] and olive trees [46].
In plants, nutrient limitation due to sink competition leading to sugar starvation is perceived as nutritional stress and generate changes in the redox status promote the synthesis of free radicals which can cause transient oxidative stress due to an increase of ROS generation [50], that can be neutralized by some adaptive mechanisms which can protect the cells from oxidative damage. The cells subjected to sugar starvation at the beginning try to adapt to this deficiency through a gradual metabolic reorganization that implies the substitution of carbohydrate metabolism by protein and lipid metabolism and that change may cause autophagy [51].
Variations in sugar levels induce changes in ROS production and sugar starvation can cause the activation of ROS production, as indicated by transcriptome profiling analysis, where sucrose starvation results in activation of oxidative stress genes, such as catalase [52]. The free polyamines could have an important physiological function in the development of flower bud abscission, which causes alternate bearing in pistachio trees [13].
It is possible that a decrease in N concentrations in plant tissues may cause a decrease in polyamines, as they can represent nitrogenous sources or as signal molecules that regulate the fruitlet abscission process in grapevine [55]. Many studies have highlighted that abscission or ethylene biosynthesis can be delayed with low levels of S-adenosylmethionine SAM.
During this phase, PAs and ethylene compete and PAs can become dominant. The low concentrations of PAs can trigger the senescence and cause abscission [56]. These references, along with our results, support the fact that polyamines could play a crucial role in the inflorescence bud abscission of pistachio.
A high level of polyamines is known to act as antisenescence agents and counteract the activity of abscisic acid and ethylene [57]. The competition between polyamines and ethylene pathways for S-adenosil methionine AdoMet or the inhibition of ACC syntase or ethylene forming enzyme EFE by polyamines can result in a mechanism that can modulate physiological events, including senescence and flower bud abscission.
Exogenous application of auxins prevented inflorescence bud abscission in pistachio [60]. In another study conducted in citrus the auxin amount is in a positive relationship with abscission by causing a delay of abscission, resulting in improvement in fruit quality and yield.
Our study shows that auxin conjugates play an important role in IAA metabolism, temporary storage reserves and inflorescence bud abscission. SnRK1 and TOR can target phosphorylation substrates to sense energy and nutrient levels and coordinate transcriptome, metabolism, cell growth and development [64]. TOR signaling plays an important role in stem and progenitor cell function and regulation that modulate proliferation and maintenance, cell-cell interactions and sink-source organ communication.
There are several studies reporting the involvement of ABA biosynthesis or ethylene perception critical for sugar signaling [65]. The ethylene signal is transmitted via a pathway that includes a transcriptional cascade, and EIN3 has been identified as a critical component within this cascade [66]. The regulation of EIN3 by ethylene and sugar indicates the cross talk between the two signaling pathways.
Interestingly, in barley, the antagonism between ABA and GA has been demonstrated to be an essential factor controlling the metabolism in aleurone cells and the PCD. In studies on abiotic stress, responses showed the involvement of polyamines in PCD through the production of hydrogen peroxide H2 O2 and Nitrogen oxide NO [68]. H2 O2 accumulation can cause the induction of PCD or stress tolerance, depending on the levels of intracellular Pas [70].
It is well known in both animals and plants that peroxisome PEX genes are induced by the universal stress signal, H2 O2 [74]. It has been found that BTS may act as an E3 ligase, which catalyzes the final step in the protein ubiquitination via the 26S proteasome [75]. Auxin related genes, on the contrary, are up-regulated, indicating a possible accumulation of this hormone, inducing cell growth and perhaps the down-regulation of TOR.
The oxidization of polyamines, such as Spd, occurs in the apoplast at a slow rate, with moderate production of H2 O2 , which activates the ROS-dependent protective pathway that does not trigger PCD or autophagy [78] Figure 6. Figure 6. Red shows the up-regulated genes and blue shows the down-regulated genes. In this situation the inflorescence buds do not occur. TOR signaling networks seem involved in cell-cell interactions, sink-source organ communication and autophagy [78].
Furthermore, in conjunction with low PA expression, down-regulation of auxin was also found resulting altogether in flower bud abscission Figure 7. Interestingly it seems that in pistachio exogenous application of PA can reduce many physiological disorders and inflorescence bud abscission [57,79], and preliminary experiments are currently being carried out in the cultivar Bianca to detect the dose and the timing of treatment. Figure 7. These results highlighted how the lack of resources carbohydrates and mineral elements in P.
This study provided further support to the theory of shoot autonomy in pistachio with regards to flower bud abscission and identified key genes and hormones associated with inflorescence bud abscission, the knowledge of which could also lead, in future, to a reduction of the inflorescence buds drop, through the development of biomarkers, and the possibility to modulate the alternate bearing.
Plant Cell, Tissue and Organ Culture —, Printed in the Netherlands. Siirt A. Siirt have been examined. Excised zygotic embryos that germinated in vitro were used as rootstocks. Current year shoot tips from mature trees of pistachio micrografted onto in vitro juvenile rootstocks, resulted in the restoration of shoot- bud proliferation. Variables tested include a size of microscion, grafting method, effects of culture medium and effects of time of the year at which shoot tips were used.
The results indicate that the easiest and most successful method for grafting was slit micrografting. High levels of micrograft take were achieved with 2—4 mm The survival rate of the shoot tips was directly related to time of the year. The best growth of microscion was obtained with the in vitro forced shoot tips rather than with shoot tips excised from tree. Slow growth and lack of axillary shoot development on the micrografts was noticeable when the micrografts were cultured on hormone-free and germination medium.
In vitro micrografted plantlets were successfully weaned and no problems were encountered with the establishment of micrografted plants in vivo. Pistacia vera L. There has been some success in widely in the Mediterranean regions of Europe and establishment of mature pistachios by direct organo- North Africa, the Middle East, China and California.
Until recently, seeds were , ; Onay, a, b. There is only one rarely obtained from controlled crosses between se- report of axillary shoot formation and plantlet re- lected clones. Since pistachio is a natural outbreeder generation from explants of mature material Onay, much genetic variation exists in seedling populations.
The first attempts to rejuvenate mature ma- Because of difficulties in rooting cuttings of pista- terials by micrografting in vitro mature scion shoot- chio cultivars Al Barazi and Schwaba, grafting tips onto juvenile P. However, very slow growth of method for vegetative propagation. Micrografting was investigated in vitro as well Hansman and Owens y de Novoa , Barghchi as in vivo by Abousalim and Mantell but re- and Alderson , Onay and Jeffree and juvenation was not reported when mature trees were Onay The initiation of cultures from ma- used as scions.
The grafted plants were cultured used to raise in vitro seedlings for rootstocks. Mature on fresh MS medium containing 2. After partial The effect of time of the year on micrografting was in- trimming of the cotyledons, the embryos were cultured vestigated for February, April, June, October, August in Magenta GA 7 vessels Chicago Corp. Scions were 4—6 mm sterile shoot tips.
Success- east Turkey. Seven to 9 weeks after tion. The regenerated shoots from the forced shoot tips micrografting, established plants were transferred to were micropropagated, and subcultured every 3 weeks soil.
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