WO2014112997A1 - Traitement des cultures à naphtalène alkyle inférieur pour modifier le cycle cellulaire et la régulation d'eau - Google Patents

Traitement des cultures à naphtalène alkyle inférieur pour modifier le cycle cellulaire et la régulation d'eau Download PDF

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Publication number
WO2014112997A1
WO2014112997A1 PCT/US2013/021740 US2013021740W WO2014112997A1 WO 2014112997 A1 WO2014112997 A1 WO 2014112997A1 US 2013021740 W US2013021740 W US 2013021740W WO 2014112997 A1 WO2014112997 A1 WO 2014112997A1
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dmn
lower alkyl
alkyl naphthalene
genes
treatment
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PCT/US2013/021740
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English (en)
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Michael Campbell
Addie Waxman
James Zalewski
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1,4 Group, Inc.
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Priority to PCT/US2013/021740 priority Critical patent/WO2014112997A1/fr
Publication of WO2014112997A1 publication Critical patent/WO2014112997A1/fr

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    • A—HUMAN NECESSITIES
    • A01—AGRICULTURE; FORESTRY; ANIMAL HUSBANDRY; HUNTING; TRAPPING; FISHING
    • A01N—PRESERVATION OF BODIES OF HUMANS OR ANIMALS OR PLANTS OR PARTS THEREOF; BIOCIDES, e.g. AS DISINFECTANTS, AS PESTICIDES OR AS HERBICIDES; PEST REPELLANTS OR ATTRACTANTS; PLANT GROWTH REGULATORS
    • A01N27/00—Biocides, pest repellants or attractants, or plant growth regulators containing hydrocarbons

Definitions

  • Potato is the fourth largest agricultural commodity on the world market and a sizable portion of the yearly harvest is placed in storage. Prolongation of the storage of harvested potato tubers is predicated on the ability to prevent sprouting of tuber meristems. Premature sprouting results in conversion of stored starch to sugars, resulting in plant material unsuitable for either the fresh market or for processing. Thus, suppression of sprouting is key to the maintenance of the potato harvest for commercial purposes.
  • potato tubers enter a state of endodormancy. This natural process of endodormancy is a quantitative trait, controlled by a suite of genes, which results in variability in the length of the dormant between potato cultivars, and at times between different harvests of the same cultivar. The onset and duration of endodormancy in potato is linked to abscisic acid (ABA) and ethylene levels in tuber tissue. Genetic analysis has previously shown that genes linked to increased ABA levels prolong the
  • Transcriptional profiling using oligo arrays developed for potato, has been previously used to ascertain gene responses during tuber initiation and in reactivated meristems following treatment with the phytohormones cytokinin and gibberellin.
  • cDNA microarrays have been used to demonstrate transcriptional changes in potato tubers meristems in response to dormancy status.
  • CIPC chlorpropham
  • the naturally occurring compound 1 ,4-dimethylnaphthalene (DMN) was isolated from potato tubers and was shown to prevent premature sprouting (Beveridge et al. 1981). DMN has demonstrated the ability to reversibly prevent sprouting, making it attractive as a sprout inhibitor on seed tubers (Pinhero et al. 2009). The mode of action of DMN is unknown but recent experiments have determined that CIPC and DMN do not function through a similar mechanism of action and neither sprout inhibitor functions by prolongation of innate dormancy (Campbell et al. 2010).
  • a particular embodiment relates to a method of maintaining hydration in crops, plants, or produce containing osmotin-expressing genes, which includes treating the crop, plant, or produce in a post-harvested state with an effective dosage of a lower alkyl naphthalene sufficient to turn-on the osmotin-expressing genes.
  • Another embodiment relates to a method of maintaining hydration of a plant (or product of said plant) that includes treatment of said plant or product thereof with an effective dosage of a lower alkyl naphthalene, wherein the plant has at least one identified gene.
  • Yet another embodiment is drawn to a method of maintaining hydration in a post-harvested, plant product, which method includes treating the plant product in a hydrated state with an effective dosage of a lower alkyl naphthalene.
  • An alternative embodiment is drawn to a method of treating plant bulbs to minimize premature sprouting in the same.
  • the method includes treating the bulbs with an effective amount of a lower alkyl naphthalene sufficient to activate genes associated with endodormancy.
  • FIG. 1 is a diagram of the treatment regimes for potato tissues treated with DMN according to a particular embodiment
  • FIG. 2 is a Venn diagram showing transcript changes between meristems isolated from various treated and control groups of tubers;
  • FIG. 3 is a QT-PCR analysis of potato meristem transcripts isolated from potatoes treated with DMN.
  • FIG. 4 is a schematic illustrating a possible mechanism for KRP1 and KRP2 in cell cycle repression following DMN exposure.
  • the instant disclosure relates to identification of certain genes in potatoes, which are activated by treatment of said potatoes with a preselected dosage of a lower alkyl naphthalene, especially 1 ,4 dimethyl naphthalene.
  • a lower alkyl naphthalene especially 1 ,4 dimethyl naphthalene.
  • the activation of these genes produces a dormancy-like state in said potatoes.
  • up-regulated genes associated with dormancy extension provides a useful technique in extending dormancy of various plants and produce containing said genes by treatment with a lower alkyl naphthalene.
  • Such plants and produce may include those intended for human or animal consumption, but also to others such as iris bulbs, tulip bulbs, gladiola bulbs, and other flowering plants such as daffodils, hyacinths and the like which are grown from bulbs. It is desirable that iris bulbs, tulip bulbs and the like do not prematurely sprout during storage and/or shipment.
  • the advantage of treatment with 1 ,4 DMN e.g., is that it does not adversely affect the later sprouting and plant growth from such treated bulbs.
  • Root vegetables such as carrots, radishes, turnips, beets, rutabaga, onions, shallots, and the like, contain the same genes, which are up-regulated in potatoes treated with 1,4 DMN, for example.
  • treatment of such root vegetables in accordance with the techniques of the instant invention will extend the period of dormancy.
  • a further advantage of such treatment is that resistance to desiccation will be improved by up-regulation of genes associated with water retention and drought assistance.
  • Potato tubers were treated with DMN.
  • Transcriptional profiling via oligo array analysis, was used to determine genes that exhibit altered expression in response to DMN.
  • Field-grown certified Russet Burbank seed tubers were obtained from a commercial grower shortly after harvest. The tubers were allowed to wound-heal for two weeks at room temperature in the dark. The tubers were then transferred to cold (3-4°C) storage. The tubers were periodically evaluated for dormancy status. Only non-dormant tubers (100% sprouting after two weeks at 20°C) were treated with 1 ,4 DMN.
  • Transcriptional profiling using oligo arrays developed for potato, has been used to examine changes in gene expression during tuber initiation (Kloosterman, et al. 2008) and in reactivated meristems following treatment with the phytohormones cytokinin and gibberellin (Hartmann, et al. 201 1).
  • cD A microarrays have been used to demonstrate transcriptional changes in potato tubers meristems in response to dormancy status (Campbell et al. 2008).
  • potato tubers were treated with DMN and transcriptional profiling, via oligo array analysis, was used to identify genes that exhibit altered expression in response to DMN.
  • DMN was shown to be effective on sprouting root and tuber crops, including but not limited to, beet, carrot, cassava, dasheen (taro), ginger, ginseng, horseradish, parsnip, pototato, sweet potato, turnip, and yam.
  • DMN was also shown to be effective on sprouting bulb crops, including but not limited to, garlic, leek, onion, and shallot.
  • DMN was also shown to be effective on sprouting ornamentals, such as flowering bulbs.
  • Field- grown certified Russet Burbank seed tubers were obtained from a commercial grower shortly after harvest. The tubers were allowed to wound-heal for two weeks at room temperature in the dark. The tubers were transferred to cold (3-4°C) storage. The tubers were periodically evaluated for dormancy status. Only non-dormant tubers (100% sprouting after two weeks at 20°C) were used in these studies. Thus, the dormancy status of these meristems can be considered to be ecodormant and held in the growth arrested state due to low temperatures. A diagram of the experimental design is found in Figure 1.
  • tubers On the day of treatment, tubers were transferred from 3° to 20°C and were immediately washed in running tap water. After drying at room temperature in the dark, the tubers were placed into 4 L airtight containers. A beaker containing about 5g of activated charcoal was placed in the chamber containing the control tubers prior to sealing (Control). DMN (0.15 mL liquid) was placed on cotton wool in a beaker and sealed in the treatment chamber (DMN3days). The chambers were incubated in the dark at 20°C for three days. At this time tubers were removed and the lateral meristems were isolated using a curette with the aid of a dissecting microscope.
  • Freshly excised meristems were washed with deionized water and groups of 20 buds were incubated on 1 mL of buffer (10 mM Mes/KOH; pH 5.7) containing 0.9 ⁇ of 3 H-thymidine (60 ⁇ / ⁇ ; American Radiolabeled Chemicals Inc., St. Louis, MO). The meristems were incubated on an oscillating shaker (100 rpm) at room temperature in the dark. After three hours, meristems were removed from the incubation media, washed extensively with deionized water followed by two washes with incubation buffer containing 5 mM unlabeled thymidine, blotted dry, frozen in liquid nitrogen, and stored at -80°C.
  • the frozen meristems were homogenized in 1 mL ice-cold 10% (w/v) TCA. After standing on ice for 3-4 hours, the samples were vigorously mixed. An aliquot was removed and placed in a scintillation vial to determine total uptake. A second equal aliquot was removed and placed on a microfiber filter (GF/C; Whatman). The filter was sequentially washed under vacuum with 2 x 20 ml 5% (w/v) TCA and 20 ml 95% (v/v) ethanol (both at 4°C). The washed filter was then placed in a scintillation vial to determine total incorporation into TCA-precipitable material (DNA).
  • DNA TCA-precipitable material
  • Total RNA (500ng) was denatured in the presence of a T7 promoter primer and cDNA was synthesized using reverse transcriptase.
  • the cDNA was used as a template for in vitro transcription by synthesis with T7RNA polymerase, which amplified target material and incorporated cyanine 3-labeled CTP.
  • the labeled cRNA was purified using spin columns and quantified using a spectrophotometer.
  • a sample of 1.65 ⁇ g of cyanine 3-labeled linearly amplified cRNA was hybridized to an Agilent 44K 60- mer-oligo microarray that was developed by the Potato Oligo Chip Initiative (POCI) (Kloosterman et al. 2008). Probe generation and hybridization to the array were conducted through a contract to Gene Togic (www.genelogic.com).
  • POCI Potato Oligo Chip Initiative
  • Analysis of the microarray was conducted in two stages; total array data was examined using a linear model and the software limma, and pathway analysis was conducted by linking the potato sequences to the Arabidopsis thaliana. Analysis of the entire microarray data set was accomplished using R and the limma package (Smyth 2005), or GeneMaths XT (Applied Maths, Inc.). The data was corrected for background and normalized between arrays using a loess method. Comparisons between treatments was established using an empirical Bayes method (Smyth 2004). The use of limma for analysis of oligo arrays was accomplished by analyzing data from a single emission channel.
  • Pathway analysis was accomplished by using tBLASTx (Altschul et al. 1990) to search the A. thaliana genome for homologs. Transcripts that exhibited an e-value of less than 1.0e-5 were assigned a function according to the described function for the A. thaliana homolog.
  • the array data was transformed to log base 2 and normalized between arrays using default parameters in GeneMathsXT software package (www.applied-maths.com). Probes had to have hybridization intensities of at least 2 standard deviations greater than background in one or more treatments to be included in the dataset.
  • Pathway Studio 8.0 www.ariadnegenomics.com was utilized to determine metabolic changes that were associated with DMN treated for three days compared to controls.
  • a set of transcripts was selected for further analysis to confirm the microarray data and to determine the possible cell cycle position of tubers treated with DMN.
  • Gene names and primers sequences used for QT-PCR are found in Table 1.
  • the oligo nucleotide sequences spotted on the array were too short for adequate primer design. Therefore, primers were determined by isolating a longer DNA sequence from the DFCI-Potato Gene Index corresponding to the microarray oligonucleotide and searching that sequence for primers using the Applied
  • cDNA template for QT-PCR was prepared from mRNA isolated from potato meristems treated with DMN for three days as described above. PCR reactions were run on a StepOneTM Real Time System and analyzed using Step One Software v2.0
  • 3H-thymidine incorporation was used to determine the effects of DMN treatment on cell division.
  • thymidine incorporation was reduced to 6.5 to 6.7% of the label taken up (Table 1).
  • the level of thymidine incorporation after DMN exposure was similar to the levels previously reported for endodormant potato meristems (Campbell et al. 1996).
  • Microarray analysis 34444 probes were considered expressed in at least one treatment; of these, 13125 were differentially expressed in at least one treatment (Benjamini Hochberg modified p value ⁇ 0.005) (Supplemental Table 1 ). These large numbers of differentially expressed genes suggest major modifications to the transcriptome for all treatments.
  • Pathway analysis highlights the similarities between meristems from eco- dormant and DMN-treated tubers in comparison to meristems incubated at 20°C for three days.
  • 294 ontologies were associated with genes that are preferentially expressed following incubation at 20°C compared to eco-dormant tissues
  • 251 ontologies were associated with genes that are preferentially expressed following incubation at 20°C compared to DMN-treated tubers. More than half of the ontologies were common between the comparisons (Supplemental Tables 2, 3).
  • QT-PCR was used to both confirm microarray data and to examine additional transcripts that were either not present on the POCI array or were eliminated from analysis after normalization and removal of systematic errors.
  • DMN suppression of transcription for the potato homologs of the cyclins CYCD1 , CYCD2, CYCD3 would limit protein components for the CDK/cyclin partners that drive initiation of cell division in potato meristems that result in sprouting (Van Leene et al. 2010).
  • CKIs cyclin-dependent kinase inhibitors
  • CDKA forms a complex with CYCD3.
  • the activity of this complex is inhibited by the CKIs KRP1 and KRP2, thereby repressing cell cycle advancement (Menges, de Jager et al. 2005).
  • KRP1 and KRP2 were found to have increased expression in tissues treated with DMN ( Figure 3). This up-regulation corresponds with the down-regulation of CYCD3, suggesting that DMN prevents sprouting by inhibiting the formation of the CDKA/CYCD3 complex, thus resulting in a Gl cell cycle block.
  • CDKB 1 indirectly increases the activity of the CDKA/CYCD3 by phosphorylation and inhibition of KRP2 (Verkes et al. 2005).
  • QT-PCR analysis indicates that DMN decreases the transcription of CDKB 1 , suggesting that the Gl block induced by DMN is also a result of the maintenance of KRP2 inhibition of the CDK/CYCD3 complex.
  • Deoxyuridine triphosphatase (dUTPase) has been shown to be an early marker of dormancy termination prior to entry into the S-phase of the cell cycle (Senning et al. 2010). Exposure to DMN resulted in a decrease of UTPase, which again suggests DMN is maintaining a cell cycle block prior to S-phase.
  • Figure 4 outlines the interaction of some of the proteins associated with cell cycle regulation.
  • the increased expression of KRP1 and KRP2 suggests two possible positions for cell cycle repression, one in the G 1 and the other in the S-phase.
  • the decreased expression of dUTPase suggests that a DMN-induced-block via KRP expression is more likely occurring during Gl because the expression of dUTPase occurs prior to S-phase entry. This hypothesis is also supported by the low levels of thymidine incorporation.
  • Nucleoside diphosphate kinase is down regulated by exposure to DMN (Table 2).
  • the function of the NDPK protein is to regulate the cellular levels of nucleotides and it is linked to organogenesis in animals (Lakso et al. 1992), cell proliferation (Keim et al. 1992), and response to salt stress in plants (Kawasaki et al. 2001).
  • NDPK expression is directly associated with response to stress and reactive oxygen and the alteration of the mitogen activated protein kinases MPK3 and MPK6 (Moon et al. 2003).
  • HOG1 is classified as a member of the mitogen-activated protein kinase (MAPK) family.
  • MAPK mitogen-activated protein kinase
  • HOG1 functions as a MAPK that interacts within the signal transduction cascade linking growth signals to cell division regulation and it also functions in the sensing system for the regulation of osmotic potential (Brewster et al. 1993; Gustin et al. 1998).
  • HOG J expression results in a shift in transcriptional profiles, which results in a large change in gene expression in response to changes in osmolarity (O'Rourke and Herskowitz 2004).
  • Exposure of potato tubers to DMN results in an increase in a HOG1 homolog.
  • the HOG1 cascade in yeast results in growth suppression by down regulation of the
  • Gl/S-phase cyclins CM and Cln2 (Clotet and Posas 2007).
  • a tBLASTx analysis of the Arabidopsis thaliana genome using the Clnl gene from Saccharomyces cerevisiae finds that the gene CYCAl is the closest homolog.
  • DMN treatment which results in elevated expression of the HOG1 homologs MPK4 and MPK6, also results in a decrease in the Clnl homolog CYCAl. This suggests that DMN growth arrest may be due to a decrease in the Gl/S-phase cyclins through the increase activity of MPK4/PK6. This information translates to many other crops and plants having the same genes wherein treatment with a lower alkyl naphthalene provides similar
  • Berries are a fragile, freshly harvested crop which maintain improved hydration and, ergo, a longer, healthier shelf life when treated with 1 ,4 DMN, for example.
  • leafy vegetables such as lettuce, cabbage, spinach and the like.
  • leafy crops such as freshly cut flowers may also be advantageously treated to maintain freshness for a longer period of time.
  • bulb-type crops such as onions, garlic, tulip, iris, gladiola and the like may be advantageously treated to extend an endodormant-like state to minimize premature sprouting and loss of turgidity.
  • Arabadopsis would contain similar genes based on the widely accepted biological principle of evolutionary conservation of genes. This further implies that many/most other plant species would respond similarly to exposure to lower alkyl napthalenes. In some instances, both types of genes are present and in other crops the presence of one group or the other may be less important for the intended result by treatment with a lower alkyl naphthalene.
  • Genes associated with endodormancy include those identified in Table 2 hereof.
  • Genes associated with drought resistance include those identified in Table 3 hereof.
  • a further advantage of the instant invention is that the effect of lower alkyl naphthalenes on the above-identified genes persists for a period of time after the chemical is no longer detectable by sophisticated chemical analysis. Thus, such chemicals are largely absent from treated edible plants on crops at the time of human consumption.
  • up-regulated transcripts and represent those that encode for proteins that function in water regulation, response to salt and water stress.
  • F11C10.33 T15N1.10, ATPSK3, UBC19, 0L12, GIF3, FASl, ATHB-2, PKL, ARP6, FAS2, MSI1,NRP2, ELOl, DDL, ABOl, UBP15, EL03, RPL5A
  • Arabidopsis genes orthologous to the potato transcripts that decrease in response to DMN treatment are a subset of down-regulated transcripts and represent those that encode for proteins that function in cell cycle regulation, DNA replication, and cell proliferation.
  • PIP2B T8P21.32, F18C1.3, AT3G06760, PIP1;4, ANNAT7, F2P16.10, SIP1, MNC17.13, F24B9.8, T24D18.16, ANN5, ATHVA22A, HIS1-3, RD19,

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Abstract

La présente invention se rapporte à l'identification de certains gènes dans les pommes de terre, qui sont activés par traitement desdites pommes de terre avec une dose présélectionnée d'un naphtalène alkyle inférieur, en particulier le 1,4 diméthyl-naphtalène. L'activation de ces gènes produit un état dormant dans lesdites pommes de terre. Des réalisations particulières ont trait à l'identification de gènes de résistance à la déshydratation dans lesdites pommes de terre, qui ont été également activé par ledit traitement avec un naphtalène alkyle inférieur, tel que le 1,4 diméthyl-naphtalène.
PCT/US2013/021740 2013-01-16 2013-01-16 Traitement des cultures à naphtalène alkyle inférieur pour modifier le cycle cellulaire et la régulation d'eau WO2014112997A1 (fr)

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CN105296443A (zh) * 2015-12-07 2016-02-03 北京市农林科学院 一种植物抗旱、耐盐相关蛋白EeSAPK7及其编码基因和应用
CN105481959A (zh) * 2016-01-27 2016-04-13 清华大学 一种通过下调PAB2和PAB4提高植物对NaCl耐受性的方法
CN107475210A (zh) * 2017-09-07 2017-12-15 四川农业大学 一种水稻白叶枯病抗性相关基因OsABA2及其应用
CN107760656A (zh) * 2016-08-23 2018-03-06 江苏省中国科学院植物研究所 玉米胁迫相关蛋白激酶sapk10在渗透胁迫中的应用
WO2019210245A1 (fr) * 2018-04-26 2019-10-31 1,4Group, Inc. Procédé de lutte contre le fusarium sur la pomme de terre à l'aide de naphtalène à alkyle inférieur
CN114672497A (zh) * 2022-04-25 2022-06-28 山东农业大学 马铃薯StPHB3基因在提高马铃薯块茎品质中的应用
CN114752604A (zh) * 2022-04-25 2022-07-15 山东农业大学 一种增强小麦生物抗性和产量的基因TaPIP2;8及其应用

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CN105296443A (zh) * 2015-12-07 2016-02-03 北京市农林科学院 一种植物抗旱、耐盐相关蛋白EeSAPK7及其编码基因和应用
CN105296443B (zh) * 2015-12-07 2019-02-05 北京市农林科学院 一种植物抗旱、耐盐相关蛋白EeSAPK7及其编码基因和应用
CN105481959A (zh) * 2016-01-27 2016-04-13 清华大学 一种通过下调PAB2和PAB4提高植物对NaCl耐受性的方法
CN105481959B (zh) * 2016-01-27 2019-02-05 清华大学 一种通过下调PAB2和PAB4提高植物对NaCl耐受性的方法
CN107760656A (zh) * 2016-08-23 2018-03-06 江苏省中国科学院植物研究所 玉米胁迫相关蛋白激酶sapk10在渗透胁迫中的应用
CN107475210A (zh) * 2017-09-07 2017-12-15 四川农业大学 一种水稻白叶枯病抗性相关基因OsABA2及其应用
WO2019210245A1 (fr) * 2018-04-26 2019-10-31 1,4Group, Inc. Procédé de lutte contre le fusarium sur la pomme de terre à l'aide de naphtalène à alkyle inférieur
CN112469276A (zh) * 2018-04-26 2021-03-09 1,4集团公司 使用低级烷基萘防治马铃薯上的镰孢菌的方法
CN114672497A (zh) * 2022-04-25 2022-06-28 山东农业大学 马铃薯StPHB3基因在提高马铃薯块茎品质中的应用
CN114752604A (zh) * 2022-04-25 2022-07-15 山东农业大学 一种增强小麦生物抗性和产量的基因TaPIP2;8及其应用
CN114752604B (zh) * 2022-04-25 2023-07-14 山东农业大学 一种增强小麦生物抗性和产量的基因TaPIP2;8及其应用

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