Ferox Seeds Research Articles

Treatment of Diabetes Nephropathy in Mice by Germinating Seeds of Euryale ferox through Improving Oxidative Stress.

Diabetes can cause severe kidney disease. Euryale ferox seeds (Gordon Euryale) have known antioxidant, hypoglycemic, and renal protection effects. Methanol extracts of Gordon Euryale were produced from ungerminated and germinated seeds. The effect of germination on polyphenol and flavonoid content was investigated by Liquid chromatography-tandem mass spectrometry (LC-MS/MS) analysis. Three doses of ungerminated seed extract (EKE) and germinated seed extract (GEKE) were administered to diabetic mice by gavage to explore the treatment-dependent improvement of oxidative stress, metabolic disorder, and kidney disease. Seed germination led to a 1.7 times increase in total phenol content in the extract, and the flavonoid content was increased by 1.9 times. Germination greatly increased the contents of 29 polyphenols and 1 terpenoid. At the same dose, GEKE more strongly improved hyperglycemia, abnormal lipid metabolism, and renal tissue lesions (as confirmed by histology) in the diabetic mice than EKE did. In diabetic mice receiving treatment, kidney microalbunminuria (ALB), blood urea nitrogen (BUN), serum creatinine (Scr), malondialdehyde (MDA), and glutathione (GSH) were all decreased, while activity of catalase (CAT), superoxide dismutase (SOD), and serum total antioxidant capacity (T-AOC) were increased. Both EKE and GEKE can improve diabetes and kidney disease by improving hyperglycemia, oxidative stress, and kidney physiological indicators and regulating the Keap1/Nrf2/HO-1 and AMPK/mTOR pathways. However, in both pathways, GEKE is more effective. The purpose of this study was to explore the effects of GEKE and EKE treatment on antioxidant defense and metabolic capacity of diabetic animals. Germination provides a suitable strategy to improve the medicinal value of these natural plant-based products.

Qualitative Phytochemical Analysis of Ethanolic Extracts of Hibiscus laevis Flowers and Euryale ferox Seeds

Qualitative Phytochemical Analysis of Ethanolic Extracts of Hibiscus laevis Flowers and Euryale ferox Seeds

Structure and physicochemical properties of low digestible Euryale ferox Salisb. seed starch.

Euryale ferox Salisb. is widely grown in China and Southeast Asia as a grain crop and medicinal plant. The composition, morphology, structure, physicochemical properties, thermal properties, and in vitro digestibility of North Euryale ferox seeds starch (NEFS), hybrid Euryale ferox seeds starch (HEFS), and South Euryale ferox seeds starch (SEFS) were studied. Of the varieties that were studied, the amylose content of NEFS (23.03%) was the highest. Starch granules of each variety were smooth, sharp, small, and had an average diameter of 2μm. All three varieties were A-type crystals with crystallinity ranging from 26.42% to 28.17%. The degree of double helix and the short-range order ranged from 1.9006 to 2.5324 and 1.4294 to 1.6006, respectively. The high proportion of C1 region in NEFS (17.74%) and HEFS (17.66%) were found. Thermodynamic properties inNorth Euryale ferox seeds included the highest onset temperature (To ) (71.43 °C), peak temperature (Tp ) (76.60 °C), conclusion temperature (Tc ) (82.77 °C), enthalpy of gelatinization (ΔH) (12.64 J g-1 ), and peak viscosity (1514 mPa·s). All three varieties maintained a low level of in vitro digestibility, with the highest resistant starch (RS) content (29.57%), the lowest rapidly digestible starch (RDS) content (27.07%), and the slowest hydrolysis kinetic constant (0.0303) in NEFS. The results revealed that the low digestibility of NEFS was attributable to compact granules, high crystallinity, high degree of order, and strong thermal stability. These digestive, physicochemical, and thermodynamic properties provide information for the future application of Euryale ferox seed starch in the food industry. © 2022 Society of Chemical Industry.

Effect of Thermal Treatment on the Physicochemical, Ultrastructural, and Antioxidant Characteristics of Euryale ferox Seeds and Flour.

Euryale ferox seeds (EFS) were less gelatinized, preventing the release of nutrients and functional compounds, resulting in limited applications in meals and the food industry. Nutraceutical importance of EFS includes starch, protein, lipids, 20 amino acids, minerals, and vitamins (C, E, and beta carotene). This study aimed to evaluate the effect of three different thermal treatments on EFS’s physicochemical and nutritional properties and expected to improve its applicability. The results showed that the bulk density, thousand-grain weight, and hardness of thermal treated EFS were significantly decreased (p < 0.05), whereas the maximum decrease was observed in the industrial infrared heating-assisted fluidized bed (IHFH) treatment. Meanwhile, there were more crevices, fissures, and heightened porous structures in EFS between the pericarp and episperm and the endosperm after heat treatment, which facilitated grinding and water absorption. Notably, EFS’s water and oil absorption capacities increased significantly (p < 0.05) with microwave and IHFH treatments. EFS ground’s solubility into powder was increased significantly with thermal treatment (p < 0.05). Furthermore, the functional properties of TPC, TFC, DPPH radical scavenging activity, and reducing power were significantly increased (p < 0.05). In general, the changes in the physicochemical properties of EFS and increased bioactivity were caused by microwave and IHFH treatments. Hence, it might improve the food value of EFS while providing valuable information to researchers and food manufacturers.

EfABI4 Transcription Factor Is Involved in the Regulation of Starch Biosynthesis in Euryale ferox Salisb Seeds.

Starch is the final product of photosynthesis and the main storage form in plants. Studies have shown that there is a close synergistic regulatory relationship between ABA signal transduction and starch biosynthesis. In this study, we employed RNA sequencing (RNA-Seq) to investigate transcriptomic changes of the Euryale ferox seeds treated by exogenous ABA. The differentially expressed genes engaged in the “Starch and sucrose” and “TCA cycle” pathway. Furthermore, the key transcription factor EfABI4 in ABA signaling pathway and the key genes of starch biosynthesis (EfDBE1, EfSBE2, EfSS1, EfSS2, EfSS3, EfSS4 and EfGBSS1) were significantly up-regulated. Further, the Euryale ferox plant was treated with ABA, it was found that the total starch content of Euryale ferox seeds at different development stages was significantly higher than that of the control, and the key genes of starch synthesis in Euryale ferox seeds were also significantly up-regulated. Finally, yeast one-hybrid and dual luciferase assay proved that EfABI4 can promote the expression of EfSS1 by directly binding to its promoter. Subcellular localization results showed that EfABI4 protein was located at the nucleus and EfSS1 protein was located in the cytomembrane. These findings revealed that ABA promotes starch synthesis and accumulation by mediating EfABI4 to directly promote EfSS1 gene expression, which is helpful for understanding starch synthesis in seeds.

Proteomic analysis of Euryale ferox Salisb seeds at different developmental stages

The development of plant seeds is accompanied by changes in their internal substances. The edible part of E. ferox is the seed, and starch and flavonoids are the storage substances and functional substances in E. ferox seeds respectively. Herein, four time points of seed development, including after flowering T10 (10 days), T20 (20 days), T30 (30 days) and T40 (40 days), were investigated by using iTRAQ technology. A total of 2809 differential proteins were identified. The enrichment analysis of differential proteins found that they were mainly enriched in starch synthesis pathways and flavonoid biosynthesis pathways. The key candidate enzymes for starch synthesis, APS (c54069), APL (c55730), SBE (c56416), SSS (c54912) and GBSS (c53181), were identified. At the same time,PAL (c50934), CHS (c49212), F3H (c35949) and ANS (c54610) may be key enzymes in flavonoid biosynthesis. In addition, the ABA signal transduction pathway was analyzed and it was identified that PYL3 (c54854) and ABI5 (c56122) are up-regulated from T10 to T40, and it is speculated that they play an important regulatory role in the development of E. ferox seeds. Together, these results reveals the dynamic changes during the development of E. ferox seeds, which will provide guidance for the study of the molecular mechanism of starch and flavonoids.

Digestive and Physicochemical Properties of Small Granular Starch from Euryale ferox Seeds Growing in Yugan of China

Digestive and Physicochemical Properties of Small Granular Starch from Euryale ferox Seeds Growing in Yugan of China

A review on Euryale Ferox: Daily dietary regime for Covid -19 patients

In recent pandemic of Covid-19 i.e. Corona Virus disease, patients who are affected by this disease shows symptoms related to different system of body in which most commonly affected parts are lungs which usually results in mortality of patients. It has also seen that older adults and those having COPD, Heart Disease, Diabetes, Chronic Kidney Disease, and obesity are at increased risk for severe COVID -19 requiring hospitalization 1. This Covid 19 can also produce abnormally high blood sugar levels, even in people without diagnosed diabetes. It also causes fluctuation in Blood Pressure by affecting ACE and RAAS system. Fox Nut or Euryale Ferox is rich in protein, fibres and carbohydrates. The Euryale ferox seeds is a grain dish good for baby and also for an old man to prolong life. It has characteristics of strengthening the spleen, nourishing the stomach, supplementing the kidney 2. Thus, makhana can be used as nutraceutical supplement for Covid -19 Patients. In a daily diet, it is also believed to increase humoral immunity and cell mediated immunity to some extent. The Euryale ferox seeds and glutinous rice are used in 1:3 ratio to prepare rice dumplings in China. These act as super natural food in diet therapy 11. Euryale ferox has many bioactive constituents which performs anti-oxidant activity, anti-diabetic activity, Anti-hyper lipidemic activity, Hepatoprotective activity, anti-cytotoxicity, cardioprotective, anti-depressant activity and much more. Thus, Euryale ferox could be an ideal supplement for Covid -19 patients.

Metabolomics and transcriptome analysis of the biosynthesis mechanism of flavonoids in the seeds of Euryale ferox Salisb at different developmental stages.

Flavonoids belong to polyphenolic compounds, which are widely distributed in plants and have rich functions. Euryale ferox Salisb is an important medicinal and edible homologous plant, and flavonoids are its main functional substances. However, the biosynthesis mechanism of flavonoids in E. ferox is still poorly understood. To explore the dynamic changes of flavonoid biosynthesis during the development of E. ferox seeds, the targeted flavonoid metabolome was determined. A total of 129 kinds of flavonoid metabolites were characterized in the seeds of E. ferox, including 11 flavanones, 8 dihydroflavanols, 16 flavanols, 29 flavones, 3 isoflavones, 12 anthocyanins, 29 flavonols, 6 flavonoid carbonosides, 3 chalcones and 13 proanthocyanidins. The relative content of flavonoid metabolites accumulated continuously during the development of E. ferox seeds, and reached the highest at T30. In transcriptome, the expression of key genes in the flavonoid pathway, such as PAL, CHS, F3H, FLS, ANS, was highest in T30, which was consistent with the trend of metabolites. Six candidate transcription factors (R2R3MYBs and bHLHs) may affect the biosynthesis of flavonoids by regulating the expression of structural genes. Furthermore, transcriptome analysis and exogenous ABA and SA treatment demonstrated that ABA (PYR1, PP2Cs, SnRK2s) and SA (NPR1) are involved in the positive regulation of flavonoid biosynthesis. This study clarified the differential changes of flavonoid metabolites during the development of E. ferox seeds, confirmed that ABA and SA promote the synthesis of flavonoids, and found key candidate genes that are involved in the regulation of ABA and SA in the positive regulation of flavonoid biosynthesis.

Transcriptome sequencing and analysis during seed growth and development in Euryale ferox Salisb

BackgroundEuryale ferox Salisb., an annual aquatic plant, is the only species in the genus Euryale in the Nymphaeaceae. Seeds of E. ferox are a nutritious food and also used in traditional Chinese medicine (Qian Shi in Mandarin). The molecular events that occurred during seed development in E. ferox have not yet been characterized. In this study, we performed transcriptomic analysis of four developmental stages (T1, T2, T3, and T4) in E. ferox seeds with three biological replicates per developmental stage to understand the physiological and biochemical processes during E. ferox seeds development.Results313,844,425 clean reads were assembled into 160,107 transcripts and 85,006 unigenes with N50 lengths of 2052 bp and 1399 bp, respectively. The unigenes were annotated using five public databases (NR, COG, Swiss-Prot, KEGG, and GO). In the KEGG database, all of the unigenes were assigned to 127 pathways, of which phenylpropanoid biosynthesis was associated with the synthesis of secondary metabolites during E. ferox seed growth and development. Phenylalanine ammonia-lyase (PAL) as the first key enzyme catalyzed the conversion of phenylalanine to trans-cinnamic acid, then was related to the synthesis of flavonoids, lignins and alkaloid. The expression of PAL1 reached its peak at T3 stage, followed by a slight decrease at T4 stage. Cytochrome P450 (P450), encoded by CYP84A1 (which also called ferulate-5-hydroxylase (F5H) in Arabidopsis), was mainly involved in the biosynthesis of lignins.ConclusionsOur study provides a transcriptomic analysis to better understand the morphological changes and the accumulation of medicinal components during E. ferox seed development. The increasing expression of PAL and P450 encoded genes in phenylpropanoid biosynthesis may promote the maturation of E. ferox seed including size, color, hardness and accumulation of medicinal components.

Physico-fracture Characteristics of Makhana (Euryale ferox) Seeds

ABSTRACTThe fracture characteristics of Makhana (Euryale ferox) seed kernels were determined at moisture contents varying from 10.28 to 15.2% (dry basis), as a function of compressive force, deformation, and energy absorbed. The force required for kernel rupture decreased as the temperature and time of roasting increased while moisture content of the seed decreased. The deformation of the seed at the point of rupture was found to be in the range of 1.69 to 2.95 mm. Energy absorbed per unit volume, sphericity and angle of repose increased linearly with increasing moisture content and were found to be in the range of 310 to 822 mJ, 0.97 to 0.98, and 46 to 50°, respectively.

La Gestión de Recursos Vegetales En Las Poblaciones Prehistóricas de Las Tierras Bajas Del Sureste Del Uruguay: Un Abordaje Multidisciplinar

This paper addresses the diversification of plant resource management practices in the lowlands of southeastern Uruguay starting in the fourth millennium B.P. Our approach includes (a) a reexamination of ethnohistorical sources; (b) a review of lithic and ceramic materials; (c) the presentation of new ethnoarchaeological information; and (d) the analysis of starch grains from grinding stones. The results highlight the importance of the palm Butia odorata as a source of food and useful materials for much of the year. Furthermore, the analysis of both macro- and micro-residues has enabled us to identify the presence of Datura ferox seeds and starch grains from edible plants. While many of these identifications confirm the use of a number of previously reported species, others are new, as is the case with Phaseolus sp., Arachis sp., and Typha domingensis. Continuity in the management of certain plant species and in environmental conditioning is interpreted as an important investment in the long-term cycles of human-environment interactions.

Seed dormancy and germination traits of an endangered aquatic plant species, Euryale ferox Salisb. (Nymphaeaceae)

Populations of Euryale ferox Salisb. have declined throughout its global range because of habitat loss and degradation. The present study aimed to evaluate the influence of storage temperature (4 and 20°C), storage period (0, 90, 180, 270, and 360 days), light condition (light and darkness), germination temperature (constant 10, 15, 20, 25, and 30°C), and seed size (two classes from 0.5cm to less than 1.2cm, and from 1.2cm to less than 1.6cm) on germination of E. ferox seeds. Our results indicated that seeds were dormant when shed and 4°C stratification promoted germination more effectively than 20°C. After stratification at 4°C, the germination frequency of the larger seeds peaked at 90 days’ stratification, whereas the smaller seeds maintained high germination frequencies up to 180 days’ stratification. The different germination responses between smaller and larger seeds may reflect a difference in the rate of induction of dormancy in the annual dormancy cycle. Seeds germinated in both light and darkness, which demonstrated that light is not involved in the regulation of E. ferox seed germination. The optimal temperature for germination was 25°C. Light condition×germination temperature interaction caused significantly higher germination frequency at 30°C in light than in darkness, and the opposite trend at 15°C.

Tolerancia a la desecación de semillas de &lt;i&gt;Prosopis ferox&lt;/i&gt; y &lt;i&gt;Pterogyne nitens&lt;/i&gt; (Fabaceae)

The high number of endemisms and species diversity together with the accelerated biodiversity loss by deforestation, especially in North Western Argentina, points out the need to work on species conservation combining ex situ and in situ strategies. The aim of this work was to study the desiccation tolerance in seeds of P ferox and P nitens for long term ex situ conservation at the Germplasm Bank of Native Species (BGEN) of the National University of Salta (Argentina). The fruits were collected from ten individuals in P ferox at the National Park Los Cardones and from two sites (Orán and Rivadavia) for P nitens. Desiccation tolerance was assessed following previous established methodologies. The moisture content (MC) of the seeds was determined by keeping them in oven at 103 degreeC and weighting the samples at different intervals till constant weight. Germination essays were carried out with two treatments (control and scarification), with different seed MC (fresh, 10-12%, 3-5%) and in desiccated seeds (3-5% MC) stored six months at -20 degreeC. The MC in P ferox seeds was 14.2% and 10% in P nitens, for both populations studied. Percentage germination in P ferox was higher in the scarification treatments (<82%). The difference between treatments increased with the reduction in MC and the storage for six months at -20 degreeC. Fresh seeds of P nitens do not need scarification treatment, but it is required with the reduction in MC and storage. Mean germination percentage of desiccated seeds stored six months at -20 degreeC was similar in both populations and greater than 82%.We concluded that both species are probably orthodox because seeds tolerated desiccation to 3-5% and storage for six months at -20 degree C.

Isolation and Identification of Compounds Responsible for Antioxidant Capacity of Euryale ferox Seeds

Euryale ferox seed is consumed medicinally or for food in China. The present study revealed it to contain significant antioxidant activity, which may be associated with its medical applications as a proteinuria inhibitor of diabetic nephropathy. This study resulted in the identification of 3 new sesquineolignans, named euryalins A-C (1-3), and 16 known compounds, which were all first isolated from this plant apart from 5,7,4-trihydroxy-flavanone. The antioxidant potential of the partial isolates was evaluated using the DPPH radical scavenging assay and mesangial cellular assay. Compounds 2, rel-(2α,3β)-7-O-methylcedrusin (4), syringylglycerol-8-O-4-(sinapyl alcohol) ether (5), and (+)-syringaresinol (7) were found to be most active on DPPH assay, whereas compounds 2, 4, 7, (1R,2R,5R,6S)-2-(3,4-dimethoxyphenyl)-6-(3,4-dihydroxyphenyl)-3,7-dioxabicyclo[3.3.0]octane, and buddlenol E could significantly inhibit high glucose-stimulated reactive oxygen species production in mesangial cells. The results suggested that E. ferox seed could be considered as an excellent source of natural antioxidants and is useful in the prevention of diabetic nephropathy.

Development of non-effervescent floating matrix tablets based on Euryale ferox seeds

Development of non-effervescent floating matrix tablets based on Euryale ferox seeds

Studies on Seed Germination, Seedling Growth, and In Vitro Shoot Induction of Aloe ferox Mill., a Commercially Important Species

A study was done to investigate the effects of some physical and chemical factors on growth and development of Aloe ferox ex vitro and in vitro. The effects of light, temperature, and smoke–water on seed germination, ex vitro seedling growth requirements, and effect of germination medium and cytokinins on shoot induction and multiplication in vitro were investigated. The highest germination percentage of A. ferox seeds was recorded between 15 and 30 °C. Seed germination was inhibited at 10 and 35 °C. There was a nonsignificant increase in germination percentage (78%) for seeds that were germinated under constant dark conditions. Smoke–water-treated seeds showed significant improvement in percentage germination (80%) over the control (66%) at 25 °C with a 16-h photoperiod. Seedlings of A. ferox subjected to alternating temperatures (30/15 °C) or being irrigated three times weekly at 25 °C showed better seedling growth. In an in vitro experiment, seedlings germinated in distilled water and one-tenth strength Murashige and Skoog medium had superior shoot induction competence compared with the other germination media. The cytokinins meta-Topolin (mT) and meta-Topolin riboside (mTR) at 5 μm gave significantly higher shoot multiplication rates compared with the control and benzyladenine (BA)-treated plants. A higher abnormality index was recorded for BA-treated plants. The findings of this study will be beneficial for commercial propagation of Aloe ferox.

The very-low-fluence and high-irradiance responses of the phytochromes have antagonistic effects on germination, mannan-degrading activities, and DfGA3ox transcript levels in Datura ferox seeds

Seed germination can be promoted by the modes of action of two of the phytochromes: the low-fluence response (LFR), which is the classical red (R)-far-red (FR) reversible response and the very-low-fluence response (VLFR) that can be saturated by extremely low levels of Pfr, which can be elicited by a saturating FR pulse. The Datura ferox seed population used in this work had acquired the capacity to germinate through a VLFR after pretreatment in a water-saturated atmosphere (WSA) at constant 25 degrees C. After 12 d in WSA germination after a FR pulse was 82%, while it was less than 10% in darkness. It was found that the VLFR of germination is associated with increments in the embryo growth potential (EGP) and in the activity of two enzymes related to the weakening of the micropylar region of the endosperm (ME); endo-beta-mannanase and beta-mannosidase. The FR pulse also significantly stimulated the expression of DfGA3ox, a GA 3beta-hydroxylase, suggesting that the promotion of germination by the VLFR is associated with an increase in the synthesis of active gibberellins. The promotive action of the VLFR on germination is reduced when the FR pulse is immediately followed by a continuous FR treatment for 24 h (FRc). The effect of FRc cannot be reproduced by hourly FR pulses during the same period, showing that the antagonistic effect of FRc is a high-irradiance response (HIR). The action of the HIR in germination is associated with a decrease of both the mannan-degrading enzyme activity and the expression of DfMan in the ME, whereas no changes in the EGP were observed. The HIR also inhibits the accumulation of DfGA3ox in embryos, indicating that its action on germination is mediated, at least in part, through the modulation of active GA contents in seeds. This is the first report of a gene that participates in the VLFR-HIR antagonism in seeds.

The Effect of Euryale Ferox (Makhana), an Herb of Aquatic Origin, on Myocardial Ischemic Reperfusion Injury

Fox nut or gorgon nut (Euryale ferox--Family Nymphaeaceae), popularly known as Makhana, has been widely used in traditional oriental medicine to cure a variety of diseases including kidney problems, chronic diarrhea, excessive leucorrhea and hypofunction of the spleen. Based on the recent studies revealing antioxidant activities of Euryale ferox and its glucosides composition, we sought to determine if Euryale ferox seeds (Makhana) could reduce myocardial ischemic reperfusion injury. Two different models were used: acute model, where isolated rat hearts were preperfused for 15 min with Krebs Henseleit bicarbonate (KHB) buffer containing three different doses of makhana (25, 125 or 250 microg/ml) followed by 30 min of ischemia and 2 h of reperfusion; and chronic model, where rats were given two different doses of makhana (250 and 500 mg/kg/day) for 21 days, after which isolated hearts were subjected to 30 min of ischemia followed by 2 h of reperfusion. In both cases, the hearts of the Makhana treated rats were resistant to ischemic reperfusion injury as evidenced by their improved post-ischemic ventricular function and reduced myocardial infarct size. Antibody array technique was used to identify the cardioprotective proteins. The Makhana-treated hearts had increased amounts of thioredoxin-1 (Trx-1) and thioredoxin-related protein-32 (TRP32) compared to the control hearts. Western blot analysis confirmed increased expression of TRP32 and thioredoxin proteins. In vitro studies revealed that Makhana extracts had potent reactive oxygen species scavenging activities. Taken together, the results of this study demonstrate cardioprotective properties of Makhana and suggest that such cardioprotective properties may be linked with the ability of makhana to induce TRP32 and Trx-1 proteins and to scavenge ROS.

A phytochrome-dependent embryonic factor modulates gibberellin responses in the embryo and micropylar endosperm of Datura ferox seeds

The promotion of germination by phytochrome is associated with extensive changes both in the embryo and in the micropylar region of the endosperm (ME) of Datura ferox seeds. These changes require de novo gibberellins (GAs) biosynthesis in the embryo, the site where the light stimulus is perceived. GAs stimulate embryo growth potential and move to ME, promoting the expression of genes related with weakening. We report here that, in addition, phytochrome stimulates the sensitivity of the seeds to gibberellic acid (GA). The phytochrome-induced signal is produced in the embryo and enhances the stimulus by GA of embryo growth potential (EGP) and the promotion of the expression of proteins thought to participate in ME weakening: endo-beta-mannanase (EC 3.2.1.78), endo-beta-mannosidase (EC 3.2.1.25) and expansin. Our results suggest that the cytokinins may be a component of the embryonic signal. Phytochrome also modulates DfPHOR and DfMYB transcript levels in ME. These genes show a high identity with components of GAs signaling identified in other species. Expression of DfPHOR in the ME is apparently regulated by phytochrome through the supply of GAs from the embryo to ME, whereas DfMYB expression is regulated by an embryonic factor with some of the characteristics of the one that modulates seed sensitivity to GAs.