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Non-Commercial Use Only International Journal of Plant Biology 2014; volume 5:4887 Proteomic analysis to horizontal controls in cotton and soybean.3,4 Paraheliotropism benefits plants by maintain- Correspondence: Savithiry Natarajan, Soybean of the pulvinus, a heliotropic ing high levels of photosynthetic quantum Genomics and Improvement Laboratory, ARS- tissue, in Glycine max yield under stressed conditions and reducing USDA, 10300 Baltimore Avenue, Beltsville, MD UVB radiation levels.5,6 20705, USA. 1 2 Tel. +1.301.504.5258 - Fax: +1.301.504.5728. Hakme Lee, Wesley M. Garrett, The pulvinus is an enlarged motor organ at E-mail: [email protected] Joseph Sullivan,1 Irwin Forseth,1 the base of leaves found in many leguminous 3 Savithiry S. Natarajan plants. It has been observed to force the move- Key words: soybean, heliotropism, nyctinasty, 1Department of Plant Sciences and ment of leaves in heliotropic, seismonastic, proteomics, LC-MS/MS. Landscape Architecture, University of and nyctinastic patterns.7,8 Unlike pulvini Maryland, College Park, MD; 2USDA-ARS, found in maize and oat that respond to gravity Acknowledgments: we thank Dr. William Kenworthy for providing the seeds for this proj- Animal Bioscience and Biotechnology by permanent growth, the movement of soy ect. This work was carried out with the support of 3 bean pulvini is reversible.8,9 In heliotropism, Laboratory, Beltsville, MD; USDA-ARS, a Cooperative Agreement between the University upon light exposure, an asymmetric turgor Soybean Genomics and Improvement of Maryland, College Park, and USDA-ARS in Laboratory, Beltsville, MD, USA gradient formed between the adaxial and abax- Beltsville, MD, USA. ial motor cells leads to leaf movement. Potassium ion influx coupled with chlorine ion Contributions: the authors contributed equally. is powered by a proton gradient and results in Abstract osmotic influx.7 In addition to heliotropism, Received for publication: 27 May 2013. the pulvinus changes turgor for nyctinastic Revision received: 19 August 2013. Accepted for publication: 23 April 2014. Certain plant species respond to light, dark, leaf folding, and is affected by alterations in and other environmental factors by leaf move- the length of the photoperiod.10 This work is licensed under a Creative Commons ment. Leguminous plants both track and avoid The structure of the pulvinus reveals its spe- Attribution NonCommercial 3.0 License (CC BY- the sun through turgor changes of the pulvi- cialized role in leaf movement. In contrast to NC 3.0). nus tissue at the base of leaves. Mechanisms the stem and petiole, the pulvinus has a rela- only ©Copyright H. Lee et al., 2014 leading to pulvinar turgor flux, particularly tively larger cortex and smaller pith. The motor 11 Licensee PAGEPress srl, Italy knowledge of the proteins involved, are not cells are part of the cortex. Two types of pulv- International Journal of Plant Biology 2014; 5:4887 well-known. In this study we used two-dimen- inar vacuoles found in many species partici- doi:10.4081/pb.2014.4887 sional gel electrophoresis and liquid chro- pate in the volume flux. Tannin-rich vacuolesuse matography-tandom mass spectrometry to sep- have been previously mentioned as a major arate and identify the proteins located in the source of cellular volume change.12 Still other relative size of its pulvinus, and ease in grow- soybean pulvinus. A total of 183 spots were studies have found the primary volume ing samples. After profiling the pulvinus pro- separated and 195 proteins from 165 spots changes to be from the type of vacuole that teome, it was compared to previously identi- were identified and functionally analyzed does not contain tannin.13 fied proteomes of nearby leaf tissue as well as using single enrichment analysis for gene A number of proteins have been linked with guard cells, which are functionally similar to ontology terms. The most significant terms pulvinar heliotropism and nyctinasty. H+- were related to proton transport. Comparison pulvini in that they also change cell shape ATPase activity increases turgor by H+ efflux with guard cell proteomes revealed similar sig- through differences in turgor. and consequent K+ influx, and H+-ATPase nificant processes but a greater number of pul- inhibitors reduce diaheliotropic response in vinus proteins are required for comparable 14 analysis. To our knowledge, this is a novel soybean pulvinus. Blue light is a deactivator report on the analysis of proteins found in soy- of H+-ATPase in Phaseolus vulgaris motor cells Materials and Methods bean pulvinus. These findings provide a better which leads to decreased turgor pressure on understanding of the proteins required for tur- the illuminated region rather than an activator Plant material gor change in the pulvinus. of H+-ATPase on the opposite region.15 Soybean (G. max cv. Clark) seeds were soaked Non-commercialFurthermore, studies on gravitropic grass pul- overnight in tap water before they were planted vini proteins have begun to identify differen- in 6-inch pots (2-3 per pot) with an LC1 soil mix- tially expressed proteins including one demon- ture (Sun Gro Horticulture, Vancouver, BC, Introduction strating MAPK-like activity.16 Canada). The plants were grown in a growth While studies have identified a number of chamber at the University of Maryland, College Plants respond to light, gravity, touch, and genes involved in a variety of tropic responses, Park, set to a 16:8 photoperiod, temperatures of other environmental signals by both temporary there remains a great dearth of knowledge on 25:20 C day:night, with a PAR of 500 mol m–2 s–1 and permanent differential growth.1,2 The the gene products and their expression pat- and 60% relative humidity. The plants were directional growth of plants as a response to an terns. The purpose of this study was to map the watered to avoid water stress and received 100 external stimulus is called tropism. The move- proteome of the soybean (Glycine max) pulvi- ppm 20:20:20 fertilizer once a week. The plants ment of leaves by sunlight, or heliotropism, nus using trichloroacetic acid/acetone extrac- were harvested after the appearance of six or can angle leaf lamina both toward (diaheliotro- tion, 2-dimensional gel electrophoresis (2-DE) seven trifoliolate leaves (between six and eight pism) and away from (paraheliotropism) the and identification by liquid chromatography- weeks). The terminal and lateral pulvini from light depending on the intensity of the irradi- tandom mass spectrometry (LC-MS/MS). This the second through sixth trifoliolate leaves were ance, circadian rhythms (nyctinasty), and map would highlight the molecular and func- separately excised with a razor one to two hours environmental stresses. Diaheliotropic move- tional characterization of the pulvinus at the into the light period and frozen in liquid nitro- ment has been shown to increase water use protein level. The soybean has been selected gen. The pulvini were stored in a −80°C freezer efficiency and carbon assimilation compared because of its importance as a food crop, the until further use. [page 8] [International Journal of Plant Biology 2014; 5:4887] Article Protein extraction excised and rinsed twice with 50% methanol, tein clusters at 100%, 90%, and 50% homology Trichloroacetic acid (TCA)/acetone precipi- ten minutes each. The gel pieces were recon- as curated by UniRef. Proteins without a name tation, described previously by Natarajan et al. stituted and subsequently dehydrated in solu- at the 50% homology level remained uncharac- was used to extract pulvinar protein.17 For tions of 25 mM ammonium bicarbonate and terized. each of three biological replicates approxi- acetonitrile respectively, by placement on a Biological process gene ontology (GO) mately 2.0 g of pulvinus were ground into a shaker for ten minutes per solution. The prior terms as listed in the Gene Ontology powder using a mortar and pestle with liquid step was repeated for a second time. The spots Annotation (GOA) Database, a collaboration nitrogen, then extracted with a 10% TCA were further dried in a speed vac concentrator with the UniProt Knowledgebase, were for about 15 minutes. Each gel piece was then (w/v)/0.07% β-mercaptoethanol (v/v) in ace- retrieved for the proteins and input into the tone mixture. Following a minimum of one reswollen with a 20 μL aliquot of 10 ng/μL agriGO version 1.2 GO analysis program hour incubation at −20°C and centrifugation porcine trypsin (sequencing grade, Promega, (China Agricultural University, Beijing, China) at 14,000 g in 4°C for 20 minutes, the super- Madison, WI, USA) in 25 mM ammonium for single enrichment analysis. For this analy- natant was discarded. The pellet was rinsed bicarbonate and refrigerated for one hour in sis, the abundance of the GO terms found in 4°C before overnight incubation at 37°C. with 0.07% β-mercaptoethanol in acetone identified pulvinar proteins were compared to The excess trypsin solution surrounding the solution followed by centrifugation at 14,000 g a background list of soybean genes. Single gel spot was transferred to new tubes and the (4°C) for 20 minutes; the rinsing and centrifu- enrichment analysis indicates the dominant remaining peptides were extracted from the gation steps were repeated until the super- biological processes in the examined tissue gel pieces by 50% acetonitrile/5% trifluo- natant was clear. After vacuum drying, the pel- against the entire plant species. It is one of roacetic acid (TFA)(v/v). 50 L of the extrac- let was resolubilized in a 7 M urea, 2 M μ several available methods to compare the sig- tion mixture was added to each gel piece and thiourea, 4% CHAPS, 1% DTT solution and nificance of each GO term. In addition, the placed in a shaker for an hour. The super- sonicated on ice for 45 minutes.
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