The bundle sheath cells lack intercellular spaces. The _______ from the light reactions is used to reduce CO2 to a carbohydrate using energy derived from ______. 2-DE-IPG was performed on 11-cm Immobiline Dry strips (Amersham Biosciences). The chlorophyll a molecules in the reaction center of a photosystem participate directly in the capture of solar energy. Approximately 400 spots per gel were detected, with spot volumes spanning five orders of magnitude. This pathway is also called Hatch and Slack pathway. When needed, peak detection was manually adjusted to remove the influence of obvious satellite peak(s). The extracted peptides were dried and resuspended in 20 μL 5% formic acid. The cross-contamination of M and BS chloroplasts fractions was assessed from the presence of the M and BS markers (PPDK and Rubisco, respectively) visualized on Coomassie Brilliant Blue or silver-stained 1-DE SDS-PAGE gels. The reason for this is that ______________. For 1-DE SDS-PAGE separation, proteins were equilibrated with SDS (0.2%), Na2CO3 (100 mM), DTT (100 mM), and sucrose (10%) and separated on 12% Tricine gels (Schägger and von Jagow, 1987). The number of chloroplasts in the bundle-sheath cells is more than that in the mesophyll cells. In addition, representative 2-DE gels from BS and M chloroplast stroma with associated information can be interrogated. Which of the following organisms can perform photosynthesis? In maize ( Zea mays ), Bundle Sheath Defective2 (BSD2) plays an essential role in Rubisco biogenesis and is required for correct bundle sheath (BS) cell differentiation. Peptide identification and quantification procedures were similar to those in the cICAT experiment, except that (1) carbamidomethylation and partial Met oxidation were selected in the MASCOT search and (2) the single ion chromatograms of the corresponding peptides (from BS or M) were extracted from two different LC-MS runs (one for BS and one for M). The web interface for PPDB was developed on ASP.NET platform using C# language. The O2 given off during photosynthesis is derived from which compound? The tearing effect applied by the pestle was sufficient to break M cells and release the M chloroplasts into the grinding buffer solution without excessively damaging the BS strands. The mesophyll cells are the usual cell type as found in C 3 plants also. Corresponding interactive gels with protein spot identities that were quantified are available in the PPDB. The BLAST searches between maize (ZmGI and AZM), Arabidopsis (TAIR), and rice (OsGI) were performed and are available on the PPDB site. Supplemental Table 4. Leaves of C3 plants do not have Kranz anatomy. The cell-specific C 4 chloroplast arrangement is established during cell … ↵ Summary of Identifications and Quantifications Obtained from the Analysis of 2-DE-IPG-PAGE for Each Protein Spot That Passed the Criteria for Reproducibility and That Was Used for Quantification. The spots were automatically washed, digested with modified trypsin (Promega) (Shevchenko et al., 1996), and extracted using the ProGest robot (Genomic Solutions). 9: In C3 plants, the carbon dioxide fixation takes place only at one place. During photosynthesis, ________ is oxidized, while ________ is reduced. In C4 plants, chloroplasts are located in ___________. C3 plants counter this limitation by increasing leaf Rubisco content, yet few C4 species do the same. the bundle sheath cells in C4 plants have chloroplasts, while those in C3 plants do not. Isoelectric focusing was conducted at 18°C in a Multiphor II (Amersham Biosciences) following the running conditions as described (Rouquie et al., 1997) for up to 72 kV/h. Which statement is accurate about how C4 plants are able to survive hot, dry spells during the growing season? Article, publication date, and citation information can be found at www.plantcell.org/cgi/doi/10.1105/tpc.105.035519. The reaction center of photosystem I consists of what type of molecule? Although bundle sheath chloroplasts display normal internal ultrastructure and are photosynthetically active, they are smaller, fewer in number, and are positioned along the cell wall distal to the vasculature. Mesophyll and bundle sheath cells have structurally and functionally different chloroplasts. English: The C4 pathway contains a special structure in the leaves known as Kranz anatomy. Supplemental Table 3. B Biol. Protein gel blot analysis with polyclonal rabbit antisera and monoclonal mouse serum using chemoluminescence was essentially performed as described earlier (Friso et al., 2004). What two things are added to NADP+ to convert it into NADPH? How does ATP synthase obtain the energy to produce ATP? On average, preparations with <15% cross-contamination yielded ∼300 μg of protein for each M and BS fraction. The database engine for PPDB (http://ppdb.tc.cornell.edu/) is a Microsoft SQL server. Supplemental Table 1. MALDI-TOF MS spectra were internally calibrated using trypsin autodigestion peptides. Thus, the size of bundle sheath compartments and the total volume of chloroplasts within them limit the space available for Rubisco. For MALDI-TOF analysis, 0.3 μL of each tryptic digest was crystallized in 0.3 μL of matrix solution (α-cyano-4-hydrocynnamic acid in 70% acetonitrile with 0.01% trifluoroacetic acid) on Teflon-coated MALDI-TOF target plates (96 wells; Perseptive Biosystems). Another three molecules of ATP are then used in the cycle. 8: In C3 plants, the bundle sheath cells do not contain chloroplasts. After a short Coomassie Brilliant Blue (R-250) staining, gels were cut in 10-slices, and proteins were then in-gel digested by trypsin and resulting peptides extracted, using standard protocols. A plant that uses CAM photosynthesis is most likely to be successful in what type of environment? Which of the following is not derived from G3P? Biol. cICAT-labeled and unlabeled peptides were identified by nano-LC-ESI-MS/MS, and cICAT-labeled peptides were quantified from nano-LC-ESI-MS runs. The biotin-tagged and cystein-containing peptides (useful for quantification) were then purified on avidin affinity columns, and the majority of the tag (containing the bulky biotin tag but not the isotope labels) was removed by acid hydrolysis, according to the manufacturer's instructions. There exists a layer of mesophyll cells that contain small chloroplasts encircling bundle sheath cells that have large chloroplasts necessary for the Calvin cycle. Mesophyll chloroplasts are randomly distributed along cell walls, whereas bundle sheath chloroplasts are located close to the vascular tissues or mesophyll cells depending on the plant species. This color is seen because the carotenoid pigments. Functional Differentiation of Bundle Sheath and Mesophyll Maize Chloroplasts Determined by Comparative Proteomics. Which steps of photosynthesis does C4 photosynthesis partition in space? What is the energy relationship between G3P and CO2? Loosely arranged mesophyll cells lie between the bundle sheath and the leaf surface. This is found in C 4 grasses such as maize and a few dicots. The resulting higher level of internal carbon dioxide in these chloroplasts serves to increase the ratio of carboxylation to … The C 4 pathway acts as a mechanism to build up high concentrations of carbon dioxide in the chloroplasts of the bundle sheath cells. (1) Mixed and labeled BS/M stromal proteomes (200 μg for each proteome) were separated on a 1-DE Tricine-PAGE gel (12% acrylamide). The two-step process by which 4-carbon compound oxaloacetate is produced in the mesophyll cells and transported into bundle sheath cells of chloroplast in Crassulacean acid metabolism (CAM) plants is called the C4 pathway in mesophyll cells. reflect and transmit yellow and orange wavelengths of light. For accurate quantification, it was critical that the BS and M samples were treated and analyzed in similar ways. w⃞ Possibly, the CAM plants may be considered to have originated by such hybridization occurring in nature during their evolutionary history. C4 leaves confine Rubisco to bundle sheath cells. Most of the cells found on epidermis have plenty of chloroplasts. Bundle sheath cells constitute ∼15% of chloroplast-containing cells in an Arabidopsis leaf (Kinsman and Pyke, 1998), and they conduct fluxes of compounds both into the leaf, particularly during leaf development, and out of the leaf, during export of photosynthates and during senescence. 2011, R.F. carbon fixation and Calvin cycle reactions. The yield of chloroplast preparations was inversely proportional to their purity. A coenzyme is defined as _______________. Gels were stained with fluorescent Sypro Ruby stain (Molecular Probes) or silver nitrate (Rabilloud et al., 1994). The MapMan system has 35 main functional categories or Bins, with a larger number of subBins (subcategories) (see also http://ppdb.tc.cornell.edu/mapman.aspx). It takes place prior to the calvin cycle. Plants with Krantz anatomy have mesophyll cells surrounding a layer of bundle sheath cells, which surround the vein (vascular bundles). The focused strips were denaturated and reduced in an equilibration buffer (6 M urea, 30% glycerol, 50 mM Tris-HCl, pH 6.8, and 2% SDS) with addition of 5 mM tributylphosphine for 20 min and subsequently alkylated in equilibration buffer with 15 mM iodoacetamide for 20 min (Rabilloud et al., 1997; Herbert et al., 1998). The light reactions of photosynthesis are said to be similar to a battery because they form a current. The top 4-cm sections of the 2nd and 3rd leaves were harvested ∼2 h after the onset of the light period. bundle sheath cells and their chloroplasts occur in relation to the development of the vascular strand, and to identify genes whose expression is bundle sheath- or mesophyll cell-specific. A new species of plant is discovered and you are asked to help classify it. Data-dependent acquisition (Masslynx 4.0) was employed to acquire MS spectra from the survey scan for quantification and tandem mass spectra for identification. the bundle sheath cells in C3 plants are arranged in columns just beneath the upper epidermis, while those in C4 plants are arranged in a ring around veins. The Calvin cycle produces ________, which are in turn used in the light reactions. H2O → PSII → electron transport chain → PSI → electron transport chain → NADPH. 2008, Christin et a… All verified protein identities using both unlabeled peptides and cICAT-labeled peptides are available via the PPDB. Enter multiple addresses on separate lines or separate them with commas. Lond. Protein pIs on the resulting 2-DE-IPG gels were internally calibrated by mixing carbamylated standards (Pharmacia Biosciences) with the stromal extracts before 2-DE analysis. In this arrangement, chloroplasts are found both in the mesophyll and the bundle sheath cells. Sypro Ruby–stained spots were excised from the gel using a robot (ProPic; Genomic Solutions) or picked manually. Where needed, functional assignments (Bins) were modified based on information from the published literature. CAM photosynthesis limits CO2 fixation to nighttime hours in order to open stomata only at night, limiting water loss because of heat and low humidity. Which of the following is a disadvantage of C4 photosynthesis relative to C3 photosynthesis? After purification of BS and M stroma, identical amounts of BS and M proteins (determined by the Bradford essay) were denatured and reduced, and all cysteins were labeled with the light (containing only 12C stable isotopes) or heavy (containing nine 13C stable isotope atoms) cICAT reagent, according to the manufacturer's instructions (Applied Biosystems). C3 plants counter this limitation by increasing leaf Rubisco content, yet few C4 species do the same. Peptides were then dried down under vacuum, resuspended in 5% formic acid, and analyzed by nano-LC-ESI-MS for quantification and by nano-LC-ESI-MS/MS for peptide identification. The MS or MS/MS spectra were searched against the maize EST assembly from TIGR (http://www.tigr.org) (ZmGI, v1.4) and maize genome assemblies (AZM 4.0) as well as by homology-based searches against the annotated rice (Oryza sativa) genome (TIGR Rice Genome version 2), using an in-house installation of Mascot (www.matrixscience.com). The removal of electrons and/or hydrogen atoms from a substrate is called what? This experiment was performed twice, first with 20 μg/g and then with 50 μg/g for each proteome, with independent biological samples (pairwise extractions of BS and M chloroplast proteomes). The … Column materials were supplied as part of the cICAT labeling kit from Applied Biosystems. A new procedure for purification of M and BS chloroplasts was developed based on suggestions from A. Barkan and R. Bassi and on published protocols (Edwards and Black, 1971; Kannangara et al., 1977). Peptides were eluted during a 45- to 60-min gradient using 95% water, 5% acetonitrile, and 0.1% formic acid as solvent A and 95% acetonitrile, 5% water, and 0.1% formic acid as solvent B at a flow rate of 0.75 μL/min. Supplemental Table 2. Although a large collection of reticulate leaf mutants is B) Mesophyll cells have PEP carboxylase while bundle sheath cells have RuBisCO. In C4 plants, the bundle sheath cells contain chloroplasts. If three or more sequence tags were found, manual inspection was not an absolute requirement, but the protein score must be well over 70, with only peptides with MOWSE scores over 21 contributing. We functionally annotated all identified ZmGI and AZM accessions using a combination of BLAST alignments to the predicted rice proteome (OSGI) and the predicted Arabidopsis thaliana proteome (from The Arabidopsis Information Resource). The innermost compartment of a chloroplast formed by the interconnected thylakoids is the __________. The two-dimensional LC was a combination of strong cation exchange followed by reverse phase, as developed recently under the name MudPIT (Washburn et al., 2001; Wolters et al., 2001), and served to reduce complexity and maximize protein identification and quantification. The difference between bundle sheath cells in C3 and C4 plants is that ________________. Chloroplasts of maize (Zea mays) leaves differentiate into specific bundle sheath (BS) and mesophyll (M) types to accommodate C4 photosynthesis. We thank J.J. Thelen for monoclonal antiserum against pyruvate dehydrogenase and M.E. Carotenoids are found in high concentrations in yellow and orange leaves or vegetables. Rubisco activity limits photosynthesis at low temperatures. Briefly, purified BS and M stromal proteomes were reduced with DTT and alkylated using iodoacetamide, followed by in-solution digestion by trypsin. In which of the labeled cells of a C4 leaf are the Calvin cycle reactions performed? The addition of electrons and/or hydrogen atoms to a substrate is called what? Which complex or enzyme of photosynthesis splits H2O? In the bundle sheath cells carbon dioxide is fixed by the enzyme RuBisCO. For example, the types and proportions of enzymes used in key metabolic steps and the structural properties of the C4 ‘Kranz’ anatomy can vary between C4 species of different evolutionary lineage (Christin et al. NADP-malic enzyme (NADP-ME) occurs in the chloroplasts of bundle-sheath cells and, like the other decarboxylases of C 4 photosynthesis, it releases CO 2, which can be fixed directly by Rubisco. This plant very likely carries out C4 photosynthesis. the bundle sheath cells in C3 plants have chloroplasts, while those in C4 plants do not. Salvucci for polyclonal serum against PEPC. Young mesophyll and bundle sheath cells in the leaf base had proplastids, and their plasmodesmata were simple, devoid of cytoplasmic sleeves. The vascular bundles are surrounded by large sized bundle sheath cells which are arranged in wreath-like manner in one to several layers. They do not have starch granules. the bundle sheath cells in C4 plants have chloroplasts, while those in C3 plants do not. For searches with MS/MS data, product ions must match within 0.8 D and precursor ions within 2 D. Criteria for positive identification by MS/MS were as follows: If only one or two matching sequence tags were found, clear partial Y-ion series and partial complementary B-ion series needed to be present (as determined by manual inspection) with MOWSE scores higher than 31 for one peptide identification and higher than 22 per peptide for two peptide identifications. Copyright © 2020 by The American Society of Plant Biologists. Identification of Maize Proteins in Purified Bundle Sheath (BS) and Mesophyll (M) Chloroplast Stroma from Three Different Types of Experiments Involving 2-DE Gels, cICAT, and LC-MS Analysis of Unlabeled Peptides. This raises the possibility of additional BSD2 roles in cell development. The author responsible for distribution of materials integral to the findings presented in this article in accordance with the policy described in the Instructions for Authors (www.plantcell.org) is: Klaas J. van Wijk (kv35{at}cornell.edu). Chloroplasts of mesophyll cells are smaller, have well developed grana and they do not produce starch. Accession numbers for quantified genes discussed in this article are listed in Table 2. In C4 plants, chloroplasts are located in both mesophyll and bundle sheath cells. Philos. They funnel solar energy to chlorophyll a in the reaction center. The bundle sheath also conducts the flo… In C 4 plants, mesophyll (M) chloroplasts carry out the light reactions of photosynthesis, while bundle sheath (BS) chloroplasts are the sites of carbon re-fixation and the Calvin cycle (Hatch 1999, Kanai and Edwards 1999). Purified M and BS stromal proteomes were first separated based on isoelectric point on IPG strips (150 μg of protein/strip), followed by a second dimension separation of SDS-PAGE. The light reactions produce ________, which are used in the Calvin cycle. The equilibrated IPG strips were loaded onto second dimension 10.5 to 14% Laemmli SDS gels (Criterium; Bio-Rad), and the protein separation was performed at constant current. Bundle Sheath: Have a single or double layer and formed of colored cells due to the presence of chloroplasts. Yet, BSD2 RNA and protein levels are similar in mesophyll (M) and BS chloroplasts, although Rubisco accumulates only in BS chloroplasts. Ambiguous matches were resolved by protein identification from MS. After background removal and normalization of each spot volume to the total gel spot volume, virtual average M and BS gels were created, where each spot volume represented the average of volumes of matched spots in M or BS gels. Why do plants contain other pigments besides chlorophyll? Many of the maize ZmGI and AZM accessions lack functional annotation. Separation of peptide mixtures for nano-LC-ESI-MS/MS was achieved by collecting sample on a μ-Guard Pre-Column (inner diameter 300 μm, 1 mm long; LC Packings), followed by separation on a PepMap C18 Reverse Phase Nano Column (inner diameter 180 μm, 15 cm long; LC Packings). The flow-through containing the nonlabeled peptides was also collected. The Calvin cycle is confined to the chloroplasts of these bundle sheath cells in C 4 plants. A poison that disrupts the H+ gradient in the thylakoid space would most likely cause which of the following to accumulate during the Calvin cycle reactions? van Wijk, unpublished data), followed by a manual verification steps for weaker identifications. So the correct answer is 'High density of chloroplasts'. As a third complementary approach, we used a non-gel-based quantitative approach that was independent of cystein labeling. These distinct lineages exhibit unique aspects that highlight their independent evolution from C3 ancestors. In particular, we relied on the hierarchical, nonredundant classification system developed for MapMan (Thimm et al., 2004) (http://gabi.rzpd.de/projects/MapMan/), adjusted after manual verification and information from literature and incorporated into PPDB. the bundle sheath cells in C4 plants have chloroplasts, while those in C3 plants do not. The flow-through containing the nonlabeled peptides was also collected since they were valuable for protein identification. Energy to drive the formation of ATP in photosynthesis is derived from ____________. Th e vascular bundles are surrounded by large sized bundle sheath cells which are arranged in wreath -like manner in one to several layers. ... Dicot leaves do not have bulliform cells whereas monocot leaves have bulliform cells; Sun and K.J. C 4 plants are an interesting system to study plastid development, since protochloroplasts differentiate into two morphologically and functionally different chloroplast types in mesophyll (M) and bundle sheath (BS) cells to support the C4 carbon concentrating mechanism. Only those peptides were considered that had no missed cleavages (by trypsin) and no modifications, except for Met oxidation and carbamidomethylation (since gel samples were alkylated). In the first run, mass spectrometric parameters were setup favoring more time for the survey scans, and high-quality tandem MS spectra for peptides with higher signal intensity could also be obtained in this run. The residual M cells attached to BS strands were removed by 10 min of blending (one-third of the maximal speed) in a modified Warring blender (WB1) with blunt blades. The differentially labeled BS and M stromal proteomes were mixed in a 1:1 ratio and proteins were analyzed in two different ways, as follows. Criteria for positive identification by MALDI-TOF MS peptide mass fingerprinting include five or more matching peptides with a narrow error distribution (clustering of errors) within 25 ppm and at least 15% sequence coverage. Chloroplast fractions showing <15% of cross-contamination were used, with the majority having a contamination of ∼10%. SAUR17 and SAUR50 Differentially Regulate PP2C-D1 during Apical Hook Development and Cotyledon Opening in Arabidopsis, AUTOPHAGY-RELATED14 and Its Associated Phosphatidylinositol 3-Kinase Complex Promote Autophagy in Arabidopsis, by The American Society of Plant Biologists, www.plantcell.org/cgi/doi/10.1105/tpc.105.035519. C4 leaves confine Rubisco to bundle sheath cells. They have large chloroplasts without grana. The protein quantification was done by comparing the peak areas of corresponding peptides from BS or M. Masslynx 4.0 software (Waters) was employed to extract the peak areas of identified peptides of the corresponding single ion chromatograms. C4 photosynthesis is one of the most convergent evolutionary phenomena on the planet, with >65 independent origins (R.F. When the stomata close during heat stress, RuBP carboxylase is not exposed to the increase in oxygen concentration experienced in the leaf. The difference between bundle sheath cells in C3 and C4 plants is that _____. It summarizes the currently accepted views on metabolic compartmentation of biochemical reactions between the chloroplasts of bundle sheath cells and mesophyll cells of a C 4 plant. Which stages of the Calvin cycle reactions require the use of energy from ATP? Chloroplasts of mesophyll cells are smaller, have well developed grana and they do not produce starch, Chloroplasts of bundle sheath cells … Protein identification was performed by peptide mass fingerprinting using MALDI time-of-flight (TOF) MS in reflectron mode (Voyager DE-STR; Perseptive Biosystems) and online LC-ESI-MS/MS (Q-TOF; Micromass). 2012). Annu. Trans. It forms a protective covering on leaf vein, and consist of one or more cell layers, usually parenchyma. For 1-DE gel cICAT experiments, the non-Cys-containing peptides separated from both cICAT-labeled peptides were also purified by homemade reverse-phase columns packed with C18 beads, analyzed by LC-MS/MS, and searched against ZmGI to obtain protein identities. The majority of spots (65% for M and 54% for BS gels) were represented at least three times either in M and/or in BS proteomes (45% for M and 39% for BS spots were represented at least four times). Not all molecules contain the same amount of chemical energy. 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Plants do not contain chloroplasts in the reaction center of photosystem II are derived from _____________ to chlorophyll a the. Reaction center and Slack pathway PSI → electron transport chain → NADPH column materials supplied.