Brief Summary of Minutes of Annual Meeting

The annual meeting of Multistate Research Project NC1168 (Regulation of Photosynthetic Processes) was hosted by the Virginia AES and held at the Peggy Lee Hahn Garden Pavilion at Virginia Tech, Blacksburg, Virginia on November 13, 2010. Attending members included Robert Aiken, Christoph Benning, Gerald Edwards, Glenda Gillaspy, Jiaxu Li, Jeff Harper, Michael E. Salvucci, Robert J. Spreitzer, and Donald P. Weeks. Attending guests included Saied Mostaghimi (Associate Dean and Director of the Virginia AES), Eva Collakova (Virginia Tech), and Jim Moroney (Louisiana State University). The meeting convened at 9:00 AM at the Hahn Garden Pavilion. Glenda Gillaspy (Virginia, AES) introduced Dr. Saied Mostaghimi, Associate Dean and Director, who summarized the organization and progress of the Virginia AES. Mike Salvucci (Arizona ARS) gave the first talk pertaining to objective 2. He described the structural basis for interactions between Rubisco and Rubisco activase (in collaboration with the Nebraska AES), and described field studies aimed at elucidating the influence of Rubisco activase on thermal tolerance. Bob Spreitzer's presentation focused on Rubisco engineering, including collaborative results from the Arizona ARS pertaining to the Rubisco structural transition. Gerry Edwards (Washington AES) described recent progress on mechanisms of single-cell C4 photosynthesis. After a short break (11:25-11:45 AM), the meeting resumed with Don Weeks summarizing recent collaborative work (with Iowa AES) in the application of TAL effector nucleases. Lunch was held at the Inn at Virginia Tech at 12:25 PM. The group returned to the Hahn Pavilion at 1:40 PM, at which time Jim Moroney (a potential new member from the Louisiana AES) brought the group up to date with regard to his research on the algal CO2 concentrating mechanism. He focused his presentation on a protein (LCI1) involved in carbon acquisition, a new protein (CIA6) that may play a role in pyrenoid formation, and various carbonic anhydrases. Christoph Benning (Michigan AES) gave the first talk pertaining to objective 3. He described progress in the use of rutabaga and algae for oil production. Eva Collakova (guest from Virginia Tech) discussed the utility of metabolic flux analysis with regard to metabolic engineering. After a short break (3:30-3:50 PM), Jeff Harper (Nevada AES) and Glenda Gillaspy (Virginia AES) described research at the interface between objectives 3 and 4. Harper discussed calcium signaling, and Gillaspy presented results on the role of inositol phosphates in carbon partitioning and signaling. In objective 4, Jiaxu Li (Mississippi AES) described histone methyltransferases involved in gene regulation. Rob Aiken (Kansas AES) described field studies focused on drought and oxidative stress. After a 15 minute break, the business meeting was called to order by the chair (Gillaspy) at 6:00 PM. Irwin Goldman (Administrative Advisor) joined the group via Skype. Goldman stressed the importance of communicating the integration and collaboration of group members in future annual reports. There was confusion about who serves as secretary for the group. It was decided/confirmed that the chair-elect is responsible for taking minutes at the annual meeting. Spreitzer voiced concern about attendance of members at the annual meeting. Because of the broad, geographical distribution of participating members, travel to the annual meeting may be difficult. Harper proposed that the group meet at a more-accessible "hub" city. After discussion, it was concluded that the host of the annual meeting would choose the location. However, Salvucci offered to serve as chair of the meeting in Phoenix (in 2013) as a more-accessible and previously-favored location. The group then considered the election of Jim Moroney as a member of NC1168. The vote of members present, and those communicating via email, was unanimous. The group discussed the renewal application of NC1168. It was agreed that the four objectives will remain the same. Benning will take the lead in preparing the application. Benning will coordinate members' input to objective 1. Spreitzer will take the lead on objective 2. Gillaspy will coordinate objective 3. Salvucci will organize members in objective 4. Goldman will provide information on important dates/deadlines for preparation of the renewal proposal. The appointment of chairs and schedule for future meetings are: 2011 Michigan AES Christoph Benning 2012 Maine AES Mary Rumpho (secretary, chair-elect in 2011) 2013 Arizona ARS Mike Salvucci (secretary, chair-elect in 2012) The meeting was adjourned at 6:45 PM. Members met as a group for dinner on both November 12 and 13 for discussions of research results and for establishing collaborative plans.

Obj. A. Plastid Function and Intracellular Communication. Photosynthesis is comprised of tightly coupled reactions requiring finely-tuned nucleocytosolic-plastid interactions and carefully executed signaling pathways. Efforts in this area center on the examination of a unique plastid model system (Maine AES) and studies of sugar signaling pathways (Ohio and Virginia AES). ME-AES continued characterization of expression of the photosynthetic mollusc, Elysia chlorotica, which harbors plastids intracellularly obtained from the algae, Vaucheria litorea. Quantitative PCR was used to compare transcript levels of both nuclear (psbO and prk) and plastid (psaA, psbA and rbcL) photosynthesis genes in E. chlorotica and V. litorea. In all cases, transcript levels were lower in the sea slug than its algal prey and they did not respond similarly to changes in light conditions over a 24-hr period. In collaboration with NC-1168 PI Wayne Loescher (MI AES), work has begun to quantify the major photosynthetic products of both organisms. Mannitol, hexoses and cyclitols have been found. Work on sugar signaling pathways seeks to understand the molecules that plants use in these pathways. OH-AES characterized the effects of a group of bZIP transcription factors and a tandem zinc finger (TZF) gene family in sugar-mediated transcriptional and post-transcriptional regulation, respectively. They have found that the expression of AtbZIP1 is repressed by sugars in a fast, sensitive, and reversible way. This sugar repression is also affected by a conserved sugar signaling component, hexokinase. VA-AES has investigated an energy sensor called SnRK1.1 which plants use to sense sugars and energy. It was found that SnRK1.1 is regulated by proteasomal turnover and proteins have been identified that participate in this process. Obj. B. Photosynthetic Capture and Photorespiratory Release of CO2. Rubisco catalyzes the rate-determining step in photosynthesis, but it does so very inefficiently. Because of this inefficiency, Rubisco presents an obvious target for increasing photosynthetic performance. The Arizona ARS and Nebraska AES have been working to understand key factors in Rubisco activity. Because the catalytic properties of Rubisco enzymes differ among various species, NE-AES has been taking a phylogenetic approach to identify those regions of the Rubisco large subunit that may account for these differences. Chlamydomonas mutant enzymes have been characterized with decreased CO2/O2 specificity, but it can be activated normally by Rubisco activase, indicating that the structural transition required for catalysis is not affected. It appears that there may be a different path to the active site that is influenced by the nature of the large/small-subunit interface. In synergistic studies, AZ-ARS has sought to understand Rubisco activase structure/function by acquiring structural information at the atomic level. In collaboration with NE-AES they have examined the effect of substituting higher plant small subunits (RBCS) on the activation of Chlamydomonas Rubisco by activase. This finding suggests that replacing native RBCS in Chlamydomonas Rubisco with a higher plant RBCS increases the stability of the binary Rubisco-activase complex without affecting the rate of Rubisco activation. Two members from the Washington AES have focused on research on the structural and biochemical diversity in mechanisms of C4 photosynthesis which evolved to concentrate CO2 around Rubisco, enhance photosynthesis, and reduce losses of CO2 due to photorespiration. These members reviewed current information on structural and biochemical diversity in C4 photosynthesis and described 25 known structural forms of Kranz anatomy and two structural forms where C4 occurs within individual photosynthetic cells without Kranz. Other efforts focused on the characterization of the mechanism of C4 in single-cell C4 species in which photosynthesis occur within individual photosynthetic cells (Chenopodiaceae). Two cytoplasmic domains develop with dimorphic chloroplasts considered to function analogous to mesophyll and bundle sheath cells in Kranz type species. WA-AES is working on the biochemical, physiological properties and molecular requirements for this form of C4. A purification protocol for the two different chloroplast types has been developed, and functional studies performed on intact chloroplasts. The results provide evidence that one domain is specialized for supporting fixation of atmospheric CO2 in the C4 cycle, and the other for accepting CO2 from decarboxylation of C4 acids and its assimilation by Rubisco in the C3 cycle. Obj. C. Mechanisms Regulating Photosynthate Partitioning. Several approaches are underway to understand how plants regulate photosynthate partitioning, and this area of work has important implications for generating better bioenergy crops. Since increasing oil biosynthesis in vegetative tissues of plants offers a way to convert carbon and increase the energy density of biomass, Michigan AES has examined converting starch to oil in the rutabaga storage organ. A transcriptomic analysis in different rutabaga tissues is underway to identify hypocotyl specific promoters and new target genes for second generation transgenics. MI-AES is also exploring the induction of triacylglycerol biosynthesis in microalgae. Synergistic studies are underway at the Nevada AES focusing on the algae Dunaliella salina. This unicellular, halophytic green alga belongs to the Chlorophyceae, and is among the most industrially important microalgae because it can produce massive amounts of ²-carotene and because of its potential as a feedstock for biofuels production. A complementary DNA (cDNA) library was constructed from D. salina cells adapted to 2.5 M NaCl and used to generate an expressed sequence tag (EST) database. ESTs were obtained for 2,831 clones representing 1,401 unique transcripts. Putative functions were assigned to 1,901 (67.2%) ESTs after comparison with protein databases. An additional 154 (5.4%) ESTs had significant similarity to known sequences whose functions are unclear and 776 (27.4%) had no similarity to known sequences. A third synergistic effort is underway by two members (Nebraska AES and Iowa AES) to develop a transformation system for algae. The ability to readily disrupt genes and/or replace gene sequences using homologous recombination has greatly speeded research with bacterial and yeast systems. When developed, this is expected to dramatically facilitate studies in algae, plants, and development of bioenergy resources. These NC-1168 members have tested a TAL effector nuclease (TALN) technology for targeted gene knockout and gene replacement in algae. To test their gene-specific TALN technology, preliminary experiments have been conducted in which naturally-occurring full-length TAL effectors were tested for the specificity of their binding to target DNA both in vitro and in vivo. The results suggest that artificial TALNs can be produced to target specific genes. A group of NC-1168 members have examined signaling components that impact photosynthate partitioning (Illinois-ARS, Nevada AES, Virginia AES). IL-ARS is examining engineering photosynthesis by a directed mutagenesis of a receptor kinase. It was found that a receptor kinase, BAK1, needs to be phosphorylated on a tyrosine residue in vivo for this. Preventing autophosphorylation by directed mutagenesis enhances plant growth. The basis for increased growth may be a higher rate of light saturated CO2 assimilation under ambient conditions, and accordingly these plants accumulate higher levels of starch and sucrose in leaves during the day. Transcriptome analysis indicated that the expression of numerous genes was altered and this may suggest new approaches to indirectly increase whole plant carbon assimilation. NV-AES has examined 14-3-3s, which are regulatory proteins that have been implicated in nitrogen and carbon metabolism by the NV-ARS. More than 124 14-3-3 clients have been identified, 103 of which have not previously been reported. Many of the newly identified clients are involved directly in metabolism, such as phosphoenol pyruvate (PEP) carboxylase. The VA-AES has investigated myo-inositol synthesis and phosphoinositide signaling molecules that appear to limit plant size. Phosphatidylinositol was found to also limit signaling by another important class of lipid signaling molecules, the sphingolipids. Three members are investigating metabolic controls on photosynthate partitioning. The Washington AES has examined the plastidic phosphorylase. The plastidic starch phosphorylase, Pho1, is essential for starch initiation at low temperatures. Although previous kinetic studies suggested that Pho1 was not involved in starch synthesis but rather starch degradation, kinetic analysis of the purified recombinant Pho1 indicates that it strongly favors the synthesis reaction. This may aid efforts to understand and modify source-sink relationships to improve crop productivity. Florida AES, and Michigan AES reported data on polyols in plants, which can be major sinks for carbon. FL-AES has found that sorbitol dehydrogenase has a role in determining the kernel number per ear in corn. In addition, the Cellulose Synthase-Like D (CSLD) genes were genetically determined to impact plant cell size. Key to this work are contributions to the UniformMu maize project which serves as a resource for sequence-indexed insertional mutants. The UniformMu population provides both seeds and sequences free of charge through the MaizeGDB. As of December 2010, 22,239 unique, germinal Mu transposon insertions have been mapped. FL-ARS and MI-AES are collaborating on the role of mannose, characterizing the impact of M6PR gene both in Arabidopsis and its original source, celery. Obj. D. Developmental and Environmental Limitations to Photosynthesis. Several NC-1168 members have pursued strategies to understand how photosynthesis can be limited by stress. Work has focused on molecular, whole plant and field responses. AZ-AES has examined differences in thermal stability between wild type Rubisco activase and structurally-altered mutant recombinant proteins in vitro and when the proteins were expressed in transgenic Arabidopsis plants. Rubisco activation studies indicated that that the thermal stability of Rubisco activase determines the heat-sensitivity of photosynthesis. In synergistic work, Kansas AES has examined the possible relationship between expression levels of Rubisco activase and plant yield in winter wheat, corn and Arabidopsis following prolonged exposure to heat stress. A significant, positive, linear correlation was found between the expression of activase and plant productivity under heat-stress conditions. In significant pilot work, the relevance of Rubisco activase to the inhibition of photosynthesis by heat stress under field conditions in Arizona was investigated. The combined effects of heat and drought under field conditions for cotton cultivars that differed in their drought tolerance were measured as part of a larger study designed to develop the capacity for high-throughput phenotyping. These studies use multispectral data spanning visible, near infrared, and thermal infrared wavelengths, along with plant canopy geometric data, to detect and evaluate plant responses to heat and drought stress at a field site in Maricopa, Arizona. Other field studies conducted by Kansas-ARS have identified genotypes in sorghum, wheat and soybean with increased stress tolerance which could enhance primary productivity and grain yield formation. Other breeding efforts by MI-AES focused on common bean to develop a better understanding of physiological parameters, quantitative trait loci and genes associated with drought tolerance in this important food crop. To investigate molecular mechanisms of stress, the NV-AES has examined the function of CAM in reactive oxygen species (ROS) alleviation using a newly isolated CAM-deficient mutant of a facultative halophyte Mesembryanthemum crystallinum L. Salinity induced nocturnal malate synthesis in the leaves of wild-type plant, but not in the mutant. These results indicated that lower levels of ROS accompanied the performance of CAM, and that CAM might alleviate oxidative stress. Another member of NV-ARS has also focused on several genes implicated in both calcium signaling and survival under extreme environmental conditions, such as cold, heat, drought, salt, and high-light stress. The calcium-dependent protein kinases, cyclic nucleotide gated channels, and P-type ATPase ion pump genes have been studied via functional genomics approaches with mutant and transgenic plants. The vacuolar calcium pumps (ACA4 and ACA11) control calcium signals that can trigger programmed cell death and limit plant productivity. Mississippi ARS reported on another set of genes, the Arabidopsis SET domain genes. One of these (SDG8) is a histone methyltransferase responsible for di- and trimethylation of the lysine 36 residue of histone 3. MI-ARS showed that knockdown mutations of this methyltransferase markedly reduce the global level of H3 trimethylation in vivo and affect rice flower and grain development. These studies suggest H3K36 trimethylation plays a key role in reproductive organ development in rice.

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