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Editing of eIF(iso)4E.c confers resistance against Turnip mosaic virus in Brassica rapa

Horticultural Plant Journal 2024年第4期10卷 1020-1034页

作者： Yujia Liu Xiaoyun Xin Peirong Li Weihong Wang Yangjun Yu Xiuyun Zhao Deshuang Zhang Jiao Wang Fenglan Zhang Shujiang Zhang Shuancang Yu Tongbing Su State Key Laboratory of Vegetable Biobreeding Beijing Vegetable Research CenterBeijing Academy of Agriculture and Forestry ScienceBeijing 100097China National Engineering Research Center for Vegetables Beijing Key Laboratory of Vegetable Germplasms ImprovementKey Laboratory of Biology and Genetics Improvement of Horticultural Crops(North China)Key Laboratory of Urban Agriculture(North)of Ministry of AgricultureBeijing Vegetable Research CenterBeijing Academy of Agriculture and Forestry ScienceBeijing 100097China institute of Vegetables and flowers Chinese Academy of Agricultural SciencesBeijing 100081China

Turnip mosaic virus(TuMV)constitutes one of the primary diseases affecting Brassica rapa,severely impacting its production and resulting in crop failures in various regions *** research has demonstrated the significance of plant translation initiation factors,specifically the eIF4E and eIF4G family genes,as essential recessive disease resistance *** our study,we conducted evolutionary and gene expression studies,leading us to identify e IF(iso)4E.c as a potential TuMV-resistant *** CRISPR/Cas9 technology,we obtained mutant *** plants with edited eIF(iso)4E.c *** confirmed eIF(iso)4E.c confers resistance against TuMV through phenotypic observations and virus content ***,we employed ribosome profiling assays on eif(iso)4e.c mutant seedlings to unravel the translation landscape in response to ***,we observed a moderate correlation between the fold changes in gene expression at the transcriptional and translational levels(R^(2)=0.729).Comparative analysis of ribosome profiling and RNA-seq data revealed that plant-pathogen interaction,and MAPK signaling pathway-plant pathways were involved in eIF(iso)4E.c-mediated TuMV *** analysis revealed that sequence features,coding sequence length,and normalized minimal free energy,influenced the translation efficiency of *** study highlights that the loss of e IF(iso)4E.c can result in a highly intricate translation mechanism,acting synergistically with transcription to confer resistance against TuMV.

关键词： Brassica rapa eIF(iso)4E.c TuMV Ribo-seq

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Non-CG DNA methylation-deficiency mutations enhance mutagenesis rates during salt adaptation in cultured Arabidopsis cells

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Stress Biology 2021年第1期1卷 144-155页

作者： Xiaohong Zhu Shaojun Xie Kai Tang Rajwant KKalia Na Liu Jinbiao Ma Ray ABressan Jian-Kang Zhu State Key Laboratory of Crop Stress Adaptation and Improvement School of Life SciencesHenan UniversityKaifengChina State Key Laboratory of Cotton Biology School of Life SciencesHenan UniversityKaifengChina Bioinformatics Core Purdue UniversityWest LafayetteIN 47907USA Department of Horticulture and landscape Architecture Purdue UniversityWest LafayetteIN 47907USA Central Arid Zone Research institute Jodhpur 342003India State Key Laboratory for Managing Biotic and Chemical Threats to the Quality and Safety of Agro-Products Institute of VegetablesZhejiang Academy of Agricultural SciencesHangzhou 310021China State Key Laboratory of Desert and Oasis Ecology Xinjiang Institute of Ecology and GeographyChinese Academy of Sciences 830011UrumqiChina Shanghai Center for Plant Stress Biology Shanghai Institutes for Biological SciencesChinese Academy of SciencesShanghai 200032China

Much has been learned about how plants acclimate to stressful environments,but the molecular basis of stress adaptation and the potential involvement of epigenetic regulation remain poorly ***,we examined if salt stress induces mutagenesis in suspension cultured plant cells and if DNA methylation affects the mutagenesis using whole genome resequencing *** generated suspension cell cultures from two Arabidopsis DNA methylation-deficient mutants and wild-type plants,and subjected the cultured cells to stepwise increases in salt stress intensity over 40 culture *** show that ddc(drm1 drm2 cmt3)mutant cells can adapt to grow in 175 mM NaCl-containing growth medium and exhibit higher adaptability compared to wild type Col-0 and nrpe1 cells,which can adapt to grow in only 125mM NaCl-containing growth *** treated nrpe1 and ddc cells but not wild type cells accumulate more mutations compared with their respective untreated *** is no enrichment of stress responsive genes in the list of mutated genes in salt treated cells compared to the list of mutated genes in untreated *** results suggest that DNA methylation prevents the induction of mutagenesis by salt stress in plant cells during stress adaptation.

关键词： Salt stress Cell suspension DNA methylation-deficient mutants Stress-induced mutagenesis Stress adaptation

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Global Lysine Acetylome Analysis of Flower Bud Development in Catalpa bungei

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Phyton-International Journal of Experimental Botany 2022年第3期91卷 507-524页

作者： Danlong Jing Nan Wang Nan Lu Guanzheng Qu Guolu Liang Wenjun Ma Shougong Zhang Junhui Wang State Key Laboratory of Tree Genetics and Breeding Key Laboratory of Tree Breeding and Cultivation of State Forestry and Grassland AdministrationResearch Institute of ForestryChinese Academy of ForestryBeijing100091China Key Laboratory of Horticulture Science for Southern Mountains Regions of Ministry of Education College of Horticulture and Landscape ArchitectureAcademy of Agricultural SciencesSouthwest UniversityChongqing400715China State Key Laboratory of Tree Genetics and Breeding Northeast Forestry UniversityHarbin150040China

Lysine acetylation,a reversible and dynamic post-translational modification,plays pivotal roles in regulating many biological processes in ***,the information of lysine acetylation during flower development of woody angiosperm remains ***,we identified a total of 667 lysine acetylation sites in 467 proteins in flower buds of Catalpa *** acetylated proteins were mainly involved in the pathways of carbon metabolism,glycometabolism and oxidative *** functional enrichment analysis,61 and 13 acetylated proteins were involved in the glycometabolism and oxidative phosphorylation pathways,*** suggests that lysine acetylated proteins play critical roles in the energy metabolism ***,the protein–protein interactions were also highly enriched in the carbon metabolic *** analysis of lysine acetylome and transcriptome showed that transcript level changes of seven key lysine-acetylated proteins/genes involved in energy metabolism were significant different during floral ***,compared with the normal flowering variety,the expression levels of phosphoribulokinase,phosphoenolpyruvate carboxykinase,ATP synthase and pyruvate dehydrogenase complex genes were significantly up-regulated at the stages of vegetative and transition buds during early flowering of *** data represent the first lysine acetylome in the *** flowers,and can serve as an essential basis for exploring the function of lysine acetylated proteins in the flower bud development of plants.

关键词： Lysine acetylome Catalpa bungei flower development energy metabolism

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Induction of Tetraploid Male Sterile Tagetes erecta by Colchicine Treatment and Its Application for Interspecific Hybridization

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Horticultural Plant Journal 2016年第5期2卷 284-292页

作者： HE Yanhong SUN Yalin ZHENG Riru AI Ye CAO Zhe BAO Manzhu Key Laboratory of Horticultural Plant Biology Ministry of Education College of Horticulture and Forestry Sciences Huazhong Agricultural University institute of Vegetable Science Wuhan Academy of Agricultural Sciences College of landscape Architecture Fujian Agriculture and Forestry University Gulf Coast Research and Education Center Institute of Food and Agricultural Sciences University of Florida

Tagetes erecta is an annual multifunctional plant which can be cultivated under a broad range of climatic conditions. Polyploidization and interspecific hybridization are applied to facilitate breeding cultivars of T. erecta with improved ornamental qualities. Colchicine treatment to the germinating seeds was proved to be a useful tool for chromosome doubling of the male sterile two-type line ‘M525AB', with the resulting frequency of polyploid seedlings ranging from 88.89%(following 0.05% w/v colchicine applied for a 3–6 h exposure period) to a maximum of100.00%(following 0.1% for 3–6 h, or 0.2% for 3 h). Morphological observation, stomatal size and density analysis, flow cytometric analysis and chromosome counting were conducted to identify the tetraploid plants. Distinctive morphological changes were observed in a notable proportion of polyploid plants. The colchicine-treated polyploid T. erecta plants showed dwarfed and more robust growth, thicker, larger and greener leaves, larger inflorescences and florets. The mutant plants identified through morphological observation all aligned as polyploid plants, thus morphological observation could be an effective method for the detection of polyploidy. The polyploid plants had significant larger stomata size over the abaxial leaf surface, whereas the density of stomata distribution was remarkably reduced. The survival rate of tetraploid cuttings(i.e. 38%)was greatly reduced compared to that of diploid plants. The fertility of tetraploid plants was also decreased, as shown by cross-pollination *** hybridizations between colchicine-induced tetraploid plants of a male sterile T. erecta line and the naturally tetraploid fully fertile Tagetes patula species resulted in hybrid progeny. Most of these hybrids displayed the dwarfed growth stature and compact, larger-flower morphology which is the typical ideotype of herbaceous flowers. Thus, polyploidization may be employed effectively as a means to facilitate in

关键词： Tagetes erecta Tagetes patula chromosome doubling colchicine tetraploid interspecific hybridization

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Recent advances in RNA structurome

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Science China(Life Sciences) 2022年第7期65卷 1285-1324页

作者： Bingbing Xu Yanda Zhu Changchang Cao Hao Chen Qiongli Jin Guangnan Li Junfeng Ma Siwy Ling Yang Jieyu Zhao Jianghui Zhu Yiliang Ding Xianyang Fang Yongfeng Jin Chun Kit Kwok Aiming Ren Yue Wan Zhiye Wang Yuanchao Xue Huakun Zhang Qiangfeng Cliff Zhang Yu Zhou MOE Laboratory of Biosystems Homeostasis&Protection Innovation Center for Cell Signaling NetworkCollege of Life SciencesZhejiang UniversityHangzhou 310058China Key Laboratory of RNA Biology Institute of BiophysicsChinese Academy of SciencesBeijing 100101China Life Sciences institute Zhejiang UniversityHangzhou 310058China State Key Laboratory of Plant Physiology and Biochemistry College of Life SciencesZhejiang UniversityHangzhou 310058China State Key Laboratory of Virology College of Life SciencesWuhan UniversityWuhan 430072China Beijing Advanced Innovation Center for Structural Biology School of Life SciencesTsinghua UniversityBeijing 100084China Stem Cell and Regenerative Biology Genome Institute of SingaporeA*STARSingaporeSingapore Department of Chemistry and State Key Laboratory of Marine PollutionCity University of Hong KongHong KongChina MOE Key Laboratory of Bioinformatics Beijing Advanced Innovation Center for Structural Biology and Frontier Research Center for Biological StructureCenter for Synthetic and Systems BiologySchool of Life SciencesTsinghua UniversityBeijing 100084China Tsinghua-Peking Center for Life Sciences Beijing 100084China Department of Cell and Developmental Biology John Innes CentreNorwich Research ParkNorwich NR47UHUnited Kingdom Shenzhen Research institute of City University of Hong Kong Shenzhen 518057China University of Chinese Academy of Sciences Beijing 100101China Key Laboratory of Molecular Epigenetics of the Ministry of Education Northeast Normal UniversityChangchun 130024China

RNA structures are essential to support RNA functions and regulation in various biological processes. Recently, a range of novel technologies have been developed to decode genome-wide RNA structures and novel modes of functionality across a wide range of species. In this review, we summarize key strategies for probing the RNA structurome and discuss the pros and cons of representative technologies. In particular, these new technologies have been applied to dissect the structural landscape of the SARS-CoV-2 RNA genome. We also summarize the functionalities of RNA structures discovered in different regulatory layers-including RNA processing, transport, localization, and mRNA translation-across viruses, bacteria, animals, and plants. We review many versatile RNA structural elements in the context of different physiological and pathological processes(e.g., cell differentiation, stress response, and viral replication). Finally, we discuss future prospects for RNA structural studies to map the RNA structurome at higher resolution and at the single-molecule and single-cell level, and to decipher novel modes of RNA structures and functions for innovative applications.

关键词： RNA structurome high-throughput techniques genome-wide 3D structure RNA secondary structure SARS-CoV-2 decoding function

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Plant Biosystems Design Research Roadmap 1.0

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BioDesign Research 2020年第1期2020卷 53-90页

作者： Xiaohan Yang June I.Medford Kasey Markel Patrick M.Shih Henrique C.De Paoli Cong T.Trinh Alistair J.McCormick Raphael Ployet Steven G.Hussey Alexander A.Myburg Poul Erik Jensen Md Mahmudul Hassan Jin Zhang Wellington Muchero Udaya C.Kalluri Hengfu Yin Renying Zhuo Paul E.Abraham Jin-Gui Chen David J.Weston Yinong Yang Degao Liu Yi Li Jessy Labbe Bing Yang Jun Hyung Lee Robert W.Cottingham Stanton Martin Mengzhu Lu Timothy J.Tschaplinski Guoliang Yuan Haiwei Lu Priya Ranjan Julie C.Mitchell Stan D.Wullschleger Gerald A.Tuskan Biosciences Division Oak Ridge National LaboratoryOak RidgeTN 37831USA The Center for Bioenergy Innovation Oak Ridge National LaboratoryOak RidgeTN 37831USA Department of Biology Colorado State UniversityFort CollinsCO 80523USA Department of Plant Biology University of CaliforniaDavisDavisCAUSA Feedstocks Division Joint BioEnergy InstituteEmeryvilleCAUSA Department of Biodesign Biological Systems and Engineering DivisionLawrence Berkeley National LaboratoryBerkeleyCAUSA Department of Chemical and Biomolecular Engineering University of TennesseeKnoxvilleTN 37996USA SynthSys and institute of Molecular Plant Sciences School of Biological SciencesUniversity of EdinburghEdinburgh EH93BFUK Department of Biochemistry Genetics and MicrobiologyForestry and Agricultural Biotechnology Institute(FABI)University of PretoriaPretoria 0002South Africa Department of Food Science University of CopenhagenRolighedsvej 26DK-1858FrederiksbergCopenhagenDenmark State Key Laboratory of Subtropical Silviculture School of Forestry and BiotechnologyZhejiang A&F UniversityHangzhouZhejiang 311300China State Key Laboratory of Tree Genetics and Breeding Research Institute of Subtropical ForestryChinese Academy of ForestryHangzhouZhejiang 311400China Department of Plant Pathology and Environmental Microbiology and the Huck institute of the Life Sciences The Pennsylvania State UniversityUniversity ParkPA 16802USA Department of Genetics Cell Biology and DevelopmentCenter for Precision Plant Genomics and Center for Genome EngineeringUniversity of MinnesotaSaint PaulMN 55108USA Department of Plant Science and landscape Architecture University of ConnecticutStorrsCT 06269USA Division of Plant Sciences Bond Life Sciences CenterUniversity of MissouriColumbiaMOUSA Donald Danforth Plant Science Center St.LouisMOUSA Environmental Sciences Division Oak Ridge National LaboratoryOak RidgeTN 37831USA

Human life intimately depends on plants for food,biomaterials,health,energy,and a sustainable *** plants have been genetically improved mostly through breeding,along with limited modification via genetic engineering,yet they are still not able to meet the ever-increasing needs,in terms of both quantity and quality,resulting from the rapid increase in world population and expected standards of living.A step change that may address these challenges would be to expand the potential of plants using biosystems design *** represents a shift in plant science research from relatively simple trial-and-error approaches to innovative strategies based on predictive models of biological *** biosystems design seeks to accelerate plant genetic improvement using genome editing and genetic circuit engineering or create novel plant systems through de novo synthesis of plant *** this perspective,we present a comprehensive roadmap of plant biosystems design covering theories,principles,and technical methods,along with potential applications in basic and applied plant biology *** highlight current challenges,future opportunities,and research priorities,along with a framework for international collaboration,towards rapid advancement of this emerging interdisciplinary area of ***,we discuss the importance of social responsibility in utilizing plant biosystems design and suggest strategies for improving public perception,trust,and acceptance.

关键词： Plant utilizing breeding

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An Arabidopsis ABC Transporter Mediates Phosphate Deficiency-Induced Remodeling of Root Architecture by Modulating Iron Homeostasis in Roots

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Molecular Plant 2017年第2期10卷 244-259页

作者： Jinsong Dong Miguel A. PiSeros Xiaoxuan Li Haibing Yang Yu Liu Angus S. Murphy Leon V. Kochian Dong Liu Ministry of Education Key Laboratory of Bioinformatics Center for Plant Biology School of Life Sciences Tsinghua University Beijing 100084 China USDA-ARS Robert Holley Center for Agriculture and Health Cornell University ithaca NY 14580 USA Department of Horticulture Purdue University West Lafayette IN 47907-2010 USA State Key Laboratory of Plant Physiology and Biochemistry College of Life Science Zhejiang University Hangzhou 310058 China Department of Plant Science and landscape Architecture University of Maryland College Park MD 20742 USA GIobal institute for Food Security University of Saskatchewan Saskatoon S7N 4J8 Canada

The remodeling of root architecture is a major developmental response of plants to phosphate （Pi） deficiency and is thought to enhance a plant＇s ability to forage for the available Pi in topsoil. The underlying mechanism controlling this response, however, is poorly understood. In this study, we identified an Arabidopsis mutant, hps 10 （hypersensitive to Pi starvation 10）, which is morphologically normal under Pi sufficient condition but shows increased inhibition of primary root growth and enhanced production of lateral roots under Pi defi- ciency, hpslO is a previously identified allele （als3-3） of the ALUMINUM SENSITIVE3 （ALS3） gene, which is involved in plant tolerance to aluminum toxicity. Our results show that ALS3 and its interacting protein AtSTAR1 form an ABC transporter complex in the tonoplast. This protein complex mediates a highly electro- genic transport in Xenopus oocytes. Under Pi deficiency, als3 accumulates higher levels of Fe3＋ in its roots than the wild type does. In Arabidopsis, LPR1 （LOW PHOSPHATE ROOT1） and LPR2 encode ferroxidases, which when mutated, reduce Fe3＋ accumulation in roots and cause root growth to be insensitive to Pi defi- ciency. Here, we provide compelling evidence showing that ALS3 cooperates with LPR1/2 to regulate Pi deficiency-induced remodeling of root architecture by modulating Fe homeostasis in roots.

关键词： phosphate deficiency root architecture iron homeostasis ABC transporter ALUMINUM SENSI-TIVE3 AtS TAR1

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Single-cell technologies:From research to application

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The Innovation 2022年第6期3卷 92-108页

作者： Lu Wen Guoqiang Li Tao Huang Wei Geng Hao Pei Jialiang Yang Miao Zhu Pengfei Zhang Rui Hou Geng Tian Wentao Su Jian Chen Dake Zhang Pingan Zhu Wei Zhang Xiuxin Zhang Ning Zhang Yunlong Zhao Xin Cao Guangdun Peng Xianwen Ren Nan Jiang Caihuan Tian Zi-Jiang Chen Biomedical Pioneering Innovation Centre(BIOPIC) Peking UniversityBeijing 100871China Bio-Med Big Data Center CAS Key Laboratory of Computational BiologyShanghai Institute of Nutrition and HealthChinese Academy of SciencesShanghai 200031China School of Chemical Engineering and Technology Sun Yat-Sen UniversityZhuhai 519082China Mozhuo Biotech(zhejiang)Co. Ltd.TongxiangJiaxing 314500China Geneis(Beijing)Co. Ltd.Beijing 100102China Guangzhou institutes of Biomedicine and Health Chinese Academy of SciencesGuangzhou 510530China Department of Medical Oncology Zhongshan HospitalFudan UniversityShanghai 200032China School of Food Science and Technology Dalian Polytechnic UniversityDalian 116034China State Key Laboratory of Transducer Technology Aerospace Information Research InstituteChinese Academy of SciencesBeijing 100190China Key Laboratory of Biomechanics and Mechanobiology Ministry of EducationBeijing Advanced Innovation Center for Biomedical EngineeringSchool of Engineering MedicineBeihang UniversityBeijing 100083China Department of Mechanical Engineering City University of Hong KongHong Kong 999077China Chongqing institute of Green and Intelligent Technology Chinese Academy of SciencesChongqing 400714China Center of Peony Institute of Vegetables and FlowersChinese Academy of Agricultural SciencesKey Laboratory of Biology and Genetic Improvement of Flower Crops(North China)Key Laboratory of Biology and Genetic Improvement of Horticultural CropsMinistry of Agriculture and Rural AffairsBeijing 100081China Department of Hepatic Surgery Fudan University Shanghai Cancer CenterShanghai 200032China Advanced Technology institute University of SurreyGuildfordSurreyGU27XHUK National Physical Laboratory TeddingtonMiddlesex TW110LWUK institute of Clinical Science Zhongshan HospitalFudan UniversityShanghai 200032China West China School of Basic Medical Sciences&Forensic Medicine Sichuan UniversityChengdu 610041China Jinfeng Laboratory Chongqing 401329China Center for Reproduc

In recent years,more and more single-cell technologies have been developed.A vast amount of single-cell omics data has been generated by large projects,such as the Human Cell Atlas,the Mouse Cell Atlas,the Mouse RNA Atlas,the Mouse ATAC Atlas,and the Plant Cell *** on these single-cell big data,thousands of bioinformatics algorithms for quality control,clustering,cell-type annotation,developmental inference,cell-cell transition,cell-cell interaction,and spatial analysis are *** powerful experimental single-cell technology and state-of-the-art big data analysis methods based on artificial intelligence,the molecular landscape at the single-cell level can be revealed.

关键词： Atlas landscape application

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Vegetation context modifies selection on flowering start and plant height in an orchid perennial herb

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Journal of Plant Ecology 2021年第5期14卷 934-944页

作者： Yun Wu Xu-Yu Duan Guang-Li Liu Yong Xiang Bo Shu Qing-Jun Li School of Civil Engineering Architecture and EnvironmentXihua UniversityChengduChina Yunnan Key Laboratory of Plant Reproductive Adaptation and Evolutionary Ecology Yunnan UniversityKunmingChina Laboratory of Ecology and Evolutionary Biology School of Ecology and Environmental ScienceYunnan UniversilyKunmingChina South-East Tibetan Plateau Station for Integrated Observation and Research of Alpine Environment Institute of Tibetan Plateau ResearchChinese Academy of SciencesBeijingChina College of landscape Architecture Sichuan Agricultural UniversityChengduChina

Aims Identifying the potential role of vegetation context(defined as the density,species identity/diversity and height of co-occurring plants)in modifying selection on floral traits is a critical step for clarifying and predicting the floral evolutionary trajectory in complex co-flowering species competition *** is also helpful to understand the variation in pollinator-mediated *** We experimentally reduced vegetation height around individual plants of Spiranthes sinensis(a bumblebee-pollinated perennial orchid herb)and estimated how vegetation context modified selection on four floral traits(flowering start,plant height,corolla size and number of flowers)through female function and pollen removal over two continuous *** randomly selected independent plants in each *** Findings We demonstrated that vegetation context modified selection for earlier flowering start and shorter plant height of *** strength of selection differed between *** addition,selection was stronger through female function than through pollen *** findings indicate the potential role of vegetation context in shaping the differentiation and diversification of flowers in angiosperms.

关键词： female fitness and pollen removal floral evolution Spiranthes sinensis temporal variation vegetation context modification selection

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Science Letters:Optimization of soybean (Glycine max (L.) Merrill) in planta ovary transformation using a linear minimal gus gene cassette

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Journal of zhejiang University-Science B(Biomedicine & Biotechnology) 2009年第12期10卷 870-876页

作者： Ming LIU Jun YANG Yun-qing CHENG Li-jia AN Department of Bioscience and Biotechnology Dalian University of Technology Dalian 116024 China institute of Eco-environmental Sciences Jilin Normal University Siping 136000 China

Soybean transformation by ovary-drip was improved by optimizing the length of the transformation pathway by cutting the styles. These modifications facilitated soybean transformation manipulation and improved transformation reproducibility and efficiency. Using a linear minimal gus gene cassette as the foreign DNA, a maximum transformation frequency of 11% was obtained in flowers of the soybean cultivar ‘Liaodou 14’ with their styles mostly removed, whereas removal of only the stigma, partial style cutting and partial ovary cutting gave transformation frequencies of 0%, 1%, and 2%, respectively. An average transformation frequency of 8.2% was obtained when 619 flowers from three soybean cultivars (‘Liaodou 14’, ‘Liaodou 13’, and ‘Tiefeng 29’) were transformed by this optimized method. Southern blotting analysis showed that the gus reporter gene (encoding β-glucuronidase) was stably inherited with a simple pattern. Reverse transcription-polymerase chain reaction (RT-PCR) and GUS staining confirmed the expression of the gus gene in transgenic plants.

关键词： Soybean Glycine max (L.) Merrill In planta ovary-drip transformation Optimization Style cutting Minimal gus renorter gene cassette

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