is involved in both dehydration- and low-temperatureresponsive gene expression has recently been analyzed extensively (for reviews see [1,5,6]).
|
|
- Dorothy Harrison
- 9 years ago
- Views:
Transcription
1 EB/CBF Molecular responses to dehydration and low temperature: differences and cross-talk between two stress signaling pathways Kazuo Shinozaki* and Kazuko Yamaguchi-Shinozaki Recently, a major transcription system that controls abscisicacid-independent gene expression in response to dehydration and low temperature has been identified. The system includes the DRE/CRT (dehydration-responsive element/c-repeat) cis-acting element and its DNA-binding protein, D (DRE-binding protein/c-repeat binding factor), which has an AP2 domain. D DREB1/CBF and DREB2, which are induced by cold and dehydration, respectively, and control the expression of various genes involved in stress tolerance. Recent studies are providing evidence of differences between dehydrationsignaling and cold-stress-signaling cascades, and of cross-talk between 8686, Japan; them. Addresses *Laboratory of Plant Molecular Biology, Tsukuba Life Science Center, Institute of Physical and Chemical Research (RIKEN), Koyadai, Tsukuba, Ibaraki , Japan; sinozaki@rtc.riken.go.jp Biological Resources Division, Japan International Research Center for Agricultural Sciences (JIRCAS), Ministry of Agriculture, Forestry and Fisheries, 2-1 Ohwashi, Tsukuba, Ibaraki kazukoys@jircas.affrc.go.jp Current Opinion in Plant Biology 2000, 3: /00/$ see front matter 2000 Elsevier Science Ltd. All rights reserved. Abbreviations ABA PSIS abscisic acid aba ABA-deficient abi ABA-responsive ABRE ABA-responsive element ATHK1 ARABIDO TWO-COMPONENT HISTIDINE KINASE bp base pairs CaMV cauliflower mosaic virus CBF C-repeat-binding factor cor cold-regulated CRT C-repeat DRE dehydration-responsive element DREB DRE-binding protein erd early responsive to dehydration EREBP ethylene-responsive element binding protein HOS HIGHEXPRESSIONOFOSMOTICALLYSENSITIVE kin cold-inducible lti low-temperature induced rd responsive to dehydration sfr sensitivity to freezing Introduction Among abiotic environmental stresses, drought and low temperature affect plant growth most seriously. Plants respond to dehydration and low temperature with a number of physiological and developmental changes. Molecular and cellular responses to these stresses have been analyzed extensively at the biochemical level: various kinds of proteins and smaller molecules, including sugars, proline, and glycine betaine, accumulate; in addition, many genes are induced by both dehydration and cold, but some respond either only to drought or only to cold. These observations suggest the existence of several cellular signal transduction pathways between the perception of stress signals and gene expression. Drought and high salinity cause plants to produce high levels of ABA; exogenous application of ABA also induces a number of genes that respond to dehydration and cold stress [1]. Nevertheless, the role of ABA in low-temperature-responsive gene expression is not clear. Several reports have described genes that are induced by dehydration and low temperature but that do not respond to exogenous abscisic acid (ABA) treatment [1 4]. It is likely, therefore, that both ABA-independent and ABA-dependent signal transduction cascades exist [1,4]. One of the transcription systems that function independently of ABA in both dehydration- and low-temperature-responsive gene expression has recently been analyzed extensively (for reviews see [1,5,6]). In 1994, we indentified a cis-acting dehydration-responsive element (DRE) [7]. A similar cis-acting element has also been reported and named C-repeat (CRT) [8] or low-temperature-responsive element [9]. The DRE/CRT element is involved in both dehydration- and low-temperatureresponsive gene expression. Our short review focuses on roles of the DRE/CRT cis-element and its DNA-binding protein, DREB/CBF (DRF-binding protein/c-repeatbinding factor), in the separation of and cross-talk between two stress signals that are involved in stress-induced gene expression in Arabidopsis. We also discuss the role of ABA in dehydration- and cold-induced gene expression. ABA plays important roles in slow and adaptive responses involving dehydration-induced gene expression. However, ABA seems not to be important in cold-induced gene expression and does not accumulate in response to low temperature. ABA does, however, have important roles in slow adaptive processes during dehydration stress. Similar genes are induced by dehydration and cold stress A variety of genes are induced by both dehydration and low temperature, and their mrna levels are subsequently reduced by release from stress conditions. This suggests that similar biochemical processes function in dehydration- and cold-stress responses. Genes induced in plants that are subjected to these stresses are thought to function not only in protecting cells by producing important metabolic proteins and cellular protectants, but also in regulating genes that are involved in transducing the stress response signal [1,2,10,11]. In Arabidopsis, these genes include rd (responsive to dehydration), erd (early responsive to dehydration), cor(cold-regulated), lti(low-temperature induced),
2 Figure 1 Signal perception Signal transduction Transcription factor Cis element Gene expression Dehydration aba MYC/MYB Osmotic change ABA abi1, abi2 era1 bzip hos5 sfr1 hos1 hos2 (a) (b) (c) (d) (e) (f) MYCR/MYBR ABRE erd1 rd22 rd29b DREB2 Low temperature Temperature change DRE/CRT esk1 Stress response and stress tolerance Current Opinion in Plant Biology CBF/DREB1 rd29a/lti78, cor15a Cellular signal transduction pathways between the initial drought-stress or cold-stress signal and gene expression in Arabidopsis. There are at least six signal transduction pathways: two (b,c) are ABA-dependent and four (a,d,e,f) are ABA-independent. Stressinducible genes rd29a/cor78/lti78, rd29b/lti65, rd22, and erd1 have been used to analyse the regulation of gene expression and the signalling process [5,40]. abi1, abi2, and era1 are involved in ABA signaling [41 44]. hos5 functions in DREB2-related dehydration signaling, and sfr6, hos1, and hos2 function in DREB1/CBF-related cold signaling [28,32 34]. esk1 is involved in responses to cold via a DRE-independent process [28 ]. Thin and thick arrows represent the minor and major signalling pathways that are involved in dehydrationresponsive gene expression, respectively. Broken arrows represent the signalling pathways that are involved in low temperature stress responses. and kin (cold-inducible). This variety of stress-inducible genes suggests that the responses of plants to dehydration and cold are complex. Some of the stress-inducible genes are overexpressed in transgenic plants that have enhanced stress tolerance, suggesting that their gene products function in stress tolerance [10 13]. Regulation of gene expression by dehydration and cold stress Most dehydration-inducible genes also respond to cold stress, and, conversely, most cold-inducible genes respond to dehydration. Analyses of the expression patterns of genes induced by both dehydration and cold have revealed broad variation in the timing of their induction and differences in their responsiveness to ABA [4]. Many of the genes that are induced by exogenous ABA treatment are also induced by cold or dehydration in ABA-deficient (aba) or ABA-insensitive (abi) Arabidopsis mutants [2]. These observations indicate that these genes are not induced by the accumulation of endogenous ABA, but respond to ABA [1,4]. Several ABA-inducible genes require protein biosynthesis for their induction by ABA [4], which suggests that at least two independent pathways signal the expression of stress-induced genes in response to endogenous ABA production. As shown in Figure 1, at least four independent signal pathways function under drought conditions [4]: two are ABA-independent and two are ABA-dependent. In DRE/CRT addition, two ABA-independent pathways are also involved in low-temperature-responsive gene expression [1]. There is a common signal transduction pathway between dehydration and cold stress involving the DRE/CRT cis-acting element, and two additional signal transduction pathways may function only in dehydration or in cold response. The role of the cis-acting element in ABA-independent gene expression In aba or abi mutants, many genes are induced by both dehydration and low temperature; this suggests that these genes do not require ABA for their expression under cold or drought conditions but that they do respond to ABA. Among these genes, the expression of two dehydrationand cold-inducible Arabidopsis genes, rd29a/lti78/cor78 and cor15a, has been analyzed in detail (for reviews see [1,5]). The transcription of rd29a in abi1 and aba1 mutants suggests that cold- and drought-regulated expression does not require ABA. DRE, a 9-base pair (bp) conserved sequence (i.e. TACCGACAT), is an essential cis-acting transcription element factors for disting the regulation of rd29a induction in the ABA-independent response to dehydration and cold [7]. Similar motifs, called CRT and low-temperature-responsive element, which include the CCGAC motif that forms the core of the DRE sequence, have been found in the promoter region of coldinducible genes [8,9]. DREB/CBF between the dehydration and low temperature stress-signaling pathways Protein factors that specifically interact with the 9-bp DRE sequence have been detected in nuclear extracts prepared from either dehydrated or adequately watered Arabidopsis plants [7]. Stockinger et al. [14] first isolated a cdna clone for a DRE/CRT-binding protein using yeast one-hybrid screening; they named this clone CBF1 (CRT-binding factor 1). In yeast, CBF1 functions as a transcription factor that upregulates DRE/CRT-dependent transcription. It contains a conserved DNA-binding motif (AP2 domain) that is also found in the EREBP (ethylene-responsive element binding protein) family and AP2 protein, which is involved in floral
3 Figure 2 A model of the induction of the rd29a/cor78/lti78 gene and cis- and transacting elements involved in stress-responsive gene expression. Two cis-acting elements, DRE/CRT and ABRE, are involved in the ABA-independent and ABA-responsive induction of rd29a, respectively. Two different DRE/CRT-binding proteins, DREB1/CBF1 and DREB2, distinguish two different signal transduction pathways in response to cold and drought stresses, respectively [17 ]. DRE/CRT-binding proteins contain an AP2 DNA-binding domain, whereas ABRE-binding proteins encode bzip transcription factors. Thick broken arrows represent a cold signalling pathway. Solid thick arrows and thin broken arrows represent an ABA-independent and an ABA-dependent signalling pathway, respectively, that are involved in the dehydration response. Signal perception Transduction Transcription Trans elements rd29a promoter Cis elements Low temperature Temperature change ABA independent DREB1/CBF genes CBF/DREB1 DREB2 genes Modification DRE/CRT ABA independent DREB2 Dehydration Osmotic change ABA biosynthesis bzip ABRE ABA signaling TATA Current Opinion in Plant Biology morphogenesis [15,16]. Independently, Liu et al. [17 ] isolated five independent cdnas for DRE/CRT-binding proteins using yeast one-hybrid screening, which they named DREBs (DRE-binding proteins). All of the DREBs also contain a conserved AP2 domain. The five cdna clones that encode DRE/CRT-binding proteins are classified into two groups, DREB1 and DREB2. The groups contain similar AP2 domains but have low sequence similarity outside that domain. There are three DREB1 proteins that are encoded by genes that lie in tandem on chromosome 4 in the order DREB1B, DREB1A, and DREB1C [18]. DREB1B is identical to CBF1. Gilmour et al. [19] also isolated two CBF1 homologues named CBF2 and CBF3. There are two DREB2 proteins, DREB2A and DREB2B [17 ]. Both DREB1A and DREB2A bind specifically to DRE/CRT and function as transcriptional activators in plant protoplasts, as well as in yeasts. Expression of the DREB1A/CBF3 gene and its two homologues (i.e. DREB1B/CBF1 and DREB1C/CBF2) is induced by lowtemperature stress, whereas expression of the two DREB2 genes is induced by dehydration. These results suggest that the DREB1 proteins are involved in cold-specific gene expression, whereas the DREB2 proteins function in dehydration-specific gene expression (Figure 2). The AP2 domain is found in many plant genes, such as EREBP, APETALA2, AINTEGUMENTA and TINY [16]. EREBPs bind to the ethylene-responsive element (i.e. the GCC box, GCCGCC), whereas DREB/CBFs bind to the DRE/CRT core sequence, PuCCGAC. DRE/CRT and the G box contain PuCCGNC as a common sequence [15,17 ]. Liu et al. [17 ] DREB/CBF showed that DREB/CBF and EREBPs have two different amino acids in the AP2 domain, which may confer different specificity for the DNA-binding of cis-acting elements [17 ]. Engineering stress tolerance of transgenics by overexpressing Jaglo-Ottosen et al. [20 ] found that overexpressing CBF1, under the control of the CaMV 35S promoter, in transgenic Arabidopsis not only induced strong expression of cor genes, but also improved freezing tolerance. The growth of these transgenics was similar to that of wild-type plants under normal growth conditions. Liu et al. [17 ] and Kasuga et al. [21 ] also observed that enhanced expression of the target cor, rd and erd genes in transgenic Arabidopsis plants that overexpress DREB1A/CBF3 (also under the control of the CaMV 35S promoter) produced dwarfed or growth-retarded phenotypes in unstressed conditions. The DREB1A transgenic plants also had enhanced freezing and dehydration tolerance. The difference in growth retardation caused by overexpression of DREB1A/CBF3 and DREB1B/CBF1 may be explained by different levels of expression of the two transgenes or the difference in the genes used. In contrast, overexpression of DREB2A cdna induced weak expression of the target genes under unstressed conditions and caused slight growth retardation of the transgenic plants [17 ]. DREB2 proteins are probably post-transcriptionally activated in dry conditions (Figure 2). These results indicate that two independent families of DREBs, DREB1/CBF and DREB2, function as trans-acting factors in two separate signal-transduction pathways under cold and dry conditions, respectively (Figure 2). As discussed above, overproduction of DREB1A/CBF3 cdnas driven by the 35S CaMV promoter in transgenic plants causes severe growth retardation under normal growth conditions [17,21 ]. Recently Kasuga et al. [21 ] found that the DREB1A cdna driven by the stress-inducible rd29a promoter was expressed at a low level in unstressed control conditions and at a high level in plants exposed to dehydra-
4 Figure 3 Low temperature CBF CRT (DREB1 DRE) system Gene expression Dehydration DREB2 DRE system ABA biosynthesis Time course bzip ABRE system Gene expression Protein synthesis Gene expression REB/CBF genes in Signal perception Rapid and emergency response Slow and adaptive response Current Opinion in Plant Biology Molecular responses to dehydration and low temperature based on stepwise gene expression. The regulation of D response to dehydration and low temperature occurs early in the stress response. DRE/CRT-dependent transcription follows the response. ABA biosynthesis is induced by dehydration and activates two regulatory ABA-dependent gene expression systems. One is the bzip/abre system; the other is the MYC/MYB system, which requires do novo production of the MYB and MYC proteins in response to ABA. tion, salt, and cold stresses. The rd29a promoter minimized the negative effects on the growth of the transgenic plants. Moreover, this stress-inducible promoter enhanced tolerance of drought, salt, and freezing to a greater extent than did the CaMV 35S promoter. The rd29a promoter::dreb1a system is a self-amplifying system that overexpresses DREB1A protein throughout exposure to stress. Greater expression of the DREB1A protein in the rd29a::dreb1a transgenics results in greater expression of the target genes involved in stress tolerance [21 ]. This system provides some promise for engineering multi-stress tolerance of transgenic crops because plants such as tobacco, Brassica, and rice, have similar transcription systems to that of Arabidopsis. ABA in dehydration and low-temperature stress response In many plants, endogenous ABA levels increase significantly in conditions of drought and high-salinity [2 4]. In Arabidopsis, however, ABA levels increase only transiently in response to low-temperature stress before returning to their basal level [1,22]. Many drought- and cold-stress-inducible genes are induced by exogenous ABA treatment. These genes contain potential ABA-responsive elements (ABREs; PyACGTGGC) in their promoter regions [1,2]. In Arabidopsis, the rd29b(or lti65) gene is induced by dehydration and high salinity, but not by cold stress [23]. rd29bdoes not contain a DRE/CRT but contains two ABREs in its promoter [7]. It is controlled downstream of the abi1 and aba1 mutations and so endogenous ABA that accumulates in response to dehydration induces its expression. During cold stress, endogenous ABA is not sufficient to induce rd29b. We therefore believe that the ABA-signaling pathway is not important in cold-stress responses. The endogenous ABA accumulation that has been observed during winter may be attributable to the dehydration of plants, which induces DRE/CRT-dependent gene expression of most cor, lti, and rdgenes to confer stress tolerance (Figure3). The rd29a promoter contains not only DRE but also an ABRE (Figure 2). An ABRE cis-acting element and bzip transcription factors function in ABA-responsive gene expression [7]. The rd29a gene is therefore controlled by three independent regulatory systems [7,17 ]. These results indicate that complex molecular responses to various environmental stresses may be mediated by both complex regulatory systems of gene expression and signal transduction, and by cross-talk between these systems. The bzip/abre system seems to function after the accumulation of endogenous ABA in drought conditions (Figure 3). The biosynthesis of novel protein factors is necessary for the expression of ABA-inducible genes in one of the two ABAdependent pathways (Figure 1). The induction of the Arabidopsis drought-inducible gene rd22 is mediated by ABA and requires protein biosynthesis for its ABA-dependent expression [24]. MYC and MYB recognition sequences are essential for the ABA- and drought-responsive expression of rd22, and ABA-inducible MYC and MYB proteins may function cooperatively in the ABA-dependent expression of rd22 [25,26]. This MYC/MYB system may also function in a slow and adaptive stress response process. The different timing of the induction of stress-inducible genes may be explained by the different regulatory systems that function in their promoters, such as DRE/CRT, ABRE, and MYB/MYC (Figure 3). Genetic analysis of signal transduction in response to dehydration and cold stress Many Arabidopsismutants that are either sensitive to or tolerant of freezing have been isolated, and their phenotypes
5 have been analyzed in detail. Warren et al. [27] isolated five freezing-sensitive (sensitivity to freezing [sfr]) mutants and mapped their positions on Arabidopsis chromosomes. Knight et al. [28 ] analyzed the expression of kin1, cor15a, and cor78/rd29a, which contain DRE/CRT in their promoters. In the sfr6 mutant, unlike wild-type Arabidopsis plants, these genes were not strongly induced in response to osmotic stress and low temperature. In contrast, ATP5CS, CBF1, CBF2, and CBF3, which do not contain DRE/CRT in their promoters, were not affected in the sfr6 mutant. The SFR6 product may be involved upstream of CBF/DREB1 and function as a positive regulator of this element (Figure 1). A transient increase in cellular calcium ion concentration in responses to dehydration and low temperature seems to stimulate cellular signalling processes. Nevertheless, evidence from calcium measurement in cold conditions suggests that calcium signalling may not be involved in sfr6 signalling. Xin and Browse [29 ] isolated a constitutively freezing-tolerant mutant named eskimo1 that, without cold acclimatization, has greater freezing tolerance than wildtype plants. The molecular mechanism of the stress tolerance of eskimo1 is not known, but proline accumulates in the eskimo1 mutant [29 ]. This proline may be involved in stress tolerance as proline functions in osmoprotection, in detoxication of active oxygen, and in protection of proteins and nucleic acids. Recently, Nanjo et al. [30] showed that the accumulation of proline in transgenics with antisense cdna for proline dehydrogenase provides strong freezing tolerance as well as salt tolerance. In eskimo1, the expression of cor15a, cor47/rd17, and cor78/rd29a remain low under normal growth conditions. This suggests that ESKIMO1 functions in a different signal transduction pathway from the DRE/CRT system (Figure 1). Genetic analysis of Arabidopsis mutants with the rd29a promoter::luciferase transgene suggests complex signaling pathways in drought-, salt-, and cold-stress responses. Ishitani et al. [31] isolated mutants that overexpressed rd29a or repressed it in response to dehydration, high salinity, cold and ABA. They propose that the ABA- and stress-signaling pathways are not independent and that the various stress-signaling pathways, including ABA-independent and ABA-dependent pathways, are not completely independent. Xiong et al. [32] isolated a hos5 mutant that has increased expression of rd29a when under osmotic stress but not when experiencing cold stress. Genetic analysis showed that HOS5 is a negative regulator of osmotic-stress-responsive gene expression. Ishitani et al. [33] and Lee et al. [34] isolated hos1 and hos2 mutants, respectively, that enhanced the expression of rd29a, cor47, cor15a, and kin1. Non-acclimatized hos1 and hos2 mutants were less cold-hardy than wild-type plants. HOS1 and HOS2 are therefore thought to function as negative regulators of a cold-specific signal transduction pathway (Figure 1). Genetic analysis of these mutants and of stressresistant or stress-sensitive mutants is likely to provide more information on stress-induced signal transduction [35]. Signal perception and signal transduction Signal transduction pathways involved in the drought-stress response have been studied in yeast and animal systems [5]. Two-component systems seem to function in sensing osmotic stress in plants as well as in bacteria and yeast [36]. Recently, Urao et al. [37 ] isolated an Arabidopsis cdna that encodes a two-component histidine kinase (ATHK1), which functions as an osmosensor in yeast. ATHK1 has a typical histidine kinase domain, a receiver domain, and two transmembrane domains in the amino-terminal domain. ATHK1 might function in signal perception during dehydration stress in Arabidopsis, but sensors for cold stress have not yet been identified. Many genes encoding factors that are involved in signaltransduction cascades are upregulated by dehydration and cold: mitogen-activated protein (MAP) kinases, calcium-dependent protein kinases, and enzymes involved in phospholipid metabolism, such as phospholipase C and phosphatidyl-4, 5-phosphate 5-kinase (PIP5) kinase [5,38,39]. These signaling factors might be involved in the amplification of stress signals and in the adaptation of plant cells to drought-stress conditions. No direct evidence has, however, been obtained of the functions of these signaling molecules. Transgenic plants that modify the expression of these genes and mutants with disrupted genes will provide more information on the function of their gene products. Conclusions and perspectives A major transcription system regulating ABA-independent gene expression in response to dehydration and cold stress includes a DRE/CRT cis-acting element and its DNA-binding protein, DREB/CBF. The DREB/CBF family of proteins contain two subclasses, DREB1/CBF and DREB2, which are induced by cold and dehydration, respectively, to express various genes involved in stress tolerance. Cross-talk between dehydration and cold occurs at the transcriptional level. ABA plays important roles in the dehydration-stress response but not, it seems, in the cold-stress response. Genetic analysis of stress-resistant or stress-sensitive mutants, and mutants with the rd29a promoter::luciferase transgene, should provide more information on signal transduction in response to dehydration and cold stress. Sequencing of the Arabidopsis genome will be completed by the end of 2000, which means that the structure of all 25,000 Arabidopsis genes will soon be determined. All stress-inducible genes can then be identified by the systematic analysis of gene expression in microarrays. In the next decade, we think it important to develop novel methods to analyze the complex networks that control the stress responses of higher plants. A reverse genetic approach, as well as classical forward genetics, will become more important for understanding not only the functions of stress-inducible genes but also the complex signaling processes of the dehydration- and cold-stress responses.
6 Acknowledgements Our work is supported in part by the Program for Promotion of Basic Research Activities for Innovative Biosciences and the Special Coordination Fund of Science and Technology Agency of Japan. References and recommended reading Papers of particular interest, published within the annual period of review, have been highlighted as: of special interest of outstanding interest 1. Thomashow MF: Plant cold acclimation: freezing tolerance genes and regulatory mechanisms. Annu Rev Plant Physiol Plant Mol Biol 1999, 50: Ingram J, Bartels D: The molecular basis of dehydration tolerance in plants. Annu Rev Plant Physiol Plant Mol Biol 1996, 47: Bray EA: Plant responses to water deficit. Trends Plant Sci 1997, 2: Shinozaki K, Yamaguchi-Shinozaki K: Gene expression and signal transduction in water-stress response. Plant Physiol 1997, 115: Shinozaki K, Yamaguchi-Shinozaki K: Molecular responses to drought stress. In Molecular Responses to Cold, Drought, Heat and Salt Stress in Higher Plants. Edited by Shinozaki K, Yamaguchi- Shinozaki Y. Austin, Texas: RG Landes; 1999: Thomashow MF, Browse J: Plant cold tolerance. In Molecular Responses to Cold, Drought, Heat and Salt Stress in Higher Plants. Edited by Shinozaki K, Yamaguchi-Shinozaki Y. Austin, Texas: RG Landes; 1999: Yamaguchi-Shinozaki K, Shinozaki K: A novel cis-acting element in an Arabidopsis gene is involved in responsiveness to drought, low-temperature, or high-salt stress. Plant Cell 1994, 6: Baker SS, Wilhelm KS, Thomashow MF: The 5 -region of Arabidopsis thaliana cor15a has cis-acting elements that confer cold-, drought- and ABA-regulated gene expression. Plant Mol Biol 1994, 24: Nordin K, Vahala T, Palva ET: Differential expression of two related low-temperature-induced genes in Arabidopsis thaliana (L.) Heynh. Plant Mol Biol 1993, 21: Steponkus PL, Uemura M, Joseph RA, Gilmour SJ, Thomashow MF: Mode of action of the COR15a gene on the freezing tolerance of Arabidopsis thaliana. Proc Natl Acad Sci USA 1998, 95: Bohnert HJ, Su H, Shen B: Molecular mechanisms of salinity tolerance. In Molecular Responses to Cold, Drought, Heat and Salt Stress in Higher Plants. Edited by Shinozaki K, Yamaguchi- Shinozaki Y. Austin, Texas: RG Landes; 1999: Holmberg N, Bülow L: Improving stress tolerance in plants by gene transfer. Trends Plant Sci 1998, 3: Thomashow MF: Role of cold-responsive genes in plant freezing tolerance. Plant Physiol 1998, 118: Stockinger EJ, Gilmour SJ, Thomashow MF: Arabidopsis thaliana CBF1 encodes an AP2 domain-containing transcription activator that binds to the C-repeat/DRE, a cis-acting DNA regulatory element that stimulates transcription in response to low temperature and water deficit. Proc Natl Acad Sci USA 1997, 94: Ohme-Takagi M, Shinshi H: Ethylene-inducible DNA binding proteins that interact with an ethylene-responsive element. Plant Cell 1995, 7: Okamuro JK, Caster B, Villarroel R, Van Montagu M, Jofuku KD: P/AP2 The AP2 DNA domain binding of APETALA2 domain, separate defines two a large cellular new signal family of DNA binding domain in Arabidopsis. Proc Natl Acad Sci USA 1997, 94: Liu Q, Sakuma Y, Abe H, Kasuga M, Miura S, Yamaguchi-Shinozaki K, Shinozaki K: Two transcription factors, DREB1 and DREB2, with an EREB transduction pathways in drought- and low temperatureresponsive gene expression, respectively, in Arabidopsis. Plant Cell 1998, 10: Two DRE-binding proteins, DREB1 and DREB2, distinguish two stress-signal-transduction pathways involved in Arabidopsis responses to cold and dehydration stress, respectively. The important role of DREB transcription factors in cross-talk between the two stress signaling pathways is demonstrated. Constitutive overexpression of DREB1A in transgenics results in acquired tolerance of drought and freezing but also causes growth retardation under non-stressful conditions. 18. Shinwari ZK, Nakashima K, Miura S, Kasuga M, Seki M, Yamaguchi Shinozaki K, Shinozaki K: An Arabidopsis gene family encoding DRE/CRT binding proteins involved in low-temperatureresponsive gene expression. Biochem Biophys Res Commun 1998, 250: Gilmour SJ, Zarka DG, Stockinger EJ, Salazar MP, Houghton JM, Thomashow MF: Low temperature regulation of the Arabidopsis CBF family of AP2 transcriptional activators as an early step in cold-induced COR gene expression. Plant J 1998, 16: Jaglo-Ottosen KR, Gilmour SJ, Zarka DG, Schabenberger O, Thomashow MF: Arabidopsis CBF1 overexpression induces cor genes and enhances freezing tolerance. Science 1998, 280: The authors report that over-expression of CBF1 under the control of the CaMV 35S promoter in Arabidopsis transgenics not only induces strong expression of cor genes but also improves freezing tolerance. This is the first proposed application of regulon biotechnology to molecular breeding for stress tolerance. 21. Kasuga M, Liu Q, Miura S, Yamaguchi-Shinozaki K, Shinozaki K: Improving plant drought, salt, and freezing tolerance by gene transfer of a single stress-inducible transcription factor. Nat Biotech 1999, 17: Overexpression of DREB1A by the rd29a stress-inducible promoter enhances tolerance to drought, salt, and freezing at a higher level than does the CaMV 35S promoter, and minimizes negative effects on the growth of the transgenic plants. This rd29a promoter::dreb1a system is useful for developing multistress-tolerant transgenics. The DREB technology is useful for molecular breeding of stress-tolerant crops. 22. Lang V, Mantyla E, Welin B, Sunberg B, Palva ET: Alterations in water status, endogenous abscisic acid content, and expression of rab18 gene during the development of freezing tolerance in Arabidopsis thaliana. Plant Physiol 1994, 104: Yamaguchi-Shinozaki K, Shinozaki K: Characterization of the expression of a desiccation-responsive rd29a gene of Arabidopsis thaliana and analysis of its promoter in transgenic plants. Mol Gen Genet 1993, 236: Yamaguchi-Shinozaki K, Shinozaki K: The plant hormone abscisic acid mediates the drought-induced expression but not the seedspecific expression of rd22, a gene responsive to dehydration stress in Arabidopsis thaliana. Mol Gen Genet 1993, 138: Abe H, Yamaguchi-Shinozaki K, Urao T, Iwasaki T, Hosokawa D, Shinozaki K: Role of Arabidopsis MYC and MYB homologs in drought- and abscisic acid-regulated gene expression. Plant Cell 1997, 9: Urao T, Yamaguchi-Shinozaki K, Urao S, Shinozaki K: An Arabidopsis myb homolog is induced by dehydration stress and its gene product binds to the conserved MYB recognition sequence. Plant Cell 1993, 5: Warren G, McKown R, Martin AL, Teutonico R: Isolation of mutations affecting the development of freezing tolerance in Arabidopsis thaliana (L.) Heynh. Plant Physiol 1996, 111: Knight H, Veale EL, Warren GJ, Knight MR: The sfr6 mutation in Arabidopsis suppresses low-temperature induction of gene dependent on the CRT/DRE sequence motif. Plant Cell 1999, 11: Genetic and molecular analysis of freezing-sensitive mutants indicated that the SFR6 product may be involved upstream of the CBF/DREB1 system and function as a positive regulator. This mutant is useful in analyzing the signaling cascade that controls the CBF/DREB1 system. 29. Xin Z, Browse J: eskimo1 mutants of Arabidopsis are constitutively freezing-tolerant. Proc Natl Acad Sci USA 1998, 95: The authors report the freezing-tolerance (without cold acclimatization) of the eskimo1-imo-mutant. ESK1 functions in a different signaling process from that of the DRE/CRT system. The accumulation of proline in eskimo1 suggests an important role for proline in freezing tolerance. 30. Nanjo T, Kobayashi M, Yoshiba Y, Kakubari Y, Yamaguchi-Shinozaki K, Shinozaki K: Antisense suppression of proline degradation improves tolerance to freezing and salinity in Arabidopsis thaliana. FEBS Lett 1999, 461:
7 31. Ishitani M, Xiong L, Stevenson B, Zhu JK: Genetic analysis of osmotic and cold stress signal transduction in Arabidopsis: interactions and convergence of abscisic acid-dependent and abscisic acid-independent pathways. Plant Cell 1997, 9: Xiong L, Ishitani M, Lee H, Zhu JK: HOS5 a negative regulator of osmotic stress-induced gene expression in Arabidopsis thaliana. Plant J 1999, 19: Ishitani M, Xiong L, Lee H, Stevenson B, Zhu JK: HOS1, a genetic locus involved in cold-responsive gene expression in Arabidopsis. Plant Cell 1998, 10: Lee H, Xiong L, Ishitani M, Stevenson B, Zhu JK: Cold-regulated gene expression and freezing tolerance in an Arabidopsis thaliana mutant. Plant J 1999, 17: Koornneef M, Peeters AJM: Genetic approaches to abiotic stress responses. In Molecular Responses to Cold, Drought, Heat and Salt Stress in Higher Plants. Edited by Shinozaki K, Yamaguchi- Shinozaki Y. Austin, Texas: RG Landes; 1999: Urao T, Yamaguchi-Shinozaki K, Shinozaki K: Two-component systems in plant signal transduction. Trends Plant Sci 2000, 5: Urao T, Yakubov B, Satoh R, Yamaguchi-Shinozaki K, Seki M, Hirayama T, Shinozaki K: A transmembrane hybrid-type histidine kinase in Arabidopsis functions as an osmosensor. Plant Cell 1999, 11: The first report of the osmosensor histidine kinase ATHK1 in Arabidopsis. ATHK1 has similar structure and functions to those of yeast osmosensor Sln1. This evidence suggests an important role for the two-component system in osmotic stress perception. 38. Mizoguchi T, Ichimura K, Shinozaki K: Environmental stress response in plants: the role of mitogen-activated protein kinases (MAPKs). Trends Biotechnol 1997, 15: Mikami K, Katagiri T, Iuchi S, Yamaguchi-Shinozaki K, Shinozaki K: A gene encoding phosphatidylinositol-4-phosphate 5-kinase is induced by water stress and abscisic acid in Arabidopsis thaliana. Plant J 1998, 15: Nakashima K, Kiyosue T, Yamaguchi-Shinozaki K, Shinozaki K: A nuclear gene, erd1, encoding a chloroplast-targeted Clp protease regulatory subunit homolog is not only induced by water stress but also developmentally up-regulated during senescence in Arabidopsis thaliana. Plant J 1997, 12: Gosti F, Beaudoin N, Serizet C, Webb AA, Vartanian N, Giraudat J: ABI1 protein phosphatase 2C is a negative regulator of abscisic acid signaling. Plant Cell 1999, 11: Pei ZM, Ghassemian M, Kwak CM, McCourt P, Schroeder JI: Role of farnesyltransferase in ABA regulation of guard cell anion channels and plant water loss. Science 1999, 282: Grill E, Himmelbach A: ABA signal transduction. Curr Opin Plant Biol 1998, 1: Bonetta D, McCourt P: Genetic analysis of ABA signal transduction pathways. Trends Plant Sci 1998, 6:
The world of non-coding RNA. Espen Enerly
The world of non-coding RNA Espen Enerly ncrna in general Different groups Small RNAs Outline mirnas and sirnas Speculations Common for all ncrna Per def.: never translated Not spurious transcripts Always/often
More informationPlant Growth & Development. Growth Stages. Differences in the Developmental Mechanisms of Plants and Animals. Development
Plant Growth & Development Plant body is unable to move. To survive and grow, plants must be able to alter its growth, development and physiology. Plants are able to produce complex, yet variable forms
More informationHow To Understand How Gene Expression Is Regulated
What makes cells different from each other? How do cells respond to information from environment? Regulation of: - Transcription - prokaryotes - eukaryotes - mrna splicing - mrna localisation and translation
More informationSite-Directed Nucleases and Cisgenesis Maria Fedorova, Ph.D.
Site-Directed Nucleases and Cisgenesis Maria Fedorova, Ph.D. Regulatory Strategy Lead Enabling Technologies DuPont-Pioneer, USA 1 New Plant Breeding Techniques 2007 New Techniques Working Group established
More informationwww.mpiz-koeln.mpg.de Abt. Entwicklungsbiologie de Pflanzen
WEB ADDRESS: www.mpiz-koeln.mpg.de Forschung Abt. Entwicklungsbiologie de Pflanzen George Coupland How is the transition from vegetative growth to flowering controlled? - How is it regulated by environmental
More informationGenetics Lecture Notes 7.03 2005. Lectures 1 2
Genetics Lecture Notes 7.03 2005 Lectures 1 2 Lecture 1 We will begin this course with the question: What is a gene? This question will take us four lectures to answer because there are actually several
More informationChapter 3. Blue Plants: Transgenic Plants With The Gus Reporter Gene
Chapter 3 Blue Plants: Transgenic Plants With The Gus Reporter Gene Susan J. Karcher Department of Biological Sciences Purdue University West Lafayette, IN 47907-1392 phone: (765) 494-8083 fax: (765) 494-0876
More informationFONCTIONS GÉNOMIQUES DE LA FAMILLE DES FACTEURS DE TRANSCRIPTION AP2 CHEZ LES CÉRÉALES
UNIVERSITÉ DU QUÉBEC À MONTRÉAL FONCTIONS GÉNOMIQUES DE LA FAMILLE DES FACTEURS DE TRANSCRIPTION AP2 CHEZ LES CÉRÉALES THÈSE PRÉSENTÉE COMME EXIGENCE PARTIELLE DU DOCTORAT EN BIOLOGIE PAR MOHAMED BADAWI
More informationSickle cell anemia: Altered beta chain Single AA change (#6 Glu to Val) Consequence: Protein polymerizes Change in RBC shape ---> phenotypes
Protein Structure Polypeptide: Protein: Therefore: Example: Single chain of amino acids 1 or more polypeptide chains All polypeptides are proteins Some proteins contain >1 polypeptide Hemoglobin (O 2 binding
More informationJust the Facts: A Basic Introduction to the Science Underlying NCBI Resources
1 of 8 11/7/2004 11:00 AM National Center for Biotechnology Information About NCBI NCBI at a Glance A Science Primer Human Genome Resources Model Organisms Guide Outreach and Education Databases and Tools
More informationName Class Date. Figure 13 1. 2. Which nucleotide in Figure 13 1 indicates the nucleic acid above is RNA? a. uracil c. cytosine b. guanine d.
13 Multiple Choice RNA and Protein Synthesis Chapter Test A Write the letter that best answers the question or completes the statement on the line provided. 1. Which of the following are found in both
More informationModule 3 Questions. 7. Chemotaxis is an example of signal transduction. Explain, with the use of diagrams.
Module 3 Questions Section 1. Essay and Short Answers. Use diagrams wherever possible 1. With the use of a diagram, provide an overview of the general regulation strategies available to a bacterial cell.
More informationControl of Gene Expression
Control of Gene Expression (Learning Objectives) Explain the role of gene expression is differentiation of function of cells which leads to the emergence of different tissues, organs, and organ systems
More informationRecombinant DNA and Biotechnology
Recombinant DNA and Biotechnology Chapter 18 Lecture Objectives What Is Recombinant DNA? How Are New Genes Inserted into Cells? What Sources of DNA Are Used in Cloning? What Other Tools Are Used to Study
More informationGENE CLONING AND RECOMBINANT DNA TECHNOLOGY
GENE CLONING AND RECOMBINANT DNA TECHNOLOGY What is recombinant DNA? DNA from 2 different sources (often from 2 different species) are combined together in vitro. Recombinant DNA forms the basis of cloning.
More informationGene Regulation -- The Lac Operon
Gene Regulation -- The Lac Operon Specific proteins are present in different tissues and some appear only at certain times during development. All cells of a higher organism have the full set of genes:
More informationModeling and Simulation of Gene Regulatory Networks
Modeling and Simulation of Gene Regulatory Networks Hidde de Jong INRIA Grenoble - Rhône-Alpes Hidde.de-Jong@inria.fr http://ibis.inrialpes.fr INRIA Grenoble - Rhône-Alpes and IBIS IBIS: systems biology
More informationCCR Biology - Chapter 9 Practice Test - Summer 2012
Name: Class: Date: CCR Biology - Chapter 9 Practice Test - Summer 2012 Multiple Choice Identify the choice that best completes the statement or answers the question. 1. Genetic engineering is possible
More informationTransfection-Transfer of non-viral genetic material into eukaryotic cells. Infection/ Transduction- Transfer of viral genetic material into cells.
Transfection Key words: Transient transfection, Stable transfection, transfection methods, vector, plasmid, origin of replication, reporter gene/ protein, cloning site, promoter and enhancer, signal peptide,
More informationThesis of Ph.D. Novel cyclin-dependent kinase inhibitor protein from Medicago truncatula. Aladár Pettkó-Szandtner
Thesis of Ph.D Novel cyclin-dependent kinase inhibitor protein from Medicago truncatula Aladár Pettkó-Szandtner Supervisors: Horváth Gábor and Dudits Dénes Institute of Plant Biology HAS, Biologcal Research
More informationAP Biology Essential Knowledge Student Diagnostic
AP Biology Essential Knowledge Student Diagnostic Background The Essential Knowledge statements provided in the AP Biology Curriculum Framework are scientific claims describing phenomenon occurring in
More informationLecture 1 MODULE 3 GENE EXPRESSION AND REGULATION OF GENE EXPRESSION. Professor Bharat Patel Office: Science 2, 2.36 Email: b.patel@griffith.edu.
Lecture 1 MODULE 3 GENE EXPRESSION AND REGULATION OF GENE EXPRESSION Professor Bharat Patel Office: Science 2, 2.36 Email: b.patel@griffith.edu.au What is Gene Expression & Gene Regulation? 1. Gene Expression
More informationSUGAR BEET (Beta vulgaris L.) PROMOTERS FOR DIRECTED TISSUE- SPECIFIC ROOT TRANSCRIPTION
SUGAR BEET (Beta vulgaris L.) PROMOTERS FOR DIRECTED TISSUE- SPECIFIC ROOT TRANSCRIPTION Senthilkumar Padmanaban, Haiyan Li, David P. Puthoff and Ann C. Smigocki USDA-ARS Molecular Plant Pathology Laboratory,
More informationArabidopsis. A Practical Approach. Edited by ZOE A. WILSON Plant Science Division, School of Biological Sciences, University of Nottingham
Arabidopsis A Practical Approach Edited by ZOE A. WILSON Plant Science Division, School of Biological Sciences, University of Nottingham OXPORD UNIVERSITY PRESS List of Contributors Abbreviations xv xvu
More informationBiochemistry. Entrance Requirements. Requirements for Honours Programs. 148 Bishop s University 2015/2016
148 Bishop s University 2015/2016 Biochemistry The Biochemistry program at Bishop s is coordinated through an interdisciplinary committee of chemists, biochemists and biologists, providing students with
More informationNO CALCULATORS OR CELL PHONES ALLOWED
Biol 205 Exam 1 TEST FORM A Spring 2008 NAME Fill out both sides of the Scantron Sheet. On Side 2 be sure to indicate that you have TEST FORM A The answers to Part I should be placed on the SCANTRON SHEET.
More informationMechanism of short-term ERK activation by electromagnetic fields at mobile phone frequencies. Biochemistry Journal. August 1, 2007 405, pp.
Mechanism of short-term ERK activation by electromagnetic fields at mobile phone frequencies 1 Biochemistry Journal August 1, 2007 405, pp. 559 568 Joseph Friedman, Sarah Kraus, Yirmi Hauptman, Yoni Schiff
More informationEngineering of Yellow Mosaic Virus Resistance (YMVR) in Blackgram. Project ID: 1 April 2000 to 31 August 2004. Project Duration:
Engineering of Yellow Mosaic Virus Resistance (YMVR) in Blackgram ID: Duration: Coordinator in Switzerland: PS1 1 April 2000 to 31 August 2004 Prof. Thomas Hohn University of Basel Botanisches Institut
More informationGene Transcription in Prokaryotes
Gene Transcription in Prokaryotes Operons: in prokaryotes, genes that encode protein participating in a common pathway are organized together. This group of genes, arranged in tandem, is called an OPERON.
More informationRegents Biology REGENTS REVIEW: PROTEIN SYNTHESIS
Period Date REGENTS REVIEW: PROTEIN SYNTHESIS 1. The diagram at the right represents a portion of a type of organic molecule present in the cells of organisms. What will most likely happen if there is
More informationActions of Hormones on Target Cells Page 1. Actions of Hormones on Target Cells Page 2. Goals/ What You Need to Know Goals What You Need to Know
Actions of Hormones on Target Cells Graphics are used with permission of: Pearson Education Inc., publishing as Benjamin Cummings (http://www.aw-bc.com) Page 1. Actions of Hormones on Target Cells Hormones
More informationLecture Series 7. From DNA to Protein. Genotype to Phenotype. Reading Assignments. A. Genes and the Synthesis of Polypeptides
Lecture Series 7 From DNA to Protein: Genotype to Phenotype Reading Assignments Read Chapter 7 From DNA to Protein A. Genes and the Synthesis of Polypeptides Genes are made up of DNA and are expressed
More informationPRESTWICK ACADEMY NATIONAL 5 BIOLOGY CELL BIOLOGY SUMMARY
Name PRESTWICK ACADEMY NATIONAL 5 BIOLOGY CELL BIOLOGY SUMMARY Cell Structure Identify animal, plant, fungal and bacterial cell ultrastructure and know the structures functions. Plant cell Animal cell
More informationTranslation Study Guide
Translation Study Guide This study guide is a written version of the material you have seen presented in the replication unit. In translation, the cell uses the genetic information contained in mrna to
More informationMUTATION, DNA REPAIR AND CANCER
MUTATION, DNA REPAIR AND CANCER 1 Mutation A heritable change in the genetic material Essential to the continuity of life Source of variation for natural selection New mutations are more likely to be harmful
More informationA Primer of Genome Science THIRD
A Primer of Genome Science THIRD EDITION GREG GIBSON-SPENCER V. MUSE North Carolina State University Sinauer Associates, Inc. Publishers Sunderland, Massachusetts USA Contents Preface xi 1 Genome Projects:
More informationThe Need for a PARP in vivo Pharmacodynamic Assay
The Need for a PARP in vivo Pharmacodynamic Assay Jay George, Ph.D., Chief Scientific Officer, Trevigen, Inc., Gaithersburg, MD For further infomation, please contact: William Booth, Ph.D. Tel: +44 (0)1235
More informationNotch 1 -dependent regulation of cell fate in colorectal cancer
Notch 1 -dependent regulation of cell fate in colorectal cancer Referees: PD Dr. Tobias Dick Prof. Dr. Wilfried Roth http://d-nb.info/1057851272 CONTENTS Summary 1 Zusammenfassung 2 1 INTRODUCTION 3 1.1
More informationHow many of you have checked out the web site on protein-dna interactions?
How many of you have checked out the web site on protein-dna interactions? Example of an approximately 40,000 probe spotted oligo microarray with enlarged inset to show detail. Find and be ready to discuss
More informationMOTHER OF FT AND TFL1 Regulates Seed Germination through a Negative Feedback Loop Modulating ABA Signaling in Arabidopsis C W
This article is a Plant Cell Advance Online Publication. The date of its first appearance online is the official date of publication. The article has been edited and the authors have corrected proofs,
More informationPublikationsliste Claudia Götz
Publikationsliste Claudia Götz 1. Reinhard,B., Götz, C., and Faillard, H.: Synthesis of N-Acetyl-9-Oacetylneuraminic acid α-p-aminophenylthioketoside and its application as ligand in the affinity chromatography
More informationActivity 7.21 Transcription factors
Purpose To consolidate understanding of protein synthesis. To explain the role of transcription factors and hormones in switching genes on and off. Play the transcription initiation complex game Regulation
More informationThe wheat wcs120 promoter is cold-inducible in both monocotyledonous and dicotyledonous species
FEBS Letters 423 (1998) 324^328 FEBS 19883 The wheat wcs120 promoter is cold-inducible in both monocotyledonous and dicotyledonous species Franc ois Ouellet, Alejandro Vazquez-Tello, Fathey Sarhan* Deèpartement
More informationMs. Campbell Protein Synthesis Practice Questions Regents L.E.
Name Student # Ms. Campbell Protein Synthesis Practice Questions Regents L.E. 1. A sequence of three nitrogenous bases in a messenger-rna molecule is known as a 1) codon 2) gene 3) polypeptide 4) nucleotide
More informationCourse Curriculum for Master Degree in Medical Laboratory Sciences/Clinical Microbiology, Immunology and Serology
Course Curriculum for Master Degree in Medical Laboratory Sciences/Clinical Microbiology, Immunology and Serology The Master Degree in Medical Laboratory Sciences / Clinical Microbiology, Immunology or
More informationControl of Gene Expression
Home Gene Regulation Is Necessary? Control of Gene Expression By switching genes off when they are not needed, cells can prevent resources from being wasted. There should be natural selection favoring
More informationAP BIOLOGY 2008 SCORING GUIDELINES
AP BIOLOGY 2008 SCORING GUIDELINES Question 1 1. The physical structure of a protein often reflects and affects its function. (a) Describe THREE types of chemical bonds/interactions found in proteins.
More informationTranscription and Translation of DNA
Transcription and Translation of DNA Genotype our genetic constitution ( makeup) is determined (controlled) by the sequence of bases in its genes Phenotype determined by the proteins synthesised when genes
More informationS1 Text. Modeling deterministic single-cell microrna-p53-mdm2 network Figure 2 Figure 2
S1 Text. Modeling deterministic single-cell microrna-p53-mdm2 network The schematic diagram of the microrna-p53-mdm2 oscillator is illustrated in Figure 2. The interaction scheme among the mrnas and the
More informationCourse Curriculum for Master Degree in Medical Laboratory Sciences/Clinical Biochemistry
Course Curriculum for Master Degree in Medical Laboratory Sciences/Clinical Biochemistry The Master Degree in Medical Laboratory Sciences /Clinical Biochemistry, is awarded by the Faculty of Graduate Studies
More informationCHAPTER 6: RECOMBINANT DNA TECHNOLOGY YEAR III PHARM.D DR. V. CHITRA
CHAPTER 6: RECOMBINANT DNA TECHNOLOGY YEAR III PHARM.D DR. V. CHITRA INTRODUCTION DNA : DNA is deoxyribose nucleic acid. It is made up of a base consisting of sugar, phosphate and one nitrogen base.the
More informationThe Steps. 1. Transcription. 2. Transferal. 3. Translation
Protein Synthesis Protein synthesis is simply the "making of proteins." Although the term itself is easy to understand, the multiple steps that a cell in a plant or animal must go through are not. In order
More informationControl of Gene Expression
Control of Gene Expression What is Gene Expression? Gene expression is the process by which informa9on from a gene is used in the synthesis of a func9onal gene product. What is Gene Expression? Figure
More informationGenetic information (DNA) determines structure of proteins DNA RNA proteins cell structure 3.11 3.15 enzymes control cell chemistry ( metabolism )
Biology 1406 Exam 3 Notes Structure of DNA Ch. 10 Genetic information (DNA) determines structure of proteins DNA RNA proteins cell structure 3.11 3.15 enzymes control cell chemistry ( metabolism ) Proteins
More informationBiosafety considerations of new dsrna molecules
Biosafety considerations of new dsrna molecules GM-free Europe Conference 2015 May 6-8 Berlin, Germany Dr. Sarah Agapito Current GMOs Plant transformed to produce new protein A G C U G G C A A U G U C
More informationNOVEL GENOME-SCALE CORRELATION BETWEEN DNA REPLICATION AND RNA TRANSCRIPTION DURING THE CELL CYCLE IN YEAST IS PREDICTED BY DATA-DRIVEN MODELS
NOVEL GENOME-SCALE CORRELATION BETWEEN DNA REPLICATION AND RNA TRANSCRIPTION DURING THE CELL CYCLE IN YEAST IS PREDICTED BY DATA-DRIVEN MODELS Orly Alter (a) *, Gene H. Golub (b), Patrick O. Brown (c)
More informationCurriculum Policy of the Graduate School of Agricultural Science, Graduate Program
Curriculum Policy of the Graduate School of Agricultural Science, Graduate Program Agricultural Science plans to conserve natural and artificial ecosystems and its ideal of "Sustainable coexistence science"
More informationUnit I: Introduction To Scientific Processes
Unit I: Introduction To Scientific Processes This unit is an introduction to the scientific process. This unit consists of a laboratory exercise where students go through the QPOE2 process step by step
More informationKeystone Review Practice Test Module A Cells and Cell Processes. 1. Which characteristic is shared by all prokaryotes and eukaryotes?
Keystone Review Practice Test Module A Cells and Cell Processes 1. Which characteristic is shared by all prokaryotes and eukaryotes? a. Ability to store hereditary information b. Use of organelles to control
More informationDNA Replication & Protein Synthesis. This isn t a baaaaaaaddd chapter!!!
DNA Replication & Protein Synthesis This isn t a baaaaaaaddd chapter!!! The Discovery of DNA s Structure Watson and Crick s discovery of DNA s structure was based on almost fifty years of research by other
More informationProtein Synthesis How Genes Become Constituent Molecules
Protein Synthesis Protein Synthesis How Genes Become Constituent Molecules Mendel and The Idea of Gene What is a Chromosome? A chromosome is a molecule of DNA 50% 50% 1. True 2. False True False Protein
More informationAntigenic variation in Plasmodium falciparum : Erythrocyte invasion and immune escape mechanisms
Antigenic variation in Plasmodium falciparum : Erythrocyte invasion and immune escape mechanisms Introduction Why does immunity to malaria take so long to develop? The parasite s survival depends on its
More informationChapter-21b: Hormones and Receptors
1 hapter-21b: Hormones and Receptors Hormone classes Hormones are classified according to the distance over which they act. 1. Autocrine hormones --- act on the same cell that released them. Interleukin-2
More informationEnzymes: Practice Questions #1
Enzymes: Practice Questions #1 1. Compound X increases the rate of the reaction below. Compound X is most likely A. an enzyme B. a lipid molecule C. an indicator D. an ADP molecule 2. The equation below
More information1. The diagram below represents a biological process
1. The diagram below represents a biological process 5. The chart below indicates the elements contained in four different molecules and the number of atoms of each element in those molecules. Which set
More informationBiotechnology and Recombinant DNA (Chapter 9) Lecture Materials for Amy Warenda Czura, Ph.D. Suffolk County Community College
Biotechnology and Recombinant DNA (Chapter 9) Lecture Materials for Amy Warenda Czura, Ph.D. Suffolk County Community College Primary Source for figures and content: Eastern Campus Tortora, G.J. Microbiology
More informationReplication Study Guide
Replication Study Guide This study guide is a written version of the material you have seen presented in the replication unit. Self-reproduction is a function of life that human-engineered systems have
More informationAppendix 2 Molecular Biology Core Curriculum. Websites and Other Resources
Appendix 2 Molecular Biology Core Curriculum Websites and Other Resources Chapter 1 - The Molecular Basis of Cancer 1. Inside Cancer http://www.insidecancer.org/ From the Dolan DNA Learning Center Cold
More informationDiabetes and Insulin Signaling
Diabetes and Insulin Signaling NATIONAL CENTER FOR CASE STUDY TEACHING IN SCIENCE by Kristy J. Wilson School of Mathematics and Sciences Marian University, Indianapolis, IN Part I Research Orientation
More informationRNA and Protein Synthesis
Name lass Date RN and Protein Synthesis Information and Heredity Q: How does information fl ow from DN to RN to direct the synthesis of proteins? 13.1 What is RN? WHT I KNOW SMPLE NSWER: RN is a nucleic
More informationStatement of ethical principles for biotechnology in Victoria
Statement of ethical principles for biotechnology in Victoria Statement of ethical principles for biotechnology in Victoria Acknowledgments Published by the Public Health Group, Rural & Regional Health
More informationRULES FOR THE EVOLUTION OF GENE CIRCUITRY
RULES FOR THE EVOLUTION OF GENE CIRCUITRY M. A. SAVAGEAU Department of Microbiology & Immunology, The University of Michigan Medical School, Ann Arbor, Michigan 48109-0629 USA Cells possess the genes required
More informationIntegrating DNA Motif Discovery and Genome-Wide Expression Analysis. Erin M. Conlon
Integrating DNA Motif Discovery and Genome-Wide Expression Analysis Department of Mathematics and Statistics University of Massachusetts Amherst Statistics in Functional Genomics Workshop Ascona, Switzerland
More informationChemical Basis of Life Module A Anchor 2
Chemical Basis of Life Module A Anchor 2 Key Concepts: - Water is a polar molecule. Therefore, it is able to form multiple hydrogen bonds, which account for many of its special properties. - Water s polarity
More informationStructure and Function of DNA
Structure and Function of DNA DNA and RNA Structure DNA and RNA are nucleic acids. They consist of chemical units called nucleotides. The nucleotides are joined by a sugar-phosphate backbone. The four
More informationBiological cell membranes
Unit 14: Cell biology. 14 2 Biological cell membranes The cell surface membrane surrounds the cell and acts as a barrier between the cell s contents and the environment. The cell membrane has multiple
More informationDepartment of Food and Nutrition
Department of Food and Nutrition Faculties Professors Lee-Kim, Yang Cha, Ph.D. (M.I.T., 1973) Nutritional biochemistry, Antioxidant vitamins, Fatty acid metabolism, Brain development, and Hyperlipidemia
More informationFACULTY OF MEDICAL SCIENCE
Doctor of Philosophy in Biochemistry FACULTY OF MEDICAL SCIENCE Naresuan University 73 Doctor of Philosophy in Biochemistry The Biochemistry Department at Naresuan University is a leader in lower northern
More informationBasic Concepts of DNA, Proteins, Genes and Genomes
Basic Concepts of DNA, Proteins, Genes and Genomes Kun-Mao Chao 1,2,3 1 Graduate Institute of Biomedical Electronics and Bioinformatics 2 Department of Computer Science and Information Engineering 3 Graduate
More information2. True or False? The sequence of nucleotides in the human genome is 90.9% identical from one person to the next. False (it s 99.
1. True or False? A typical chromosome can contain several hundred to several thousand genes, arranged in linear order along the DNA molecule present in the chromosome. True 2. True or False? The sequence
More information13.4 Gene Regulation and Expression
13.4 Gene Regulation and Expression Lesson Objectives Describe gene regulation in prokaryotes. Explain how most eukaryotic genes are regulated. Relate gene regulation to development in multicellular organisms.
More informationHuman Genome and Human Genome Project. Louxin Zhang
Human Genome and Human Genome Project Louxin Zhang A Primer to Genomics Cells are the fundamental working units of every living systems. DNA is made of 4 nucleotide bases. The DNA sequence is the particular
More informationBiotechnology: DNA Technology & Genomics
Chapter 20. Biotechnology: DNA Technology & Genomics 2003-2004 The BIG Questions How can we use our knowledge of DNA to: diagnose disease or defect? cure disease or defect? change/improve organisms? What
More informationLecture 8. Protein Trafficking/Targeting. Protein targeting is necessary for proteins that are destined to work outside the cytoplasm.
Protein Trafficking/Targeting (8.1) Lecture 8 Protein Trafficking/Targeting Protein targeting is necessary for proteins that are destined to work outside the cytoplasm. Protein targeting is more complex
More informationGenetics Test Biology I
Genetics Test Biology I Multiple Choice Identify the choice that best completes the statement or answers the question. 1. Avery s experiments showed that bacteria are transformed by a. RNA. c. proteins.
More informationSpecial report. Chronic Lymphocytic Leukemia (CLL) Genomic Biology 3020 April 20, 2006
Special report Chronic Lymphocytic Leukemia (CLL) Genomic Biology 3020 April 20, 2006 Gene And Protein The gene that causes the mutation is CCND1 and the protein NP_444284 The mutation deals with the cell
More informationby the PCR-mediated method (Krawchuk and Wahls, 1999). The construction of Ams2-null and conditional ams2-shut-off strains was previously described
Table S1. Fission yeast strains used in this study. Gene disruption was performed by the PCR-mediated method (Krawchuk and Wahls, 1999). The construction of Ams2-null and conditional ams2-shut-off strains
More information2.1.2 Characterization of antiviral effect of cytokine expression on HBV replication in transduced mouse hepatocytes line
i 1 INTRODUCTION 1.1 Human Hepatitis B virus (HBV) 1 1.1.1 Pathogenesis of Hepatitis B 1 1.1.2 Genome organization of HBV 3 1.1.3 Structure of HBV virion 5 1.1.4 HBV life cycle 5 1.1.5 Experimental models
More informationEnzyme Response Profiling: Integrating proteomics and genomics with xenobiotic metabolism and cytotoxicity
Enzyme Response Profiling: Integrating proteomics and genomics with xenobiotic metabolism and cytotoxicity Proteomics high throughput separation and characterization of expressed proteins in tissue and
More informationOutline. interfering RNA - What is dat? Brief history of RNA interference. What does it do? How does it work?
Outline Outline interfering RNA - What is dat? Brief history of RNA interference. What does it do? How does it work? What is RNA interference? Recently discovered regulatory level. Genome immune system.
More informationChapter 8. Summary and Perspectives
Chapter 8 Summary and Perspectives 131 Chapter 8 Summary Overexpression of the multidrug resistance protein MRP1 confer multidrug resistance (MDR) to cancer cells. The contents of this thesis describe
More informationGuidance for Industry
Guidance for Industry Interpreting Sameness of Monoclonal Antibody Products Under the Orphan Drug Regulations U.S. Department of Health and Human Services Food and Drug Administration Center for Drug Evaluation
More informationISTEP+: Biology I End-of-Course Assessment Released Items and Scoring Notes
ISTEP+: Biology I End-of-Course Assessment Released Items and Scoring Notes Page 1 of 22 Introduction Indiana students enrolled in Biology I participated in the ISTEP+: Biology I Graduation Examination
More informationComplex multicellular organisms are produced by cells that switch genes on and off during development.
Home Control of Gene Expression Gene Regulation Is Necessary? By switching genes off when they are not needed, cells can prevent resources from being wasted. There should be natural selection favoring
More informationBiochemistry Major Talk 2014-15. Welcome!!!!!!!!!!!!!!
Biochemistry Major Talk 2014-15 August 14, 2015 Department of Biochemistry The University of Hong Kong Welcome!!!!!!!!!!!!!! Introduction to Biochemistry A four-minute video: http://www.youtube.com/watch?v=tpbamzq_pue&l
More informationGenetomic Promototypes
Genetomic Promototypes Mirkó Palla and Dana Pe er Department of Mechanical Engineering Clarkson University Potsdam, New York and Department of Genetics Harvard Medical School 77 Avenue Louis Pasteur Boston,
More informationGENE REGULATION. Teacher Packet
AP * BIOLOGY GENE REGULATION Teacher Packet AP* is a trademark of the College Entrance Examination Board. The College Entrance Examination Board was not involved in the production of this material. Pictures
More informationHormones & Chemical Signaling
Hormones & Chemical Signaling Part 2 modulation of signal pathways and hormone classification & function How are these pathways controlled? Receptors are proteins! Subject to Specificity of binding Competition
More information8/20/2012 H C OH H R. Proteins
Proteins Rubisco monomer = amino acids 20 different amino acids polymer = polypeptide protein can be one or more polypeptide chains folded & bonded together large & complex 3-D shape hemoglobin Amino acids
More informationCarbohydrates, proteins and lipids
Carbohydrates, proteins and lipids Chapter 3 MACROMOLECULES Macromolecules: polymers with molecular weights >1,000 Functional groups THE FOUR MACROMOLECULES IN LIFE Molecules in living organisms: proteins,
More information