AP-1 function and regulation.

OXIDATIVE STRESS . . . . . . . . . . . . . .. . . . . .... . . . . .. . . . . .. . . . ... .. . . . . .... . . .. .. . . . . . ... . 84 RESPONSE OF SOD TO ENVIRONMENTAL STRESS . . . . . . . . . . . . . . . .. . . . .... . . .... . . . . . . 87 Photoinhibition . . . . . . . . . . .. . ... . . .. . . . .. . . . . ..... . . . , ... ", , ... ,' , ... . ,., . . . "" . .. . ,'.' . . ,' . . . . , ., 87 Paraquat and Other Herbicides . . .. , . . .... , ..... , . . . . ... . . . .. . . . . . . . . . . . . ,.. 91 Atmospheric Pollutants . . " .... , .... , " .... " , , . . , .... , ' 94 Waterlogging and Drought .. ", . . ... , . . . . . . , .. "", . . ", . . . . . ", . . . .. ", . . .. , . . . ... """", . . " 97 The Defense Response to Pathogens . ... . . . . "" ... "" .... " ...... " ,."" .... . , 98 The Phenomenon of Cross-Tolerance . . . , , .. . . . . ...... . . , . . . . . . , . . .... , .. . . . . . . . .. .. 101 The Mechanism of SOD Regulation ........ , , ...... , " 102 GENETIC ENGINEERING OF SOD IN PLANTS .. ' ...... ' . . . . . . . . . . . . . . . . . . . . . . . .. . . . , .... ' 104 Cu/ZnSOD and MnSOD Overexpression . . . . . .. . . . . . , . . . . . . .. . . . . . .. . . . . . , .... . . . . .. . . ,.... . . 105 Prospects for Stress Tolerance through Genetic Engineering of SOD . . . . ,,, .. , . . . . . . . 106

Superoxide dismutase and stress tolerance

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1. Light and Energy. 2. Organization and Structure of Photosynthetic Systems. 3. History and Development of Photosynthesis. 4. Photosynthetic Pigments-Structure and Spectroscopy. 5. Antenna Complexes and Energy Transfer Processes. 6. Reaction Center Complexes. 7. Electron Transfer Pathways and Components. 8. Chemiosmotic Coupling and ATP Synthesis. 9. Carbon Metabolism. 10. Genetics, Assembly and Regulation of Photosynthetic. Systems. 11. Origin and Evolution of Photosynthesis. Appendix 1. Light, Energy and Kinetics

Molecular mechanisms of photosynthesis

Auxin: regulation, action, and interaction.

Many auxin responses are dependent on redistribution and/or polar transport of indoleacetic acid. Polar transport of auxin can be inhibited through the application of phytotropins such as 1-naphthylphthalamic acid (NPA). When Arabidopsis thaliana seedlings were grown in the light on medium containing 1.0 microM NPA, hypocotyl and root elongation and gravitropism were strongly inhibited. When grown in darkness, however, NPA disrupted the gravity response but did not affect elongation. The extent of inhibition of hypocotyl elongation by NPA increased in a fluence-rate-dependent manner to a maximum of about 75% inhibition at 50 mumol m-2 s-1 of white light. Plants grown under continuous blue or far-red light showed NPA-induced hypocotyl inhibition similar to that of white-light-grown plants. Plants grown under continuous red light showed less NPA-induced inhibition. Analysis of photoreceptor mutants indicates the involvement of phytochrome and cryptochrome in mediating this NPA response. Hypocotyls of some auxin-resistant mutants had decreased sensitivity to NPA in the light, but etiolated seedlings of these mutants were similar in length to the wild type. These results indicate that light has a significant effect on NPA-induced inhibition in Arabidopsis, and suggest that auxin has a more important role in elongation responses in light-grown than in dark-grown seedlings.

http://www.plantphysiol.org/content/plantphysiol/116/2/455.full.pdf

Auxin Transport Is Required for Hypocotyl Elongation in Light-Grown but Not Dark-Grown Arabidopsis

Flowering in Arabidopsis thaliana is promoted by longday (LD) photoperiods such that plants grown in LD flower earlier, and after the production of fewer leaves, than plants grown in short-day (SD) photoperiods The early-flowering 3 (elf3) mutant of Arabidopsis, which is insensitive to photoperiod with regard to floral initiation has been characterized elf3 mutants are also altered in several aspects of vegetative photomorphogenesis, including hypocotyl elongation When inhibition of hypocotyl elongation was measured, elf3 mutant seedlings were less responsive than wild-type to all wavelengths of light, and most notably defective in blue and green light-mediated inhibition When analyzed for the flowering-time phenotype, elf3 was epistatic to mutant alleles of the blue-light receptor encoding gene, HY4 However, when elf3 mutants were made deficient for functional phytochrome by the introduction of hy2 mutant alleles, the elf3 hy2 double mutants displayed the novel phenotype of flowering earlier than either single mutant while still exhibiting photoperiod insensitivity, indicating that a phytochrome-mediated pathway regulating floral initiation remains functional in elf3 single mutants In addition, the inflorescences of one allelic combination of elf3 hy2 double mutants form a terminal flower similar to the structure produced by tfk1 single mutants These results suggest that one of the signal transduction pathways controlling photoperiodism in Arabidopsis is regulated, at least in part, by photoreceptors other than phytochrome, and that the activity of the Arabidopsis inflorescence and floral meristem identity genes may be regulated by this same pathway

The Arabidopsis ELF3 gene regulates vegetative photomorphogenesis and the photoperiodic induction of flowering

Plants have evolved highly sensitive and selective mechanisms that detect and respond to various aspects of their environment. As a plant develops, it integrates the environmental information perceived by all of its sensory systems and adapts its growth to the prevailing environmental conditions. Light is of critical importance because plants depend on it for energy and, thus, survival. The quantity, quality and direction of light are perceived by several different photosensory systems that together regulate nearly all stages of plant development, presumably in order to maintain photosynthetic efficiency. Gravity provides an almost constant stimulus that is the source of critical spatial information about its surroundings and provides important cues for orientating plant growth. Gravity plays a particularly important role during the early stages of seedling growth by stimulating a negative gravitropic response in the primary shoot that orientates it towards the source of light, and a positive gravitropic response in the primary root that causes it to grow down into the soil, providing support and nutrient acquisition. Gravity also influences plant form during later stages of development through its effect on lateral organs and supporting structures. Thus, the final form of a plant depends on the cumulative effects of light, gravity and other environmental sensory inputs on endogenous developmental programs. This article is focused on developmental interactions modulated by light and gravity.

Gravity, light and plant form

Recent studies of auxin response have focused on the functions of three sets of proteins: the auxin (Aux) response factors (ARFs), the Aux/IAAs, and the F-box protein TIR1. The ARF proteins bind DNA and directly activate or repress transcription of target genes while the Aux/IAA proteins repress ARF function. TIR1 is part of a ubiquitin protein ligase required for degradation of Aux/IAA proteins. Here we report the isolation and characterization of a novel mutant of Arabidopsis called axr5-1. Mutant plants are resistant to auxin and display a variety of auxin-related growth defects including defects in root and shoot tropisms. Further, the axr5-1 mutation results in a decrease in auxin-regulated transcription. The molecular cloning of AXR5 revealed that the gene encodes the IAA1 protein, a member of the Aux/IAA family of proteins. AXR5 is expressed throughout plant development consistent with the pleiotropic mutant phenotype. The axr5-1 mutation results in an amino acid substitution in conserved domain II of the protein, similar to gain-of-function mutations recovered in other members of this gene family. Biochemical studies show that IAA1/AXR5 interacts with TIR1 in an auxin-dependent manner. The mutation prevents this interaction suggesting that the mutant phenotype is caused by the accumulation of IAA1/AXR5. Our results provide further support for a model in which most members of the Aux/IAA family are targeted for degradation by SCFTIR1 in response to auxin.

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The IAA1 protein is encoded by AXR5 and is a substrate of SCF(TIR1).

The interaction of tropisms is important in determining the final growth form of the plant body. In roots, gravitropism is the predominant tropistic response, but phototropism also plays a role in the oriented growth of roots in flowering plants. In blue or white light, roots exhibit negative phototropism that is mediated by the phototropin family of photoreceptors. In contrast, red light induces a positive phototropism in Arabidopsis roots. Because this red-light-induced response is weak relative to both gravitropism and negative phototropism, we used a novel device to study phototropism without the complications of a counteracting gravitational stimulus. This device is based on a computer-controlled system using real-time image analysis of root growth and a feedback-regulated rotatable stage. Our data show that this system is useful to study root phototropism in response to red light, because in wild-type roots, the maximal curvature detected with this apparatus is 30 degrees to 40 degrees, compared with 5 degrees to 10 degrees without the feedback system. In positive root phototropism, sensing of red light occurs in the root itself and is not dependent on shoot-derived signals resulting from light perception. Phytochrome (Phy)A and phyB were severely impaired in red-light-induced phototropism, whereas the phyD and phyE mutants were normal in this response. Thus, PHYA and PHYB play a key role in mediating red-light-dependent positive phototropism in roots. Although phytochrome has been shown to mediate phototropism in some lower plant groups, this is one of the few reports indicating a phytochrome-dependent phototropism in flowering plants.

Roger P. Hangarter

Papers

Auxin Transport Is Required for Hypocotyl Elongation in Light-Grown but Not Dark-Grown Arabidopsis

The Arabidopsis ELF3 gene regulates vegetative photomorphogenesis and the photoperiodic induction of flowering

Gravity, light and plant form

The IAA1 protein is encoded by AXR5 and is a substrate of SCF(TIR1).

Phytochromes A and B Mediate Red-Light-Induced Positive Phototropism in Roots