▪ Abstract In eukaryotic cells, most proteins in the cytosol and nucleus are degraded via the ubiquitin-proteasome pathway. The 26S proteasome is a 2.5-MDa molecular machine built from ∼31 different subunits, which catalyzes protein degradation. It contains a barrel-shaped proteolytic core complex (the 20S proteasome), capped at one or both ends by 19S regulatory complexes, which recognize ubiquitinated proteins. The regulatory complexes are also implicated in unfolding and translocation of ubiquitinated targets into the interior of the 20S complex, where they are degraded to oligopeptides. Structure, assembly and enzymatic mechanism of the 20S complex have been elucidated, but the functional organization of the 19S complex is less well understood. Most subunits of the 19S complex have been identified, however, specific functions have been assigned to only a few. A low-resolution structure of the 26S proteasome has been obtained by electron microscopy, but the precise arrangement of subunits in the 19S co...

The 26S Proteasome: A Molecular Machine Designed for Controlled Proteolysis

Bioactive gibberellins (GAs) are diterpene plant hormones that are biosynthesized through complex pathways and control diverse aspects of growth and development. Biochemical, genetic, and genomic approaches have led to the identification of the majority of the genes that encode GA biosynthesis and deactivation enzymes. Recent studies have highlighted the occurrence of previously unrecognized deactivation mechanisms. It is now clear that both GA biosynthesis and deactivation pathways are tightly regulated by developmental, hormonal, and environmental signals, consistent with the role of GAs as key growth regulators. In some cases, the molecular mechanisms for fine-tuning the hormone levels are beginning to be uncovered. In this review, I summarize our current understanding of the GA biosynthesis and deactivation pathways in plants and fungi, and discuss how GA concentrations in plant tissues are regulated during development and in response to environmental stimuli.

Gibberellin Metabolism and its Regulation

Summary) The abstract should follow the structure of the article (relevance, degree of exploration of the problem, the goal, the main results, conclusion) and characterize the theoretical and practical significance of the study results. The abstract should not contain wording echoing the title, cumbersome grammatical structures and abbreviations. The text should be written in scientific style. The volume of abstracts (summaries) depends on the content of the article, but should not be less than 250 words. All abbreviations must be disclosed in the summary (in spite of the fact that they will be disclosed in the main text of the article), references to the numbers of publications from reference list should not be made. The sentences of the abstract should constitute an integral text, which can be made by use of the words “consequently”, “for example”, “as a result”. Avoid the use of unnecessary introductory phrases (eg, “the author of the article considers...”, “The article presents...” and so on.)

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Ministry of Education and Science of the Russian Federation

Seed Dormancy and Germination

The power of molecular genetics has dramatically advanced our understanding of all aspects of gibberellin (GA) signaling. Many genes encoding GA response pathway components have been identified using Arabidopsis and cereal mutants, and more elaborate genetic screens are producing additional mutants

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Gibberellin signaling: biosynthesis, catabolism, and response pathways.

It is perhaps not surprising that plants have evolved a mechanism to sense the light environment about them and to modify growth for optimal use of the available life-giving' light. Green plants, and ultimately all forms of life, depend on the energy of sunlight fixed during photosynthesis. Unlike animals that use behaviour to find food, sedentary plants use physiology to optimize their growth and development for light absorption. By appreciating the quality, quantity, direction and duration of light, plants can control such complex processes as germination, growth and flowering. To perceive the light environment several receptor pigments have evolved, including the red/far-red reversible phytochrome and the blue/UV-absorbing photoreceptors (Part 1). The quantification of light (Part 2) and importance of instrumentation for photomorphogenesis research are introduced in Part 3. Isolation and characterization of phytochrome is a classic example of how photobiological techniques can predict the nature of an unknown photoreceptor. Current knowledge of the phytochrome photoreceptor family is given in Part 4 and that of blue/UV receptors in Part 5. Part 6 deals with the coaction of photoreceptors. The light environment and its perception is addressed in Part 7. Molecular and genetic approaches and the photoregulation of gene expression compose Part 8. Part 9 contains further selected topics: photomodulation of growth phototropism, photobiology of stomatal movements, photomovement, photocontrol of flavonoid biosynthesis, photobiology of fungi and photobiology of ferns.The 28 chapters written by leading experts from Europe, Israel, Japan and the USA, provide an advanced treatise on the excitingand rapidly developing field of plant photomorphogenesis.

Photomorphogenesis in plants

A novel cDNA sequence homologous to a phytochrome B (phyB) gene that was isolated in a library from tobacco tissue has been used in an Escherichia coli expression system to raise anti-phytochrome B (anti-PHYB) polypeptide-specific monoclonal antibodies. The specificity of these antibodies has been tested by cross-reactivity against purified pea light-labile type 1 and light-stable type 2 phytochromes, with some antibodies reacting with the type 2 and none with the type 1 phytochromes. One such antibody, monoclonal mAT1, has been employed to analyze the phytochrome molecular species present in a photomorphogenic long hypocotyl (lh) mutant of cucumber. The results indicated that the mutant contains wild-type levels of the light-labile type 1 phytochrome polypeptide (PHYA), which has an apparent molecular mass of approximately 120 kD, but shows less than 1% (detection limit) of a light-stable polypeptide recognized by mAT1 in wild-type seedlings. This protein, not detectable in the lh mutant, has the properties of light-stable type 2 phytochrome, has an apparent molecular mass of 116 to 117 kD, and remains at constant levels under continuous low-fluence-rate red light. Therefore, we conclude that the lh mutant lacks at least one type 2 phytochrome-like polypeptide, most probably a phyB gene product. The correlation between the lack of this protein and the deficiency or absence of physiological responses to a light-stable phytochrome species in this mutant helps to identify the physiological roles played by the products of different subfamilies within the phytochrome gene family.

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The cucumber long hypocotyl mutant lacks a light-stable PHYB-like phytochrome.

The aurea (au) and yellow-green-2 (yg-2) mutants of tomato (Solanum lycopersicum L.) are unable to synthesize the linear tetrapyrrole chromophore of phytochrome, resulting in plants with a yellow-green phenotype. To understand the basis of this phenotype, we investigated the consequences of the au and yg-2 mutations on tetrapyrrole metabolism. Dark-grown seedlings of both mutants have reduced levels of protochlorophyllide (Pchlide) due to an inhibition of Pchlide synthesis. Feeding experiments with the tetrapyrrole precursor 5-aminolevulinic acid (ALA) demonstrate that the pathway between ALA and Pchlide is intact in au and yg-2 and suggest that the reduction in Pchlide is a result of the inhibition of ALA synthesis. This inhibition was independent of any deficiency in seed phytochrome, and experiments using an iron chelator to block heme synthesis demonstrated that both mutations inhibited the degradation of the physiologically active heme pool, suggesting that the reduction in Pchlide synthesis is a consequence of feedback inhibition by heme. We discuss the significance of these results in understanding the chlorophyll-deficient phenotype of the au and yg-2 mutants.

Feedback inhibition of chlorophyll synthesis in the phytochrome chromophore-deficient aurea and yellow-green-2 mutants of tomato

Photomorphogenic responses of long hypocotyl mutants of tomato

We have selected two recessive mutants of tomato with slightly longer hypocotyls than the wild type, one under low fluence rate (3 μmol/m2/s) red light (R) and the other under low fluence rate blue light. These two mutants were shown to be allelic and further analysis revealed that hypocotyl growth was totally insensitive to far-red light (FR). We propose the gene symbol fri (far-red light insensitive) for this locus and have mapped it on chromosome 10. Immunochemically detectable phytochrome A polypeptide is essentially absent in the fri mutants as is the bulk spectrophotometrically detectable labile phytochrome pool in etiolated seedlings. A phytochrome B-like polypeptide is present in normal amounts and a small stable phytochrome pool can be readily detected by spectrophotometry in the fri mutants. Inhibition of hypocotyl growth by a R pulse given every 4 h is quantitatively similar in the fri mutants and wild type and the effect is to a large extent reversible if R pulses are followed immediately by a FR pulse. After 7 days in darkness, both fri mutants and the wild type become green on transfer to white light, but after 7 days in FR, the wild-type seedlings that have expanded their cotyledons lose their capacity to green in white light, while the fri mutants de-etiolate. Adult plants of the fri mutants show retarded growth and are prone to wilting, but exhibit a normal elongation response to FR given at the end of the daily photoperiod. The inhibition of seed germination by continuous FR exhibited by the wild type is normal in the fri mutants. It is proposed that these fri mutants are putative phytochrome A mutants which have normal pools of other phytochromes.

Richard E. Kendrick

Papers

Photomorphogenesis in plants

The cucumber long hypocotyl mutant lacks a light-stable PHYB-like phytochrome.

Feedback inhibition of chlorophyll synthesis in the phytochrome chromophore-deficient aurea and yellow-green-2 mutants of tomato

Photomorphogenic responses of long hypocotyl mutants of tomato

Far-red light-insensitive, phytochrome A-deficient mutants of tomato.