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From flour to flower: how Polycomb group proteins influence multiple aspects of plant development.

Cell identity and differentiation are determined by patterns of regulatory gene expression. Spatially and temporally regulated homeotic gene expression defines segment identities along the anterior-posterior axis of animal embryos. Polycomb group (PcG) proteins form a cellular memory system that maintains the repressed state of homeotic gene expression. Conserved PcG proteins control multiple aspects of Arabidopsis development and maintain homeotic gene repression. In animals, PcG proteins repress their target genes by modifying histone tails through deacetylation and methylation, generating a PcG-specific histone code that recruits other chromatin remodeling proteins to establish a stable, heritable mechanism of epigenetic expression control. Plant PcG proteins might function through a similar biochemical mechanism owing to their conserved structural and functional relationship to animal PcG proteins.

Arabidopsis↗

Control of plant development and gene expression by sugar signaling.

Coordination of development with the availability of nutrients, such as soluble sugars, may help ensure an adequate supply of building materials and energy with which to carry out specific developmental programs. For example, in-vivo and in-vitro experiments suggest that increasing sugar levels delay seed germination and stimulate the induction of flowering and senescence in at least some plant species. Higher sugar concentrations can also increase the number of tubers formed by potatoes and can stimulate the formation of adventitious roots by Arabidopsis. New insights into the mechanisms by which sugar-response pathways interact with other response pathways have been provided by microarray experiments examining sugar-regulated gene expression under different light and nitrogen conditions.

Carbohydrates↗

A FUSCA gene of Arabidopsis encodes a novel protein essential for plant development.

Arabidopsis fusca mutants display striking purple coloration due to anthocyanin accumulation in their cotyledons. We describe six recessive fusca mutants isolated from Agrobacterium-transformed Arabidopsis families. These mutants first become defective during embryogenesis and exhibit limited seedling development. Double mutant constructs revealed that developmental defects were not simply a consequence of anthocyanin accumulation. fusca seedlings showed altered responses to several environmental and endogenous factors. Allelism tests established that three fusca loci are represented by mutants previously described as defective in light-regulated responses. To study the molecular basis of the fusca phenotype, we cloned the FUS6 gene. FUS6 encodes a novel protein that is hydrophilic, alpha-helical, and contains potential protein kinase C phosphorylation sites. The FUSCA proteins appear to act in a network of signal transduction pathways critical for plant development.

Alleles↗

A new family of plasma membrane polypeptides differentially regulated during plant development.

Two cDNAs encoding polypeptides identified in a tobacco leaf plasma membrane fraction prepared by phase partitioning were cloned. The deduced polypeptides, P16 and P17, exhibit a striking primary structure, similar to that of P19, a previously cloned plasma membrane polypeptide. Antibodies raised to the recombinant proteins were used to probe the cellular location of P16, P17 and P19 by means of western blotting of sucrose density-gradient fractions; all three polypeptides were found to be located solely at the plasma membrane. Furthermore, P19 antigen accumulated transiently at the time of floral induction while P16 and P17 antigens accumulated towards the end of the life-cycle. These results together with sequence database searches and multiple-sequence alignments suggest that we have identified a new family of plasma membrane polypeptides that (i) are putatively plant specific and (ii) are differentially regulated during plant development. These polypeptides are termed DREPPs for developmentally regulated plasma membrane polypeptides.

Amino Acid Sequence↗

Low-temperature tolerance and genetic potential in wheat (Triticum aestivum L.): response to photoperiod, vernalization, and plant development.

It is frequently observed that winter habit types are more low-temperature (LT) tolerant than spring habit types. This raises the question of whether this is due to pleiotropic effects of the vernalization loci or to the linkage of LT-tolerance genes to these vernalization loci. Reciprocal near-isogenic lines (NILs) for alleles at the Vrn-A1 locus, Vrn-A1 and vrn-A1, determining spring and winter habit respectively, in two diverse genetic backgrounds of wheat (Triticum aestivum L.) were used to separate the effects of vernalization, photoperiod, and development on identical, or near identical, genetic backgrounds. The vrn-A1 allele in the winter lines allowed full expression of genotype dependent LT tolerance potential. The winter allele (vrn-A1) in a very cold tolerant genetic background resulted in 11 degrees C, or a 2.4-fold, greater LT tolerance compared to the spring allele. Similarly, the delay in development caused by short-day (SD) versus long-day (LD) photoperiod in the identical spring habit NIL resulted in an 8.5 degrees C or 2.1-fold, increase in LT tolerance. The duration of time in early developmental stages was shown to underlie full expression of genetic LT-tolerance potential. Therefore, pleiotropic effects of the vernalization loci can explain the association of LT tolerance and winter habit irrespective of either the proposed closely linked Fr-A1 or the more distant Fr-A2 LT-tolerance QTLs. Plant development progressively reduced LT-acclimation ability, particularly after the main shoot meristem had advanced to the double ridge reproductive growth stage. The Vrn-1 genes, or other members of the flowering induction pathway, are discussed as possible candidates for involvement in LT-tolerance repression.

Acclimatization↗

AtBXL1, a novel higher plant (Arabidopsis thaliana) putative beta-xylosidase gene, is involved in secondary cell wall metabolism and plant development.

To investigate mechanisms involved in cell wall development, an Arabidopsis T-DNA insertion mutant collection was screened to identify mutants with beta-glucuronidase fusion gene expression in tissues undergoing secondary cell wall thickening. This promoter-trapping strategy allowed the isolation of a transformant containing the GUS coding sequence inserted 700 bp upstream of the ATG of a putative beta-xylosidase gene. The transformant has no phenotype as the expression of the gene was not disrupted by the insertion. The analysis of the predicted protein, AtBXL1, suggests its targeting to the extracellular matrix and its involvement in cell wall metabolism through a putative activity towards xylans. The 2-kb promoter sequence of AtBXL1 was fused to the GUS coding sequence and introduced into wild-type Arabidopsis thaliana. GUS expression was shown to be restricted to tissues undergoing secondary cell wall formation. Beta-xylosidase activity was associated with the cell wall-enriched fraction of different organs of wild-type plants. The level of activity correlates with transcript accumulation of AtBXL1 and other AtBXL1-related genes. Transgenic plants expressing the AtBXL1 cDNA in antisense orientation were generated. Lines exhibiting the highest decrease in AtBXL1 transcript accumulation and beta-xylosidase activity had phenotypic alterations. This newly identified gene is proposed to be involved in secondary cell wall hemicellulose metabolism and plant development.

Amino Acid Sequence↗

Plant development: regulation by protein degradation.

Many aspects of eukaryotic development depend on regulated protein degradation by the ubiquitin-proteasome pathway. This highly conserved pathway promotes covalent attachment of ubiquitin to protein substrates through the sequential action of three enzymes called a ubiquitin-activating enzyme (E1), a ubiquitin-conjugating enzyme (E2), and a ubiquitin-protein ligase (E3). Most ubiquitinated proteins are then targeted for degradation by the 26S proteasome. Recent studies have also shown that the ubiquitin-related protein RUB/Nedd8 and the proteasome-related COP9 signalosome complex cooperate with the ubiquitin-proteasome pathway to promote protein degradation. Most of these components are conserved in all three eukaryotic kingdoms. However, the known targets of the pathway in plants, and the developmental processes they regulate, are specific to the plant kingdom.

COP9 Signalosome Complex↗

Unraveling the knots in plant development.

Homeobox genes, first discovered from studies of homeotic mutations in Drosophila, have recently been found in plants. The proteins encoded by homeobox genes thus join the ranks of other animal transcription factors that have plant developmental counterparts, suggesting that even though plant and animal development are very different, regulatory mechanisms that direct development may be shared among all higher eukaryotes. The role of homeobox genes in plants remains elusive; nonetheless, gain-of-function mutations of one homeobox gene, Knotted, profoundly affect development. The phenotype suggests that ectopic expression of Knotted in leaves causes cells to take on alternative fates.

Animals↗

Epigenetic control of plant development by Polycomb-group proteins.

Recent genetic studies indicate that the plant Polycomb-group genes play much broader roles in development than was initially apparent from their single mutant phenotypes. At the mechanistic level, evidence is accumulating that their protein products act together in complexes that direct changes in histone methylation patterns. We discuss recent studies that give clues as to how these epigenetic changes are propagated through mitosis, how they are interpreted, and how they might be reset.

Gene Expression Regulation, Developmental↗

Monitoring of plant development in controlled environment with machine vision.

Information acquisition is the foremost requirement for the control and continued operation of any complex system. This is especially true when a plant production system is used as a major component in a sustainable life support system. The plant production system not only provides food and fiber but is a means of providing critically needed life supporting elements such as O2 and purified H2O. The success of the plant production system relies on close monitoring and control of the production system. Machine vision technology was evaluated for the monitoring of plant health and development and showed promising results. Spectral and morphological characteristics of a model plant were studied under various artificially induced stress conditions. From the spectroscopic studies, it was found that the stresses can be determined from visual and non-visual symptoms. The development of the plant can also be quantified using a video image analysis base approach. The correlations between the qualities of the model plant and machine vision measured spectral features were established. The success of the research has shown a great potential in building an automated, closed-loop plant production system in controlled environments.

Ecological Systems, Closed↗

Homeoboxes in plant development.

The homeobox is a 180 bp consensus DNA sequence present in a number of genes involved in developmental processes. This review focuses on the structure and function of plant homeobox genes and of the proteins they encode. Plant homeobox genes have been identified in studies using mutants, degenerate oligonucleotides deduced from conserved sequences, differential screening or binding to known promoters. According to sequence conservation, plant homeoboxes can be subdivided into different families, each comprising several members. Evolutionary studies indicate that the different families have diverged prior to the separation of the branches leading to animals, plants and fungi. Accordingly, members of different families show characteristic structural and functional properties. As an example, kn1-like genes seem to be involved in different aspects of the control of cell fate determination in the shoot meristem; HD-Zip genes, which encode proteins containing a leucine zipper motif adjacent to the homeodomain, are believed to operate at later stages of development; and gl2-like genes are involved in epidermal cell differentiation. Future studies should be oriented to discern the precise function of the many homeobox genes present in plant genomes, and to evaluate their use as modifiers of plant development.

Gene Expression Regulation, Developmental↗

[The role of plastids and assimilate transport system in the control of plant development].

Phylogenetic and ontogenetic relationships between the plastids, cell endoplasmic reticulum, and plant transport communication have been reviewed. The initiating role of plastids (endosymbionts) in the origin of endoplasmic reticulum (buffer zone of endosymbiogenesis) has been shown, as well as a similar role of endoplasmic reticulum in the development of transport communication of xylem and phloem. Plastids, sugars and transport system for their distribution can be interpreted as leading sections in the mechanism of developmental control: gene expression of nuclear genome and genome of organelles, cell and tissue differentiation, and plant morphogenesis. The conflict between the bulk of plant genome and low percentage of its realization is explained as a result of limitation of the nuclear genome realization by plastid genome. The concept of development as applied to plant ontogenesis has been critically analyzed. The possibilities of the concept correction by with the help of symbiogenetic hypothesis are discussed.

Biological Transport↗

Overexpression of a soybean gene encoding cytosolic glutamine synthetase in shoots of transgenic Lotus corniculatus L. plants triggers changes in ammonium assimilation and plant development.

A soybean cytosolic glutamine synthetase gene (GS15) was fused with the constitutive 35S cauliflower mosaic virus (CaMV) promoter in order to direct overexpression in Lotus corniculatus L. plants. Following transformation with Agrobacterium rhizogenes, eight independent Lotus transformants were obtained which synthesized additional cytosolic glutamine synthetase (GS) in the shoots. To eliminate any interference caused by the T-DNA from the Ri plasmid, three primary transformants were crossed with untransformed plants and progeny devoid of TL- and TR-DNA sequences were chosen for further analyses. These plants had a 50-80% increase in total leaf GS activity. Plants were grown under different nitrogen regimes (4 or 12 mM NH4+) and aspects of carbon and nitrogen metabolism were examined. In roots, an increase in free amino acids and ammonium was accompanied by a decrease in soluble carbohydrates in the transgenic plants cultivated with 12 mM NH4+ in comparison to the wild type grown under the same conditions. Labelling experiments using 15NH4+ were carried out in order to monitor the influx of ammonium and its subsequent incorporation into amino acids. This experiment showed that both ammonium uptake in the roots and the subsequent translocation of amino acids to the shoots was lower in plants overexpressing GS. It was concluded that the build up of ammonium and the increase in amino acid concentration in the roots was the result of shoot protein degradation. Moreover, following three weeks of hydroponic culture early floral development was observed in the transformed plants. As all these properties are characteristic of senescent plants, these findings suggest that expression of cytosolic GS in the shoots may accelerate plant development, leading to early senescence and premature flowering when plants are grown on an ammonium-rich medium.

Cytosol↗

A novel cell-ablation strategy for studying plant development.

The processes controlling the differentiation of plant cells are not well understood. Two alternative, but not mutually exclusive, mechanisms probably play a major role in plant-cell differentiation. One mechanism utilizes a position-independent, cell-autonomous differentiation process. The other, employs a position-dependent, cell-cell interaction process that requires signals from neighbouring cells. Cell ablation studies can be used to distinguish between these two models of plant-cell differentiation. In this article we outline a new cell-ablation strategy that utilizes promoters with distinct, but overlapping, cell specificities that are fused with cytotoxic and anticytotoxic structural genes. We present preliminary observations on how this strategy can be used to dissect the events controlling anther development.

Cell Differentiation↗

Foliar contamination of Phaseolus vulgaris with aerosols of 137Cs, 85Sr, 133Ba and 123mTe: influence of plant development stage upon contamination and rain.

As part of a requirement to improve the assessment of the impact of radioactive fallout on consumed agricultural products, bean plants at four development stages (seedlings, preflowering, late flowering and mature plants) were contaminated by dry deposition of (137)Cs, (85)Sr, (133)Ba and (123m)Te aerosols. The influence of two rain scenarios and of the development stage upon contamination on interception, retention, and translocation to pods was studied. Interception of the four radionuclides was almost identical and varied from 30 to 60% with increasing development stage. The most important rain parameter was the time which elapsed between contamination and the first rain. Whatever the development stage, rain washed off more cesium from the leaves when it occurred 2 days after the deposit (37% at the seedling stage, for example) rather than later on (6 days, 27%), due to rapid migration of Cs in the plant. The first rain washed off nearly 40% of Ba whatever the scenario. For later stages, Sr and Ba were more washed off by heavy weekly rains than by weak twice-a-week rains, perhaps because of the Sr/Ba-contaminated material loss associated with wash off (desquamation of cuticles). Te showed little wash off (less than 5%). Wash off decreased with an older development stage for a weak rain intensity, due to the superimposition of leaves. Heavy rains removed this shelter effect. At harvest, rain effect was no longer detectable as foliar activity was similar for both rain scenarios. Translocation factors (TF) for strontium and barium increased from 6 x 10(-3) to 1 x 10(-1) with the plant development stage upon contamination, whereas those for cesium remained almost unchanged between 2 x 10(-1) and 4 x 10(-1). Flowering is the most critical stage towards residual contamination in pods at harvest, with the exception of direct deposit on pods at the mature stage (TF values are one order of magnitude higher). TF value for Te was 6.5 x 10(-2) and was due to direct deposit. Modelling reflected the trends, through the differential values of the wash off and absorption coefficients, of what was reported for experimental results.

Aerosols↗

Down-regulation of the 26S proteasome subunit RPN9 inhibits viral systemic transport and alters plant vascular development.

Plant viruses utilize the vascular system for systemic movement. The plant vascular network also transports water, photosynthates, and signaling molecules and is essential for plant growth. However, the molecular mechanisms governing vascular development and patterning are still largely unknown. From viral transport suppressor screening using virus-induced gene silencing, we identified a 26S proteasome subunit, RPN9, which is required for broad-spectrum viral systemic transport. Silencing of RPN9 in Nicotiana benthamiana inhibits systemic spread of two taxonomically distinct viruses, Tobacco mosaic virus and Turnip mosaic virus. The 26S proteasome is a highly conserved eukaryotic protease complex controlling many fundamental biochemical processes, but the functions of many 26S proteasome regulatory subunits, especially in plants, are still poorly understood. We demonstrate that the inhibition of viral systemic transport after RPN9 silencing is largely due to alterations in the vascular tissue. RPN9-silenced plants display extra leaf vein formation with increased xylem and decreased phloem. We further illustrate that RPN9 functions at least in part through regulation of auxin transport and brassinosteroid signaling, two processes that are crucial for vascular formation. We propose that RPN9 regulates vascular formation by targeting a subset of regulatory proteins for degradation. The brassinosteroid-signaling protein BZR1 is one of the targets.

Arabidopsis↗

The Arabidopsis major intrinsic protein NIP5;1 is essential for efficient boron uptake and plant development under boron limitation.

Boron (B) is essential in plants but often present at low concentrations in the environment. To investigate how plants survive under conditions of B limitation, we conducted a transcriptome analysis and identified NIP5;1, a member of the major intrinsic protein family, as a gene upregulated in B-deficient roots of Arabidopsis thaliana. Promoter-beta-glucuronidase fusions indicated that NIP5;1 is strongly upregulated in the root elongation zone and the root hair zone under B limitation, and green fluorescent protein-tagged NIP5;1 proteins localized to the plasma membrane. Expression in Xenopus laevis oocytes demonstrated that NIP5;1 facilitated the transport of boric acid in addition to water. Importantly, two T-DNA insertion lines of NIP5;1 displayed lower boric acid uptake into roots, lower biomass production, and increased sensitivity of root and shoot development to B deficiency. These results identify NIP5;1 as a major plasma membrane boric acid channel crucial for the B uptake required for plant growth and development under B limitation.

Animals↗