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Yellow-green variegation leaf phenotype adds more value to ornamental plants, but it is regarded as an undesirable trait in crop plants, affecting their yields. Until recently, the underlying mechanism regulating the yellow-green variegation phenotype has remained largely unexplored in soybean. In the present study, we indentified four Glycine max leaf yellow/green variegation mutants, Gmvar1 , Gmvar2 , Gmvar3 , and Gmvar4 , from artificial mutagenesis populations. Map-based cloning, together with the allelic identification test and CRISPR-based gene knockout, proved that mutated GmCS1 controls yellow-green variegation phenotype of the Gmvar mutants. GmCS1 encodes a chorismate synthase in soybean. The content of Phe, Tyr, and Trp were dramatically decreased in Gmcs1 mutants. Exogenous supply of three aromatic amino acid mixtures, or only Phe to Gmvar mutants, leads to recovery of the mutant phenotype. The various biological processes and signalling pathways related to metabolism and biosynthesis were altered in Gmvar mutants. Collectively, our findings provide new insights about the molecular regulatory network of yellow-green variegation leaf phenotype in soybean. [ABSTRACT FROM AUTHOR]

We here study the level of albedo variegation on the nucleus of Comet 67P/Churyumov–Gerasimenko. This is done by fitting the parameters of a standard photometric phase function model to disc–average radiance factor data in images acquired by the Rosetta /OSIRIS Narrow Angle Camera in the orange filter. Local discrepancies between the observed radiance factor and the disc–average solution are interpreted as a proxy |$\mathcal {W}$| of the local single–scattering albedo. We find a wide range |$0.02 \lesssim \mathcal {W}\lesssim 0.09$| around an average of |$\mathcal {W}=0.055$|⁠. The observed albedo variegation is strongly correlated with nucleus morphology – smooth terrain is brighter, and consolidated terrain is darker, than average. Furthermore, we find that smooth terrain darken prior to morphological changes, and that stratigraphically low terrain (with respect to the centre of each nucleus lobe) is brighter than stratigraphically high terrain. We propose that the observed albedo variegation is due to differences in porosity and the coherent effect: compaction causes small brighter particles to act collectively as larger optically effective particles that are darker. Accordingly, we consider the dark consolidated terrain materials more compacted than smooth terrain materials, and darkening of the latter is due to subsidence. [ABSTRACT FROM AUTHOR]

The world region is missing from financialisation analysis of emerging economies (EEs) with little attention given to regional commonalities or comparative analysis across regions. This article sets out to identify regional commonalities in financialisation experiences across EEs, rooted in domestic institutions and countries' varying integration into the global financial system. Bringing commonalities within and differences between emerging regions to the fore will help us understand the specificities of their political economies and current capitalist experiences. Based on existing research and data availability, six financialisation indicators across five macroeconomic aggregates are identified. Considering 2008–17, a ranking emerges with the most affected region being Central Easter Europe (CEE), followed by Latin America. Emerging Asia takes an intermediate position while EEs in the Middle East and North Africa (MENA) region and Africa show limited signs of financialisation. Our analysis identifies distinct regional features. Financialisation in CEE and Latin America is strongly driven by external forces, the key difference being the role that large domestic capitalists play in the process. Across Emerging Asia, financialisation has mainly unfolded in the private sector while state authorities could to some extent insulate public policy. In MENA, much of the private sector appears outside of the reach of financialisation because of the economic and political power of regional conglomerates. In emerging Africa, the phenomenon is concentrated in very few, if intensely affected, financialisation centres. From a theoretical perspective, we find financialisation is driven by a mix of external factors and domestically influential capitalist elites pursuing their interests. [ABSTRACT FROM AUTHOR]

Changes in chromatin state may drive changes in gene expression, and it is of growing interest to understand the population genetic forces that drive differences in chromatin state. Here, we use the phenomenon of position effect variegation (PEV), a well-studied proxy for chromatin state, to survey variation in PEV among a naturally derived population. Further, we explore the genetic architecture of natural variation in factors that modify PEV. While previous mutation screens have identified over 150 suppressors and enhancers of PEV, it remains unknown to what extent allelic variation in these modifiers mediate interindividual variation in PEV. Is natural variation in PEV mediated by segregating genetic variation in known Su(var) and E(var) genes, or is the trait polygenic, with many variants mapping elsewhere in the genome? We designed a dominant mapping study that directly answers this question and suggests that the bulk of the variance in PEV does not map to genes with prior annotated impact to PEV. Instead, we find enrichment of top P-value ranked associations that suggest impact to active promoter and transcription start site proximal regions. This work highlights extensive variation in PEV within a population, and provides a quantitative view of the role naturally segregating autosomal variants play in modifying PEV-a phenomenon that continues to shape our understanding of chromatin state and epigenetics. [ABSTRACT FROM AUTHOR]

In eukaryotic cells, the organization of genomic DNA into chromatin regulates many biological processes, from the control of gene expression to the regulation of chromosome segregation. The proper maintenance of this structure upon cell division is therefore of prime importance during development for the maintenance of cell identity and genome stability. The chromatin assembly factor 1 (CAF-1) is involved in the assembly of H3-H4 histone dimers on newly synthesized DNA and in the maintenance of a higher order structure, the heterochromatin, through an interaction of its large subunit with the heterochromatin protein HP1a. We identify here a conserved domain in the large subunit of the CAF-1 complex required for its interaction with HP1a in the Drosophila fruit fly. Functional analysis reveals that this domain is dispensable for viability but participates in two processes involving heterochromatin: position-effect variegation and long range chromosomal interactions during meiotic prophase. Importantly, the identification in the large subunit of CAF-1 of a domain required for its interaction with HP1 allows the separation of its functions in heterochromatinrelated processes from its function in the assembly of H3-H4 dimers onto newly synthesized DNA. [ABSTRACT FROM AUTHOR]

The ornamental peach cultivar 'Hongbaihuatao (HBH)' can simultaneously bear pink, red, and variegated flowers on a single tree. Anthocyanin content in pink flowers is extremely low, being only 10% that of a red flower. Surprisingly, the expression of anthocyanin structural and potential regulatory genes in white flowers was not significantly lower than that in both pink and red flowers. However, proteomic analysis revealed a GST encoded by a gene--regulator involved in anthocyanin transport (Riant)--which is expressed in the red flower, but almost undetectable in the variegated flower. The Riant gene contains an insertion-deletion (indel) polymorphism in exon 3. In white flowers, the Riant gene is interrupted by a 2-bp insertion in the last exon, which causes a frameshift and a premature stop codon. In contrast, both pink and red flowers that arise from bud sports are heterozygous for the Riant locus, with one functional allele due to the 2-bp deletion or a novel 1 -bp insertion. Southern blot analysis indicated that the Riant gene occurs in a single copy in the peach genome and it is not interrupted by a transposon. The function of the Riant gene was confirmed by its ectopic expression in the Arabidopsis tt19 mutant, where it complements the anthocyanin phenotype, but not the proanthocyanidin pigmentation in seed coat. Collectively,these results indicate that a small indel mutation in the Riant gene, which is not the result of a transposon insertion or excision, causes variegated colouration of peach flowers. [ABSTRACT FROM AUTHOR]

Background and Aims Foliar variegation is recognized as arising from two major mechanisms: leaf structure and pigment-related variegation. Begonia has species with a variety of natural foliar variegation patterns, providing diverse examples of this phenomenon. The aims of this work are to elucidate the mechanisms underlying different foliar variegation patterns in Begonia and to determine their physiological consequences. Methods Six species and one cultivar of Begonia were investigated. Light and electron microscopy revealed the leaf structure and ultrastructure of chloroplasts in green and light areas of variegated leaves. Maximum quantum yields of photosystem II were measured by chlorophyll fluorescence. Comparison with a cultivar of Ficus revealed key features distinguishing variegation mechanisms. Key Results Intercellular space above the chlorenchyma is the mechanism of variegation in these Begonia. This intercellular space can be located (a) below the adaxial epidermis or (b) below the adaxial water storage tissue (the first report for any taxa), creating light areas on a leaf. In addition, chlorenchyma cell shape and chloroplast distribution within chlorenchyma cells differ between light and green areas. Chloroplasts from both areas showed dense stacking of grana and stroma thylakoid membranes. The maximum quantum yield did not differ significantly between these areas, suggesting minimal loss of function with variegation. However, the absence of chloroplasts in light areas of leaves in the Ficus cultivar led to an extremely low quantum yield. Conclusions Variegation in these Begonia is structural, where light areas are created by internal reflection between air spaces and cells in a leaf. Two forms of air space structural variegation occur, distinguished by the location of the air spaces. Both forms may have a common origin in development where dermal tissue becomes loosely connected to mesophyll. Photosynthetic functioning is retained in light areas, and these areas do not include primary veins, potentially limiting the costs of variegation. [ABSTRACT FROM PUBLISHER]

Mutants lacking a thylakoid membrane-bound metalloprotease, FtsH, are known to cause leaf variegation in Arabidopsis. However, the effect of reduced FtsH levels on leaf variegation has scarcely been examined in other plants. In this study, we performed RNA interference (RNAi) by which FtsH expression was suppressed in tobacco. The resulting FtsH knock-down tobacco plants showed variegation in their leaves, and a negative correlation between the degree of variegation and the level of FtsH, which supported earlier observations in Arabidopsis. A decrease of NtFtsH2 as well as NtFtsH1 suggested that these are the two major isoforms comprising the FtsH complex in tobacco chloroplasts. The RNAi tobacco lines also showed photoinhibition-vulnerable phenotypes, as evidenced by high-light-sensitive PSII activity and retarded degradation of D1 protein. Interestingly, the formation of variegated sectors during leaf development appeared to differ between Arabidopsis and tobacco. In contrast to the formation of variegation in Arabidopsis, the yellow sectors in FtsH RNAi tobacco emerged from green leaves at a late stage of leaf development. A series of cytological observations implied that thylakoid membranes were dismantled after development had already occurred. Late formation of variegation in FtsH RNAi tobacco suggested that the heteromeric FtsH complex is important for maintaining thylakoid membranes. [ABSTRACT FROM AUTHOR]

VAR2 is an integral thylakoid membrane protein and a member of the versatile FtsH class of metalloproteases in prokaryotes and eukaryotes. Recessive mutations in the VAR2 locus give rise to variegated plants (var2) that contain white sectors with abnormal plastids and green sectors with normal-appearing chloroplasts. In a continuing effort to isolate second-site suppressors of var2 variegation, we characterize in this report ems2505, a suppressor strain that has a virescent phenotype due to a missense mutation in At4g28590, the gene for a pioneer protein. We designated this gene SVR4 (for SUPPRESSOR OF VARIEGATION4) and the mutant allele in ems2505 as svr4-1. We demonstrate that SVR4 is located in chloroplasts and that svr4-1 single mutants are normal with respect to chloroplast anatomy and thylakoid membrane protein accumulation. However, they are modestly impaired in several aspects of photochemistry and have enhanced non-photochemical quenching (NPQ) capacity. A T-DNA insertion allele of SVR4, svr4-2, is seedling-lethal due to an early blockage of chloroplast development. We conclude that SVR4 is essential for chloroplast biogenesis, and hypothesize that SVR4 mediates some aspect of thylakoid structure or function that controls NPQ. We propose that in the suppressor strain, photoinhibitory pressure caused by a lack of VAR2 is ameliorated early in chloroplast development by enhanced NPQ capacity caused by reduced SVR4 activity. This would result in an increase in the number of chloroplasts that are able to surmount a threshold necessary to avoid photo-damage and thereby develop into functional chloroplasts. [ABSTRACT FROM PUBLISHER]

The Arabidopsis (Arabidopsis thaliana) yellow variegated2 (var2) mutant has green- and white-sectored leaves due to loss of VAR2, a subunit of the chloroplast FtsH protease/chaperone complex. Suppressor screens are a valuable tool to gain insight into VAR2 function and the mechanism of var2 variegation. Here, we report the molecular characterization of 004-003, a line in which var2 variegation is suppressed. We found that the suppression phenotype in this line is caused by lack of a chloroplast pentatricopeptide repeat (PPR) protein that we named SUPPRESSOR OF VARIEGATION7 (SVR7). PPR proteins contain tandemly repeated PPR motifs that bind specific RNAs, and they are thought to be central regulators of chloroplast and mitochondrial nucleic acid metabolism in plants. The svr7 mutant has defects in chloroplast ribosomal RNA (rRNA) processing that are different from those in other svr mutants, and these defects are correlated with reductions in the accumulation of some chloroplast proteins, directly or indirectly. We also found that whereas var2 displays a leaf variegation phenotype at 22°C, it has a pronounced chlorosis phenotype at 8°C that is correlated with defects in chloroplast rRNA processing and a drastic reduction in chloroplast protein accumulation. Surprisingly, the cold-induced phenotype of var2 cannot be suppressed by svr7. Our results strengthen the previously established linkage between var2 variegation and chloroplast rRNA processing/chloroplast translation, and they also point toward the possibility that VAR2 mediates different activities in chloroplast biogenesis at normal and chilling temperatures. [ABSTRACT FROM AUTHOR]

FtsH is an ATP-dependent metalloprotease present as a hexameric heterocomplex in thylakoid membranes. Encoded in the Arabidopsis thaliana YELLOW VARIEGATED2 (VAR2) locus, FtsH2 is one isoform among major Type A (FtsH1/5) and Type B (FtsH2/8) isomers. Mutants lacking FtsH2 (var2) and FtsH5 (var1) are characterized by a typical leaf-variegated phenotype. The functional importance of the catalytic center (comprised by the zinc binding domain) in FtsH2 was assessed in this study by generating transgenic plants that ectopically expressed FtsH2(488), a proteolytically inactive version of FtsH2. The resulting amino acid substitution inhibited FtsH protease activity in vivo when introduced into Escherichia coli FtsH. By contrast, expression of FtsH2(488) rescued not only leaf variegation in var2 but also seedling lethality in var2 ftsh8 , suggesting that the protease activity of Type B isomers is completely dispensable, which implies that the chloroplastic FtsH complex has protease sites in excess and that they act redundantly rather than coordinately. However, expression of FtsH2(488) did not fully rescue leaf variegation in var1 var2 because the overall FtsH levels were reduced under this background. Applying an inducible promoter to our complementation analysis revealed that rescue of leaf variegation indeed depends on the overall amount of FtsH. Our results elucidate protein activity and its amount as important factors for the function of FtsH heterocomplexes that are composed of multiple isoforms in the thylakoid membrane. [ABSTRACT FROM AUTHOR]

Leaf variegation is seen in many ornamental plants and is often caused by a cell-lineage type formation of white sectors lacking functional chloroplasts. A mutant showing such leaf variegation is viable and is therefore suitable for studying chloroplast development. In this study, the formation of white sectors was temporally investigated in the Arabidopsis leaf-variegated mutant var2. Green sectors were found to emerge from white sectors after the formation of the first true leaf. Transmission electron microscopic examination of plastid ultrastructures confirmed that the peripheral zone in the var2 shoot meristem contained proplastids but lacked developing chloroplasts that were normally detected in wild type. These data suggest that chloroplast development proceeds very slowly in var2 variegated leaves. A notable feature in var2 is that the plastids in white sectors contain remarkable globular vacuolated membranes and prolamellar body-like structures. Although defective plastids were hardly observed in shoot meristems, they began to accumulate during early leaf development. Consistent with these observations, large plastid nucleoids detected in white sectors by DNA-specific fluorescent dyes were characteristic of those found in proplastids and were clearly distinguished from those in chloroplasts. These results strongly imply that in white sectors, differentiation of plastids into chloroplasts is arrested at the early stage of thylakoid development. Interestingly, large plastid nucleoids were detected in variegated sectors from species other than Arabidopsis. Thus, plastids in variegated leaves appear to share a common feature and represent a novel plastid type. [ABSTRACT FROM PUBLISHER]

We hypothesized that chloroplast energy imbalance sensed through alterations in the redox state of the photosynthetic electron transport chain, measured as excitation pressure, governs the extent of variegation in the immutans mutant of Arabidopsis thaliana. To test this hypothesis, we developed a nondestructive imaging technique and used it to quantify the extent of variegation in vivo as a function of growth temperature and irradiance. The extent of variegation was positively correlated (R² = 0.750) with an increase in excitation pressure irrespective of whether high light, low temperature, or continuous illumination was used to induce increased excitation pressure. Similar trends were observed with the variegated mutants spotty, var1, and var2. Measurements of greening of etiolated wild-type and immutans cotyledons indicated that the absence of IMMUTANS increased excitation pressure twofold during the first 6 to 12 h of greening, which led to impaired biogenesis of thylakoid membranes. In contrast with IMMUTANS, the expression of its mitochondrial analog, AOXla, was transiently upregulated in the wild type but permanently upregulated in immutans, indicating that the effects of excitation pressure during greening were also detectable in mitochondria. We conclude that mutations involving components of the photosynthetic electron transport chain, such as those present in immutans, spotty, var1, and var2, predispose Arabidopsis chloroplasts to photooxidation under high excitation pressure, resulting in the variegated phenotype. e [ABSTRACT FROM AUTHOR]

The Arabidopsis thaliana yellow variegated2 (var2) mutant is variegated due to lack of a chloroplast FtsH-like metal-Ioprotease (FtsH2/VAR2). We have generated suppressors of var2 variegation to gain insight into factors and pathways that interact with VAR2 during chloroplast biogenesis. Here, we describe two such suppressors. Suppression of variegation in the first line, TAG-FN, was caused by disruption of the nuclear gene (SUPPRESSOR OF VARIEGATION1 [SVR1]) for a chloroplast-localized homolog of pseudouridine (Ψ) synthase, which isomerizes uridine to ψ in noncoding RNAs. svr1 single mutants were epistatic to var2, and they displayed a phenotypic syndrome that included defects in chloroplast rRNA processing, reduced chloroplast translation, reduced chloroplast protein accumulation, and elevated chloroplast mRNA levels. In the second line (TAG-IE), suppression of variegation was caused by a lesion in SVR2, the gene for the ClpR1 subunit of the chloroplast ClpP/R protease. Like svr1, svr2 was epistatic to var2, and clpR1 mutants had a phenotype that resembled svr1. We propose that an impairment of chloroplast translation in TAG-FN and TAG-IE decreased the demand for VAR2 activity during chloroplast biogenesis and that this resulted in the suppression of var2 variegation. Consistent with this hypothesis, var2 variegation was repressed by chemical inhibitors of chloroplast translation. In planta mutagenesis revealed that SVR1 not only played a role in uridine isomerization but that its physical presence was necessary for proper chloroplast rRNA processing. Our data indicate that defects in chloroplast rRNA processing are a common, but not universal, molecular phenotype associated with suppression of var2 variegation. [ABSTRACT FROM AUTHOR]

The yellow variegated2 (var2) is one of the best-characterized Arabidopsis (Arabidopsis thaliana) mutants showing leaf variegation. Leaf variegation of var2 results from the loss of an ATP-dependent metalloprotease, FtsH2, which is a major component of the FtsH heterocomplex in thylakoid membranes. While the functional role of FtsH2 in protein quality control has been extensively studied, the physiological state of plastids in white tissues of the var2 is not well characterized. Here we show that the white tissue in var2 is neither the result of photobleaching nor enhanced senescence. Visualization of plastids by plastid-targeted green fluorescent protein revealed that plastids in the white sector are distinct and have undifferentiated characteristics. The plastids are also distinct in that they contain large nucleoids, a complex structure of plastid DNA and proteins, that are typically found in undifferentiated plastids. Comparative analyses of protein profiles from green and white tissues suggested that the difference was observed in the proteins related to photosynthesis but not due to proteins of other organelles. Thus, cells in the white tissue are viable and their defect is limited to plastid function. The plastid accumulates normal levels of chloroplast transcripts, whereas a substantial repression of nuclear-encoded photosynthetic genes was evident in the white sector. Based upon these results, we inferred that the white sectors in var2 are made by viable cells that have plastids arrested in thylakoid formation. A proposed model to form the variegated sector in var2 is provided. [ABSTRACT FROM AUTHOR]

In this article we show that hypomorphic loss-of-function alleles of the JIL-1 histone H3S10 kinase are strong suppressors of position effect variegation (PEV) of the wm4 allele and that lack of JTL-1 activity can counteract the effect of the dominant enhancer E(var)2-1 on PEV. [ABSTRACT FROM AUTHOR]

Plant variegations are characterized by the presence of white sectors in normally green tissues and organs. Whereas the white sectors contain defective plastids that lack coloured pigments, the green sectors contain morphologically normal chloroplasts. Variegation mutants are defective in chloroplast developmental processes and arise due to mutations in nuclear or organellar genes. Despite their widespread occurrence in nature, only a few variegations have been studied at the molecular level. In this review, recent progress toward understanding two Arabidopsis variegations, immutans (im) and var2 is summarized. Both im and var2 are caused by nuclear recessive mutations and the responsible genes have been cloned and characterized. IMMUTANS functions as a chloroplast terminal oxidase that transfers electrons from the plastoquinol pool to oxygen. It appears to be a versatile electron sink, especially early in chloroplast development, when its function is crucial for carotenoid biosynthesis, and in excess light, when it serves as a 'safety valve'. IM also probably functions in chlororespiration. VAR2 encodes a chloroplast FtsH metalloprotease (termed AtFtsH2). Along with other AtFtsH proteins (AtFtsH1, 5 and 8), it forms complexes in the thylakoid membrane that are probably involved in the process of PSII repair during photoinhibition. A model has been proposed to explain the mechanism of var2 variegation, which suggests that threshold levels of FtsH complexes are required for green sector formation. It is concluded that studies on im and var2 have provided novel insights into nuclear--chloroplast interactions and, especially, into mechanisms of photoprotection. [ABSTRACT FROM AUTHOR]

We studied the patterns of sexual dichromatism and seasonal variation in plumage color in the Diademed Tanager (Stephanophorus diadematus), a species previously considered devoid of variation in adult plumage. The general coloration of this species is dark blue-violet, with a white-blue and red crown. Plumage reflectance of seven body regions from 33 study skins belonging to adults of both sexes was measured. Reflectance values were used in a principal components analysis (PCA) and hue, short-wave chroma, and UV chroma were also measured directly on the spectra. Both PCA factor scores and these latter variables were subjected to two- way ANCOVAs with sex and season as main factors and the year of capture as a covariate. We found that crowns of males were significantly brighter than those of females. In addition, the nape, chest, and belly showed significant differences in spectral shape, with relatively greater short-wave reflectance and less long-wave reflectance in males than in females. Although sexes were alike in hue, they differed in chroma in almost all body regions. Brightness also differed between seasons, and contrary to our expectation nonbreeding birds were brighter than breeding ones. This result may be a consequence of the particular molt program of tanagers that includes only a complete post-reproductive molt. Despite finding seasonal differences in spectral shape in various body regions, no significant changes in hue, short-wave chroma, or (IV chroma were evident. To our knowledge, this is the first report of variation in adult plumage color for the Diademed Tanager, and we suggest that dichromatism in tanagers may be even more pervasive than is currently recognized. [ABSTRACT FROM AUTHOR]

Bonus, a Drosophila TIF1 homolog, is a nuclear receptor cofactor required for viability, molting, and numerous morphological events. Here we establish a role for Bonus in the modulation of chromatin structure. We show that weak loss-of-function alleles of bonus have a more deleterious effect on males than on females. This male-enhanced lethality is not due to a defect in dosage compensation or somatic sex differentiation, but to the presence of the Y chromosome. Additionally, we show that bonus acts as both an enhancer and a suppressor of position-effect variegation. By immunostaining, we demonstrate that Bonus is associated with both interphase and prophase chromosomes and through chromatin immunopre- cipitation show that two of these sites correspond to the histone gene cluster and the Stellate locus. [ABSTRACT FROM AUTHOR]

• Background and Aims Sections leaves of Ficus rubiginosa ‘Variegata’ show that it is a chimera with a chlorophyll deficiency in the second layer of the leaf meristem (GWG structure). Like other Ficus species, it has a multiseriate epidermis on the adaxial and abaxial sides of the leaf, formed by periclinal cell divisions as well as anticlinal divisions. The upper and lower laminae of the leaf often exhibit small dark and light green patches of tissue overlying internal leaf tissue.• Methods The distribution of chlorophyll in transverse sections of typical leaves was determined by fluorescence microscopy.• Key Results Patches of dark and light green tissue which arise in the otherwise colourless palisade and spongy mesophyll tissue in the entire leaf are due to further cell divisions arising from the bundle sheath which is associated with major vascular bundles or from the green multiseriate epidermis. Leaves produced in winter exhibit more patches of green tissue than leaves which expand in mid‐summer. Many leaves produced in summer have no spotting and appear like a typical GWG chimera. There is a strong relationship between the number of patches on the adaxial side of leaves and the number on the abaxial side, showing that the cell division in upper and lower layers of leaves is strongly coordinated. In both winter and summer, there are fewer patches on the abaxial side of leaves compared with the adaxial side, indicating that periclinal and anticlinal cell divisions from the outer meristematic layer are less frequent in the lower layers of leaf tissue. Most of the patches are small (<1 mm in longest dimension) and thus the cell divisions which form them occur late in leaf development. Leaves which exhibit large patches generally have them on both sides of the leaves.• Conclusion In this cultivar, the outer meristematic layer appears to form vascular bundle sheaths and associated internal leaf tissue in the entire leaf lamina. [ABSTRACT FROM AUTHOR]

Arabidopsis YELLOW VARIEGATED1 (VAR1) and VAR2 are separate loci that encode similar chloroplast FtsH proteases. To date, FtsH is the best-characterized protease in thylakoid membranes involved in the turnover of photosynthetic protein complexes. It comprises a protein family that is encoded by 12 different nuclear genes in Arabidopsis. We show here that nine FtsH proteins are located in the chloroplasts. Mutations in either VAR1 or VAR2 cause typical leaf variegation and sensitivity to photoinhibition. By contrast, none of these phenotypes was observed in T-DNA insertion mutants in other ftsH genes (ftsh1, ftsh6, and ftsh8) closely related to VAR1 and VAR2. This finding suggests that VAR1 and VAR2 play a predominant role in the photosystem II repair cycle in thylakoid membranes. By generating VAR1- and VAR2-specific antibodies, we found that loss of either VAR1 or VAR2 results in the decreased accumulation of the other. Thus, the genetic nonredundancy between VAR1 and VAR2 could be attributed to their coordinated regulation at the protein level. These observations led us to examine whether VAR1 and VAR2 form a complex. Sucrose density gradient and gel filtration analyses revealed a complex of ∼400 to 450 kD, probably representing a hexamer. Furthermore, VAR1 and VAR2 were shown to coprecipitate by immunoprecipitation using VAR1- and VAR2-specific antibodies. The majority of VAR1 appears to exist as heterocomplexes with VAR2, whereas VAR2 may be present as homocomplexes as well. Based on these results, we conclude that VAR1 and VAR2 are the major components of an FtsH complex involved in the repair of photodamaged proteins in thylakoid membranes. [ABSTRACT FROM AUTHOR]

Both defensive and nutritional hypotheses have been used to explain the improved performance of sucking insects on variegated plants. With portions of leaves unable to photosynthesize, variegated plants are likely to have less carbon assimilate to allocate to the production of defensive compounds than green plants. Alternatively, like plants grown in the shade, improved response of sucking insects on variegated plants may be caused by greater mobility of amino acid nitrogen and carbohydrates in the phloem to fuel compensatory production of leaves. We examined amino acid and sucrose content of stem exudate collected from red-variegated, yellow-variegated, and green-leafed coleus plants, Solenostemon scutellarioides (L.) Codd, and their relation to life history characteristics of citrus mealybug, Planococcus citri (Risso), to test these defensive and nutritional hypotheses. Although higher concentrations of both total amino acids and sucrose were found in green- and red-leaved plants, these quantitative measures were uncorrelated with life history parameters of mealybugs. In contrast, our analysis of qualitative measures of amino acid composition supported both defensive and nutritional hypotheses. Green-leafed plants had the greatest levels of shikimic acid precursors and lowest levels of other nonessential amino acids. Life history characteristics of citrus mealybugs were correlated negatively with increasing proportions of shikimie acid precursors and positively with increasing proportions of other nonessential amino acids. Thus, our studies of life history and stem chemistry indicate that improved suitability of variegated plants is consistent with both carbon- and nutrient-based theories of plant defense. [ABSTRACT FROM AUTHOR]

Examines the multiple effects of genetic background on variegated transgene expression in mice. Discussion on ovine &Beta-lactoglobulin transgene during lactation; Evaluation on inbreeding depression for selectively downregulating transgene expression; Description on the defense system to inactivate foreign sequences.

Examines the modifiers of terminal deficiency-associated position effect of variegation (PEV) in Drosophila. Suppression of PEV by the presence of second minichromosomes; Characterization of mutation; Analysis of mitotic chromosomes from homozygous larvae.

Examines the genetic recombination effect on suppressor of position-effect variegation in Drosophila. Induction of gene silencing in position-effect variegation; Suppression of data by heterochromatin frequencies in the genome; Effects of mutation in proximal euchromatin.

Describes pitkin[sup dominant] (ptn[sup D]), the gain-of-function enhancer of position-effect variegation (PEV) in Drosophila. Characteristics of modifier genes of PEV in Drosophila; Enhancing capabilities of ptn[sup D]; Role played by ptn gene in chromatin regulation.

Describes a dicistronic transcription unit that evolved by the fusion of two genes, the suppressor of position-effect variegation Su(var)3-9 and the gene encoding the gamma subunit of translation initiation factor eIF-2. Su(var)3-9/eIF-2 gamma dicistronic transcription unit of holometabolic insects; Characterization of eIF-2 gamma gene structure in arthropods.

Explores genetic and molecular complexity of the position effect variegation modifier mod(mdg4) in Drosophila. Need for gene regulation in spatial and temporal expression of multicellular organism development; Identification of chromatin proteins involved in structural changes of transcriptional regulation; Details on the characterization of deduced proteins.

Examines the significance of size and internal heterochromatic block structure in determining variegation in Drosophila melanogaster. Transposition of the 1A-2E euchromatic segment to the X heterochromatin (Xh) vacinity; Effect of In(1LR)pn2a irridiation on position effect variegation (PEV); Correlation of wapl PEV with Xh centromere-carrying blocks.

Studies the inheritance, karyotype and temperature effects of a variegation mutant phenotype called Mosaic (Mos.). Origin and phenotype of Mos; Genetic analysis of Mos; Materials and methods used in the study; Results and discussion.

Examines the factors influencing the spreading of variegation in the Drosophila melanogaster's white-roughest region. Gene inactivation from heterochromatin into adjacent euchromatin in the Drosophila melanogaster; Silencing of the white and roughest genes in the variegating rearrangements.

Analyzes genomic imprinting and position-effect variegation in a mini-X chromosome in Drosophila melanogaster. Difference in imprinting when genes are transmitted by male or female parent; Effect of factors that modify position-effect variegation on somatic expression; Importance of chromatin structure in maintenance of genomic imprint.

Studies changes in the eye color of Drosophila resulting from a euchromatic-heterochromatin fusion gene. Isolation of the dominant mutant gene pugilist (pugD); Nature of pugD mutation; Mechanism of pugD action; Patterned pteridine pigmentation in pugD; Variegated pigmentation in pugD.

Studies the ability of mutant genes of ribosomal protein PO to modify gene expression and variegation in the fly Drosophila. Isolation and cloning of P-element insertion mutant l(3)01544; Experimental procedure; Effect of mutant allele PO l(3)01544 on the transcript abundance of unrelated genes.

Reports that an enhancer of position-effect variegation in Drosophila melanogaster encodes a putative ribonucleic acid (RNA) helicase, which binds chromosomes and is regulated by the cell cycle. Information on position-effect variegation in Drosophila melanogaster; Reference to the importance of the product of the white gene of Drosophila.

Investigates the classical position-effect variegation (PEV) of a Drosophila brown transgene affected by a heterochromatic junction. Enhancement of PEV when the transgene was locally duplicated using P transposase; Enhancement as revertable; Selection for enhanced variegation yielding transgene duplications; Effect of orientation on susceptibility to PEV.

Examines the results of a study on the inheritance of a variegated seed coat color mutation and its linkage relationship with the high palmitic acid mutation. Detailed description of the methodology used in the study; Discussion on its results.