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Reversing transmembrane electron flow: the DsbD and DsbB protein families.

DsbD and DsbB are two proteins that in Escherichia coli catalyze transmembrane electron flow in opposite directions, thereby allowing reversible oxidoreduction of periplasmic dithiol/disulfide-containing proteins. We have identified all recognizable homologues of these two proteins in the databases and have conducted structural and phylogenetic analyses of the two families. The larger DsbD family is more diverse in sequence, topology, function and organismal distribution than the smaller DsbB family. DsbB homologues are rarely found outside of the proteobacteria, although DsbD homologues are found in many bacterial kingdoms as well as archaea and plant chloroplasts. Few organisms with a fully sequenced genome and a DsbB homologue lack a DsbD homologue, and most of these DsbD homologues fall within two clusters in the DsbD tree, exhibiting phylogenetic relationships that are the same as those observed for the DsbB proteins. These observations suggest that a subset of the DsbD homologues evolved in parallel with the DsbB family to perform a single unified function involving reversible extracytoplasmic protein dithiol-disulfide bond interchange. DsbD family proteins are shown to have arisen by an internal gene duplication event, and this observation leads to prediction of the pathway taken for the evolutionary appearance of the different protein topological types found within this family.

Archaea↗

The yfhQ gene of Escherichia coli encodes a tRNA:Cm32/Um32 methyltransferase.

BACKGROUND: Naturally occurring tRNAs contain numerous modified nucleosides. They are formed by enzymatic modification of the primary transcripts during the complex RNA maturation process. In model organisms Escherichia coli and Saccharomyces cerevisiae most enzymes involved in this process have been identified. Interestingly, it was found that tRNA methylation, one of the most common modifications, can be introduced by S-adenosyl-L-methionine (AdoMet)-dependent methyltransferases (MTases) that belong to two structurally and phylogenetically unrelated protein superfamilies: RFM and SPOUT. RESULTS: As a part of a large-scale project aiming at characterization of a complete set of RNA modification enzymes of model organisms, we have studied the Escherichia coli proteins YibK, LasT, YfhQ, and YbeA for their ability to introduce the last unassigned methylations of ribose at positions 32 and 34 of the tRNA anticodon loop. We found that YfhQ catalyzes the AdoMet-dependent formation of Cm32 or Um32 in tRNASer1 and tRNAGln2 and that an E. coli strain with a disrupted yfhQ gene lacks the tRNA:Cm32/Um32 methyltransferase activity. Thus, we propose to rename YfhQ as TrMet(Xm32) according to the recently proposed, uniform nomenclature for all RNA modification enzymes, or TrmJ, according to the traditional nomenclature for bacterial tRNA MTases. CONCLUSION: Our results reveal that methylation at position 32 is carried out by completely unrelated TrMet(Xm32) enzymes in eukaryota and prokaryota (RFM superfamily member Trm7 and SPOUT superfamily member TrmJ, respectively), mirroring the scenario observed in the case of the m1G37 modification (introduced by the RFM member Trm5 in eukaryota and archaea, and by the SPOUT member TrmD in bacteria).

Escherichia coli Proteins↗

JEvTrace: refinement and variations of the evolutionary trace in JAVA.

BACKGROUND: Details of functional speciation within gene families can be difficult to identify using standard multiple sequence alignment (MSA) methods. The evolutionary trace (ET) was developed as a visualization tool to combine MSA, phylogenetic and structural data for identification of functional sites in proteins. The method has been successful in extracting evolutionary details of functional surfaces in a number of biological systems and modifications of the method are useful in creating hypotheses about the function of previously unannotated genes. We wish to facilitate the graphical interpretation of disparate data types through the creation of flexible software implementations. RESULTS: We have implemented the ET method in a JAVA graphical interface, JEvTrace. Users can analyze and visualize ET input and output with respect to protein phylogeny, sequence and structure. Function discovery with JEvTrace is demonstrated on two proteins with recently determined crystal structures: YlxR from Streptococcus pneumoniae with a predicted RNA-binding function, and a Haemophilus influenzae protein of unknown function, YbaK. To facilitate analysis and storage of results we propose a MSA coloring data structure. The sequence coloring format readily captures evolutionary, biological, functional and structural features of MSAs. CONCLUSIONS: Protein families and phylogeny represent complex data with statistical outliers and special cases. The JEvTrace implementation of the ET method allows detailed mining and graphical visualization of evolutionary sequence relationships.

Amino Acid Sequence↗

Evidence for dual pathways of Tc1/mariner domestication in Drosophila.

BACKGROUND: The domestication of transposable elements is a key source of evolutionary innovation, yet the pathways by which their functional modules are repurposed by the host remain poorly understood. The Tc1/mariner superfamily is a widespread group of DNA transposons, but the prevalence and patterns of their domestication are underexplored. RESULTS: We performed a systematic genomic screen across 43 drosophilid species using stringent criteria for molecular domestication. This analysis identified five high-confidence, evolutionarily conserved genes derived from Tc1/mariner transposases. Phylogenetic and structural analyses suggest domestication via two distinct molecular pathways: co-option of the DNA-binding module and co-option of the catalytic domain. The DNA-binding module pathway includes CG4570, the previously known genes cag and toy (the latter fused with a homeodomain), and a lineage-restricted gene in the Drosophila obscura group that exhibits signatures of recent domestication. In contrast, the catalytic domain pathway is represented solely by CG14478. Structural modeling reveals that CG14478 protein preserves a canonical DDE endonuclease fold. Co-expression network analysis suggests potential cellular roles of these genes: CG14478 is linked to RNA/chromatin-related processes, CG4570 to cell cycle/chromosome functions, cag to ciliary and nuclear functions, and toy to neuronal development. CONCLUSIONS: This study establishes a stringent framework for identifying domesticated TEs, demonstrating that Tc1/mariner elements are co-opted via two distinct pathways: retention of either catalytic or DNA-binding modules. Our findings suggest that domestication is a dynamic continuum, ranging from recent, lineage-specific events to ancient, conserved genes, and underscore how genomic conflict with TEs can drive eukaryotic evolution and regulatory complexity.

Animals↗

Structural, biochemical, and expression analysis of two distinct insulin-like growth factor I receptors and their ligands in zebrafish.

We have cloned and characterized cDNAs encoding the zebrafish IGF ligands and receptors. Sequence comparison showed that the primary structures of zebrafish IGF-I, IGF-II, and IGF-I receptors (IGF-IRs) have been highly conserved in vertebrates. In contrast to the presence of a single IGF-IR gene in mammals, two distinct IGF-IR genes, termed igf-1ra and igf-1rb, were found in zebrafish. Structural and phylogenetic analyses indicated that both genes are orthologous to the human igf-1r gene. Immunoprecipitation studies with specific antibodies showed that both IGF-IR genes are expressed and both receptors bind to IGFs and des(1-3)IGF-I, but not to insulin. The spatio-temporal expression patterns of the two IGF-IRs and their ligands were determined using a combination of RT-PCR, whole mount in situ hybridization, and immunocytochemistry. Transcripts for both IGF-I and -II mRNAs were found throughout embryogenesis in a ubiquitous manner. In adult tissues, IGF-I mRNA was more abundant in liver and testis, and its level was increased after GH treatment, whereas IGF-II mRNA was not regulated by GH. IGF-IRa and IGF-IRb mRNAs and proteins were expressed in overlapping spatial domains, but exhibited distinct temporal expression patterns. In particular, the relative level of IGF-IRa mRNA was low during early embryogenesis and increased in the hatched larva, whereas the situation was reversed for IGF-IRb mRNA. In adult zebrafish, the overall tissue distribution patterns of the two IGF-IRs were similar, but there were differences in their cellular localization and relative abundance in defined cells/regions. The differential expression pattern of IGF-IRa and IGF-IRb suggest that they may play distinct roles in regulating the growth and development of zebrafish.

Amino Acid Sequence↗

Evolution of novel functions: cryptocyanin helps build new exoskeleton in Cancer magister.

Hemocyanin, the blue blood protein of many arthropods and molluscs, reversibly binds oxygen at its highly conserved copper-oxygen-binding sites and supplies tissues with oxygen. Cryptocyanin, closely related structurally and phylogenetically to arthropod hemocyanin, lacks several of the six critical copper-binding histidines, however, and has lost the ability to bind oxygen. Despite this loss of function, cryptocyanin continues to be synthesized, an indication that it has been exploited to carry out new functions. Here, we show that cryptocyanin is present in extremely high concentrations in the hemolymph of the crab during the premolt portion of the molt cycle. Both proteins are specifically expressed in the same type of cell in the hepatopancreas and secreted into the hemolymph, but cryptocyanin plays a major role in forming the new exoskeleton, while hemocyanin functions in oxygen transport. A cessation in cryptocyanin, but not hemocyanin, synthesis after eyestalk ablation supports our hypothesis that cryptocyanin is closely regulated by molting hormones. The contrasts between the two gene products illustrate how a gene duplication of a copper-oxygen protein and its subsequent mutation may work in concert with the evolution of new regulatory mechanisms, leading to the assumption of new functions.

Animals↗

Novel cysteine proteinase inhibitors homologous to the proregions of cysteine proteinases.

Propeptides of papain-like cysteine proteinases such as papain, cathepsins B, L and S are potent inhibitors of their cognate cysteine proteinases with Ki values in the nanomolar range, and they exhibit highest inhibition selectivity for enzymes from which they originate. Recent studies have identified novel inhibitor proteins that are homologous to the proregions of papain-like cysteine proteinases. Mouse activated T-lymphocytes express cytotoxic T-lymphocyte antigen (CTLA-2), which is homologous to the proregion of mouse cathepsin L. CTLA-2 exhibits inhibitory activities to several cysteine proteinases. We have also identified a similar propeptide-like cysteine proteinase inhibitor, Bombyx cysteine proteinase inhibitor (BCPI), in the silkmoth Bombyx mori. BCPI is a slow and tight binding inhibitor of cathepsin L-like cysteine proteinases with Ki values in picomolar range, and the inhibition is highly selective towards these proteinases just like the propeptides. Recent genome analyses have shown the expression of similar propeptide-like proteins in Drosophila and rat, suggesting the presence of a novel class of cysteine proteinase inhibitors in a variety of organisms. Studies of the gene structures and phylogenetic analysis have shown that genes of the propeptide-like cysteine proteinase inhibitors have emerged from ancestor genes of their parental enzymes.

Amino Acid Sequence↗

Genome-Wide Characterization of the Apple HD-Zip IV Gene Family and Functional Validation of MdHDZIV3 Under PEG-Induced Osmotic Stress.

The homeodomain-leucine zipper IV (HD-Zip IV) transcription factor subfamily plays essential roles in epidermal development, cuticle formation, lipid metabolism, and environmental adaptation in plants. Despite its biological importance, the HD-Zip IV family has not been systematically characterized in apple (Malus domestica). Here, we identified 17 apple HD-Zip IV genes and named them MdHDZIV1-MdHDZIV17 based on their locations on the chromosomes. The 17 genes showed a nonuniform distribution on eight chromosomes, while the occurrence of both tandem and segmental duplications indicated that family expansion involved more than one duplication mechanism. All MdHDZIV proteins contained the conserved HD, LZ, START, and SAD domains but lacked the MEKHLA domain, consistent with typical HD-Zip IV structural features. Phylogenetic analysis classified MdHDZIV proteins into five groups together with HD-Zip IV members from Arabidopsis thaliana and rice, indicating evolutionary conservation of this subfamily. Collinearity and Ka/Ks analyses revealed that duplicated MdHDZIV gene pairs were mainly subjected to purifying selection. Promoter scanning revealed diverse cis-regulatory motifs associated with hormonal signaling, environmental stress, light response, and epidermal regulation, including ABRE, ARE, W-box, MYC, G-box, and L1-box motifs. Integration of transcriptomic profiling with qRT-PCR validation revealed pronounced tissue-dependent differences in the expression of MdHDZIV genes in leaf, fruit skin, and branch bark. Under PEG6000-induced osmotic stress and NaCl-induced salt stress, 10 candidate MdHDZIV genes displayed gene-specific and stress type-specific expression patterns, with MdHDZIV3 showing strong induction under PEG6000 treatment. Functional validation in apple calli showed that MdHDZIV3 overexpression enhanced PEG tolerance, increased fresh weight, elevated SOD and POD activities, and reduced MDA accumulation under osmotic stress. These findings provide a genome-wide framework for understanding the apple HD-Zip IV gene family.

abiotic stress↗

Synthetic TLR agonists reveal functional differences between human TLR7 and TLR8.

Although TLR7 and TLR8 are phylogenetically and structurally related, their relative functions are largely unknown. The role of TLR7 has been established using TLR7-deficient mice and small molecule TLR7 agonists. The absence of TLR8-selective agonists has hampered our understanding of the role of TLR8. In this study TLR agonists selective for TLR7 or TLR8 were used to determine the repertoire of human innate immune cells that are activated through these TLRs. We found that TLR7 agonists directly activated purified plasmacytoid dendritic cells and, to a lesser extent, monocytes. Conversely, TLR8 agonists directly activated purified myeloid dendritic cells, monocytes, and monocyte-derived dendritic cells (GM-CSF/IL-4/TGF-beta). Accordingly, TLR7-selective agonists were more effective than TLR8-selective agonists at inducing IFN-alpha- and IFN-regulated chemokines such as IFN-inducible protein and IFN-inducible T cell alpha chemoattractant from human PBMC. In contrast, TLR8 agonists were more effective than TLR7 agonists at inducing proinflammatory cytokines and chemokines, such as TNF-alpha, IL-12, and MIP-1alpha. Thus, this study demonstrated that TLR7 and TLR8 agonists differ in their target cell selectivity and cytokine induction profile.

Aminoquinolines↗

Identification and characterization of a FcR homolog in an ectothermic vertebrate, the channel catfish (Ictalurus punctatus).

An FcR homolog (IpFcRI), representing the first such receptor from an ectothermic vertebrate, has been identified in the channel catfish (Ictalurus punctatus). Mining of the catfish expressed sequence tag databases using mammalian FcR sequences for CD16, CD32, and CD64 resulted in the identification of a teleost Ig-binding receptor. IpFcRI is encoded by a single-copy gene containing three Ig C2-like domains, but lacking a transmembrane segment and cytoplasmic tail. The encoded Ig domains of IpFcRI are phylogenetically and structurally related to mammalian FcR and the presence of a putative Fc-binding region appears to be conserved. IpFcRI-related genomic sequences are also present in both pufferfish and rainbow trout, indicating the likely presence of a soluble FcR in other fish species. Northern blot and qualitative PCR analyses demonstrated that IpFcRI is primarily expressed in IgM-negative leukocytes derived from the lymphoid kidney tissues and PBL. Significantly lower levels of IpFcRI expression were detected in catfish clonal leukocyte cell lines. Using the native leader, IpFcRI was secreted when transfected into insect cells and importantly the native IpFcRI glycoprotein was detected in catfish plasma using a polyclonal Ab. Recombinant IpFcRI binds catfish IgM as assessed by both coimmunoprecipation and cell transfection studies and it is presumed that it functions as a secreted FcR akin to the soluble FcR found in mammals. The identification of an FcR homolog in an ectothermic vertebrate is an important first step toward understanding the evolutionary history and functional importance of vertebrate Ig-binding receptors.

Amino Acid Sequence↗

The cisterna magna septa: vestigial remnants of Blake's pouch and a potential new marker for normal development of the rhombencephalon.

OBJECTIVE: The purpose of this study was to show the normal sonographic embryologic anatomy of the cisterna magna septa, fourth ventricle, and cerebellar vallecula at various stages of development and our experience with their variable appearance in multiple planes and to discuss the probable relationship between the cisterna magna septa, Dandy-Walker continuum, mega cisterna magna, and persistent Blake's pouch. METHODS: Retrospective and prospective selection of examples of cisterna magna septa was performed over approximately a 12-month period. Standard and nonstandard imaging planes were adopted as necessary. RESULTS: The septa are typically seen inferoposterior to the cerebellar vermis, usually straight and parallel, arising at the cerebellovermian angle and coursing posteriorly to the occipital bone. The cisterna magna septa become contiguous with the roof of the fourth ventricle inferior to the cerebellar vermis. The cerebrospinal fluid space enclosed between the cisterna magna septa is in direct contiguity with the fourth ventricle via the vallecula and is always completely anechoic because it develops intra- and not extra-axially. CONCLUSIONS: We propose that the cisterna magna septa represent the walls of Blake's pouch, a phylogenetic vestigial structure observed during ontogeny. Additionally, our observations support current opinion that a persistent Blake's pouch and mega cisterna magna represent (less severe) abnormalities within the Dandy-Walker continuum. The cisterna magna septa therefore are a marker of normal development of the roof of the rhombencephalon. Deviation from their normal appearances should prompt a closer assessment for associated abnormalities of the cerebellum, vermis, and brain stem by additional imaging in orthogonal planes with either sonography or magnetic resonance imaging.

Cisterna Magna↗

[Heliobacterium sulfidophilum sp. Nov. and Heliobacterium undosum sp. Nov.: sulfide-oxidizing Heliobacteria from thermal sulfidic springs].

Two new species of heliobacteria isolated from cyanobacterial mats of two alkaline sulfidic hot springs are formally described. Strains BR4 and BG29 are assigned to anoxygenic phototrophic bacteria of the family Heliobacteriaceae, since they possess the unique properties of this taxon: strict anaerobiosis, formation of bacteriochlorophyll g, the lack of extensive intracytoplasmic membranes and chlorosomes, an unusual cell wall structure, and phylogenetic relatedness to the low G + C gram-positive eubacteria. Based on the 16S rDNA sequence similarity, strains BR4 and BG29 are assigned to the genus Heliobacterium and described as two new species of this genus: Heliobacterium sulfidophilum sp. nov. and Heliobacterium undosum sp. nov. The G + C content of the DNA is 51.3 mol % in Hbt. sulfidophilum and 57.2-57.7 mol % in Hbt. undosum. The cells of Hbt. sulfidophilum are rods, and the cells of Hbt. undosum are slightly twisted spirilla or short rods. Both new bacteria are motile by peritrichous flagella. Hbt. sulfidophilum produces endospores. The new bacteria are strict anaerobes growing photoheterotrophically on a limited range of organic compounds. In the dark, they can switch from photosynthesis to the slow fermentation of pyruvate. Biotin is required as a growth factor. Both species are highly tolerant to sulfide (up to 2 mM at pH 7.5) and oxidize it photoheterotrophically to elemental sulfur; photoautotrophic growth was not observed. The temperature optimal for growth of Hbt. sulfidophilum and Hbt. undosum is 30-35 degrees C, and the optimal pH is 7-8.

Bacteria↗

[Quantitative analysis of genetic parameters in populations of European (Capreolus capreolus L.) and Siberian (Capreolus pygargus Pall.) roe deer with RAPD markers].

A technique for obtaining unbiased estimates of genetic parameters (allelic frequencies of RAPD loci, heterozygosity (H), Wright's F statistic, and Nei's genetic distances) in populations of the European (Capreolus capreolus L.) and Siberian (Capreolus pygargus Pall.) roe deer is presented. The technique employs jackknifing and multiple comparative analysis based on a modified Holmes's procedure for Bonferroni's test. It was demonstrated that samples from local groups of roe deer in the Trans-Ural region did not differ significantly in allelic frequencies (0.8, 0.81, and 0.78; P > 0.447) or Nei's genetic distances (0.0056, 0.0273, and 0.0218; P = 0.26), but they could be differentiated based on Wright's F statistic (0.0346, 0.0519, and 0.0450; P = 10(-9)). The parameters of intrapopulation heterozygosity (from 0.18 to 0.042) formed a gradient from the east to the west. Calibration estimates of molecular evolution rate in the family Cervidae obtained based on published data and Jukes-Cantor genetic distances estimated in this study demonstrated that the Siberian roe deer has split into two subspecies, C. pygargus pygargus Pall. and C. pygargus tianschanicus Satunin in the interval between 229 and 462.3 thousand years ago. The species formation of the Siberian and European roe deer was dated between 1.375 and 2.75 Myr ago. Based on the results obtained we recommend the approaches used in the study for analysis of population genetic structure and phylogenetic relationships between populations, subspecies, species, and higher taxa.

Animals↗

Neurophysiological mechanisms of induction of meditation: a hypothetico-deductive approach.

A detailed analysis of methods of induction of meditation and meditative experience encountered therein implicates involvement of several mechanisms in inducing 'meditative effect'. 'Efferent attenuation', 'sensory attenuation' and 'cognitive restructuring' appear three possible mechanisms employed in varying degree of combinations to produce the 'meditative effect' during different types of meditations. Using hypothetico-deductive approach, it is possible to generate a neural model for explaining the 'meditative effect'. Primarily, the meditation is produced by disengaged association cortices driven by thalamus or other older group of reticular nuclei. Secondarily, there may be involvement of some more phylogenetically older structures depending upon depth and types of meditation. This model explains induction, maintenance and long-term effects of meditation.

Attention↗

[Separation and characterization of mitochondrial COI pseudogenes in Scylla paramamosain].

Mitochondrial DNA marker has been applied in genetic structure and phylogenetic analysis, but the differences between nuclear mitochondrial pseudogenes (Numts) and mitochondrial DNA complicate phylogenetic analysis. Using the universal and special primers, 34 pseudogene sequences (Numts) and 5 real mtDNA COI sequences were amplified respectively from 34 individuals of Scylla paramamosain from the coast of southeast China. Twenty-nine haplotypes were identified and two groups (Group I and Group II) were determined according to the homology of the 34 pseudogene sequences. Two insertions and an 8-bp deletion were found in some sequences of Group I, but no indel (insertion and deletion) was found in Group II. Compared to mtDNA COI sequences, the two groups had their conservative sites respectively. At least two independent nuclear transfer events are required to explain the codon position biases revealed through the pairwise comparisons of the 34 Numts, one for Group I and one for Group II. The results showed the necessity of great care in interpreting PCR-generated mtDNA sequences using universal primers when used in population or evolutionary studies in genus Scylla.

Animals↗

Morphological evolution of subterranean mammals: integrating structural, functional, and ecological perspectives.

The widely recognized convergence of subterranean mammals offers unique opportunities for the study of patterns, causes and consequences of morphological integration. For example, behavioral and biomechanical observations have revealed a diversity of digging modes among subterranean mammals. In rodents, scratch-digging and chisel-tooth digging, alone or combined, are widespread. This dual nature of the digging apparatus must be understood to avoid erroneous assessments of specializations based on forelimb modifications only. Additionally, bulldozing and disposal of loose soil is performed by head-lifting, by means of the forelimbs, or with the hindlimbs. Insectivores are known to resort to scratch-digging, sand-swimming, or to a unique system known as humeral rotation. Semi-fossorial species among rodents and insectivores are scratch-diggers. In contrast, fully subterranean taxa show further specializations for scratch-digging and/or resort to some of the other digging modes. The ecological correlates of these morphological trends are still poorly known; work in geomyids suggests that scratch-digging specializations are efficient in friable soils, whereas tooth-digging allows the utilization of a much broader spectrum of soil types. This indicates that alternative pathways of morphological specialization are not equivalent in their ecological potentialities. Important morphological features, such as incisor procumbency among tooth-diggers, are shown to be constrained by structural, allometric, and mechanical factors. In geomyids, the same procumbent morphologies may be acquired as a byproduct of size increases, by means of adaptive shifts independent from size, or by a combination of both. Thus, geometric similarity may not be indicative of functional similarity. Further constraints result from the integration of different functions. For instance, it is suggested that mastication imposes limitations upon potential modifications of jaw morphology for tooth-digging. Scratch-digging appears to be less constrained by locomotion, but the effects of integration of various functions in fore- and hind-limbs are largely unexplored. Multiple approaches, combining behavioral, functional, ecological, structural, and phylogenetic data, are necessary for the study of morphological evolution among subterranean mammals.

Animals↗

Fine structure and cytochemistry of the mitotic plaques of Trypanosoma cruzi and Crithidia fasciculata.

The mitotic plaques are double, electron-dense structures which are located at the equator of the nucleus during the equatorial (metaphase) stage of mitosis in Trypanosoma cruzi, Crithidia fasciculata and other trypanosomatids. Each part of the equatorial plaques separates from the other and becomes an hemiplaque at the beginning of nuclear elongation. Variations of size of the plaques in different species of Trypanosomatidae are restricted to a limited range (less than 30% of the average thickness). At least two different components are found in the plaques with cytochemical methods: a) a basic protein with a high affinity for ethanolic-phosphotungstic acid, which is located in a narrow band towards the cleavage plane in each hemiplaque; and b) an osmiophilic component (possibly a protein) with a low affinity for uranyl acetate and which is located throughout the body of the plaque. The affinity for uranyl acetate can be abolished by methylation and acetylation, but remains after extraction with cold perchloric acid. No cytochemical evidence for the presence of DNA in the plaques is found. However, electron microscopy and cytochemical observations show that the PTA-affine band of the plaques is associated at its sides with chromatin fibers. Thus plaques, as the outer layer of kinetochores in higher eukaryots, have a component with high affinity for phosphotungstic acid, strengthening the hypothesis that these structures are phylogenetically related.

Acetylation↗