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PubMed1990

Minimal sequence and factor requirements for the initiation of transcription from an atypical, TATATAA box-containing housekeeping promoter.

We have established the minimal sequence and factor requirements for both constitutive and viral-induced transcription from an atypical, TATATAA box-containing human housekeeping promoter. Utilizing a transient cotransfection protocol, we have found that efficient transactivation of triosephosphate isomerase (TPI) gene transcription by the immediate early proteins of adenovirus and pseudorabies virus is dependent upon the same assembly of sequence elements that collectively confer minimal TPI promoter function in the absence of viral protein. These elements span TPI promoter positions -65 and -6 (where +1 is the transcription initiation site) and include not only a TFIID-responsive TATATAA box (-27 to -21) but a single GC box (-53 to -48) that binds Spl, and a novel cap proximal element (-18 to -6) that binds a 110-kDa nuclear factor that is present in HeLa cells. We demonstrate that these elements function in an interdependent fashion; deleting either GC box 1 or the cap proximal element completely or nearly abolished both basal transcription and viral transactivation. Therefore, these elements and their cognate factors represent the basal transcription initiation complex through which the immediate early protein of adenovirus or pseudorabies virus mediates the stimulation of TPI gene transcription. We discuss the implications of these data for both constitutive and viral-induced transcription.

Base Sequence↗

PubMed1987

Intracranial injuries resulting from boxing: a review (1918-1985).

A review of the available records indicates that there have been a substantial number of fatalities in boxers due to intracranial injuries sustained in the ring in comparison to the numbers engaged at both amateur and professional levels. The number of such fatalities has decreased steadily in recent years owing to different measures taken by boxing authorities to decrease the physical hazards in the ring and to improve monitoring of boxers during bouts by referees and physicians. The considerable concern about the long-term effects of repeated brain injury as the result of boxing in producing chronic encephalopathy is adequately justified by the many studies of live boxers and pathologic examinations of brains of former boxers made and recorded over the years since 1952. These indicate clearly a significant relationship between the numbers of bouts fought and the presence of severity of chronic encephalopathy. Because it is unlikely that currently adopted protective measures will substantially reduce these hazards, even though they have apparently reduced direct fatalities, a reasonable approach to prevention would be to reduce the number and severity of blows to the head. This might be done by making any blow to the head in boxing a foul and a reason for disqualification. Another approach would be the elimination of boxing.

Athletic Injuries↗

PubMed1996

[Acute subdural hematoma caused by professional boxing].

Knockout in boxing entails deliberate production of the state of unconsciousness. Acute subdural hematoma which is the most common acute brain injury in boxing, accounts for 75% of all acute brain injuries and is the leading cause of boxing fatalities. The aim of this study is to evaluate acute subdural hematoma caused by professional boxing by analyzing the content of bouts, the level of consciousness on admission, CT scan, therapy and outcome 3 months after admission. Fifteen boxers who had suffered from acute subdural hematoma were classified into three groups according to the pattern of loss of consciousness. Transient unconsciousness type (Transient type): boxers who had returned to alertness within an hour from the time of injury. Lucid interval type: neurological deterioration appeared with a lucid interval from ten minutes to an hour after knockout. Deterioration of consciousness type (Deterioration type): A state of unconsciousness appeared and worsened from a few minutes after knockout. Analyzing the number of rounds in bouts indicated that the hematoma occurred most frequently in bouts of 10 rounds. All of our subjects presented subdural hematomas without cerebral contusions on CT scan. With regard to the location of the hematomas, 9 hematomas involved the left side, 3 the right, 2 the suboccipit and 1 the interhemisphere. Transient type was found in 7 patients who had GCS scores of 14, 15 on admission. Since CT scan revealed thin subdural hematoma with or without mild midline shift, conservative therapy was carried out in all patients. All patients had a good recovery. Five patients of lucid interval type with an admission GCS score of 4, 6 and 7 demonstrated thicker hematoma compared to that presented by the transient type with significant midline shift on CT scan. All patients required surgery. Outcome of this type was good recovery (n = 2), moderate disability (n = 1), persistent vegetative state (n = 1), death (n = 1). Three patients of deterioration type had GCS scores of 5, 6. Because of subdural hematoma with remarkable midline shift on CT scan, all patients underwent surgery. Outcome was good recovery (n = 1), moderate disability (n = 1), persistent vegetative state (n = 1). Overall outcome was good recovery 66.7%, moderate disability 13.3%, persistent vegetative state 13.3%, death 6.7%. Furthermore, 8 patients who underwent surgery with a GCS score of less than 8 exhibited good recovery 37.5%, moderate disability 25%, persistent vegetative state 25%, death 12.5%. CT scan of lucid interval and deterioration type showed a tendency to show thick subdural hematoma and remarkable midline shift compared to transient type. Outcomes of lucid interval and deterioration type were worse than those of transient type. This result suggests that the influence of repeated head injury and diffuse brain injury might make a difference between these groups. Repeated head injury means that further impacts repeatedly damaged the injured brain after bleeding in the bouts. Overall outcome was better than that published in previous reports and also than that observed in other head injuries, for example, traffic accident and fall. The reasons for this could be that the patients were younger, that there was immediate surgical treatment, and that brain injury without cerebral contusion had contributed to better outcome. Finally, the best medical management intervention seems to be to diagnose and treat the lesions as early as possible after occurrence of subdural hematoma.

Acute Disease↗

PubMed1997

Apolipoprotein E epsilon4 associated with chronic traumatic brain injury in boxing.

CONTEXT: Given the similarities between Alzheimer disease and dementia pugilistica, we evaluated the relationship between apolipoprotein E (APOE) genotype and chronic traumatic brain injury (CTBI) in boxers to determine whether there is a genetic susceptibility to the effects of head trauma. OBJECTIVE: To assess the relationship between CTBI and APOE genotype in boxers. DESIGN AND SETTING: Clinical characterization of 24 volunteer and 6 referred boxers in an outpatient setting. PARTICIPANTS: Thirty professional boxers aged 23 to 76 years underwent neurologic and behavioral assessment in conjunction with APOE genotyping. MAIN OUTCOME MEASURES: Apolipoprotein E genotype was examined in relationship to measures of CTBI. A 10-point clinical rating scale (0-9), the Chronic Brain Injury (CBI) scale, was devised to assess the severity of traumatic encephalopathy associated with boxing. Boxers with abnormal CTBI scores were further classified on the basis of whether their impairments were possibly or probably related to boxing. Scores were analyzed in relation to boxing exposure (number of bouts) and APOE genotype. RESULTS: Among the 30 boxers, 11 were found to be normal (CBI score=0), 12 showed mild deficits (CBI score=1-2), 4 were moderately impaired (CBI score=3-4), and 3 showed signs of severe impairment (CBI score > 4). High-exposure boxers (ie, those with > or = 12 professional bouts) had significantly higher CBI scores (mean [SD], 2.6 [1.9]) than low-exposure boxers (mean [SD], 0.3 [0.7]) (P<.001), indicating that neurologic impairment as measured by the CBI scale seems related to boxing exposure. The APOE genotype frequencies of the study population were approximately the same as those found in the general population. Boxers with low exposure had mean CBI scores of 0.33, irrespective of APOE genotype. However, high-exposure boxers with an APOE epsilon4 allele had significantly greater CBI scores (mean [SD], 3.9 [2.3]) than high-exposure boxers without APOE epsilon4 (mean [SD], 1.8 [1.2]) (P=.04). All boxers with severe impairment possessed at least 1 APOE epsilon4 allele. The tendency for greater CTBI among those with both high exposure and an epsilon4 allele was statistically significant at the P<.001 level. CONCLUSIONS: These preliminary findings suggest that possession of an APOE epsilon4 allele may be associated with increased severity of chronic neurologic deficits in high-exposure boxers.

Adult↗

PubMed1997

[Present views on boxing--Henner's legacy].

Fifty years ago academician Henner formulated clearly the negative view of neurologists on boxing as practised in his time. It was his merit that a protective helmet was introduced, many of his demands concerning the regime after a KO and the importance of medical examinations were no yet adequately appreciated. At the opportunity of the 100th anniversary of Henner's birth and the fiftieth anniversary of the Clinic he founded in the premises of saint Catherine the author discusses contemporary views on boxing, the risk of repeated injuries for the development of encephalopathia pugilistica and their relationship to Alzheimer's disease. Contemporary boxing is a spectacle which is a continuation of historical fights of gladiators. If the main objective is to hurt the adversary, this activity does not deserve the name of "sport" Sport should promote and maintain physical and mental health which is not the case in professional boxing.

Boxing↗

PubMed1998

TATA-box mutant in the promoter of the uridine diphosphate glucuronosyltransferase gene in Italian patients with Gilbert's syndrome.

BACKGROUND: A mutation in the promoter of the uridine diphosphate glucuronosyltransferase gene has been described in patients with Gilbert's syndrome from Northern Europe. AIMS: To assess the frequency of this mutation in Italian patients with Gilbert's syndrome and in normal controls, in order to establish the molecular basis and molecular epidemiology of the syndrome in Italy. PATIENTS: Forty-six patients with a clinical diagnosis of Gilbert's syndrome and 44 individuals from the general population unselected for bilirubin levels. METHODS: Polymerase chain reaction amplification of the TATA-box element in the promoter of uridine diphosphate glucuronosyltransferase and identification of wild-type and variant alleles by high-resolution polyacrylamide gel electrophoresis. RESULTS: A TATA-box variant in the promoter of uridine diphosphate glucuronosyltransferase was found on 93% of chromosomes from patients with Gilbert's syndrome. The same variant was present on 44% of chromosomes from controls, unselected for bilirubin levels. Only 55% of controls homozygous for the TATA-box variant, however, had increased bilirubin levels. CONCLUSIONS: The TATA-box variant in the promoter of uridine diphosphate glucuronosyltransferase is strongly associated with the phenotype of Gilbert's syndrome in Italy. The incomplete penetrance of the mutation observed in controls indicates that other acquired or inherited conditions affecting bilirubin production, uptake, cellular transport or excretion may contribute to the hyperbilirubinaemia of Gilbert's syndrome.

Adolescent↗

PubMed1999

OsMADS13, a novel rice MADS-box gene expressed during ovule development.

MADS-box genes have been shown to play a major role in defining plant architecture. Recently, several MADS-box genes have been reported that are highly expressed in the ovule. However, only for the Petunia genes FBP7 and FBP11 has a function in defining ovule identity been shown. We have isolated a rice MADS-box gene named OsMADS13. Expression analysis has shown that this gene is highly expressed in developing ovules. In order to facilitate a detailed characterization of rice ovule-expressed genes, a comprehensive morphological description of ovule development in rice has been performed. The predicted amino acid sequence of OsMADS13 shows significant homology with ZAG2, a maize MADS-box gene, which is also expressed mainly in the ovule. Mapping of the gene in the rice genome showed that it is located on chromosome 12, which is syntenic to two maize regions where ZAG2 and its paralogous gene ZMM1 have been mapped. Our results suggest that OsMADS13 is the ortholog of ZAG2 and ZMM1 and might play a role in rice ovule and seed development.

Amino Acid Sequence↗

PubMed2000

Expression screening of factors binding to the osteocalcin bone-specific promoter element OC box I: isolation of a novel osteoblast differentiation-specific factor.

Contributing to bone-specific expression of the osteocalcin gene is the promoter element OC Box I (-99 to -76), which binds both Hox proteins and another nonhomeodomain factor (designated OCBP for osteocalcin-box binding protein) (Hoffmann et al. [1996] J Cell Biochem 61:310-324). OCBP correlates with increased promoter activity and may, therefore, be important to development or maintenance of the osteoblast phenotype. To identify OCBP candidates, we used a multimerized OC Box I sequence to screen a gammagt11 cDNA expression library, constructed from the rat osteosarcoma osteoblastic ROS 17/2.8 cell line, for cDNA clones encoding factors that recognize this element. Mutant OC Box I sequences that do not bind OCBP and/or homeodomain proteins were used to counterscreen the cDNA isolates. Clones showing binding specificity were sequenced and further characterized for patterns of expression in different tissues and cell lines. Among the characterized nonhomeodomain-related isolates, we identified a nucleolin, a clone encoding rCAP2 that is related to myogenic differentiation and more importantly, a cDNA clone containing a previously uncharacterized gene that has been designated as a cell differentiation-related factor (DRF). DRF mRNA is highly expressed in ROS 17/2.8 cells and in a developmentally regulated pattern during osteoblast differentiation, being upregulated at the postproliferative maturation transition and coinciding with the induction of osteocalcin gene expression. The 7.7-kb transcript encoded by clone 44 is abundantly expressed in osteoblasts, but the mRNA was not present at detectable levels in bone and soft tissues by Northern blot analysis. However, related expressed sequence tags were recently reported in cDNA libraries of rat lung and mouse sympathetic ganglion. The identification of DRF represents a novel osteoblast differentiation-specific marker related to osteocalcin expression. The identification of DRF may further facilitate definition of gene regulatory mechanisms mediating the final stages of bone cell differentiation

Animals↗

PubMed1986

Isolation of a mouse cDNA coding for a developmentally regulated, testis-specific transcript containing homeo box homology.

A clone, pHBT-1, containing sequences homologous to Drosophila homeo boxes has been isolated from a mouse testis cDNA library. The sequence is 80% homologous at the DNA level and 88% homologous at the amino acid level to the homeo box sequence of the Antennapedia gene of Drosophila. Sequences flanking the 3' end of the homeo box are highly diverged from other murine homeo box-containing genes characterized to date. RNA blot hybridization analysis of mouse testis poly(A)+ RNA revealed transcripts of approximately 1.4 kb in length. Within the limits of sensitivity of detection of Northern blot analysis, no transcripts were seen in any of the adult somatic tissues examined. Other tissues that contain stem cells, namely those of the hemopoietic system, also lacked detectable amounts of HBT-1 transcripts. HBT-1 transcripts were limited to male germ cell-containing tissues, since RNAs from juvenile and adult ovaries did not contain detectable amounts of the 1.4-kb transcripts. Expression of the HBT-1 gene was not detected in embryonic testes, nor in tests of neonatal animals which contain germ cells up to the Type B stage of spermatogonial development. A role for the expression of the HBT-1 gene in the meiotic stages of male germ cell differentiation is postulated.

Animals↗

PubMed1987

Hox-1.6: a mouse homeo-box-containing gene member of the Hox-1 complex.

Hox-1.6, a mouse homeo-box-containing gene member of the Hox-1 complex, is described. The Hox-1.6 homeo-box shows more divergence than the other members of the complex with the Drosophila Antennapedia-like homeo-box class. This previously undescribed gene was studied with respect to its transcription pattern and was found to be expressed during mouse fetal development in an intestine-specific manner in adults, and in tumours or cell types exhibiting early endodermal-like differentiation. The study of embryonic partial Hox-1.6 cDNA clones revealed structural features common to other Drosophila and vertebrate homeo-box-containing genes, but also indicated that Hox-1.6 transcripts might display splicing patterns more complex than those known for other vertebrate homeo-genes. One of these cDNA clones contains a rather short open reading frame which would encode a protein of approximately 14.5 kd. The use of this clone as a probe for S1 nuclease mapping confirmed that different Hox-1.6 transcripts were present both in embryonic total RNA and in embryonal carcinoma cell cytoplasmic RNA. These various transcripts are probably generated by an alternative splicing mechanism and may thus encode a set of related proteins.

Amino Acid Sequence↗

PubMed1993

Structure of the HMG box motif in the B-domain of HMG1.

The conserved, abundant chromosomal protein HMG1 consists of two highly homologous, folded, basic DNA-binding domains, each of approximately 80 amino acid residues, and an acidic C-terminal tail. Each folded domain represents an 'HMG box', a sequence motif recently recognized in certain sequence-specific DNA-binding proteins and which also occurs in abundant HMG1-like proteins that bind to DNA without sequence specificity. The HMG box is defined by a set of highly conserved residues (most distinctively aromatic and basic) and appears to define a novel DNA-binding structural motif. We have expressed the HMG box region of the B-domain of rat HMG1 (residues 88-164 of the intact protein) in Escherichia coli and we describe here the determination of its structure by 2D 1H-NMR spectroscopy. There are three alpha-helices (residues 13-29, 34-48 and 50-74), which together account for approximately 75% of the total residues and contain many of the conserved basic and aromatic residues. Strikingly, the molecule is L-shaped, the angle of approximately 80 degrees between the two arms being defined by a cluster of conserved, predominantly aromatic, residues. The distinctive shape of the HMG box motif, which is distinct from hitherto characterized DNA-binding motifs, may be significant in relation to its recognition of four-way DNA junctions.

Amino Acid Sequence↗

PubMed1993

Sox-4, an Sry-like HMG box protein, is a transcriptional activator in lymphocytes.

Previous studies in lymphocytes have described two DNA-binding HMG box proteins, TCF-1 and LEF-1, with affinity for the A/TA/TCAAAG motif found in several T cell-specific enhancers. Evaluation of cotransfection experiments in non-T cells and the observed inactivity of an AACAAAG concatamer in the TCF-1/LEF-1-expressing T cell line BW5147, led us to conclude that these two proteins did not mediate the observed enhancer effect. We therefore searched for additional HMG box proteins. By a PCR-aided strategy, we cloned Sox-4, a gene with homology to the HMG box region of the sex determining gene SRY. Sox-4 was expressed in T and pre-B lymphocyte lines and in the murine thymus. Significantly, BW5147 T cells did not express Sox-4. Recombinant Sox-4 bound with high affinity (Kd 3 x 10(-11) M) to the minor groove of the AACAAAG motif, most likely contacting all seven base pairs. In contrast with observations on TCF-1 and LEF-1, cotransfection with Sox-4 unveiled a transactivating capacity, which mapped to its serine-rich C terminus. This region remained functional upon grafting onto a GAL4 DNA-binding domain. Sox-4 is thus the first 'classical' transcription factor in the Sox gene family with separable DNA-binding and transactivation domains. Our observations indicate that a detailed understanding of T cell-specific gene control must integrate the concerted activity of at least three tissue-specific HMG box genes.

Amino Acid Sequence↗

PubMed1995

CCAAT-box contributions to human thymidine kinase mRNA expression.

In order to examine the role of two inverted CCAAT boxes near the start of transcription of the human thymidine kinase (TK) gene, a series of constructs were prepared in which one or both CCAAT boxes were deleted or mutated. These altered promoters (1.2 kb of 5'-flanking sequence) were used to express a TK minigene containing the first two exons and introns followed by the remainder of the cDNA. RNA blots were prepared from stable cell lines of ts13 cells containing these constructs under three conditions: 1) serum deprived cells, 2) serum stimulated cells, and 3) cells that had been stimulated with serum, but were arrested in the G1 phase of the cell cycle by the temperature sensitive mutation carried by these cells. TK mRNA expression from each construct was suppressed by the temperature sensitive block to cell cycle progression. Measurement of protein expression from the various altered TK promoters indicated that both CCAAT boxes contribute to promoter strength. These experiments also suggested that the two CCAAT boxes were not equivalent and that the distal CCAAT could substitute for the proximal CCAAT, but the converse was not true.

Animals↗

PubMed1995

Farnesylation and proteolysis are sequential, but distinct steps in the CaaX box modification pathway.

Membrane localization of Ras proteins requires posttranslational modification of a conserved C-terminal sequence motif known as the CaaX box (C is Cys, a is any aliphatic amino acid, and X is the carboxyl terminal residue). The modification steps include farnesylation, removal of the three C-terminal amino acids, carboxyl-methylation, and palmitoylation. In yeast, the farnesyltransferase (FTase) is encoded by the RAM1(DPR1) and RAM2 genes, and the methyltransferase is the product of STE14. The gene encoding the protease(s) that is responsible for modification of the CaaX has not been identified. We have used in vitro-synthesized Ras2p and synthetic peptide substrates to investigate the relationship between farnesylation and proteolysis. Addition of yeast cytosolic extracts to rabbit reticulocyte extracts programmed to synthesize Ras2p led to prenylation of Ras2p and a change in electrophoretic mobility similar to that observed during Ras maturation in vivo. However, it was not possible to determine if the mobility shift is the result of prenylation, proteolysis or a combination of both steps. Therefore, we examined the relationship between farnesylation and proteolysis directly using extracts prepared from bacteria overexpressing the genes for the yeast FTase (RAM1 and RAM2) and synthetic CaaX box peptides. Extracts from bacteria expressing RAM1/RAM2 efficiently prenylate CaaX box peptides, but do not proteolyze the -aaX residues. However, addition of yeast extracts from wild type, ram1, or ste14 mutants resulted in the removal of the -aaX residues from prenylated CaaX box peptides.

Alkyl and Aryl Transferases↗

PubMed1996

The serum response factor (SRF) is needed for muscle-specific activation of CArG boxes.

CArG boxes, whose consensus sequence is CC(A/T)6GG, are involved in two very different types of transcriptional responses: response of immediate early genes to serum, mediated by so-called Serum Response Elements (SRE), and transcriptional activation of muscle-specific genes during muscle differentiation. Although previous studies have shown that the Serum Response Factor (SRF) binds to muscular CArG boxes, the role of such a binding in muscle-specific activation of CArG box-dependent genes was not directly demonstrated. Here, by transient co-transfection experiments, we demonstrate that intact SRF is required for muscle-specific transcriptional activation through CArG boxes.

Actins↗

PubMed1999

The E-box of the human glycophorin B promoter is involved in the erythroid-specific expression of the GPB gene.

Previous studies performed on the glycophorin B (GPB) expression demonstrated that this gene is expressed in erythroid cells only and that the ubiquitous factor Ku70 is involved in the process. Here, we investigated the contribution of the -70 E-box sequence toward the GPB promoter expression. We found that the E-box bound two factors, the USF1/USF2 protein and an unidentified ubiquitous protein which was named factor U. Site-directed mutagenesis performed on the -70 E-box showed that the USF factor had an activating effect in CAT assays. Conversely, mutation of the -70 E-box that impaired the binding of factor U led to a positive CAT activity in nonerythroid cells and thus to the loss of the erythroid-specific expression of the GPB gene. This indicates that, in addition to the Ku70 factor, the extinction of the GPB promoter expression in nonerythroid cells depends also on the repressing effect of the factor U.

Antigens, Nuclear↗

PubMed1999

An E box comprises a positional sensor for regional differences in skeletal muscle gene expression and methylation.

To dissect the molecular mechanisms conferring positional information in skeletal muscles, we characterized the control elements responsible for the positionally restricted expression patterns of a muscle-specific transgene reporter, driven by regulatory sequences from the MLC1/3 locus. These sequences have previously been shown to generate graded transgene expression in the segmented axial muscles and their myotomal precursors, fortuitously marking their positional address. An evolutionarily conserved E box in the MLC enhancer core, not recognized by MyoD, is a target for a nuclear protein complex, present in a variety of tissues, which includes Hox proteins and Zbu1, a DNA-binding member of the SW12/SNF2 gene family. Mutation of this E box in the MLC enhancer has only a modest positive effect on linked CAT gene expression in transfected muscle cells, but when introduced into transgenic mice the same mutation elevates CAT transgene expression in skeletal muscles, specifically releasing the rostral restriction on MLC-CAT transgene expression in the segmented axial musculature. Increased transgene activity resulting from the E box mutation in the MLC enhancer correlates with reduced DNA methylation of the distal transgenic MLC1 promoter as well as in the enhancer itself. These results identify an E box and the proteins that bind to it as a positional sensor responsible for regional differences in axial skeletal muscle gene expression and accessibility.

Animals↗

PubMed1993

Activation of gene expression via CArG boxes during myogenic differentiation.

We have studied gene activation via CArG boxes in the context of myogenesis. The proximal CArG box of the human cardiac actin gene (HCA1) stimulates transcription from the herpes simplex virus thymidine kinase (TK) promoter in a tissue-specific fashion. Thus in transient transfection assays, when the expression of chloramphenicol acetyltransferase (CAT) from p(HCA1)4 TKCAT is compared to that derived from p(M1)4 TKCAT which contains an inactive mutated version (M1) of the HCA1 element, high levels of expression are seen in C2 mouse myoblasts and myotubes, and in the T4 myoblast cell line derived from the C3H10T1/2 cell line by 5-azacytidine treatment, whereas only low levels of expression are seen in the mouse L fibroblast cell line. The parental C3H10T1/2 cell line shows intermediate levels of expression. A similar situation is seen in stably transfected cell lines. Gene activation via CArG boxes was also analyzed in the course of myogenic conversion of C3H10T1/2 cells treated with 5-azacytidine. Our results indicate that activation of the CAT gene from the HCA1 element is slightly posterior to the appearance of the first MyoD1 and myogenin transcripts, concomitant with the appearance of cardiac alpha-actin transcripts, but clearly precedes the accumulation of myosin light-chain 1a transcripts and the appearance of troponin T-positive cells. These results further establish that CArG boxes can be seen as muscle-specific cis-acting regulatory element prior to terminal differentiation.

Actins↗