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PubMed2003

Transforming growth factor-beta-inducible gene-h3 (beta(ig)-h3) promotes cell adhesion of human astrocytoma cells in vitro: implication of alpha6beta4 integrin.

Beta(ig)-h3 is a secretory protein that is induced by transforming growth factor (TGF)-beta. We have recently found that beta(ig)-h3 expression is induced in cultured astrocytes by TGF-beta1 and in rat cerebral cortex by stab wound. The purpose of this study was to examine the effect of the secreted beta(ig)-h3 on cell adhesion of astrocytes and the underlying mechanisms. When U87 human astrocytoma cells were seeded on dishes coated with recombinant beta(ig)-h3, cell adhesion was significantly enhanced. Blocking experiments using various antibodies to the integrin subunit suggested alpha6beta4 integrin could be involved in the beta(ig)-h3-mediated astrocyte cell adhesion. Cell adhesion to beta(ig)-h3 substrate was substantially blocked by preincubation with the inhibitor to the src kinase. When cells were plated on beta(ig)-h3-coated dishes, tyrosine phosphorylation of focal adhesion kinase was prominently increased within 20 min in a beta4 integrin-dependent manner. The results suggest that alpha6beta4 integrin-mediated interactions of astrocytes with beta(ig)-h3 transduce intracellular signals through the focal adhesion proteins, which may regulate certain aspects of astrocyte response to brain injury.

Astrocytoma↗
PubMed2000

Integrin expression during epithelial migration and restratification in the tenascin-C-deficient mouse cornea.

In the unwounded cornea, tenascin-C localizes to a short stretch of the basement membrane zone at the corneoscleral junction or limbus. To determine whether the function of the limbus is affected by the absence of tenascin-C, mice possessing a deletion of tenascin-C and strain-matched wild-type mice are used in corneal debridement wounding experiments. The expression of integrins (alpha3, alpha9, and beta4) in the tenascin-C knockout corneas is evaluated by producing polyclonal cytoplasmic domain antipeptide sera and performing immunofluorescence microscopy. In addition, we evaluate the localization of several other proteins involved in wound healing, including fibronectin, laminin beta1, nidogen/entactin, and VCAM-1, in both the tenascin knockout and wild-type mice. There are no differences in healing rate, scarring, or neovascularization after corneal debridement wounds. alpha9 integrin is expressed at the limbal border of unwounded tenascin-C knockout animals and is upregulated during migration only after the larger wounds. At 8 weeks after larger wounds, the localization of alpha9 again becomes restricted to the limbal border. Results show that tenascin-C is not required for development or maintenance of the corneal limbus or for normal re-epithelialization of corneal epithelial cells after debridement wounding.

Animals↗
PubMed2000

Effects of sulfur mustard on the basal cell adhesion complex.

Among the most intriguing questions about sulfur mustard (di(2-chloroethyl) sulfide) is why basal cells are the primary targets of its vesicating lesions. To investigate this problem, replicate cultures of human epidermal keratinocytes (HEK) were grown from normal skin and exposed to 400 microM sulfur mustard (HD) for 5 min. Using fluorescein isothiocyanate (FITC)-conjugated antibodies, confocal laser microscopy and image analyses, we found that in early passages, sham-treated HEK maintained in a 0.15 mM Ca2+ medium continued to express keratins K5 and K14 as well as alpha6beta4-integrin. Both K5 and K14 are intermediate filaments characteristic of basal cells and linked with attachment mechanisms effecting epidermolysis bullosa simplex, a family of blistering skin diseases. Acute exposure to HD caused a statistically significant (P < 0.01) 30.74% decrease in K14 fluorescence within 1 h of exposure. Within 2 h of exposure, K14 fluorescence decreased to near-zero values. The loss in expression of K14 was progressive and occurred well before the expected appearance of in vivo blisters, which have a dose-dependent, clinical latent phase of 8-24 h. Acute exposure to HD also caused a statistically significant (P < 0.002) decrease in expression of beta4, an integrin which has been associated with junctional epidermolysis bullosa (JEB). Disruption of K14 and alpha6beta4-integrin may be early events in the HD injury pathway; however, they had no immediate or obvious effect on cell to substrate attachment.

Antibodies↗
PubMed1999

Integrin alpha6beta1 plays a significant role in the attachment of hepatoma cells to laminin.

BACKGROUND/AIMS: Tumor invasion and metastasis consist of a series of complex events. During this process, the ability of tumor cells to adhere to laminin, a major component of basement membranes, is required at various steps. The expression of laminin-binding integrins and the extent of tumor metastasis and progression appear to be related. In hepatocellular carcinoma, increased expression of laminin-binding integrins is observed. However, little is known concerning the possible functional interactions between laminin-binding integrins and laminin. Therefore, we investigated the participation of laminin-binding integrins in the attachment of hepatoma cells to laminin. METHODS: Human hepatoma cell lines (KIM-1, KYN-1, 2) were used. We investigated the expression of integrin alpha1, alpha2, alpha3, alpha6, beta1, and beta4 subunits on hepatoma cells by immunocytochemical and flow cytometric analysis. Participation of these integrin subunits in the attachment of hepatoma cells to laminin was evaluated by an inhibition of cell adhesion assay. RESULTS: Integrin alpha1, alpha2, alpha3, alpha6 and beta1 subunits were expressed at the marginal areas of hepatoma cells, while the integrin beta4 subunit was scarcely detected. Laminin promoted the attachment of hepatoma cells in a dose-dependent manner. Although anti-integrin alpha1, alpha2, beta3 and beta4 subunit antibodies did not inhibit cell attachment to laminin, anti-integrin alpha6 and beta1 subunit antibodies inhibited the attachment by 50% or more. CONCLUSIONS: These findings indicate that integrin alpha6beta1 is very important in the attachment of hepatoma cells to laminin, suggesting the participation of this integrin in metastasis and invasion of hepatoma cells.

Carcinoma, Hepatocellular↗
PubMed1998

Characterization of a tight molecular complex between integrin alpha 6 beta 4 and laminin-5 extracellular matrix.

In many adult epithelia, e.g., epidermis or intestine, adhesion of epithelial cells to basement membrane requires the integrin alpha6 beta4 and laminin-5 (Ln-5). In the absence of one or the other, extensive blistering and exfoliation occur. While alpha6 beta4 was reported to be a receptor for Ln-5, this interaction is poorly understood. We characterize complexes between alpha6 beta4 and Ln-5 in cell-free preparations of extracellular matrix (ECM) from the epithelial cell line, 804G. By microsequencing, Ln-5 and alpha6 beta4 were the major proteins in this ECM and were likely engaged in receptor/ligand complexes because, by immunofluorescence, alpha6 beta4 was colocalized with Ln-5 both in cell monolayers and in cell-free ECM preparations, but they disappeared after preincubation of the monolayers with alpha6 beta4 or Ln-5 function-blocking antibodies. The alpha6 beta4/Ln-5 complexes were resistant to dissociation by extreme pH, urea, chaotropes, eDTA, non-ionic detergents, and b-mercaptoethanol. They were only dissociated by strong anionic detergents, e.g., 1% SDS, suggesting receptor/ligand interactions based on high affinity or avidity. We propose that these alpha6 beta4/Ln-5 complexes may provide links between plasma membrane and basement membrane that resist mechanical stress and support epithelial integrity.

Amino Acid Sequence↗
PubMed2002

Acute hyperglycaemia induces changes in the transcription levels of 4 major genes in human endothelial cells: macroarrays-based expression analysis.

Hyperglycaemia, in insulin-dependent or independent diabetes mellitus, promotes endothelial cell (EC) dysfunction and is a major factor in the development of macro- or microvascular diseases. The mechanisms and the disease-related genes in vascular diseases resulting from hyperglycaemia are poorly understood. Macroarrays. bearing a total of 588 cDNA known genes, were used to analyze HUVEC gene transcription subjected to 25 or 5-mM glucose for 24 h. Isolated mRNA derived from treated first passage HUVEC were reverse transcribed, 32P labeled, and hybridized to the cDNA macroarrays. Results show that acute hyperglycaemia induces an up-regulation of seven major genes, four of which were not previously reported in the literature. Northern blot analyses, performed on these 4 genes, confirm macroarrays results for alphav, beta4, c-myc, and MUC18. Moreover, time course analysis (0, 2, 4, 8, 2, 16, 24 h) of alphav, beta4 c-myc, and MUC18 mRNA expression, observed by northern blot assays, showed a peak at time points situated between 2 to 8 h. The 3 other genes (ICAM-1, beta1, and IL-8), were shown by others to be significantly upregulated after glucose stimulation. Furthermore, ELISA assays performed on the supernatant of HUVEC culture medium showed a significant increase of IL-8 for cells treated with 25-mM compared to 5-mM glucose. Identified genes, upregulated in endothelial cells as a result of acute hyperglycaemia, may serve as therapeutic or diagnostic targets in vascular lesions present in diabetic patients. These results also demonstrate the use of cDNA macroarrays as an effective approach in identifying genes implicated in a diseased cell.

Acute Disease↗
PubMed1996

Expression of the beta 4 integrin subunit in the mouse heart during embryonic development: retinoic acid advances beta 4 expression.

Using immunohistochemical techniques as well as in situ hybridization we were able to elicit the expression pattern of the beta 4 integrin subunit in the murine heart during development. We show that beta 4 is not expressed in the heart before E13 and is afterwards restricted to the endocardium of the atrioventricular canal, the outflow tract, and the venous valves in the right atrium. As these are all sites of high shear stress in the heart, we propose a role for alpha 6 beta 4 in the tight adhesion of the endocardial cells to their basement membranes in these segments. Moreover, mouse embryos were treated with all-trans retinoic acid, which was previously shown to induce congenital malformations, among which malformations of the heart. We show an advanced expression without ectopic localization of cardiac beta 4 after the administration of retinoic acid. This advanced appearance of beta 4 was also shown in extracardiac tissue like migrating neural crest cells. Several hypotheses on the mechanism of beta 4 up-regulation and a possible role for alpha 6 beta 4 in the development of heart malformations after the administration of retinoic acid are discussed.

Animals↗
PubMed1997

Plasma surface modification of artificial corneas for optimal epithelialization.

We have demonstrated that the optimal surface treatment of a polyvinylalcoholcopolymer hydrogel for epithelial cell migration and proliferation is an argon radio frequency (rf) plasma treatment. The surface chemistry of the material was determined prior to each cellular evaluation, allowing us to compare the biological response with a known surface chemistry. The cellular response was carried out in a consistent manner a minimum of three separate runs. We found that the optimal conditions required culturing the cells under constant rotation. Cells became confluent on argon-plasma-treated surfaces coated under several different reactions pressures, and after 2 weeks they became multilayered. Our experiments demonstrated that cells proliferated and extracellular matrix and adhesion proteins were present only when the surface was treated with an argon rf plasma; acetone- and ammonia-treated surfaces did not yield the desired results. Organ culture experiments further demonstrated the efficacy of the argon-treated surfaces. In these experiments, intact keratoprosthetic devices with modified hydrogel surfaces were implanted into rabbit corneas. The excised corneas containing the devices were cultured, and 3 weeks later, using confocal laser scanning microscopy, confluent epithelium was detected on the modified hydrogel surface. This is the first demonstration that rabbit limbal epithelial cells can migrate onto a synthetic cornea containing a modified hydrogel-treated surface and form a confluent surface of epithelium.

Animals↗
PubMed1997

The expression of integrin subunits alpha 6 and beta 4 by corneal epithelial cells on modified hydrogel surfaces.

Our goal was to quantitate the expression and localization of integrin subunits alpha 6 and beta 4 by corneal epithelial cells on defined synthetic substrates. Previously we demonstrated that the cytoplasmic pH and translocation of the alpha 6 integrin subunit to the cell membrane was modified by ionic interactions. These results suggest that changes in the ionic interactions at the cell-substrate interface not only alter the intracellular milieu but ultimately affect the expression of adhesion proteins. To test this hypothesis, hydroxyethylmethacrylate (hema) hydrogels were modified by the addition of amines (N,N-dimethylaminoethylmethacrylate) or carboxyl moieties (methacrylic acid). Changes in the distribution of mRNA and protein were monitored using confocal laser scanning microscopy. The steady state level of integrin mRNA was evaluated, and the results indicate that while the plating efficiency was identical on all surfaces, the expression and localization of integrin subunits was surface dependent. Alpha 6 and beta 4 proteins were localized along the basal surface of nonpermeabilized cells cultured on laminin, on surfaces with amine moieties, and on those with amine and carboxyl moieties. The level of diffuse cytoplasmic staining increased with the presence of carboxyl moieties. Alpha 6 and beta 4 integrin subunits were negligible when the surfaces contained carboxyl moieties alone. The expression of alpha 6 and beta 4 mRNA was higher on surfaces containing amine moieties than on surfaces containing only carboxyl moieties. These results indicate that the characteristics of the substrate and the resulting cell-matrix interaction alter protein and mRNA expression of integrin subunits.

Animals↗
PubMed1996

Beta 4 integrin and other Schwann cell markers in axonal neuropathy.

Schwann cell gene expression is dynamically regulated after peripheral nerve injury and during regeneration. We hypothesized that the changes in protein expression described after rat peripheral nerve injury could be used to identify single Schwann cell-axon units in human axonal neuropathy. Therefore, we performed immuno-fluorescence staining on sections of injured rat sciatic nerves compared with sections of neuropathic human sural nerves. We chose the markers beta 4 integrin, P0 glycoprotein, and glial fibrillary acidic protein (GFAP) to characterize Schwann cells, and neurofilament-heavy (NF-H) to recognize axons. Normal rat or human myelin-forming units demonstrated a sharp ring of beta 4 staining at their outer surface, P0 staining in the myelin sheath, and NF-H staining in the axon. Acutely denervated rat units transited from broken rings of beta 4 and P0 staining, to diffuse beta 4 and absent P0 and NF-H staining. Chronically denervated rat Schwann cells re-expressed beta 4 more highly, but in a diffuse, non-polarized pattern. In contrast, regenerating units re-expressed beta 4, P0, and NF-H; beta 4 staining was polarized to the outer surface of Schwann cells. Finally, GFAP staining increased progressively after injury and decreased during regeneration in the distal nerve stump. In neuropathic human sural nerves, we identified units exhibiting each of these beta 4, P0, and NF-H staining patterns; the proportion of each pattern correlated best with the extent and chronicity of axonal injury. Thus, synchronous injury of rat sciatic nerve predicts patterns of Schwann cell marker expression in human axonal neuropathy. In addition, the unique changes in the polarity of beta 4 integrin expression, in combination with changes in P0 and NF-H expression, may distinguish normal from denervated or reinnervated myelin-forming Schwann cells in human sural nerve biopsies.

Adult↗
PubMed2003

Identification of the antigens predominantly reacted with serum from patients with hepatocellular carcinoma.

BACKGROUND: To identify antigens specifically recognized by the immune surveillance system in patients with hepatocellular carcinoma (HCC), the authors examined two complementary DNA (cDNA) libraries of moderately differentiated HCC by serologic analysis of recombinant cDNA expression libraries (SEREX). METHODS: The libraries were screened with autologous patients' sera, and sequences of the reacted clones were determined. To study the immunoreactivity of the antigens, sera from 20 patients with HCC, from 20 healthy volunteers, and from 16 patients with chronic viral hepatitis were examined. RESULTS: Twenty-seven antigens were identified. They included SART1, p57Kip2, ROCK-1, gamma-catenin, and heat shock proteins, which are classified as tumor-associated genes. Three of 27 antigens-Tat-binding protein-1 (TBP-1), beta4 integrin-binding protein (p27[BBP]), and ribosomal protein L30 (rpL30)-were reacted predominantly with sera from patients with HCC (55% of patients, 45% of patients, and 20% of patients, respectively). Patients in the control group had no antibodies against these three antigens. Seventy percent of patients with HCC had the antibody against at least one of these antigens. CONCLUSIONS: Disease specific humoral immune response against TBP-1, p27(BBP), and rpL30 was induced in patients with HCC, and the antibodies against these antigens also may be used as tumor markers.

Aged↗
PubMed2001

alpha(6) Integrin is the main receptor of human papillomavirus type 16 VLP.

The present study was performed to determine the specific receptor of type HPV-16 using recombinant human papillomavirus-like particle (HPV-16 L1-VLP). The expression levels of alpha(6), beta(1), and beta(4) integrins were determined and compared with the amount of HPV-VLP binding in ten cell lines by flow cytometry. Our results show that the amount of VLP binding and the expression level of alpha(6) integrin are correlated, which was confirmed by an inhibition experiment using antibodies and by immunocytochemistry. Both the expression level of alpha(6) integrin and the amount of HPV-VLP binding were high in cervical cancer cell lines, as the type HPV-16 is the main cause of cervical cancer. The degree of binding of HPV-VLP matched the alpha(6) integrin expression level in cell lines but was not correlated with beta(1) and beta(4) levels, which suggests that alpha(6) integrin is the main receptor of HPV type 16.

Animals↗
PubMed2001

In vitro organ culture model for mucous membrane pemphigoid.

Mucous membrane pemphigoid (MMP), also known as cicatricial pemphigoid, is a rare vesiculobullous disease of mucosal tissues, which involves the oral, ocular, and other mucous membranes. We have studied a group of patients with histologically and immunopathologically proven pemphigoid disease involving predominantly the conjunctiva and oral mucosa in addition to other mucosae. The purpose of our study was to (i) demonstrate the specific binding of autoantibodies present in the sera of patients with MMP to normal human oral mucosa by indirect immunofluorescence (IIF) and (ii) to study the role of these autoantibodies in the pathogenesis of subepithelial blister formation using normal human buccal mucosa in organ culture. Serum and IgG fractions from MMP patients showed homogeneous smooth linear binding along the basement membrane zone (BMZ) of the normal buccal mucosa on IIF. Serum from pemphigus vulgaris patients showed intercellular or keratinocyte cell surface staining. BMZ separation developed at 48 h after incubation of normal human buccal mucosa in organ culture, with serum or IgG from patients with MMP but not after addition of normal human serum. Addition of pemphigus vulgaris serum to the in vitro culture of normal human buccal mucosa showed acantholysis. This preliminary report suggests that circulating autoantibodies may have an important role in the pathogenesis of MMP. This in vitro organ culture model will facilitate enhancing our understanding of various molecular events during the process of blister formation in MMP and in the study of other mucosal diseases.

Acantholysis↗
PubMed1996

Molecular complexity of the cutaneous basement membrane zone.

Ultrastructural examination of the cutaneous basement membrane zone (BMZ) reveals the presence of several attachment structures, which are critical for integrity of the stable association of epidermis and dermis. These include hemidesmosomes which extend from the intracellular compartment of the basal keratinocyte to the underlying basement membrane where they complex with anchoring filaments, thread-like structures traversing the lamina lucida. At the lower portion of dermal-epidermal attachment zone, anchoring fibrils extend from the lamina densa to the papillary dermis, where they associate with basement membrane-like structures, known as anchoring plaques. Molecular cloning of the cutaneous BMZ components has allowed elucidation of the structural features of the proteins which constitute these attachment structures. Specifically, hemidesmosomes have been shown to consist of at least four distinct proteins. The intracellular hemidesmosomal inner plaque is comprised of the 230-kD bullous pemphigoid antigen (BPAG1), and plectin, a high-molecular weight cytomatrix protein, encoded by the corresponding gene, PLEC1. The transmembrane component of the hemidesmosomes consists of the 180-kD bullous pemphigoid antigen (BPAG2), a collagenous protein also known as type XVII collagen (COL17A1), as well as of the basal keratinocyte-specific integrin alpha 6 beta 4. The anchoring filaments consist predominantly of laminin 5 with three constitutive subunit polypeptides, the alpha 3, beta 3 and gamma 2 chains, which is associated with laminin 6 with the chain composition alpha 3, beta 1 and gamma 1. Also associated with anchoring filaments is a novel protein, ladinin, which serves as autoantigen in the linear IgA disease, and the corresponding gene, LAD1, has been mapped to human chromosome 1. Finally, the major, if not the exclusive, component of anchoring fibrils is type VII collagen, encoded by the gene (COL7A1) which consists of 118 distinct exons, the largest number of exons in any gene published thus far. Collectively, the cutaneous basement membrane zone is a complex continuum of macromolecules which form a network providing the stable association of the epidermis to the underlying dermis. Thus, genetic lesions resulting in abnormalities in any part of this network could result in a blistering skin disease, such as epidermolysis bullosa.

Antigens, CD↗
PubMed2002

Crucial role of fibroblasts in regulating epidermal morphogenesis.

Epidermis reconstructed on de-epidermized dermis (DED) was used to investigate whether fibroblasts can substitute growth factors needed for generation of a fully differentiated epidermis. For this purpose, a centrifugal seeding method was developed to reproducibly incorporate different fibroblast numbers into DED. Using (immuno)histochemical techniques, we could demonstrate that in the absence of fibroblasts the formed epidermis consisted only of two to three viable cell layers with a very thin stratum corneum layer. However, in the presence of fibroblasts keratinocyte proliferation and migration was stimulated and epidermal morphology markedly improved. The stimulatory effect of fibroblasts showed a biphasic character: keratinocyte proliferation increased in the initial phase but decreased in later stages of cell culture. After 3 weeks culture at the air-liquid interface, the proliferation index decreased irrespective of the number of fibroblasts present within the dermal matrix to levels observed also in native epidermis. Keratin 10 was localized in all viable suprabasal cell layers irrespective of the absence or presence of fibroblasts. Keratin 6 was downregulated with increasing numbers of fibroblasts, and keratins 16 and 17 were absent in fibroblast-populated matrices. The expression of involucrin or transglutaminase 1 showed a similar pattern as for the keratins. Irrespective of the number of fibroblasts incorporated into DED, the expression of alpha(3), alpha(6), beta(1), and beta(4) integrin subunits was upregulated. In fibroblast-free DED matrices normalization of epidermal differentiation was only achieved when the culture medium was supplemented by keratinocyte growth factor. The results of this study indicate that normalization of epidermal differentiation can be achieved using a non-contractile dermal matrix populated with fibroblasts.

Cell Division↗
PubMed2000

Tyrosine phosphorylation: a critical component in the formation of hemidesmosomes.

Our goal was to evaluate the role of tyrosine phosphorylation in the complete formation of hemidesmosomes that occurs during development or during remodeling after injury. A corneal organ culture system was used to study hemidesmosome formation as it would occur in an intact tissue. Phosphorylation of the integrin subunit beta 4 and bullous pemphigoid antigen-1 (BPAG-1) was examined, as these proteins are known to play a role in linking the electron-dense plaques along the basal surface with the intermediate filaments to complete the formation of hemidesmosomes. Corneal epithelial sheets were placed on substrata that contained an intact basal lamina or basal laminae that had been either modified or removed. These constructs were incubated for up to 18 h, and hemidesmosome formation was evaluated by using transmission electron microscopy. When epithelial sheets were placed on intact basal laminae and incubated in the presence of the tyrosine kinase inhibitor genistein (200 microM), hemidesmosome formation was impaired. The formation of electron-dense regions was delayed, and no association of intermediate filaments was detected. Results were confirmed by biochemical studies. When the epithelium and underlying proteins were extracted and immunoprecipitated with beta 4 or BPAG-1, tyrosine phosphorylation decreased in the presence of genistein. In addition, the phosphorylation of beta 4 decreased when epithelial sheets were incubated on substrata from which the basal lamina had been removed or altered. Thus, a reduction in phosphorylation of tyrosine residues impairs the formation of mature hemidesmosomes, and substrata that fail to support hemidesmosome formation also demonstrate decreased phosphorylation of tyrosine residues.

Alkalies↗
PubMed2001

Characterization of cultured human prostatic epithelial cells by cluster designation antigen expression.

Cultured prostatic epithelial cells have been extensively studied as a model of prostate biology. What is the lineage relationship of the cultured cells to the epithelial cell types in tissue? How different are cultured cells derived from tumor tissue to those derived from benign tissue? Expression of cluster designation (CD) cell surface molecules has been shown to be useful in characterizing cells according to lineage. A CD profile was therefore generated for cultured human prostatic epithelial cells and compared with those previously established for basal and luminal epithelial cells in the prostate. Presence of CD44, CD49b, CD49f, and CD104 and absence of CD57 suggests that cultured cells were derived from basal cells of prostatic tissues. However, expression of certain CD antigens characteristic of luminal epithelial cells was also observed in subpopulations of cultured cells. The pattern of CD antigens in cultured cells reflects a phenotype similar to that of transit-amplifying cells that have been described in the prostate. Several CD antigens were found expressed by both cultured prostatic epithelial and stromal cells, and are probably associated with cell proliferation. The CD profiles of cultured epithelial cell strains derived from normal compared with malignant tissues were notably similar to each other and to that of the prostate cancer cell line PC-3. We conclude that cells in culture retain expression of certain lineage-characteristic CD antigens. Furthermore, CD antigens can define subpopulations of cells with differential gene expression.

Aged↗