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PubMed1977

Further observations on the site of bone prospective areas in the chick embryo wing bud.

The portion of the pre-axial region of the wing bud anterior to the cranial boundary of the skeletogenous territory of the stylo-zeugopod, according to Stark and Searls' recent maps, was surgically isolated in chicken embryos of the stages 18-25, and autoplastically grafted to the dorsal surface of the hind-limb bud or to the trunk. In other embryos of the same stages the cranial half or the cranial two thirds of the pre-axial region mentioned were isolated and heterotopically implanted. The experimental results consistently showed that the isolate contained a variously large portion of the presumptive radius and a significant part of the proximal half of the prospective humerus. In fact, the skeletal parts mentioned did not develop in situ in the donor wing while they differentiated, in general, in the implantation site. In this regard, the present research confirms data from previous experiments showing that Stark and Searls' maps do not offer an accurate representation of the position and the cranio-caudal width of the presumptive areas of the skeletal pieces of the wing. Our findings seem also to indicate that the major (or longitudinal) axis of the future radius is arranged along the proximodistal axis of the wing bud throughout the developmental stages mentioned; instead, the major axis of the future humerus undergoes a gradual shift, being first (stages 18-20) arranged nearly parallel to the bud base and getting more and more parallel to the proximo-distal axis of the wing bud through stages 21-24. Besides, between stage 18 and 25 the cranio-caudal width of the presumptive skeletogenous territory of the zeugopod, and to a lesser extent of the stylopod, seems to undergo a slight relative reduction with respect to the cranio-caudal thickness of the whole wing bud; this might depend on a process of aggregation of the prospective skeletogenous cells into gradually denser and more defined precartilaginous blastemes.

Animals↗

PubMed2004

The bud scar-based screening system for hunting human genes extending life span.

We developed a high-throughput screening system that allows identification of genes prolonging life span in the budding yeast Saccharomyces cerevisiae. The method is based on isolating yeast mother cells with an extended number of cell divisions as indicated by the increased number of bud scars on their surface. Fluorescently labeled wheat germ agglutinin (WGA) was used for specific staining of bud scars. Screening of a human HepG2 cDNA expression library in yeast resulted in the isolation of several yeast transformants with a potentially prolonged life span. The budding yeast S. cerevisiae, one of the favorite models used to study aging, has been studied extensively for the better understanding of the mechanisms of human aging. Because human disease genes often have yeast counterparts, they can be studied efficiently in this organism. One interesting example is the WRN gene, the human DNA helicase, which participates in the DNA repair pathway. The mutation of the WRN gene causes Werner syndrome showing premature-aging phenotype. Budding yeast contains WRN homologue, SGS1, and its mutation results in shortening yeast life span. The knowledge gained from the studies of budding yeast will benefit studies in humans for better understanding of aging and aging-related disease.

Aging↗

PubMed1988

Characterization of retinoid metabolism in the developing chick limb bud.

Retinoids (vitamin A derivatives) have been shown to have striking effects on developing and regenerating vertebrate limbs. In the developing chick limb, retinoic acid is a candidate morphogen that may coordinate the pattern of cellular differentiation along the anteroposterior limb axis. We describe a series of investigations of the metabolic pathway of retinoids in the chick limb bud system. To study retinoid metabolism in the bud, all-trans-[3H]retinol, all-trans-[3H]retinal and all-trans-[3H]retinoic acid were released into the posterior region of the limb anlage, the area that contains the zone of polarizing activity, a tissue possibly involved in limb pattern formation. We found that the locally applied [3H]retinol is primarily converted to [3H]retinal, [3H]retinoic acid and a yet unidentified metabolite. When [3H]retinal is locally applied, it is either oxidized to [3H]retinoic acid or reduced to [3H]retinol. In contrast, local delivery of retinoic acid to the bud yields neither retinal nor retinol nor the unknown metabolite. This flow of metabolites agrees with the biochemical pathway of retinoids that has previously been elucidated in a number of other animal systems. To find out whether metabolism takes place directly in the treated limb bud, we have compared the amount of [3H]retinoid present in each of the four limb anlagen following local treatment of the right wing bud. The data suggest that retinoid metabolism takes place mostly in the treated limb bud. This local metabolism could provide a simple mechanism to generate in a controlled fashion the biologically active all-trans-retinoic acid from its abundant biosynthetic precursor retinol. In addition, local metabolism supports the hypothesis that retinoids are local chemical mediators involved in pattern formation.

Animals↗

PubMed1993

Retroviral expression of FGF-2 (bFGF) affects patterning in chick limb bud.

To investigate the role of fibroblast growth factor-2 (basic fibroblast growth factor) in chick limb development, we constructed a replication-defective spleen necrosis virus to ectopically express fibroblast growth factor-2 in stage 20-22 chick limb bud. Because infecting cells in vivo proved to be inefficient, limb bud cells were dissociated, infected in vitro, and then grafted back into host limbs. This procedure caused duplications of anterior skeletal elements, including proximal humerus, distal radius, and digits 2 and 3. Eighty-nine percent of host wings receiving infected grafts at their anterior borders had duplications of one or more of these elements. The frequency of duplication declined dramatically when infected cells were grafted to progressively more posterior sites of host limb buds, and grafting to the posterior border had no effect at all. Several techniques were used to determine the role of infected tissue in forming skeletal duplications. First, staining with an fibroblast growth factor-2 specific monoclonal antibody showed higher than endogenous levels of fibroblast growth factor-2 expression associated with extra elements. Second, the host/donor composition of duplicated elements was determined by simultaneously infecting donor cells with viruses encoding fibroblast growth factor-2 or beta-galactosidase; donor tissue was then visualized by X-gal staining. Patterns of ectopic fibroblast growth factor-2 expression and X-gal staining confirmed the presence of infected donor tissue near duplicated structures, but the duplicated skeletal elements themselves showed very little staining. Similar results were obtained in duplications caused by infected quail wing bud cells grafted to the chick wing bud. These observations suggest that fibroblast growth factor-2-expressing donor tissue induced host tissue to form normally patterned extra elements. In support of this conclusion, implanting beads containing fibroblast growth factor-2 caused partial duplications of digit 2. These data provide the first direct evidence that fibroblast growth factor-2 plays a role in patterning in the limb bud.

Animals↗

PubMed1993

FGF-4 maintains polarizing activity of posterior limb bud cells in vivo and in vitro.

The polarizing region is a major signalling tissue involved in patterning the tissues of the vertebrate limb. The polarizing region is located at the posterior margin of the limb bud and can be recognized by its ability to induce additional digits when grafted to the anterior margin of a chick limb bud. The signal from the polarizing region operates at the tip of the bud in the progress zone, a zone of undifferentiated mesenchymal cells, maintained by interactions with the apical ectodermal ridge. A number of observations have pointed to a link between the apical ectodermal ridge and signalling by the polarizing region. To test this possibility, we removed the posterior apical ectodermal ridge of chick wing buds and assayed posterior mesenchyme for polarizing activity. When the apical ectodermal ridge is removed, there is a marked decrease in polarizing activity of posterior cells. The posterior apical ectodermal ridge is known to express FGF-4 and we show that the decrease in polarizing activity of posterior cells of wing buds that normally follows ridge removal can be prevented by implanting a FGF-4-soaked bead. Furthermore, we show that both ectoderm and FGF-4 maintain polarizing activity of limb bud cells in culture.

Animals↗

PubMed1999

FGF-7 modulates ureteric bud growth and nephron number in the developing kidney.

The importance of proportioning kidney size to body volume was established by clinical studies which demonstrated that in-born defecits of nephron number predispose the kidney to disease. As the kidney develops, the expanding ureteric bud or renal collecting system induces surrounding metanephric mesenchyme to proliferate and differentiate into nephrons. Thus, it is likely that nephron number is related to ureteric bud growth. The expression patterns of mRNAs encoding Fibroblast Growth Factor-7 (FGF-7) and its high affinity receptor suggested that FGF-7 signaling may play a role in regulating ureteric bud growth. To test this hypothesis we examined kidneys from FGF-7-null and wild-type mice. Results of these studies demonstrate that the developing ureteric bud and mature collecting system of FGF-7-null kidneys is markedly smaller than wild type. Furthermore, morphometric analyses indicate that mature FGF-7-null kidneys have 30+/-6% fewer nephrons than wild-type kidneys. In vitro experiments demonstrate that elevated levels of FGF-7 augment ureteric bud growth and increase the number of nephrons that form in rodent metanephric kidney organ cultures. Collectively, these results demonstrate that FGF-7 levels modulate the extent of ureteric bud growth during development and the number of nephrons that eventually form in the kidney.

Animals↗

PubMed2001

Induced repatterning of type XVIII collagen expression in ureter bud from kidney to lung type: association with sonic hedgehog and ectopic surfactant protein C.

Epithelial-mesenchymal tissue interactions regulate the formation of signaling centers that play a role in the coordination of organogenesis, but it is not clear how their activity leads to differences in organogenesis. We report that type XVIII collagen, which contains both a frizzled and an endostatin domain, is expressed throughout the respective epithelial bud at the initiation of lung and kidney organogenesis. It becomes localized to the epithelial tips in the lung during the early stages of epithelial branching, while its expression in the kidney is confined to the epithelial stalk region and is lost from the nearly formed ureter tips, thus displaying the reverse pattern to that in the lung. In recombinants, between ureter bud and lung mesenchyme, type XVIII collagen expression pattern in the ureter bud shifts from the kidney to the lung type, accompanied by a shift in sonic hedgehog expression in the epithelium. The lung mesenchyme is also sufficient to induce ectopic lung surfactant protein C expression in the ureter bud. Moreover, the shift in type XVIII collagen expression is associated with changes in ureter development, thus resembling aspects of early lung type epigenesis in the recombinants. Respecification of collagen is necessary for the repatterning process, as type XVIII collagen antibody blocking had no effect on ureter development in the intact kidney, whereas it reduced the number of epithelial tips in the lung and completely blocked ureter development with lung mesenchyme. Type XVIII collagen antibody blocking also led to a notable reduction in the expression of Wnt2, which is expressed in the lung mesenchyme but not in that of the kidney, suggesting a regulatory interaction between this collagen and Wnt2. Respecification also occurred in a chimeric organ containing the ureter bud and both kidney and lung mesenchymes, indicating that the epithelial tips can integrate the morphogenetic signals independently. A glial cell line-derived neurotrophic factor signal induces loss of type XVIII collagen from the ureter tips and renders the ureter bud competent for repatterning by lung mesenchyme-derived signals. Our data suggest that differential organ morphogenesis is regulated by an intra-organ patterning process that involves coordination between inductive signals and matrix molecules, such as type XVIII collagen.

Animals↗

PubMed2005

Novel regulators of kidney development from the tips of the ureteric bud.

Mammalian nephrogenesis depends on the interaction between the ureteric bud and the metanephric mesenchyme. As the ureteric bud undergoes branching and segmentation, the stalks differentiate into the collecting system of the mature kidney, while the tip cells interact with the adjacent cells of the metanephric mesenchyme, inducing their conversion into nephrons. This induction is mediated by secreted factors. For identifying novel mediators, the tips of the ureteric tree were isolated and microarray analyses were performed using manually refined, multistep gene ontology annotations. For identifying conserved factors, two databases were developed, one from mouse E12.5 and one from rat E13.5 ureteric buds. The overlap of mouse and rat data sets yielded 20 different transcripts that were enriched in the ureteric bud compared with metanephric mesenchyme and predicted to code for secreted proteins. Real-time reverse transcriptase-PCR and in situ hybridization confirmed these identifications. One of the genes that was highly specific to the ureteric bud tip was cytokine-like factor 1 (CLF-1). Recombinant CLF-1 in complex with its physiologic ligand, cardiotrophin-like cytokine (CLC), triggered phosphorylation of signal transducer and activator of transcription 3 in mesenchyme, a pathway characteristic of mesenchymal-to-epithelial conversion. Indeed, when applied to isolated rat metanephric mesenchyme, CLF-1/CLC (3 nM) induced mature nephron structures expressing glomerular and tubular markers. These results underline the power of this first comprehensive gene expression analysis of the ureteric bud tip to identify bioactive molecules.

Animals↗

PubMed2005

Ras and the Rho effector Cla4 collaborate to target and anchor Lte1 at the bud cortex.

Lte1, a protein important for exit from mitosis, localizes to the bud cortex as soon as the bud forms and remains there until cells exit from mitosis. Ras, the Rho GTPase Cdc42 and its effector the protein kinase Cla4 are required for Lte1's association with the bud cortex. Here we investigate how Ras, and the Cdc42 effector Cla4 regulate the localization of Lte1. We find that Ras2 and Lte1 associate in stages of the cell cycle when Lte1 is phosphorylated and associated with the bud cortex and that this association requires CLA4. Additionally, RAS1 and RAS2 are required for CLA4-dependent Lte1 phosphorylation. Our findings suggest that Cla4-dependent phosphorylation promotes the initial association of Lte1 with Ras at the bud cortex and that Ras is required to stabilize phosphorylated forms of Lte1 at the bud cortex. Our results also raise the interesting possibility that the localization of Lte1 affects the protein's ability to promote mitotic exit.

Adenylyl Cyclases↗

PubMed1976

[Contribution of somite cells to the development of posterior limb buds in mice].

1. The structural changes accompanying the early genesis of the mouse hindlimb bud have been studied in serial Epon thick sections at five successive somitic stages collected during the 10th developmental day. 2. As soon as the 14-somite stage, increased proliferative activities occurring in a limited area of the embryonic somatopleure close to the caudal end of the coelomic cavity point out the onset of limb morphogenesis. From its very beginning on, up to its ceasing at the 27-somite stage, the somatopleural proliferation is more active in the caudal part of the prospective limb territory and in the whole area, it appears stronger ventrally than dorsally. 3. Discrete structural changes occur during this period in the mesoderm interposed between the hindlimb bud territory and the overlying unsegmented sheet of somitic mesoderm, in relation with the genesis of the mesonephritic constituents and of the posterior cardinal vein. A detailed analysis of this area in serial sections reveals that the unsegmented somitic mesoderm does not provide any cell to the limb bud mesoderm. 4. As soon as the somitic mesoderm becomes organized into segmented somites, obvious histological changes located on the lateral aspect of the ventro-lateral somitic edge reveal that numerous undifferentiated somitic cells invade the proximal area of the limb bud mesoderm. Starting at the 27-somite stage in the first two metameres adjacent to the precordonal portion of the limb bud, this processus extends to the five somites of the limb territory at the 33-somite stage. In each metamere, the area of cell migration takes place near the caudal border of the somite and, from one somite to the other, the number of migrating cells increases in a cephalo-caudal direction. No sign of a similar cell migration was observed in the somites located cranially or caudally to the hindlimb bud. 5. The results are discussed in relation with the various experimental and descriptive evidences obtained in non-mammalian Vertebrates about the contribution of the somites to limb morphogenesis and about the factors involved in the onset of cell proliferation in the prospective limb mesoderm.

Animals↗

PubMed1999

The protein phosphatase inhibitor cantharidin induces head and foot formation in buds of Cassiopea andromeda (Rhizostomae, Scyphozoa).

The polyps of Cassiopea andromeda produce spindle shaped, freely swimming buds which do not develop a head (a mouth opening surrounded by tentacles) and a foot (a sticky plate at the opposite end) until settlement to a suited substrate. The buds, therewith, look very similar to the planula larvae produced in sexual reproduction. With respect to both, buds and planulae, several peptides and the phorbolester TPA have been found to induce the transformation into a polyp. Here it is shown that cantharidin, a serine/threonine protein phosphatase inhibitor, induces head and foot formation in buds very efficiently in a 30 min treatment, the shortest yet known efficient treatment. Some resultant polyps show malformations which indicate that a bud is ordinary polyp tissue in which preparatory steps of head and foot formation mutually block each other from proceeding. Various compounds related to the transfer of methyl groups have been shown to affect head and foot formation in larvae of the hydrozoon Hydractinia echinata. These compounds including methionine, homocysteine, trigonelline, nicotinic acid and cycloleucine are shown to also interfere with the initiation of the processes which finally lead to head and foot formation in buds of Cassiopea andromeda.

Alkaloids↗

PubMed2000

Time-lapse observation of branching morphogenesis of the lung bud epithelium in mesenchyme-free culture and its relationship with the localization of actin filaments.

It has been shown that branching morphogenesis of the lung bud is mediated by epithelial-mesenchymal interaction via such molecules as FGF10, BMP4 and Shh. However, a recent study showed that the isolated lung epithelium still undergoes branching morphogenesis in vitro even in the absence of mesenchyme (Nogawa and Ito, 1995). In the present study, we observed in vitro the dynamic movement of the isolated lung epithelium of the fetal mouse using time-lapse recording, and investigatedthe roles of actinfilaments in branching of the lung bud. First, time-lapse observation of the initial phase of lung branching morphogenesis revealed that at the sites of cleft formation, the epithelial surface was retracted inward from its original position. From this observation we assumed that there should be some structures which exert a physical force on the epithelium, and the localization and arrangement of actin fibers in the cultured lung epithelium were examined at various stages of branching morphogenesis. At the prebudding (6 h) and onset-budding (24 h) stages, no specific localization of actin filaments was observed in the lung bud epithelium, but at the postbudding stage (48 h) they were localized densely in the cells at the tip of the branched lung epithelium. The cell density was not different between the tip and cleft regions of the lung bud epithelium. When cultured with FGF-soaked beads, an actin-rich region was induced at the tip of the lung bud which was growing toward an FGF-soaked bead. These results indicate that actin fibers do not play a significant part in cleft formation but can be secondarily induced by FGF in the surrounding matrix and play some roles at later shaping of the branch in lung morphogenesis.

Actins↗

PubMed2001

Tumour budding intensity in relation to cathepsin D expression and some clinicopathological features of colorectal cancer.

Although it has been suggested that tumour budding at the invasive edge of colorectal cancer is an important prognostic factor its biological significance for tumour progression is still to be evaluated. The aim of the study was to correlate tumour budding intensity with cathepsin D expression and some other clinicopathological variables of presumed or established prognostic value. 48 patients with colorectal cancer at pT3 stage, G2 grade of histological differentiation and tumour budding at the invasive edge were evaluated. Colorectal tumours were investigated for cathepsin D expression by immunohistochemistry of formalin-fixed and paraffin embedded tissues. There was no statistically significant relationships between tumour budding intensity grade and primary tumour cathepsin D expression, stromal cell cathepsin D expression and histochemical immunostaining of cathepsin D in rumour budding at its invasive edge. The tumour budding intensity was not associated with lymph node status, tumour site, peritumoral inflammatory response as well as the patient's age and sex. The results of this study suggest that intensity of tumour budds formation at the invasive margin of colorectal cancer is not associated with presumed or established prognostic factors such as lymph node metastases, and peritumoural inflammatory reaction as well as cathepsin D expression.

Adenocarcinoma↗

PubMed2002

Exogenous retinoic acid induces a stage-specific, transient and progressive extension of Sonic hedgehog expression across the pectoral fin bud of zebrafish.

We have performed a time course analysis of the expression of Sonichedgehog (shh) and patched1 (ptc1) in response to exogenous retinoic acid (RA) application to get some insight into the mechanism(s) underlying the formation of a mirror-image duplication of shh and ptc1 domains of expression in the pectoral fin buds of zebrafish. We have shown that RA exposure during the early stages of pectoral fin development first results in a rapid decrease or complete loss of shh/ptc1 expression. This is followed by reappearance of transcripts in the normal posterior domain, then by a stage-dependent and progressive expansion of the shh domain from the ZPA towards the anterior margin of the bud. Shh transcripts are induced in mesenchymal cells underlying the ventral ectoderm at the base of the bud. Once shh expression is activated in the most anterior cells, the number of shh-expressing cells increases in this region, possibly through an amplification mechanism involving signals from the apical ectodermal ridge. At this time, shh expression disappears from cells centrally located in the bud, resulting in the formation of the two distinct domains. An anterior extension of shh expression is also obtained in syu mutants with impaired shh function, suggesting that shh induction across the fin bud is independent of shh signaling. This study suggests the existence of complex mechanisms controlling the spatial and temporal expression of shh in the developing fin bud.

Animals↗

PubMedDate not supplied

Laminin-5 gamma 2 chain and matrix metalloproteinase-2 may trigger colorectal carcinoma invasiveness through formation of budding tumor cells.

UNLABELLED: Tumor budding has been reported to be significantly associated with lymph node or distant metastasis in colorectal carcinomas. We attempted to elucidate the contributing factors for the formation of budding tumor cells. PATIENTS AND METHODS: Fifty T1 colorectal carcinomas were immunostained using several antibodies against adhesion-related molecules, and the associations between tumor budding and clinicopathological factors were examined. RESULTS: Univariate analysis revealed that predominant histology, lymphatic permeation, depth of invasion, width of invasion, CD44 variant 6 expression, laminin-5 gamma 2 chain expression and stromal MMP-2 expression were associated with tumor budding. Multivariate analysis showed that lymphatic permeation, laminin-5 gamma 2 chain expression and stromal MMP-2 expression were independently and significantly associated with tumor budding. Furthermore, laminin-5 gamma 2 chain and stromal MMP-2 expression patterns were significantly associated with locoregional failure. CONCLUSION: These results suggest that laminin-5 gamma 2 chain and MMP-2 may contribute synergistically to the formation of budding tumor cells.

Aged↗

PubMed2004

[Research of the amounts of flavonoids accumulated in the buds of single-styled hawthorn].

OBJECTIVE: To determine the amounts of flavonoids accumulated in the buds of single-styled hawthorn (Crataegus monogyna Jacq.); and to establish the possibility of usage of hawthorn buds in pharmacy practice. MATERIAL AND METHODS: Different examples of hawthorn buds collected in 2001, 2002 and 2003 from some Lithuanian regions: Akmene (V1, V2, V3), Kedainiai (S1, S2, S3), Klaipeda (M1), Lazdijai (D1, D2, D3) and Vilnius (VL1, VL2) from branches of lower storey. Examples were extracted with ethanol and flavonoids were measured by spectroscopic method and by high performance liquid chromatography. RESULTS: From 0.72 to 1.89% of flavonoids are found in the hawthorn buds. The dominating flavonoid is vitexin-2-O-rhamnosid (6.72-10.91 milligrams in one gram of dried crude drug). Other flavonoids are: vitexin: 0.88-6.53 milligrams/gram, hyperosid: 0.85-2.70 milligrams/gram, rutin: 0.72-2.10 milligrams/gram and quercitrin: 0.82-1.01 milligrams/gram. CONCLUSION: The marks of phytochemical compounds of Lithuanian single-styled hawthorn buds corroborated the theory of possibility to use hawthorn buds as pharmaceutical crude drug and form sufficient ground for planning phytochemical and pharmacological researches of new hawthorn crude drug.

Apigenin↗

PubMed2006

[The changes of three endogenous hormones during flower bud differentiation of Schisandga chinensis].

OBJECTIVE: To reveal the relation between endogenous hormones and the flower bud differentiation in Schisandga chinensis. METHOD: Top buds of extremely short branch and axillary buds of long branch in the same plant of S. chinensis were used as material and the contents of endogenous hormones were measured during different periods of the flower bud differentiation with HPLC. RESULT: The result showed that flower bud differentiation and the formation of female flower were inhibited by high concentration of GA3 and were promoted by high concentration of ABA or ZT. Low ratio of GA3/ABA has the same result. CONCLUSION: There was a correlation between endogenous hormones and the flower bud differentiation of S. chinensis.

Abscisic Acid↗

PubMedDate not supplied

MMP-9 and cathepsin B expression in tumor budding as an indicator of a more aggressive phenotype of colorectal cancer (CRC).

BACKGROUND: The aim of this study was to clarify the relationship between the expression of MMP-9 and Cathepsin B (CathB) at the main tumor mass and lymph node metastases and the presence of tumor budding and vascular invasion at the invasion front, as well as lymph node involvement. PATIENTS AND METHODS: The immunohistochemical expressions of MMP-9 and CathB were studied in 55 specimens of colorectal adenocarcinomas (pT3, G2). A standard avidin-biotin immunoperoxidase staining method (ABCu-NCL) was performed on 4-microm paraffin-embedded tissue sections with a mouse anti-human MMP-9 and CathB monoclonal antibody. RESULTS: Positive immunostaining for MMP-9 in the primary tumor was observed in 30/55 (54.6%) cases and in 29 cases (52.7%) for CathB. A statistically significant association was found between the expressions of MMP-9 and CathB in the primary tumor and lymph node involvement (p<0.01). The expression of CathB in the primary tumor was associated with the presence of tumor budding and vascular invasion (p<0.01); but no such association was found for MMP-9. The expressions of MMP-9 and CathB in buds were strongly associated with lymph node involvement (p<0.01). However, the presence of vascular invasion was significantly associated only with positive expression of CathB in the buds (p<0.01). CONCLUSION: If the presence of budding in the colorectal cancer invasive front indicates higher invasive potential, the present results seem to suggest the existence of a strong relationship between MMP-9 and CathB expressions in the buds and a more aggressive tumor phenotype.

Adenocarcinoma↗