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PubMed2003

Structure, regulation, and function of mammalian membrane guanylyl cyclase receptors, with a focus on guanylyl cyclase-A.

Besides soluble guanylyl cyclase (GC), the receptor for NO, there are at least seven plasma membrane enzymes that synthesize the second-messenger cGMP. All membrane GCs (GC-A through GC-G) share a basic topology, which consists of an extracellular ligand binding domain, a short transmembrane region, and an intracellular domain that contains the catalytic (GC) region. Although the presence of the extracellular domain suggests that all these enzymes function as receptors, specific ligands have been identified for only three of them (GC-A through GC-C). GC-A mediates the endocrine effects of atrial and B-type natriuretic peptides regulating arterial blood pressure and volume homeostasis and also local antihypertrophic actions in the heart. GC-B is a specific receptor for C-type natriuretic peptide, having more of a paracrine function in vascular regeneration and endochondral ossification. GC-C mediates the effects of guanylin and uroguanylin on intestinal electrolyte and water transport and on epithelial cell growth and differentiation. GC-E and GC-F are colocalized within the same photoreceptor cells of the retina and have an important role in phototransduction. Finally, the functions of GC-D (located in the olfactory neuroepithelium) and GC-G (expressed in highest amounts in lung, intestine, and skeletal muscle) are completely unknown. This review discusses the structure and functions of membrane GCs, with special emphasis on the physiological endocrine and cardiac functions of GC-A, the regulation of hormone-dependent GC-A activity, and the relevance of alterations of the atrial natriuretic peptide/GC-A system to cardiovascular diseases.

Animals↗

PubMed1986

Cartilage grafting in facial reconstruction with special consideration of irradiated grafts.

The search for the perfect facial implant for reconstruction of the face continues. Cartilage, once thought to be an undesirable graft material because of its propensity for absorption, has regained popularity in the past decade. Various preparation techniques have been employed to ensure graft sterility and diminished absorption. An improved understanding of cartilage structure and physiology has shed considerable light on the host-graft relationship. Gamma irradiation is a time-honored method of preservation. An experiment was undertaken to investigate the physiology of irradiated cartilage grafts following prolonged implantation on the facial skeleton of sheep and dog. Merthiolate preserved grafts were used as controls. Direct observation, histochemical techniques, autoradiography, and transmission electron micrography were used to determine chondrocyte viability and matrix composition. It was surprising to note that following implantation of 16 to 72 months, complete resorption was seen in 87.7% of the irradiated grafts and in 43.8% of the Merthiolate stored controls. Many of the grafts acquired chondrocytes from the host and produced new proteoglycan matrix as well as undergoing some degree of ossification. A comparison to the clinical situation in humans is made.

Animals↗

PubMed1997

[Gray scale histographic analysis of the acetabular cartilage for quantifying hip ultrasound images].

AIM: Congenital dislocated hips are classified in ultrasound studies using the angle measuring system of Graf and evaluating the typical changes in acetabular form and structure. We attempted to quantify physiological and pathological changes in the acetabular cartilage using B-mode scan ultrasound studies. METHODS: Gray-level histograms were performed on 894 infant hips; 833 were classified by the Graf scheme as Type IIa and 61 as Type IIIa/b. We compared the histological structure of the cartilage of both the acetabulum and the head of the femur using objective parameters. RESULTS: We could show that gray-level histograms of the acetabular cartilage are an ideal way to study objectively ultrasound evaluations of the same area. Type IIa hips with histographically identified ossification zones show significantly higher healing rates than those without such areas. Type IIIb hips with histographically demonstrated pathological changes showed a significantly longer healing rate and a higher incidence of persistent dislocations that did Type IIa hips with normal cartilage. CONCLUSION: Structural changes in the acetabular cartilage can be evaluated not only qualitatively but also quantitatively to provide prognostic information in infants with congenially dislocated hips. Gray-level histograms nicely supplement ultrasound examination.

Acetabulum↗

PubMed2004

Evaluation of fetal growth and estimation of fetal age based on skeletal growth in Hokkaido sika deer (Cervus nippon yesoensis Heude, 1884).

We investigated fetal development and the estimation of fetal age of 127 Hokkaido sika deer fetuses, categorizing them into three groups according to the nutritional condition of populations. The order and time of the appearance of ossification centers were clarified, and fetal age was determined based on bone length and the appearance of ossification centers. Then we observed the differences in fetal growth among three populations, and discussed the effect of poor nutrition on the fetal growth. The results suggest that fetal diaphysial length of the femur was affected very little by nutritional conditions, whereas conception dates were delayed and fetal weight was restricted as the nutritional condition became poorer. Although it is impossible to know the exact accurate fetal age in wild populations, it was possible to create a standard to estimate fetal age more precisely by the method described in this study. Both the bone length and the appearance of ossification centers are reliable indices to estimate fetal age precisely in measurements available from fetuses of unknown age, and can be applied to estimate the fetal age of other populations of sika deer, whereas estimation of fetal age based on weight is prone to great errors.

Age Determination by Skeleton↗

PubMed2006

The physiology of bone blood flow: a review.

INTRODUCTION: Bone circulation plays an important role in bone physiology, but has been relatively poorly studied, because most techniques of circulatory research are difficult to apply to bone. This article summarizes briefly some of the important aspects of the physiology of bone blood flow most relevant to orthopaedics. METHODS: The gold standard for experimental measurement of bone blood flow is the radioactive microsphere technique, though advances are being made in other techniques, such as positron emission tomography, laser and ultra-sound Doppler velocimetry, and near infrared spectroscopy, that may provide useful clinical measurement in the future. RESULTS: Multiple vascular pathways contribute to an adaptive response to traumatic disruption of bone circulation. The microcirculation is not merely a passive conduit for blood flow, but plays an active role in controlling bone processes such as osteochondral ossification. DISCUSSION: The pathophysiology of bone circulation has been associated with osteonecrosis, but more and more evidence is pointing to the importance of bone circulation in fracture repair and osteoporosis, both of which are potentially very exciting areas for future studies.

Bone and Bones↗

PubMed1996

Physiological roles for parathyroid hormone-related protein: lessons from gene knockout mice.

A null mutation in the PTHrP gene produces profound abnormalities in endochondral bone formation in vivo. The role of PTHrP as a modulator of the chondrocytic proliferation and differentiation program is evident in the alterations that occur in its absence in the temporal and spatial sequence of chondrocyte development and subsequent endochondral bone formation that is necessary for normal bone elongation. These actions of PTHrP are probably responsible for the delay in chondrocyte development seen in Jansen osteochondrodystrophy, a disease caused by ligand-independent activation of the PTH-PTHrP receptor (Schipani et al., 1995). Furthermore, these conclusions have been corroborated by the observation that chondrocyte-specific overexpression of PTHrP causes a profound delay in the developmental program of chondrocyte differentiation and endochondral ossification (Weir et al., 1995). The morphological abnormalities in the knockout mice were limited to the skeletal system, despite the widespread production of PTHrP during fetal development. At this point, one can only speculate about the limited tissue distribution of the abnormalities. It is possible, for example, that other gene products, such as PTH, can compensate for the loss of PTHrP in some tissues. Alternatively, possible abnormalities in proliferation and differentiation may be present but morphologically subtle. A molecular assessment of these possible actions may well reveal more widespread effects of PTHrP.

Animals↗

PubMed2001

Extrinsic tension results in FGF-2 release, membrane permeability change, and intracellular Ca++ increase in immature cranial sutures.

There are numerous studies cataloging the temporal profiles of the various growth factors during the morphogenesis of cranial sutures. There are also many clearly documented mutations of the receptors of some of these growth factors such as fibroblast growth factor (FGF)R-2 and FGFR-3 in clinical craniosynostosis. It is obvious, and often concluded, that growth factors play a role or are involved in craniofacial development. However, precisely what that role is, what causes the changes in the growth factor levels, and why these changes occur in the particular temporal and spatial patterns observed remains elusive. Using simple physics, we applied a plasma membrane disruption model and the principles of complex adaptive systems to arrive at a conjecture of calvarial morphogenesis. The purpose of this article is to introduce the concept of complex adaptive systems, to propose our conjecture, and to provide experimental proof of some key steps in this conjecture: tension induces rapid and demonstrable physiological responses in some cells within the immature cranial sutures. These responses include increases of intracellular Ca++, plasma membrane permeability, and the release of growth factors, e.g., FGF-2. Paired coronal sutures from 1-week-old Sprague-Dawley rat pups were subjected to either 0.59 N of tensile force or no force for 5 minutes in a protein-free medium. FGF-2 levels in the media were measured by slot blot analysis. Western blot analysis was used to determine FGF-2 levels in the sutures. To determine cell membrane permeability changes, fluorescein-conjugated dextran, with a molecular weight of 10 kd, was added to the media during the 5 minutes with or without tensile force. Laser confocal microscopy was used to compare the amount of entry of this impermeant tracer and the pattern of permeability change at the tissue level. To determine the intracellular pCa++, the sutures were first loaded with a calcium indictor, FURA-2 AM, and then subjected isotonically to 0.059 N of tension. The intracellular pCa++ was expressed as ratio of Ca++-bound FURA-2 to Ca++-free FURA-2. The experimental findings were as follows: 1) Sutures, in response to tension, release FGF-2. 2) Sutures contain higher levels of FGF-2 when strained. 3) There is an increase in the sutural cell membrane permeability as a result of tensile strain. 4) The cells along the leading edges of the ossification fronts (at the insertion sites of Sharpey's fibers) demonstrated the maximum permeability increase. 5) There was an immediate (within seconds) increase in intracellular Ca++. and 6) This increase in intracellular Ca++ caused by tension was reversible and independent of the extracellular Ca++ ion availability. In summary, these data support, in part, the conjecture that growth of the brain places strain on the cells within the immature sutures, which causes the iteration of a set of cellular subroutines. These subroutines integrate to generate the emergent property of directed cranial expansion with dissipation of the initiating strains.

Adaptation, Physiological↗

PubMed2000

A role for smad6 in development and homeostasis of the cardiovascular system.

Smad proteins are intracellular mediators of signalling initiated by Tgf-betasuperfamily ligands (Tgf-betas, activins and bone morphogenetic proteins (Bmps)). Smads 1, 2, 3, 5 and 8 are activated upon phosphorylation by specific type I receptors, and associate with the common partner Smad4 to trigger transcriptional responses. The inhibitory Smads (6 and 7) are transcriptionally induced in cultured cells treated with Tgf-beta superfamily ligands, and downregulate signalling in in vitro assays. Gene disruption in mice has begun to reveal specific developmental and physiological functions of the signal-transducing Smads. Here we explore the role of an inhibitory Smad in vivo by targeted mutation of Madh6 (which encodes the Smad6 protein). Targeted insertion of a LacZ reporter demonstrated that Smad6 expression is largely restricted to the heart and blood vessels, and that Madh6 mutants have multiple cardiovascular abnormalities. Hyperplasia of the cardiac valves and outflow tract septation defects indicate a function for Smad6 in the regulation of endocardial cushion transformation. The role of Smad6 in the homeostasis of the adult cardiovascular system is indicated by the development of aortic ossification and elevated blood pressure in viable mutants. These defects highlight the importance of Smad6 in the tissue-specific modulation of Tgf-beta superfamily signalling pathways in vivo.

Animals↗

PubMed1981

Postnatal morphogenesis of the anconeal process of the ulna of swine.

The proximal ulnae of 79 swine ranging in age from 1 day to 2 years were radiographed and those of 46 of the swine were studied histologically. The anconeal process was a hyaline cartilaginous model at birth but ossified by 6 months of age. Endochondral ossification began at the base of the anconeal process and progressed in a pyramidal manner toward its apex. Secondary center of ossification was not observed in porcine anconeal processes examined. A lesion in the anconeal process of a 2-year-old animal was interpreted as a fracture of the anconeal process, and not a fracture of an anconeal physis. Implications of the development of the anconeal process as related to osteochondrosis in swine were discussed.

Animals↗

PubMed2003

[The hyoid bone and vertical dimension].

Located at the center of the neck and the pharyngeal column and, thanks to its attachments, tied to the base of the skull, to the mandible, to the tongue, to the larynx, and to the scapular belt, the hyoid bone is intimately related to an extensive tendino-muscular complex. From this strategic position, the hyoid bone participates in all functional and nutritional activities of the oro-facial complex. Heavily involved in the physiology of the pharyngeal column, it appears to enjoy a remarkable stability in relation to its surrounding structures as it participates in the critical process of vertical ontogenetic development. Deceptively simple in its shape and location it is anything but. While it would be easy to scorn it as a vestigial organ of little importance, the hyoid bone is actually an ancient witness of the evolution of the first human vertebrae and a vestige of the 2nd and 3rd brachial arches. Like a toy buffeted by powerful groups of muscles and tendons, the hyoid bone occupies a crossroads position where it is solicited by many vital functions, the most critical of which is respiration. Its functional behavior seems to have evolved considerably over the course of phylogeny. By piecing together the conceptions of many authors, we can conclude that the similarities--and, later, the differences--existing between the human fetus and those of other primates, and, then between young children and young primates, are related to an essentially different status quo, which modifies the delicate equilibrium of the pharyngeal region. In metamorphosing from a sagittal equilibrium to one that is vertical, this very status quo maintains an open respiratory airway during each time frame, continuously permitting the articulation and modulation of sounds, particularly during the vertical period. The hyoid bone is unpaired, in a mid-line position, symmetrical, and located just below the mandible with which it shares, to an astonishing extent, certain morphological characteristics. Its maturation and its ossification occur slowly and late even though it migrates much more rapidly to a vertical position than does the bulk of the cervical complex. True articulations between the body and the horns of the hyoid bone persist for a very long time; and so the junction between the small horns doesn't fully calcify until about the age of 50; and, in spite of its appearance, the hyoid bone is extremely supple, a quality that allows it to make substantial contributions to functional activity. If the hyoid bone itself scarcely moves during normal respiration, modern research attributes an increasing role to it in the maintenance of the equilibrium of the pharyngeal column. It seems to be called upon to respond to a number of demands that it manages to satisfy as it helps to maintain the permeability of the pharyngeal column and thus make respiration possible. In so doing, the hyoid bone adjusts its positioning, and, perhaps most important, its orientation to the physiological requirements imposed by pharyngeal obstruction and mouth breathing. Intimately connected to the larynx, the hyoid bone plays a part in phonation that has long been recognized. Recent studies now also show that the contribution the hyoid bone makes to respiratory equilibrium is far more important than its small size would suggest.

Adaptation, Biological↗

PubMed2003

Growth hormone modulates cholecalciferol metabolism with moderate effects on intestinal mineral absorption and specific effects on bone formation in growing dogs raised on balanced food.

The aim of the study was to investigate the influence of growth hormone (GH) on Vitamin D3 metabolism and the subsequent effects on calcium (Ca) homeostasis and skeletal growth in growing dogs. A group of Miniature Poodles received supraphysiological doses of GH (GH group; n = 6; 0.5 IU GH per kg body per day) from 12 to 21 weeks of age and was compared with a control placebo-treated group (n = 8). Biologic activity of GH in the GH compared to the control group was indicated by (a) the 2.5- to 3.5-fold increase in the plasma concentrations of insulin-like growth factor I (IGF-I), (b) the increased production of 1,25-dihydroxycholecalciferol as indicated by the significantly increased plasma 1,25-dihydroxycholecalciferol concentrations and the 12.9-fold increase in renal 1alpha-hydroxylase gene expression, and (c) the inhibited production of 24,25-dihydroxycholecalciferol as indicated by the significantly lower plasma 24,25-dihydroxycholecalciferol concentrations and the similar levels of renal 24-hydroxylase gene expression. Despite the distinct effects on Vitamin D(3) metabolism in the GH group, there were only moderate effects on the intestine, i.e. at 20 weeks of age there was a significant increase of 14.4 and 5.6% in fractional absorption of Ca and phosphate (Pi), respectively, compared to the control group. GH administration resulted in significantly elevated glomerular filtration rate, with no differences in Pi urine excretion as a result of a concomitant increase in the tubular reabsorption of Pi. GH had only limited disturbing effects on endochondral ossification as indicated by the maintenance of the regularity of the growth plates. However, GH had specific anabolic effects on bone formation without concomitant effect on bone resorption that may result in disorders of skeletal remodeling and manifestation of enostosis.

Analysis of Variance↗

PubMed1998

Distinct and overlapping patterns of localization of bone morphogenetic protein (BMP) family members and a BMP type II receptor during fracture healing in rats.

Bone morphogenetic proteins (BMPs) and their receptors (BMPRs) are thought to play an important role in bone morphogenesis. The purpose of this study was to determine the locations of BMP-2/-4, osteogenic protein-1 (OP-1, also termed BMP-7), and BMP type II receptor (BMPR-II) during rat fracture healing by immunostaining, and thereby elucidate the possible roles of the BMPs and BMPR-II in intramembranous ossification and endochondral ossification. In the early stage of fracture repair, the expression of BMP-2/-4 and OP-1 was strongly induced in the thickened periosteum near the fracture ends, and coincided with an enhanced expression of BMPR-II. On day 7 after fracture, staining for BMP-2/-4 and OP-1 immunostaining was increased in various types of chondrocytes, and was strong in fibroblast-like spindle cells and proliferating chondrocytes in endochondral bone. On day 14 after fracture, staining with OP-1 antibody disappeared in proliferating and mature chondrocytes, while BMP-2/-4 staining continued in various types of chondrocytes until the late stage. In the newly formed trabecular bone, BMP-2/-4 and OP-1 were present at various levels. BMPR-II was actively expressed in both intramembranous ossification and endochondral ossification. Additionally, immunostaining for BMP-2/-4 and OP-1 was observed in multinucleated osteoclast-like cells on the newly formed trabecular bone, along with BMPR-II. In reference to our previous study of BMP type I receptors (BMPR-IA and BMPR-IB), BMPR-II was found to be co-localized with BMPR-IA and BMPR-IB. BMP-2/-4 and OP-1 antibodies exhibited distinct and overlapping immunostaining patterns during fracture repair. OP-1 may act predominantly in the initial phase of endochondral ossification, while BMP-2/-4 acts throughout this process. Thus, these findings suggested that BMPs acting through their BMP receptors may play major roles in modulating the sequential events leading to bone formation.

Animals↗

PubMed1981

[Boundaries between the physiological and the beginnings of pathology in the development of the proximal femoral epiphysis (author's transl)].

This article describes the physiological development of the proximal femoral epiphysis and its variable aspects. It also includes a discussion of the transition to the pathological changes characteristic of dysplasia of the head of the femur and of Legg-Calvé-Perthes' disease, basing on a latent ischaemic phase in the development of the proximal femoral epiphysis, resulting from hypoplastic vascular development.

Child↗

PubMed1996

Chondrocytes synthesize type I collagen and accumulate the protein in the matrix during development of rat tibial articular cartilage.

The present study was designed to investigate whether or not chondrocytes in articular cartilage express type I collagen in vivo under physiological conditions. Expressions of the gene and the phenotype of type I collagen were examined in rat tibial articular cartilage in the knee joint during development. Knee joints of Wistar rats at 1, 5, and 11 weeks postnatal were fixed in 4% paraformaldehyde with or without 0.5% glutaraldehyde and decalcified in 10% EDTA. After the specimens were embedded in paraffin and serial sections made, adjacent sections were processed for immunohistochemistry and in situ hybridization for type I collagen. The epiphysis of the tibia was composed of cartilage in week- 1 rats. Formation of articular cartilage was in progress in week 5 as endochondral ossification proceeded and was completed in week 11. Anti-type I collagen antibody stained only the superficial area of the epiphysis in week 1, but the immunoreactivity was expanded into the deeper region of the articular cartilage with development in weeks 5 and 11. Hybridization signals for pro-alpha 1 (I) collagen were seen in some of chondrocytes in the epiphysis of the week-1 tibia. The most intense signals were identified in chondrocytes in week 5 and the signals appeared weaker in week 11. The present study demonstrated that chondrocytes synthesize type I collagen and accumulate the protein in the matrix during development of the articular cartilage.

Age Factors↗

PubMed1996

Mechanical factors in bone growth and development.

Mechanobiologic factors strongly influence skeletal ossification and regulate changes in bone geometry and apparent density during ontogeny. We have developed computer models that implement a simple mathematical rule relating cyclic tissue stresses to bone apposition and resorption. Beginning at the fetal stages of the femoral anlage, these models successfully predict the appositional bone growth and modeling observed in the development of the diaphyseal cross section. The same basic mechanobiologic rule can also predict the architectural construction of the proximal cancellous bone formed in regions of endochondral ossification. Geometry and density changes in adult diaphyseal and cancellous bone as a result of changes in physical activity can be simulated by invoking the same rule used during development. Future clinical and experimental work is needed to provide more quantitative data for mechanobiologic rules and elucidate the interactions between chemical and mechanical factors influencing bone biology.

Adaptation, Physiological↗

PubMed2002

Effects of transforming growth factor-beta deficiency on bone development: a Fourier transform-infrared imaging analysis.

Transforming growth factor-beta 1 (TGF-beta1) is a cytokine member of the TGF-beta superfamily involved in the control of proliferation and differentiation of various cell types. TGF-beta1 plays an important role in bone formation and resorption. To determine the effect of TGF-beta1 deficiency on bone mineral and matrix, tibias from mice in which TGF-beta1 expression had been ablated (TGF-beta1 null) were analyzed and compared with background- and age-matched wild-type (WT) control animals by Fourier transform-infrared imaging (FTIRI) and histochemistry. FTIRI allows the characterization of nondemineralized thin tissue sections at the ultrastructural level with a spatial resolution of approximately 7 microm. The spectroscopic parameters calculated were: mineral-to-matrix ratio (previously shown to correspond to ash weight); mineral crystallinity (related to the crystallographically determined crystallite size and perfection in the apatite c-axis direction); and collagen maturity (related to the ratio of pyridinoline:deH-DHLNL collagen cross-links). Several fields were selected to represent different stages of bone development within the same specimen from the secondary ossification center to the distal diaphysis. Anatomically equivalent areas were compared as a function of age and genotype. The spectroscopic results were expressed both as color-coded images and as pixel population distributions for each of the three parameters monitored. Based on comparisons of histochemistry and FTIRI, there were distinctive age and genotype variations. At all ages examined, in the TGF-beta1 null mice growth plates, alkaline phosphatase (ALP) activity and collagen maturity were reduced, but no effect on mineral content or crystallinity was noted. In the TGF-beta1 null mice metaphyses, there was a persistence of trabeculae, but no significant alterations in mineral content or crystallinity. In contrast, mineral content, mineral crystallinity, and collagen maturity were reduced in the secondary ossification center and cortical bone of the TGF-beta1 null mice. These results, consistent with a mechanism of impaired bone maturation in the TGF-beta1 null mice, may be directly related to TGF-beta1 deficiency and indirectly to increased expression of inflammatory cytokines in the TGFbeta1 null mice.

Animals↗

PubMed2002

Angiogenesis and mineralization during distraction osteogenesis.

Distraction osteogenesis is currently a standard method of bone lengthening. It is a viable method for the treatment of short extremities as well as extensive bone defects, because large amounts of bone can be regenerated in the distraction gap. Mechanical stimulation by distraction induces biological responses of skeletal regeneration that is accomplished by a cascade of biologic processes that may include differentiation of pluripotential tissue, angiogenesis, mineralization, and remodeling. There are complex interactions between bone-forming osteoblasts and other cells present within the bone microenvironment, particularly vascular endothelial cells that may be pivotal members of a complex interactive communication network in bone. Regenerate bone forms by three modes of ossification, which include intramembranous, enchondral, and transchondroid ossifications, although intramembraneous bone formation is the predominant mechanism of ossification. In this review we discussed the coupling between angiogenesis and mineralization, the biological and mechanical factors affecting them, the cellular and molecular events occurring during distraction osteogenesis, and the emerging modalities to accelerate regenerate bone healing and remodeling.

Animals↗

PubMed2002

Mechanical effects on skeletal growth.

The growth (i.e. increase of external dimensions) of long bones and vertebrae occurs longitudinally by endochondral ossification at the growth plates, and radially by apposition of bone at the periosteum. It is thought that mechanical loading influences the rate of longitudinal growth. The 'Hueter-Volkmann Law' proposes that growth is retarded by increased mechanical compression, and accelerated by reduced loading in comparison with normal values. The present understanding of this mechanism of bone growth modulation comes from a combination of clinical observation (where altered loading and growth is implicated in some skeletal deformities) and animal experiments in which growth plates of growing animals have been loaded. The gross effect of growth modulation has been demonstrated qualitatively and semi-quantitatively. Sustained compression of physiological magnitude inhibits growth by 40% or more. Distraction increases growth rate by a much smaller amount. Experimental studies are underway to determine how data from animal studies can be scaled to other growth plates. Variables include: differing sizes of growth plate, different anatomical locations, different species and variable growth rate at different stages of skeletal maturity. The two major determinants of longitudinal growth are the rate of chondrocytic proliferation and the amount of chondrocytic enlargement (hypertrophy) in the growth direction. It is largely unknown what are the relative changes in these key variables in mechanically modulated growth, and what are the signaling pathways that produce these changes.

Journal Article↗