Abstract: The concept of functional adapatation of articular cartilage during maturation has emerged from earlier biochemical research. However, articular cartilage has principally a biomechanical function governed by joint loading. Objective: To verify whether the concept of functional adaptation can be confirmed by direct measurement of biomechanical properties of cartilage. Objective: Fetuses have homogeneous (i.e. site-independent) cartilage with regard to biomechanical properties. During growth and development to maturity, the biomechanical characteristics adapt according to functional (loading) demands, leading to distinct, site-dependent biomechanical heterogeneity of articular cartilage. Methods: Osteochondral plugs were drilled out of the surface at 2 differently loaded sites (Site 1: intermittent impact-loading during locomotion, Site 2: low-level constant loading during weightbearing) of the proximal articular cartilage surface of the proximal phalanx in the forelimb from stillborn foals (n = 8), horses of age 5 (n = 9) and 18 months (n = 9) and mature horses (n = 13). Cartilage thickness was measured using ultrasonic, optical and needle-probe techniques. The osteochondral samples were biomechanically tested in indentation geometry. Young's modulus at equilibrium, dynamic modulus at 1 Hz and the ratios of these moduli values between Sites 1 and 2 were calculated. Age and site effects were evaluated statistically using ANOVA tests. The level of significance was set at P<0.05. Results: Fetal cartilage was significantly thicker compared to the other ages with no further age-dependent differences in cartilage thickness from age 5 months onwards. Young's modulus stayed constant at Site 1, whereas at Site 2 there was a gradual, statistically significant increase in modulus during maturation. Values of dynamic modulus at both Sites 1 and 2 were significantly higher in the fetus and decreased after birth. Values for both moduli were significantly different between Sites 1 and 2 from age 18 months onwards. The ratio of values between Sites 1 and 2 for Young's modulus and dynamic modulus showed a gradual decrease from approximately 1.0 at birth to 0.5-0.6 in the mature horse. At age 18 months, all values were comparable to those in the mature horse. Conclusions: In line with the concept of functional adaptation, the neonate is born with biomechanically 'blank' or homogeneous cartilage. Functional adaptation of biomechanical properties takes place early in life, resulting in cartilage with a distinct heterogeneity in functional characteristics. At age 18 months, functional adaptation, as assessed by the biomechanical characteristics, has progressed to a level comparable to the mature horse and, after this age, no major adaptations seem to occur. Conclusions: Throughout life, different areas of articular cartilage are subjected to different types of loading. Differences in loading can adequately be met only when the tissue is biomechanically adapted to withstand these different loading conditions without injury. This process of functional adaptation starts immediately after birth and is completed well before maturity. This makes the factor of loading at a young age a crucial variable, and emphasises the necessity to optimise joint loading during early life in order to create an optimal biomechanical quality of articular cartilage, which may well turn out to be the best prevention for joint injury later in life.
The Equine Research Bank provides access to a large database of publicly available scientific literature. Inclusion in the Research Bank does not imply endorsement of study methods or findings by Mad Barn.
This research summary has been generated with artificial intelligence and may contain errors and omissions. Refer to the original study to confirm details provided. Submit correction.
The research looks into how articular cartilage, which plays a key role in joint function, adapts to different loading conditions from birth to maturity, suggesting that this process completes well before maturity and could be instrumental in preventing joint injuries later in life.
Study Design and Methodology
The study’s aim was to confirm the concept of functional adaptation of articular cartilage during maturation by directly measuring its biomechanical properties.
The researchers hypothesized that fetuses have homogeneous cartilage, meaning its properties are consistent across the site, and that these properties adapt to functional demands (loading conditions) as the organism matures, leading to distinct, site-dependent biomechanical attributes.
To test this, osteochondral plugs were drilled from the surface at two sites of different loading. One experienced intermittent loading during locomotion and the other low-level consistent loading during weight-bearing, from the proximal phalanx in the forelimb of stillborn foals, horses aged 5 and 18 months, and adult horses.
Cartilage thickness measurements were made using ultrasonic, optical, and needle-probe techniques, and the samples underwent biomechanical testing in indentation geometry. Young’s modulus at equilibrium, dynamic modulus at 1 Hz, and ratios between the two sites were calculated.
Findings and Interpretation
Fetal cartilage was found to be significantly thicker compared with other ages, with no further age-dependent differences in thickness from 5 months onwards.
Young’s modulus, a measure of stiffness, remained constant at Site 1, but at Site 2 there was a gradual, statistically significant increase during maturation, suggesting site-specific adaptations.
Values for both Young’s and dynamic moduli showed significant differences between the two sites from 18 months onwards; the ratio between the two sites decreased from around 1.0 at birth to 0.5-0.6 in adult horses.
The study concluded that the neonate is born with biomechanically ‘blank’ cartilage that adapts to functional demands early in life, completing by around 18 months in horses. After this age, no major adaptations appeared to occur.
Implications and Conclusion
The concept of functional adaptation suggests that articular cartilage is subjected to different types of loads throughout life. These different loading conditions can only be optimally countered when the cartilage is biomechanically adapted to resist them without injury.
The study stresses the importance of optimising joint loading during early life to achieve optimal biomechanical quality of cartilage. This process, which completes well before maturity, could prevent joint injuries later in life.
Overall, the research underlines the necessity of healthy loading conditions for the functional adaptation and health of joint cartilage.
Cite This Article
APA
Brommer H, Brama PA, Laasanen MS, Helminen HJ, van Weeren PR, Jurvelin JS.
(2005).
Functional adaptation of articular cartilage from birth to maturity under the influence of loading: a biomechanical analysis.
Equine Vet J, 37(2), 148-154.
https://doi.org/10.2746/0425164054223769
Department of Equine Sciences, Faculty of Veterinary Medicine, Utrecht University, Yalelaan 12, NL-3584 CM Utrecht, The Netherlands.
Brama, P A J
Laasanen, M S
Helminen, H J
van Weeren, P R
Jurvelin, J S
MeSH Terms
Adaptation, Physiological
Age Factors
Aging / physiology
Analysis of Variance
Animals
Animals, Newborn
Biomechanical Phenomena
Cadaver
Cartilage, Articular / anatomy & histology
Cartilage, Articular / growth & development
Cartilage, Articular / physiology
Fetus
Horses / anatomy & histology
Horses / growth & development
Horses / physiology
Weight-Bearing
Citations
This article has been cited 31 times.
Batool S, Hammami M, Mantebea H, Badar F, Xia Y. Location-Specific Study of Young Rabbit Femoral Cartilage by Quantitative µMRI and Polarized Light Microscopy. Cartilage 2022 Jan-Mar;13(1):19476035221085143.
Lee J, Jang S, Kwon J, Oh TI, Lee E. Comparative Evaluation of Synovial Multipotent Stem Cells and Meniscal Chondrocytes for Capability of Fibrocartilage Reconstruction. Cartilage 2021 Dec;13(2_suppl):980S-990S.
Gruber BL, Mienaltowski MJ, MacLeod JN, Schittny J, Kasper S, Flück M. Tenascin-C expression controls the maturation of articular cartilage in mice. BMC Res Notes 2020 Feb 17;13(1):78.
Vail DJ, Somoza RA, Caplan AI, Khalil AM. Transcriptome dynamics of long noncoding RNAs and transcription factors demarcate human neonatal, adult, and human mesenchymal stem cell-derived engineered cartilage. J Tissue Eng Regen Med 2020 Jan;14(1):29-44.
Jessop ZM, Zhang Y, Simoes IN, Al-Sabah A, Badiei N, Gazze SA, Francis L, Whitaker IS. Morphological and biomechanical characterization of immature and mature nasoseptal cartilage. Sci Rep 2019 Aug 28;9(1):12464.
Ren P, Niu H, Gong H, Zhang R, Fan Y. Morphological, biochemical and mechanical properties of articular cartilage and subchondral bone in rat tibial plateau are age related. J Anat 2018 Mar;232(3):457-471.
Chan DD, Cai L, Butz KD, Nauman EA, Dickerson DA, Jonkers I, Neu CP. Functional MRI can detect changes in intratissue strains in a full thickness and critical sized ovine cartilage defect model. J Biomech 2018 Jan 3;66:18-25.
Drobnic M, Perdisa F, Kon E, Cefalì F, Marcacci M, Filardo G. Implant strategy affects scaffold stability and integrity in cartilage treatment. Knee Surg Sports Traumatol Arthrosc 2018 Sep;26(9):2774-2783.
Al-Himdani S, Jessop ZM, Al-Sabah A, Combellack E, Ibrahim A, Doak SH, Hart AM, Archer CW, Thornton CA, Whitaker IS. Tissue-Engineered Solutions in Plastic and Reconstructive Surgery: Principles and Practice. Front Surg 2017;4:4.
Jessop ZM, Javed M, Otto IA, Combellack EJ, Morgan S, Breugem CC, Archer CW, Khan IM, Lineaweaver WC, Kon M, Malda J, Whitaker IS. Combining regenerative medicine strategies to provide durable reconstructive options: auricular cartilage tissue engineering. Stem Cell Res Ther 2016 Jan 28;7:19.
Chan DD, Cai L, Butz KD, Trippel SB, Nauman EA, Neu CP. In vivo articular cartilage deformation: noninvasive quantification of intratissue strain during joint contact in the human knee. Sci Rep 2016 Jan 11;6:19220.
Malda J, de Grauw JC, Benders KE, Kik MJ, van de Lest CH, Creemers LB, Dhert WJ, van Weeren PR. Of mice, men and elephants: the relation between articular cartilage thickness and body mass. PLoS One 2013;8(2):e57683.
Hamann N, Zaucke F, Dayakli M, Brüggemann GP, Niehoff A. Growth-related structural, biochemical, and mechanical properties of the functional bone-cartilage unit. J Anat 2013 Feb;222(2):248-59.
Chan EF, Harjanto R, Asahara H, Inoue N, Masuda K, Bugbee WD, Firestein GS, Hosalkar HS, Lotz MK, Sah RL. Structural and functional maturation of distal femoral cartilage and bone during postnatal development and growth in humans and mice. Orthop Clin North Am 2012 Apr;43(2):173-85, v.
Mahmoodian R, Leasure J, Philip P, Pleshko N, Capaldi F, Siegler S. Changes in mechanics and composition of human talar cartilage anlagen during fetal development. Osteoarthritis Cartilage 2011 Oct;19(10):1199-209.
van Turnhout MC, Schipper H, van Lagen B, Zuilhof H, Kranenbarg S, van Leeuwen JL. Postnatal development of depth-dependent collagen density in ovine articular cartilage. BMC Dev Biol 2010 Oct 22;10:108.
van Turnhout MC, Schipper H, Engel B, Buist W, Kranenbarg S, van Leeuwen JL. Postnatal development of collagen structure in ovine articular cartilage. BMC Dev Biol 2010 Jun 7;10:62.
Price C, Li W, Novotny JE, Wang L. An in-situ fluorescence-based optical extensometry system for imaging mechanically loaded bone. J Orthop Res 2010 Jun;28(6):805-11.
Księżarczyk MM, IJsseldijk LL, van Weeren PR, Levato R, Malda J. Age-dependent development and microarchitecture of the osteochondral unit of the humeral head in harbour porpoises (Phocoena phocoena). Sci Rep 2026 Feb 12;16(1).