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Degeneration of Articular Cartilage in Digit and Knee Joints\nOutside of the ankle region, little or no evidence was seen for expansion of the synovial membrane. Instead, mutant mice showed histological signs of osteoarthritis, such as fibrillation of the articular surface (Figure 7). As previously seen in 1- and 2-wk-old animals, Safranin O staining and Agg and Col10 expression were all reduced in mutant articular regions of the forefeet and hindfeet by 7 wk of age, and the beginning signs of cartilage loss were observed (unpublished data). By 9 mo of age, many regions of articular cartilage were completely missing or extremely fibrillated, leaving regions of exposed bone on the surface (Figure 7A–7D). No alterations were seen in the expression of Osteocalcin, Col1a1, or matrix metalloprotease-13 at either 7 wk or 9 mo.\nFigure 7 Loss of Bmpr1a Signaling Leads to Articular Cartilage Fibrillation and Degeneration in Digits and Knees of Aging Mice\n(A–D) Near adjacent sections of metatarsal-phalangeal joints from 9 month old mice. Articular cartilage of controls is complete and stains strongly with Safranin O (A, orange stain). In contrast, articular cells of mutants are severely fibrillated or absent with much reduced staining of Safranin O (C, arrowheads). LACZ expression confirms Cre-mediated recombination has occurred in articular cells (B and D).\n(E–P) Sagittal sections through knee joints of 7-wk- (E–J) or 9-mo-old animals (K–P); fe, femur; ti, tibia; gp, growth plate. Seven weeks after birth, the height of the tibial epiphysis is reduced in mutants (E and H, bars), and their articular layer stains poorly with Safranin O, is fibrillated, and is strikingly thinner (F and I, black arrowhead, and brackets). Near adjacent sections with LACZ staining confirm Cre-mediated recombination has occurred in articular cells (G and J). Note that in mutants, LACZ is absent in cells adjacent to those that do stain with Safranin O, suggesting Bmpr1a may act cell autonomously (I and J, white arrowheads). At 9 mo old, the mutant tibial epiphysis is extremely thin (K and N, bars), and the articular layer is completely absent, leaving bone to rub directly on bone (L and O, bracket). LACZ staining shows Cre-mediated recombination occurred in articular cells of controls (M) and in some remaining skeletal tissue of mutants (P). Also note aberrantly formed meniscal cartilage in mutants (E, H, K, and N, arrows), and increased sclerosis in mutant epiphyses (E, H, K, and N, asterisks).\n(A and K) Scale bar = 50 μm; (I) scale bar = 300 μm. The major weight-bearing joint of the hindlimb, the knee, showed changes that closely paralleled that seen in the foot joints. All markers of cartilage matrix looked similar to controls at E16.5, suggesting that early stages of joint formation were not disrupted (unpublished data). By postnatal day 7, Safranin O staining and Col2a1 and Agg expression were clearly reduced in the mutant, despite continued expression of Sox9 (unpublished data). The overall shape of mutant knee skeletal elements appeared similar to controls, although the fibrocartilaginous meniscus that resides between the femur and tibia appeared much less dense in mutants at E16.5. Some cartilage formed in the meniscus region, but the size of these elements was greatly reduced and contained abundant cells with fibrous, noncartilaginous appearance (unpublished data). This reduction of the meniscus can also be seen in sections from 7-wk- and 9-mo-old animals (Figure 7E, 7H, 7K, and 7N, arrows).\nAt 7 wk of age the normally domed tibial epiphysis was flattened and depressed in the knees of mutant animals, markedly reducing the distance between the growth plate and articular surface (Figure 7E and 7H, vertical bar). Articular cartilage was also thinner than in control animals, showed nearly complete absence of Safranin O staining, and was either acellular or beginning to fibrillate in many regions (Figure 7F and 7I). The few large Safranin O-stained cells still apparent in mutant articular regions appeared to correspond in position to rare LACZ-negative cells in adjacent sections, suggesting that Bmpr1a is required cell-autonomously in articular cartilage (Figure 7I and 7J, white arrowheads). By 9 mo, large areas of mutant knees were devoid of articular cells, and the bones of the femur and tibia appeared to rub directly against each other. Furthermore, the epiphysis of the tibia was extremely depressed, to the point that growth plate cartilage was almost exposed through the surface of the bone (Figure 7K, 7L, 7N, and 7O). In addition, mutants at 7 wk and 9 mo showed subchondral sclerosis, especially in the epiphysis of the femur (Figure 7E, 7H, 7K, and 7N, asterisks). While subchondral sclerosis is commonly seen in cases of osteoarthritis, it is unclear in this case whether the sclerosis is mainly a response of bone formation to compensate for decreased articular cartilage, or whether it is the effect of loss of Bmpr1a signaling in some LACZ-positive cells that are also observed in these regions (unpublished data).\nThe histological signs of joint arthritis were accompanied by functional impairments in both grasping ability and range of motion in mutant animals. Gdf5-Cre/Bmpr1afloxP mutant animals showed a highly significantly reduced ability to grasp and remain suspended on a slender rod (mean suspension time: controls 38 ± 6 s, n = 39; mutants 6 ± 3 s, n = 11; p \u003c 0.0001). Mutant mice also showed a clear decrease in the maximum range of mobility of two different joints in the digits, as assayed by passive manipulation (MT/P1 joint: controls 100 ± 0°, n = 26; mutants 82 ± 3°, n = 8; p \u003c 0.0003; P1/P2 joint: controls 152 ± 1°, n = 23; mutants 140 ± 5°, n = 6; p \u003c 0.05). The structural, histological, marker gene expression, and functional changes in mutant mice demonstrate that BMPR1A is required for normal postnatal maintenance of articular cartilage."}

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Degeneration of Articular Cartilage in Digit and Knee Joints\nOutside of the ankle region, little or no evidence was seen for expansion of the synovial membrane. Instead, mutant mice showed histological signs of osteoarthritis, such as fibrillation of the articular surface (Figure 7). As previously seen in 1- and 2-wk-old animals, Safranin O staining and Agg and Col10 expression were all reduced in mutant articular regions of the forefeet and hindfeet by 7 wk of age, and the beginning signs of cartilage loss were observed (unpublished data). By 9 mo of age, many regions of articular cartilage were completely missing or extremely fibrillated, leaving regions of exposed bone on the surface (Figure 7A–7D). No alterations were seen in the expression of Osteocalcin, Col1a1, or matrix metalloprotease-13 at either 7 wk or 9 mo.\nFigure 7 Loss of Bmpr1a Signaling Leads to Articular Cartilage Fibrillation and Degeneration in Digits and Knees of Aging Mice\n(A–D) Near adjacent sections of metatarsal-phalangeal joints from 9 month old mice. Articular cartilage of controls is complete and stains strongly with Safranin O (A, orange stain). In contrast, articular cells of mutants are severely fibrillated or absent with much reduced staining of Safranin O (C, arrowheads). LACZ expression confirms Cre-mediated recombination has occurred in articular cells (B and D).\n(E–P) Sagittal sections through knee joints of 7-wk- (E–J) or 9-mo-old animals (K–P); fe, femur; ti, tibia; gp, growth plate. Seven weeks after birth, the height of the tibial epiphysis is reduced in mutants (E and H, bars), and their articular layer stains poorly with Safranin O, is fibrillated, and is strikingly thinner (F and I, black arrowhead, and brackets). Near adjacent sections with LACZ staining confirm Cre-mediated recombination has occurred in articular cells (G and J). Note that in mutants, LACZ is absent in cells adjacent to those that do stain with Safranin O, suggesting Bmpr1a may act cell autonomously (I and J, white arrowheads). At 9 mo old, the mutant tibial epiphysis is extremely thin (K and N, bars), and the articular layer is completely absent, leaving bone to rub directly on bone (L and O, bracket). LACZ staining shows Cre-mediated recombination occurred in articular cells of controls (M) and in some remaining skeletal tissue of mutants (P). Also note aberrantly formed meniscal cartilage in mutants (E, H, K, and N, arrows), and increased sclerosis in mutant epiphyses (E, H, K, and N, asterisks).\n(A and K) Scale bar = 50 μm; (I) scale bar = 300 μm. The major weight-bearing joint of the hindlimb, the knee, showed changes that closely paralleled that seen in the foot joints. All markers of cartilage matrix looked similar to controls at E16.5, suggesting that early stages of joint formation were not disrupted (unpublished data). By postnatal day 7, Safranin O staining and Col2a1 and Agg expression were clearly reduced in the mutant, despite continued expression of Sox9 (unpublished data). The overall shape of mutant knee skeletal elements appeared similar to controls, although the fibrocartilaginous meniscus that resides between the femur and tibia appeared much less dense in mutants at E16.5. Some cartilage formed in the meniscus region, but the size of these elements was greatly reduced and contained abundant cells with fibrous, noncartilaginous appearance (unpublished data). This reduction of the meniscus can also be seen in sections from 7-wk- and 9-mo-old animals (Figure 7E, 7H, 7K, and 7N, arrows).\nAt 7 wk of age the normally domed tibial epiphysis was flattened and depressed in the knees of mutant animals, markedly reducing the distance between the growth plate and articular surface (Figure 7E and 7H, vertical bar). Articular cartilage was also thinner than in control animals, showed nearly complete absence of Safranin O staining, and was either acellular or beginning to fibrillate in many regions (Figure 7F and 7I). The few large Safranin O-stained cells still apparent in mutant articular regions appeared to correspond in position to rare LACZ-negative cells in adjacent sections, suggesting that Bmpr1a is required cell-autonomously in articular cartilage (Figure 7I and 7J, white arrowheads). By 9 mo, large areas of mutant knees were devoid of articular cells, and the bones of the femur and tibia appeared to rub directly against each other. Furthermore, the epiphysis of the tibia was extremely depressed, to the point that growth plate cartilage was almost exposed through the surface of the bone (Figure 7K, 7L, 7N, and 7O). In addition, mutants at 7 wk and 9 mo showed subchondral sclerosis, especially in the epiphysis of the femur (Figure 7E, 7H, 7K, and 7N, asterisks). While subchondral sclerosis is commonly seen in cases of osteoarthritis, it is unclear in this case whether the sclerosis is mainly a response of bone formation to compensate for decreased articular cartilage, or whether it is the effect of loss of Bmpr1a signaling in some LACZ-positive cells that are also observed in these regions (unpublished data).\nThe histological signs of joint arthritis were accompanied by functional impairments in both grasping ability and range of motion in mutant animals. Gdf5-Cre/Bmpr1afloxP mutant animals showed a highly significantly reduced ability to grasp and remain suspended on a slender rod (mean suspension time: controls 38 ± 6 s, n = 39; mutants 6 ± 3 s, n = 11; p \u003c 0.0001). Mutant mice also showed a clear decrease in the maximum range of mobility of two different joints in the digits, as assayed by passive manipulation (MT/P1 joint: controls 100 ± 0°, n = 26; mutants 82 ± 3°, n = 8; p \u003c 0.0003; P1/P2 joint: controls 152 ± 1°, n = 23; mutants 140 ± 5°, n = 6; p \u003c 0.05). The structural, histological, marker gene expression, and functional changes in mutant mice demonstrate that BMPR1A is required for normal postnatal maintenance of articular cartilage."}

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Degeneration of Articular Cartilage in Digit and Knee Joints\nOutside of the ankle region, little or no evidence was seen for expansion of the synovial membrane. Instead, mutant mice showed histological signs of osteoarthritis, such as fibrillation of the articular surface (Figure 7). As previously seen in 1- and 2-wk-old animals, Safranin O staining and Agg and Col10 expression were all reduced in mutant articular regions of the forefeet and hindfeet by 7 wk of age, and the beginning signs of cartilage loss were observed (unpublished data). By 9 mo of age, many regions of articular cartilage were completely missing or extremely fibrillated, leaving regions of exposed bone on the surface (Figure 7A–7D). No alterations were seen in the expression of Osteocalcin, Col1a1, or matrix metalloprotease-13 at either 7 wk or 9 mo.\nFigure 7 Loss of Bmpr1a Signaling Leads to Articular Cartilage Fibrillation and Degeneration in Digits and Knees of Aging Mice\n(A–D) Near adjacent sections of metatarsal-phalangeal joints from 9 month old mice. Articular cartilage of controls is complete and stains strongly with Safranin O (A, orange stain). In contrast, articular cells of mutants are severely fibrillated or absent with much reduced staining of Safranin O (C, arrowheads). LACZ expression confirms Cre-mediated recombination has occurred in articular cells (B and D).\n(E–P) Sagittal sections through knee joints of 7-wk- (E–J) or 9-mo-old animals (K–P); fe, femur; ti, tibia; gp, growth plate. Seven weeks after birth, the height of the tibial epiphysis is reduced in mutants (E and H, bars), and their articular layer stains poorly with Safranin O, is fibrillated, and is strikingly thinner (F and I, black arrowhead, and brackets). Near adjacent sections with LACZ staining confirm Cre-mediated recombination has occurred in articular cells (G and J). Note that in mutants, LACZ is absent in cells adjacent to those that do stain with Safranin O, suggesting Bmpr1a may act cell autonomously (I and J, white arrowheads). At 9 mo old, the mutant tibial epiphysis is extremely thin (K and N, bars), and the articular layer is completely absent, leaving bone to rub directly on bone (L and O, bracket). LACZ staining shows Cre-mediated recombination occurred in articular cells of controls (M) and in some remaining skeletal tissue of mutants (P). Also note aberrantly formed meniscal cartilage in mutants (E, H, K, and N, arrows), and increased sclerosis in mutant epiphyses (E, H, K, and N, asterisks).\n(A and K) Scale bar = 50 μm; (I) scale bar = 300 μm. The major weight-bearing joint of the hindlimb, the knee, showed changes that closely paralleled that seen in the foot joints. All markers of cartilage matrix looked similar to controls at E16.5, suggesting that early stages of joint formation were not disrupted (unpublished data). By postnatal day 7, Safranin O staining and Col2a1 and Agg expression were clearly reduced in the mutant, despite continued expression of Sox9 (unpublished data). The overall shape of mutant knee skeletal elements appeared similar to controls, although the fibrocartilaginous meniscus that resides between the femur and tibia appeared much less dense in mutants at E16.5. Some cartilage formed in the meniscus region, but the size of these elements was greatly reduced and contained abundant cells with fibrous, noncartilaginous appearance (unpublished data). This reduction of the meniscus can also be seen in sections from 7-wk- and 9-mo-old animals (Figure 7E, 7H, 7K, and 7N, arrows).\nAt 7 wk of age the normally domed tibial epiphysis was flattened and depressed in the knees of mutant animals, markedly reducing the distance between the growth plate and articular surface (Figure 7E and 7H, vertical bar). Articular cartilage was also thinner than in control animals, showed nearly complete absence of Safranin O staining, and was either acellular or beginning to fibrillate in many regions (Figure 7F and 7I). The few large Safranin O-stained cells still apparent in mutant articular regions appeared to correspond in position to rare LACZ-negative cells in adjacent sections, suggesting that Bmpr1a is required cell-autonomously in articular cartilage (Figure 7I and 7J, white arrowheads). By 9 mo, large areas of mutant knees were devoid of articular cells, and the bones of the femur and tibia appeared to rub directly against each other. Furthermore, the epiphysis of the tibia was extremely depressed, to the point that growth plate cartilage was almost exposed through the surface of the bone (Figure 7K, 7L, 7N, and 7O). In addition, mutants at 7 wk and 9 mo showed subchondral sclerosis, especially in the epiphysis of the femur (Figure 7E, 7H, 7K, and 7N, asterisks). While subchondral sclerosis is commonly seen in cases of osteoarthritis, it is unclear in this case whether the sclerosis is mainly a response of bone formation to compensate for decreased articular cartilage, or whether it is the effect of loss of Bmpr1a signaling in some LACZ-positive cells that are also observed in these regions (unpublished data).\nThe histological signs of joint arthritis were accompanied by functional impairments in both grasping ability and range of motion in mutant animals. Gdf5-Cre/Bmpr1afloxP mutant animals showed a highly significantly reduced ability to grasp and remain suspended on a slender rod (mean suspension time: controls 38 ± 6 s, n = 39; mutants 6 ± 3 s, n = 11; p \u003c 0.0001). Mutant mice also showed a clear decrease in the maximum range of mobility of two different joints in the digits, as assayed by passive manipulation (MT/P1 joint: controls 100 ± 0°, n = 26; mutants 82 ± 3°, n = 8; p \u003c 0.0003; P1/P2 joint: controls 152 ± 1°, n = 23; mutants 140 ± 5°, n = 6; p \u003c 0.05). The structural, histological, marker gene expression, and functional changes in mutant mice demonstrate that BMPR1A is required for normal postnatal maintenance of articular cartilage."}