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earliest step of limb development in which BMP signaling has been implicated is the establishment of the anterior-posterior limb axis. Differences in anterior-posterior pattern are instructed as a graded response to Sonic Hedghog (SHH) signaling emanating from the posterior margin of the limb bud [5]. It has remained controversial, however, whether this response is direct or indirect. If indeed the long-range effects of SHH are indirectly mediated by local production of secondary signals, the leading candidates have been two members of the BMP family, BMP2 and BMP7. Both are expressed in a broader domain than SHH in the early posterior limb bud mesenchyme [6,7], although BMP7 also has a second, weaker domain of expression in the anterior limb bud mesenchyme. BMP2 [8] and BMP7 [7] can be induced by ectopic SHH and their expression is greatly diminished in the absence of SHH activity [9]. BMP2 and BMP7 are thus secondary signals produced in response to SHH activity. Moreover, BMP signaling has a weak ability to posteriorly polarize the limb in ectopic grafting experiments [10], an activity enhanced by prior low-level exposure to SHH [11]. It remains unclear, however, whether BMP2 and BMP7 activity is required endogenously for anterior-posterior limb patterning by SHH. Bmp2 mutant embryos die too early to assess their limb phenotypes. A targeted deletion of Bmp7 has been made, and Bmp7-deficient embryos display hindlimb polydactyly with incomplete penetrance but otherwise phenotypically normal limbs [12,13]. Nonetheless, Bmp7 knockout mice do not show any defect in limb polarity. However, a redundant function in anterior-posterior patterning with BMP2 remains a possibility. In addition to BMP2 and BMP7, a third member of this family that is closely related to BMP2, BMP4, is also expressed in the early limb bud. Like BMP7, it is expressed in both the anterior and posterior margins of the limb bud mesenchyme [4,14]; however, it does not appear to be induced by SHH signaling, nor does its expression change in SHH-deficient limb buds. Thus, BMP4 is not a candidate for a secondary signal downstream of SHH in early patterning. However, all three of these molecules, BMP2, BMP4, and BMP7, have been suggested to act in a second distinct phase of limb patterning, when digit identities are established downstream of earlier patterning events. In the vertebrate limb, each digit can be uniquely identified based on its size, length, number of phalanges, and location within the autopod. As a consequence of the initial establishment of anterior-posterior positioned information within the limb by SHH and/or BMP signaling, the interdigital mesenchyme of the hand plate becomes specified in a graded manner. Grafting and extirpation experiments have shown that it is this polarized interdigital tissue which directs digit morphology [15]. BMP2, BMP4, and BMP7 are all expressed in the interdigital mesenchyme [4,14], and experiments inhibiting BMP signaling in this tissue suggested that differential levels of BMPs in the interdigital mesenchyme might be the relevant factor directing digit morphology [15]. As noted above, BMP7-deficient embryos produce mostly normal limbs, and mice harboring a conditional deletion of Bmp4 in the limb bud mesenchyme also do not show evidence of anterior digit transformation [16], as would have been predicted if BMP4 acted as an interdigital regulator of digit morphology. In the light of these observations, it would seem unlikely that quantitative differences in levels of interdigital BMP signal are responsible for establishing digit identity, although redundancy between BMP2, BMP4, and BMP7 in this role remains a possibility."}

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earliest step of limb development in which BMP signaling has been implicated is the establishment of the anterior-posterior limb axis. Differences in anterior-posterior pattern are instructed as a graded response to Sonic Hedghog (SHH) signaling emanating from the posterior margin of the limb bud [5]. It has remained controversial, however, whether this response is direct or indirect. If indeed the long-range effects of SHH are indirectly mediated by local production of secondary signals, the leading candidates have been two members of the BMP family, BMP2 and BMP7. Both are expressed in a broader domain than SHH in the early posterior limb bud mesenchyme [6,7], although BMP7 also has a second, weaker domain of expression in the anterior limb bud mesenchyme. BMP2 [8] and BMP7 [7] can be induced by ectopic SHH and their expression is greatly diminished in the absence of SHH activity [9]. BMP2 and BMP7 are thus secondary signals produced in response to SHH activity. Moreover, BMP signaling has a weak ability to posteriorly polarize the limb in ectopic grafting experiments [10], an activity enhanced by prior low-level exposure to SHH [11]. It remains unclear, however, whether BMP2 and BMP7 activity is required endogenously for anterior-posterior limb patterning by SHH. Bmp2 mutant embryos die too early to assess their limb phenotypes. A targeted deletion of Bmp7 has been made, and Bmp7-deficient embryos display hindlimb polydactyly with incomplete penetrance but otherwise phenotypically normal limbs [12,13]. Nonetheless, Bmp7 knockout mice do not show any defect in limb polarity. However, a redundant function in anterior-posterior patterning with BMP2 remains a possibility. In addition to BMP2 and BMP7, a third member of this family that is closely related to BMP2, BMP4, is also expressed in the early limb bud. Like BMP7, it is expressed in both the anterior and posterior margins of the limb bud mesenchyme [4,14]; however, it does not appear to be induced by SHH signaling, nor does its expression change in SHH-deficient limb buds. Thus, BMP4 is not a candidate for a secondary signal downstream of SHH in early patterning. However, all three of these molecules, BMP2, BMP4, and BMP7, have been suggested to act in a second distinct phase of limb patterning, when digit identities are established downstream of earlier patterning events. In the vertebrate limb, each digit can be uniquely identified based on its size, length, number of phalanges, and location within the autopod. As a consequence of the initial establishment of anterior-posterior positioned information within the limb by SHH and/or BMP signaling, the interdigital mesenchyme of the hand plate becomes specified in a graded manner. Grafting and extirpation experiments have shown that it is this polarized interdigital tissue which directs digit morphology [15]. BMP2, BMP4, and BMP7 are all expressed in the interdigital mesenchyme [4,14], and experiments inhibiting BMP signaling in this tissue suggested that differential levels of BMPs in the interdigital mesenchyme might be the relevant factor directing digit morphology [15]. As noted above, BMP7-deficient embryos produce mostly normal limbs, and mice harboring a conditional deletion of Bmp4 in the limb bud mesenchyme also do not show evidence of anterior digit transformation [16], as would have been predicted if BMP4 acted as an interdigital regulator of digit morphology. In the light of these observations, it would seem unlikely that quantitative differences in levels of interdigital BMP signal are responsible for establishing digit identity, although redundancy between BMP2, BMP4, and BMP7 in this role remains a possibility."}

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Differences in anterior-posterior pattern are instructed as a graded response to Sonic Hedghog (SHH) signaling emanating from the posterior margin of the limb bud [5]. It has remained controversial, however, whether this response is direct or indirect. If indeed the long-range effects of SHH are indirectly mediated by local production of secondary signals, the leading candidates have been two members of the BMP family, BMP2 and BMP7. Both are expressed in a broader domain than SHH in the early posterior limb bud mesenchyme [6,7], although BMP7 also has a second, weaker domain of expression in the anterior limb bud mesenchyme. BMP2 [8] and BMP7 [7] can be induced by ectopic SHH and their expression is greatly diminished in the absence of SHH activity [9]. BMP2 and BMP7 are thus secondary signals produced in response to SHH activity. Moreover, BMP signaling has a weak ability to posteriorly polarize the limb in ectopic grafting experiments [10], an activity enhanced by prior low-level exposure to SHH [11]. It remains unclear, however, whether BMP2 and BMP7 activity is required endogenously for anterior-posterior limb patterning by SHH. Bmp2 mutant embryos die too early to assess their limb phenotypes. A targeted deletion of Bmp7 has been made, and Bmp7-deficient embryos display hindlimb polydactyly with incomplete penetrance but otherwise phenotypically normal limbs [12,13]. Nonetheless, Bmp7 knockout mice do not show any defect in limb polarity. However, a redundant function in anterior-posterior patterning with BMP2 remains a possibility. In addition to BMP2 and BMP7, a third member of this family that is closely related to BMP2, BMP4, is also expressed in the early limb bud. Like BMP7, it is expressed in both the anterior and posterior margins of the limb bud mesenchyme [4,14]; however, it does not appear to be induced by SHH signaling, nor does its expression change in SHH-deficient limb buds. Thus, BMP4 is not a candidate for a secondary signal downstream of SHH in early patterning. However, all three of these molecules, BMP2, BMP4, and BMP7, have been suggested to act in a second distinct phase of limb patterning, when digit identities are established downstream of earlier patterning events. In the vertebrate limb, each digit can be uniquely identified based on its size, length, number of phalanges, and location within the autopod. As a consequence of the initial establishment of anterior-posterior positioned information within the limb by SHH and/or BMP signaling, the interdigital mesenchyme of the hand plate becomes specified in a graded manner. Grafting and extirpation experiments have shown that it is this polarized interdigital tissue which directs digit morphology [15]. BMP2, BMP4, and BMP7 are all expressed in the interdigital mesenchyme [4,14], and experiments inhibiting BMP signaling in this tissue suggested that differential levels of BMPs in the interdigital mesenchyme might be the relevant factor directing digit morphology [15]. As noted above, BMP7-deficient embryos produce mostly normal limbs, and mice harboring a conditional deletion of Bmp4 in the limb bud mesenchyme also do not show evidence of anterior digit transformation [16], as would have been predicted if BMP4 acted as an interdigital regulator of digit morphology. In the light of these observations, it would seem unlikely that quantitative differences in levels of interdigital BMP signal are responsible for establishing digit identity, although redundancy between BMP2, BMP4, and BMP7 in this role remains a possibility."}

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earliest step of limb development in which BMP signaling has been implicated is the establishment of the anterior-posterior limb axis. Differences in anterior-posterior pattern are instructed as a graded response to Sonic Hedghog (SHH) signaling emanating from the posterior margin of the limb bud [5]. It has remained controversial, however, whether this response is direct or indirect. If indeed the long-range effects of SHH are indirectly mediated by local production of secondary signals, the leading candidates have been two members of the BMP family, BMP2 and BMP7. Both are expressed in a broader domain than SHH in the early posterior limb bud mesenchyme [6,7], although BMP7 also has a second, weaker domain of expression in the anterior limb bud mesenchyme. BMP2 [8] and BMP7 [7] can be induced by ectopic SHH and their expression is greatly diminished in the absence of SHH activity [9]. BMP2 and BMP7 are thus secondary signals produced in response to SHH activity. Moreover, BMP signaling has a weak ability to posteriorly polarize the limb in ectopic grafting experiments [10], an activity enhanced by prior low-level exposure to SHH [11]. It remains unclear, however, whether BMP2 and BMP7 activity is required endogenously for anterior-posterior limb patterning by SHH. Bmp2 mutant embryos die too early to assess their limb phenotypes. A targeted deletion of Bmp7 has been made, and Bmp7-deficient embryos display hindlimb polydactyly with incomplete penetrance but otherwise phenotypically normal limbs [12,13]. Nonetheless, Bmp7 knockout mice do not show any defect in limb polarity. However, a redundant function in anterior-posterior patterning with BMP2 remains a possibility. In addition to BMP2 and BMP7, a third member of this family that is closely related to BMP2, BMP4, is also expressed in the early limb bud. Like BMP7, it is expressed in both the anterior and posterior margins of the limb bud mesenchyme [4,14]; however, it does not appear to be induced by SHH signaling, nor does its expression change in SHH-deficient limb buds. Thus, BMP4 is not a candidate for a secondary signal downstream of SHH in early patterning. However, all three of these molecules, BMP2, BMP4, and BMP7, have been suggested to act in a second distinct phase of limb patterning, when digit identities are established downstream of earlier patterning events. In the vertebrate limb, each digit can be uniquely identified based on its size, length, number of phalanges, and location within the autopod. As a consequence of the initial establishment of anterior-posterior positioned information within the limb by SHH and/or BMP signaling, the interdigital mesenchyme of the hand plate becomes specified in a graded manner. Grafting and extirpation experiments have shown that it is this polarized interdigital tissue which directs digit morphology [15]. BMP2, BMP4, and BMP7 are all expressed in the interdigital mesenchyme [4,14], and experiments inhibiting BMP signaling in this tissue suggested that differential levels of BMPs in the interdigital mesenchyme might be the relevant factor directing digit morphology [15]. As noted above, BMP7-deficient embryos produce mostly normal limbs, and mice harboring a conditional deletion of Bmp4 in the limb bud mesenchyme also do not show evidence of anterior digit transformation [16], as would have been predicted if BMP4 acted as an interdigital regulator of digit morphology. In the light of these observations, it would seem unlikely that quantitative differences in levels of interdigital BMP signal are responsible for establishing digit identity, although redundancy between BMP2, BMP4, and BMP7 in this role remains a possibility."}