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the present study, we examined the function of ADAM22 by generating mice that lacked the Adam22 gene. ADAM22-deficient mice exhibited severe ataxia and premature death. In contrast, heterozygous mutants were healthy, fertile and survived more than 1 year without obvious abnormalities. The cause of death in the homozygous mutants is not clear; however, it is likely to be due to multiple seizures. This is because convulsive seizures were occasionally observed in homozygotes, after which the suffered mice appeared exhausted and, in most cases, died on the next day. It is known that cortical dysplasia resulting from aberrant brain development causes seizure syndrome [22]. For example, mice lacking the neuron-specific activator of cyclin 5, p35, exhibit seizures and a severe neuronal migration defect [23]. Because ADAM22 protein is expressed on the cell surface and is likely to be involved in cell-cell or cell-matrix interactions, we hypothesized that depletion of ADAM22 would generate aberrant neuronal migration. Histochemical analysis was performed to verify this hypothesis. However, no marked abnormalities were observed in the mutant mouse brain, ruling out a critical role for ADAM22 in neuronal migration. Another possible explanation is that the premature death in first 2 weeks of life is caused by the dysfunction of the autonomic nervous system. Especially, nerves which control the breathing would be very important, because the respiration system undergoes significant maturation in the first 2–3 weeks after birth. For example, the myelin-deficient (MD) rat, which carries a point mutation in the proteolipid protein (PLP) gene, exhibits ataxia, tremor, dysmyelination and dies at approximately postnatal day 21. In MD rat, early death is caused by the dysfunction of the brain stem which is essential for autonomic control of respiration during hypoxia [24]. Further study is needed to verify the cause of death in ADAM22-deficient mice.\nAnother interesting phenotype observed in ADAM22-deficient mice was hypomyelinogenesis. Histopathological analysis revealed marked hypomyelination in the sciatic and trigeminal nerves. However, myelin formation in the spinal cord and the brain was normal. It was therefore evident that hypomyelinogenesis occurred specifically in the peripheral nerves in the knockout mice, suggesting Schwann cell dysfunction. In the mutant sciatic nerve, the density of pro-myelinating Schwann cells (S100-positive, MBP-negative) was markedly higher than that seen in heterozygotes. Higher pro-myelinating Schwann cell density and marked delay in myelin formation indicate that ADAM22 plays an important role in Schwann cell differentiation, but not in proliferation.\nInvestigations on why hypomyelinogenesis occurred in ADAM22-deficient mice centre on integrin. This is because several ADAMs have been shown to interact with integrins via disintegrin domains [9], and it has been suggested that alpha-6 beta-1 integrin on the Schwann cell plays an important role in myelin formation [25]. In addition, laminin-2 is a key player in peripheral myelin formation, because laminin-2 deficiency causes dysmyelination in mice and humans, and its receptors, alpha-6 beta-1 integrin and dystroglycan, are also shown to be deeply involved in myelin formation [26,27]. The conditional knockout studies of beta-1 integrin and laminin gamma-1 in Schwann cell suggested that both laminin-2 and integrins are essential for segregation of large bundles of axons in the early stage of myelin formation [25,28]. In contrast, the phenotype seen in the ADAM22-deficient mice was quite different: for example, unsorted bundles of axons were not seen in nerves and roots. We assume that ADAM22 partially modulates integrin-mediated signals or is involved at a later stage of myelin formation."}

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the present study, we examined the function of ADAM22 by generating mice that lacked the Adam22 gene. ADAM22-deficient mice exhibited severe ataxia and premature death. In contrast, heterozygous mutants were healthy, fertile and survived more than 1 year without obvious abnormalities. The cause of death in the homozygous mutants is not clear; however, it is likely to be due to multiple seizures. This is because convulsive seizures were occasionally observed in homozygotes, after which the suffered mice appeared exhausted and, in most cases, died on the next day. It is known that cortical dysplasia resulting from aberrant brain development causes seizure syndrome [22]. For example, mice lacking the neuron-specific activator of cyclin 5, p35, exhibit seizures and a severe neuronal migration defect [23]. Because ADAM22 protein is expressed on the cell surface and is likely to be involved in cell-cell or cell-matrix interactions, we hypothesized that depletion of ADAM22 would generate aberrant neuronal migration. Histochemical analysis was performed to verify this hypothesis. However, no marked abnormalities were observed in the mutant mouse brain, ruling out a critical role for ADAM22 in neuronal migration. Another possible explanation is that the premature death in first 2 weeks of life is caused by the dysfunction of the autonomic nervous system. Especially, nerves which control the breathing would be very important, because the respiration system undergoes significant maturation in the first 2–3 weeks after birth. For example, the myelin-deficient (MD) rat, which carries a point mutation in the proteolipid protein (PLP) gene, exhibits ataxia, tremor, dysmyelination and dies at approximately postnatal day 21. In MD rat, early death is caused by the dysfunction of the brain stem which is essential for autonomic control of respiration during hypoxia [24]. Further study is needed to verify the cause of death in ADAM22-deficient mice.\nAnother interesting phenotype observed in ADAM22-deficient mice was hypomyelinogenesis. Histopathological analysis revealed marked hypomyelination in the sciatic and trigeminal nerves. However, myelin formation in the spinal cord and the brain was normal. It was therefore evident that hypomyelinogenesis occurred specifically in the peripheral nerves in the knockout mice, suggesting Schwann cell dysfunction. In the mutant sciatic nerve, the density of pro-myelinating Schwann cells (S100-positive, MBP-negative) was markedly higher than that seen in heterozygotes. Higher pro-myelinating Schwann cell density and marked delay in myelin formation indicate that ADAM22 plays an important role in Schwann cell differentiation, but not in proliferation.\nInvestigations on why hypomyelinogenesis occurred in ADAM22-deficient mice centre on integrin. This is because several ADAMs have been shown to interact with integrins via disintegrin domains [9], and it has been suggested that alpha-6 beta-1 integrin on the Schwann cell plays an important role in myelin formation [25]. In addition, laminin-2 is a key player in peripheral myelin formation, because laminin-2 deficiency causes dysmyelination in mice and humans, and its receptors, alpha-6 beta-1 integrin and dystroglycan, are also shown to be deeply involved in myelin formation [26,27]. The conditional knockout studies of beta-1 integrin and laminin gamma-1 in Schwann cell suggested that both laminin-2 and integrins are essential for segregation of large bundles of axons in the early stage of myelin formation [25,28]. In contrast, the phenotype seen in the ADAM22-deficient mice was quite different: for example, unsorted bundles of axons were not seen in nerves and roots. We assume that ADAM22 partially modulates integrin-mediated signals or is involved at a later stage of myelin formation."}

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the present study, we examined the function of ADAM22 by generating mice that lacked the Adam22 gene. ADAM22-deficient mice exhibited severe ataxia and premature death. In contrast, heterozygous mutants were healthy, fertile and survived more than 1 year without obvious abnormalities. The cause of death in the homozygous mutants is not clear; however, it is likely to be due to multiple seizures. This is because convulsive seizures were occasionally observed in homozygotes, after which the suffered mice appeared exhausted and, in most cases, died on the next day. It is known that cortical dysplasia resulting from aberrant brain development causes seizure syndrome [22]. For example, mice lacking the neuron-specific activator of cyclin 5, p35, exhibit seizures and a severe neuronal migration defect [23]. Because ADAM22 protein is expressed on the cell surface and is likely to be involved in cell-cell or cell-matrix interactions, we hypothesized that depletion of ADAM22 would generate aberrant neuronal migration. Histochemical analysis was performed to verify this hypothesis. However, no marked abnormalities were observed in the mutant mouse brain, ruling out a critical role for ADAM22 in neuronal migration. Another possible explanation is that the premature death in first 2 weeks of life is caused by the dysfunction of the autonomic nervous system. Especially, nerves which control the breathing would be very important, because the respiration system undergoes significant maturation in the first 2–3 weeks after birth. For example, the myelin-deficient (MD) rat, which carries a point mutation in the proteolipid protein (PLP) gene, exhibits ataxia, tremor, dysmyelination and dies at approximately postnatal day 21. In MD rat, early death is caused by the dysfunction of the brain stem which is essential for autonomic control of respiration during hypoxia [24]. Further study is needed to verify the cause of death in ADAM22-deficient mice.\nAnother interesting phenotype observed in ADAM22-deficient mice was hypomyelinogenesis. Histopathological analysis revealed marked hypomyelination in the sciatic and trigeminal nerves. However, myelin formation in the spinal cord and the brain was normal. It was therefore evident that hypomyelinogenesis occurred specifically in the peripheral nerves in the knockout mice, suggesting Schwann cell dysfunction. In the mutant sciatic nerve, the density of pro-myelinating Schwann cells (S100-positive, MBP-negative) was markedly higher than that seen in heterozygotes. Higher pro-myelinating Schwann cell density and marked delay in myelin formation indicate that ADAM22 plays an important role in Schwann cell differentiation, but not in proliferation.\nInvestigations on why hypomyelinogenesis occurred in ADAM22-deficient mice centre on integrin. This is because several ADAMs have been shown to interact with integrins via disintegrin domains [9], and it has been suggested that alpha-6 beta-1 integrin on the Schwann cell plays an important role in myelin formation [25]. In addition, laminin-2 is a key player in peripheral myelin formation, because laminin-2 deficiency causes dysmyelination in mice and humans, and its receptors, alpha-6 beta-1 integrin and dystroglycan, are also shown to be deeply involved in myelin formation [26,27]. The conditional knockout studies of beta-1 integrin and laminin gamma-1 in Schwann cell suggested that both laminin-2 and integrins are essential for segregation of large bundles of axons in the early stage of myelin formation [25,28]. In contrast, the phenotype seen in the ADAM22-deficient mice was quite different: for example, unsorted bundles of axons were not seen in nerves and roots. We assume that ADAM22 partially modulates integrin-mediated signals or is involved at a later stage of myelin formation."}