PMC:548520 / 1245-6397 JSONTXT

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These sensory neurons are a shared element in all organisms capable of sensing light. In humans, genetic diseases of the eye are common and the primary site of disease is most frequently photoreceptors (for review see [1-3]).\nPhotoreceptors have been well studied, particularly with respect to the biochemistry and physiology of phototransduction. Insight into the development of vertebrate photoreceptors, however, has lagged behind our understanding of function. Only recently have the first molecular mechanisms regulating photoreceptor development been identified (for review see, [2,4]). Crx (cone-rod homeobox) is an otx-family homeobox gene expressed predominantly in photoreceptors, from early in their development through to the adult ages [5-7]. Crx gene expression is critically dependent upon Otx2, another member of the same homeobox family which is expressed in early photoreceptor cells [8]. In rod photoreceptors, Crx appears to work in concert with Nrl, a leucine zipper protein that is also restricted in its expression in the retina to rod photoreceptors [9]. Many photoreceptor-specific genes have putative Crx-binding elements in their regulatory regions [10], including rhodopsin [11] and arrestin [12]. Mutations in Crx have been associated with several human diseases that lead to blindness, including cone-rod dystrophy 2 [6,13,14], retinitis pigmentosa [14], and LCA [14-16]. Based on these data, Crx was hypothesized to play a critical role in the differentiation and maintainance of photoreceptor cells [5,7].\nLCA is the most severe genetic disease of photoreceptors (see [17], for recent review). Affected infants exhibit a complete or near complete absence of vision from birth. Mutations in retinal specific genes, such as Crx, have been associated with LCA [14,15], as well as GUCY2D [18], RPE65 [19], AIPL-1 [20], CRB-1 [21], and RPGRIP-1 [22]. There also may be as many as three additional genetically linked loci where genes have not been identified [23]. Crx mutations in LCA are varied, and include a putative dominant mutation that is proposed to encode a dominant-negative form of Crx [14,15]. Recessive mutations also have been reported and at least one allele encodes a protein with decreased DNA-binding activity [16]. Histopathological and ultrastructural studies of LCA should enable a better understanding of the disease process, and the design of suitable therapies. Few such studies exist for human LCA (reviewed in [17]) and the majority of such studies examine the globes of adults with LCA, after the tissue has undergone secondary changes. Only a single study exists where the developing eye of an infant was examined [24]. Animal models for LCA have recently been reported and have already served to broaden our understanding of the pathology of this disease [25-28]. Since LCA is a clinically and genetically heterogeneous disorder, additional mouse models are in order to allow a full understanding of the many ways in which photoreceptor development can go awry.\nIn addition to their importance as a locus of disease, photoreceptor cells serve as an excellent model for studies in neuronal differentiation. Photoreceptor cells are highly polarized. At their apex, these neurons have a membranous outer segment, which contains proteins involved in the phototransduction cascade. Loss of function mutations in rhodopsin [29], or the structural protein, peripherin [30], result in an inability to form outer segments. At the other extremity, photoreceptors terminate with synaptic endings that make contact with the processes of horizontal and bipolar cells [31,32]. Rod spherules establish an invaginating synapse with rod bipolar dendrites and axonal endings of horizontal cells. This synapse is characterized by the presence of a ribbon in the presynaptic cytoplasm. Cone pedicles make invaginating synapses with the dendrites of on-cone bipolar cells and horizontal cells and basal junctions with the dendrites of off-cone bipolar cells. The factors regulating the formation of the photoreceptor synapses are completely unknown. At least one photoreceptor synaptic protein, HRG4, contains a potential Crx target sequence in its transcriptional regulatory sequence [33].\nFew studies of LCA animal models have extended their examination of retinal pathology to the ultrastructural level. Certain features of neuronal differentiation, such as synapse formation, can be detected definitively at this level of analysis. With the hope of understanding the neuropathology of LCA in greater detail, we have analyzed the differentiation of the outer retina in Crx-/- mice at the ultrastructural level. These retinas exhibit several prominent defects. Crx-/- photoreceptors demonstrate a complete block in outer segment formation at the elongation stage. Further, these cells exhibit abnormal synaptic morphology, thereby broadening the function of Crx to photoreceptor synaptogenesis. This represents the first report strongly implicating the process of synapse formation in LCA."}

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cells play a primary role in vision by capturing light energy and converting it into neural stimuli. These sensory neurons are a shared element in all organisms capable of sensing light. In humans, genetic diseases of the eye are common and the primary site of disease is most frequently photoreceptors (for review see [1-3]).\nPhotoreceptors have been well studied, particularly with respect to the biochemistry and physiology of phototransduction. Insight into the development of vertebrate photoreceptors, however, has lagged behind our understanding of function. Only recently have the first molecular mechanisms regulating photoreceptor development been identified (for review see, [2,4]). Crx (cone-rod homeobox) is an otx-family homeobox gene expressed predominantly in photoreceptors, from early in their development through to the adult ages [5-7]. Crx gene expression is critically dependent upon Otx2, another member of the same homeobox family which is expressed in early photoreceptor cells [8]. In rod photoreceptors, Crx appears to work in concert with Nrl, a leucine zipper protein that is also restricted in its expression in the retina to rod photoreceptors [9]. Many photoreceptor-specific genes have putative Crx-binding elements in their regulatory regions [10], including rhodopsin [11] and arrestin [12]. Mutations in Crx have been associated with several human diseases that lead to blindness, including cone-rod dystrophy 2 [6,13,14], retinitis pigmentosa [14], and LCA [14-16]. Based on these data, Crx was hypothesized to play a critical role in the differentiation and maintainance of photoreceptor cells [5,7].\nLCA is the most severe genetic disease of photoreceptors (see [17], for recent review). Affected infants exhibit a complete or near complete absence of vision from birth. Mutations in retinal specific genes, such as Crx, have been associated with LCA [14,15], as well as GUCY2D [18], RPE65 [19], AIPL-1 [20], CRB-1 [21], and RPGRIP-1 [22]. There also may be as many as three additional genetically linked loci where genes have not been identified [23]. Crx mutations in LCA are varied, and include a putative dominant mutation that is proposed to encode a dominant-negative form of Crx [14,15]. Recessive mutations also have been reported and at least one allele encodes a protein with decreased DNA-binding activity [16]. Histopathological and ultrastructural studies of LCA should enable a better understanding of the disease process, and the design of suitable therapies. Few such studies exist for human LCA (reviewed in [17]) and the majority of such studies examine the globes of adults with LCA, after the tissue has undergone secondary changes. Only a single study exists where the developing eye of an infant was examined [24]. Animal models for LCA have recently been reported and have already served to broaden our understanding of the pathology of this disease [25-28]. Since LCA is a clinically and genetically heterogeneous disorder, additional mouse models are in order to allow a full understanding of the many ways in which photoreceptor development can go awry.\nIn addition to their importance as a locus of disease, photoreceptor cells serve as an excellent model for studies in neuronal differentiation. Photoreceptor cells are highly polarized. At their apex, these neurons have a membranous outer segment, which contains proteins involved in the phototransduction cascade. Loss of function mutations in rhodopsin [29], or the structural protein, peripherin [30], result in an inability to form outer segments. At the other extremity, photoreceptors terminate with synaptic endings that make contact with the processes of horizontal and bipolar cells [31,32]. Rod spherules establish an invaginating synapse with rod bipolar dendrites and axonal endings of horizontal cells. This synapse is characterized by the presence of a ribbon in the presynaptic cytoplasm. Cone pedicles make invaginating synapses with the dendrites of on-cone bipolar cells and horizontal cells and basal junctions with the dendrites of off-cone bipolar cells. The factors regulating the formation of the photoreceptor synapses are completely unknown. At least one photoreceptor synaptic protein, HRG4, contains a potential Crx target sequence in its transcriptional regulatory sequence [33].\nFew studies of LCA animal models have extended their examination of retinal pathology to the ultrastructural level. Certain features of neuronal differentiation, such as synapse formation, can be detected definitively at this level of analysis. With the hope of understanding the neuropathology of LCA in greater detail, we have analyzed the differentiation of the outer retina in Crx-/- mice at the ultrastructural level. These retinas exhibit several prominent defects. Crx-/- photoreceptors demonstrate a complete block in outer segment formation at the elongation stage. Further, these cells exhibit abnormal synaptic morphology, thereby broadening the function of Crx to photoreceptor synaptogenesis. This represents the first report strongly implicating the process of synapse formation in LCA."}

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cells play a primary role in vision by capturing light energy and converting it into neural stimuli. These sensory neurons are a shared element in all organisms capable of sensing light. In humans, genetic diseases of the eye are common and the primary site of disease is most frequently photoreceptors (for review see [1-3]).\nPhotoreceptors have been well studied, particularly with respect to the biochemistry and physiology of phototransduction. Insight into the development of vertebrate photoreceptors, however, has lagged behind our understanding of function. Only recently have the first molecular mechanisms regulating photoreceptor development been identified (for review see, [2,4]). Crx (cone-rod homeobox) is an otx-family homeobox gene expressed predominantly in photoreceptors, from early in their development through to the adult ages [5-7]. Crx gene expression is critically dependent upon Otx2, another member of the same homeobox family which is expressed in early photoreceptor cells [8]. In rod photoreceptors, Crx appears to work in concert with Nrl, a leucine zipper protein that is also restricted in its expression in the retina to rod photoreceptors [9]. Many photoreceptor-specific genes have putative Crx-binding elements in their regulatory regions [10], including rhodopsin [11] and arrestin [12]. Mutations in Crx have been associated with several human diseases that lead to blindness, including cone-rod dystrophy 2 [6,13,14], retinitis pigmentosa [14], and LCA [14-16]. Based on these data, Crx was hypothesized to play a critical role in the differentiation and maintainance of photoreceptor cells [5,7].\nLCA is the most severe genetic disease of photoreceptors (see [17], for recent review). Affected infants exhibit a complete or near complete absence of vision from birth. Mutations in retinal specific genes, such as Crx, have been associated with LCA [14,15], as well as GUCY2D [18], RPE65 [19], AIPL-1 [20], CRB-1 [21], and RPGRIP-1 [22]. There also may be as many as three additional genetically linked loci where genes have not been identified [23]. Crx mutations in LCA are varied, and include a putative dominant mutation that is proposed to encode a dominant-negative form of Crx [14,15]. Recessive mutations also have been reported and at least one allele encodes a protein with decreased DNA-binding activity [16]. Histopathological and ultrastructural studies of LCA should enable a better understanding of the disease process, and the design of suitable therapies. Few such studies exist for human LCA (reviewed in [17]) and the majority of such studies examine the globes of adults with LCA, after the tissue has undergone secondary changes. Only a single study exists where the developing eye of an infant was examined [24]. Animal models for LCA have recently been reported and have already served to broaden our understanding of the pathology of this disease [25-28]. Since LCA is a clinically and genetically heterogeneous disorder, additional mouse models are in order to allow a full understanding of the many ways in which photoreceptor development can go awry.\nIn addition to their importance as a locus of disease, photoreceptor cells serve as an excellent model for studies in neuronal differentiation. Photoreceptor cells are highly polarized. At their apex, these neurons have a membranous outer segment, which contains proteins involved in the phototransduction cascade. Loss of function mutations in rhodopsin [29], or the structural protein, peripherin [30], result in an inability to form outer segments. At the other extremity, photoreceptors terminate with synaptic endings that make contact with the processes of horizontal and bipolar cells [31,32]. Rod spherules establish an invaginating synapse with rod bipolar dendrites and axonal endings of horizontal cells. This synapse is characterized by the presence of a ribbon in the presynaptic cytoplasm. Cone pedicles make invaginating synapses with the dendrites of on-cone bipolar cells and horizontal cells and basal junctions with the dendrites of off-cone bipolar cells. The factors regulating the formation of the photoreceptor synapses are completely unknown. At least one photoreceptor synaptic protein, HRG4, contains a potential Crx target sequence in its transcriptional regulatory sequence [33].\nFew studies of LCA animal models have extended their examination of retinal pathology to the ultrastructural level. Certain features of neuronal differentiation, such as synapse formation, can be detected definitively at this level of analysis. With the hope of understanding the neuropathology of LCA in greater detail, we have analyzed the differentiation of the outer retina in Crx-/- mice at the ultrastructural level. These retinas exhibit several prominent defects. 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cells play a primary role in vision by capturing light energy and converting it into neural stimuli. These sensory neurons are a shared element in all organisms capable of sensing light. In humans, genetic diseases of the eye are common and the primary site of disease is most frequently photoreceptors (for review see [1-3]).\nPhotoreceptors have been well studied, particularly with respect to the biochemistry and physiology of phototransduction. Insight into the development of vertebrate photoreceptors, however, has lagged behind our understanding of function. Only recently have the first molecular mechanisms regulating photoreceptor development been identified (for review see, [2,4]). Crx (cone-rod homeobox) is an otx-family homeobox gene expressed predominantly in photoreceptors, from early in their development through to the adult ages [5-7]. Crx gene expression is critically dependent upon Otx2, another member of the same homeobox family which is expressed in early photoreceptor cells [8]. In rod photoreceptors, Crx appears to work in concert with Nrl, a leucine zipper protein that is also restricted in its expression in the retina to rod photoreceptors [9]. Many photoreceptor-specific genes have putative Crx-binding elements in their regulatory regions [10], including rhodopsin [11] and arrestin [12]. Mutations in Crx have been associated with several human diseases that lead to blindness, including cone-rod dystrophy 2 [6,13,14], retinitis pigmentosa [14], and LCA [14-16]. Based on these data, Crx was hypothesized to play a critical role in the differentiation and maintainance of photoreceptor cells [5,7].\nLCA is the most severe genetic disease of photoreceptors (see [17], for recent review). Affected infants exhibit a complete or near complete absence of vision from birth. Mutations in retinal specific genes, such as Crx, have been associated with LCA [14,15], as well as GUCY2D [18], RPE65 [19], AIPL-1 [20], CRB-1 [21], and RPGRIP-1 [22]. There also may be as many as three additional genetically linked loci where genes have not been identified [23]. Crx mutations in LCA are varied, and include a putative dominant mutation that is proposed to encode a dominant-negative form of Crx [14,15]. Recessive mutations also have been reported and at least one allele encodes a protein with decreased DNA-binding activity [16]. Histopathological and ultrastructural studies of LCA should enable a better understanding of the disease process, and the design of suitable therapies. Few such studies exist for human LCA (reviewed in [17]) and the majority of such studies examine the globes of adults with LCA, after the tissue has undergone secondary changes. Only a single study exists where the developing eye of an infant was examined [24]. Animal models for LCA have recently been reported and have already served to broaden our understanding of the pathology of this disease [25-28]. Since LCA is a clinically and genetically heterogeneous disorder, additional mouse models are in order to allow a full understanding of the many ways in which photoreceptor development can go awry.\nIn addition to their importance as a locus of disease, photoreceptor cells serve as an excellent model for studies in neuronal differentiation. Photoreceptor cells are highly polarized. At their apex, these neurons have a membranous outer segment, which contains proteins involved in the phototransduction cascade. Loss of function mutations in rhodopsin [29], or the structural protein, peripherin [30], result in an inability to form outer segments. At the other extremity, photoreceptors terminate with synaptic endings that make contact with the processes of horizontal and bipolar cells [31,32]. Rod spherules establish an invaginating synapse with rod bipolar dendrites and axonal endings of horizontal cells. This synapse is characterized by the presence of a ribbon in the presynaptic cytoplasm. Cone pedicles make invaginating synapses with the dendrites of on-cone bipolar cells and horizontal cells and basal junctions with the dendrites of off-cone bipolar cells. The factors regulating the formation of the photoreceptor synapses are completely unknown. At least one photoreceptor synaptic protein, HRG4, contains a potential Crx target sequence in its transcriptional regulatory sequence [33].\nFew studies of LCA animal models have extended their examination of retinal pathology to the ultrastructural level. Certain features of neuronal differentiation, such as synapse formation, can be detected definitively at this level of analysis. With the hope of understanding the neuropathology of LCA in greater detail, we have analyzed the differentiation of the outer retina in Crx-/- mice at the ultrastructural level. These retinas exhibit several prominent defects. Crx-/- photoreceptors demonstrate a complete block in outer segment formation at the elongation stage. Further, these cells exhibit abnormal synaptic morphology, thereby broadening the function of Crx to photoreceptor synaptogenesis. This represents the first report strongly implicating the process of synapse formation in LCA."}