Drosophila Importin-α2 Is Involved in Synapse, Axon and Muscle Development
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{"title"=>"Drosophila importin-α2 is involved in synapse, axon and muscle development", "type"=>"journal", "authors"=>[{"first_name"=>"Timothy J.", "last_name"=>"Mosca", "scopus_author_id"=>"8237160700"}, {"first_name"=>"Thomas L.", "last_name"=>"Schwarz", "scopus_author_id"=>"35325739900"}], "year"=>2010, "source"=>"PLoS ONE", "identifiers"=>{"issn"=>"19326203", "pui"=>"360158260", "sgr"=>"78650145173", "doi"=>"10.1371/journal.pone.0015223", "scopus"=>"2-s2.0-78650145173", "isbn"=>"1932-6203 (Electronic)\\r1932-6203 (Linking)", "pmid"=>"21151903"}, "id"=>"02ca7108-f6d2-3583-af1b-9b9428888b38", "abstract"=>"Nuclear import is required for communication between the cytoplasm and the nucleus and to enact lasting changes in gene transcription following stimuli. Binding to an Importin-α molecule in the cytoplasm is often required to mediate nuclear entry of a signaling protein. As multiple isoforms of Importin-α exist, some may be responsible for the entry of distinct cargoes rather than general nuclear import. Indeed, in neuronal systems, Importin-α isoforms can mediate very specific processes such as axonal tiling and communication of an injury signal. To study nuclear import during development, we examined the expression and function of Importin-α2 in Drosophila melanogaster. We found that Importin-α2 was expressed in the nervous system where it was required for normal active zone density at the NMJ and axonal commissure formation in the central nervous system. Other aspects of synaptic morphology at the NMJ and the localization of other synaptic markers appeared normal in importin-α2 mutants. Importin-α2 also functioned in development of the body wall musculature. Mutants in importin-α2 exhibited errors in muscle patterning and organization that could be alleviated by restoring muscle expression of Importin-α2. Thus, Importin-α2 is needed for some processes in the development of both the nervous system and the larval musculature.", "link"=>"http://www.mendeley.com/research/drosophila-importin%CE%B12-involved-synapse-axon-muscle-development", "reader_count"=>27, "reader_count_by_academic_status"=>{"Professor > Associate Professor"=>1, "Researcher"=>10, "Student > Ph. D. Student"=>11, "Student > Postgraduate"=>1, "Student > Master"=>2, "Other"=>1, "Student > Bachelor"=>1}, "reader_count_by_user_role"=>{"Professor > Associate Professor"=>1, "Researcher"=>10, "Student > Ph. D. Student"=>11, "Student > Postgraduate"=>1, "Student > Master"=>2, "Other"=>1, "Student > Bachelor"=>1}, "reader_count_by_subject_area"=>{"Unspecified"=>1, "Agricultural and Biological Sciences"=>21, "Medicine and Dentistry"=>1, "Neuroscience"=>3, "Computer Science"=>1}, "reader_count_by_subdiscipline"=>{"Medicine and Dentistry"=>{"Medicine and Dentistry"=>1}, "Neuroscience"=>{"Neuroscience"=>3}, "Agricultural and Biological Sciences"=>{"Agricultural and Biological Sciences"=>21}, "Computer Science"=>{"Computer Science"=>1}, "Unspecified"=>{"Unspecified"=>1}}, "reader_count_by_country"=>{"United States"=>2, "Israel"=>1, "France"=>1, "Germany"=>1}, "group_count"=>3}

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Figshare

  • {"files"=>["https://ndownloader.figshare.com/files/814037"], "description"=>"<p>(A) Diagram of the Importin-α2 protein reflecting the Importin-β binding domain (grey), eight Armadillo (ARM) repeats (green) and C-terminal SAR domain (blue). The extent of the D14 deletion <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0015223#pone.0015223-Gorjanacz1\" target=\"_blank\">[20]</a> and the region against which polyclonal antibodies were raised <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0015223#pone.0015223-Giarre1\" target=\"_blank\">[19]</a> are indicated. (B,C) Representative confocal z-stacks of the ventral nerve cord (VNC) from wild-type (<i>y,w; FRT42D; +; +</i>) and <i>importin-a2</i> mutant (<i>y,w; imp-α2<sup>D14</sup>; +; +</i>) third-instar larvae stained with antibodies to Importin-α2 (magenta). The antibody labels a population of wild-type cell bodies within the brain lobes and at the midline of the VNC. (D–F) High magnification images of the midline of the VNC in wild-type larvae stained with antibodies to Importin-α2 (magenta) and the glial marker Repo (green). Repo does not overlap with Importin-α2 staining. (G–H) In high magnification images of the brain lobes of wild-type larvae, Importin-α2 (magenta) and Repo (green) immunoreactivities do not overlap. (I) A larval imaginal disc in a wild-type larva stained with antibodies to Importin-α2 (magenta). In all panels, scale bar = 10 µm.</p>", "links"=>[], "tags"=>["larval"], "article_id"=>484411, "categories"=>["Neuroscience", "Genetics", "Developmental Biology"], "users"=>["Timothy J. Mosca", "Thomas L. Schwarz"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0015223.g001", "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Importin_945_2_is_Expressed_in_the_Larval_Nervous_System_/484411", "title"=>"Importin-α2 is Expressed in the Larval Nervous System.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-12-06 01:13:31"}
  • {"files"=>["https://ndownloader.figshare.com/files/814335"], "description"=>"<p>(A–C) Representative confocal images of wild-type (<i>y,w; FRT42D; +; +</i>) larval NMJs from muscle 4 stained with antibodies against Bruchpilot (magenta), GluRIIC (green) and HRP (blue). Insets show high magnification images of individual boutons. Each Brp punctum is apposed by a corresponding GluRIIC punctum. (D–F) Representative confocal images stained as above in <i>importin-α2</i> mutants (<i>y,w; imp-α2<sup>D14</sup>; +; +</i>). No defects in apposition are observed, but the density of both Brp and GluRIIC puncta is increased. Scale bar = 10 µm for full NMJ, 4 µm for high magnification images. (G–H) Quantification of Brp and GluRIIC puncta in wild-type larvae, <i>importin-α2</i> mutants, and <i>importin-α2</i> mutants with restored neuronal expression of Importin-α2 (<i>y,w; imp-α2<sup>D14</sup>; elav-GAL4/UASt-Importin-α2; +</i>). In <i>importin-α2</i> mutants, the density of Brp and GluRIIC puncta per µm<sup>2</sup> is increased by 40% and restored to wild-type levels by neuronal restoration of Importin-α2 expression. *** <i>p</i><0.0001. (I–J) The synaptic area (as determined by anti-HRP staining of terminals) and the ratio of Brp to GluRIIC puncta are unchanged in the mutants. Error bars represent S.E.M. For all cases, <i>n</i>≥2 NMJs from 6 larvae.</p>", "links"=>[], "tags"=>["neuronal", "increases"], "article_id"=>484711, "categories"=>["Neuroscience", "Genetics", "Developmental Biology"], "users"=>["Timothy J. Mosca", "Thomas L. Schwarz"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0015223.g003", "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Loss_of_Neuronal_Importin_945_2_Increases_Active_Zone_Density_/484711", "title"=>"Loss of Neuronal Importin-α2 Increases Active Zone Density.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-12-06 01:18:31"}
  • {"files"=>["https://ndownloader.figshare.com/files/814988"], "description"=>"<p>Quantification of muscle patterning errors from third-instar larvae stained with TxRed-conjugated phalloidin. Percents are the number of errors observed/total hemisegments scored</p><p><i>n</i>>8 animals for all genotypes.</p>", "links"=>[], "tags"=>["patterning", "errors", "mutant"], "article_id"=>485359, "categories"=>["Neuroscience", "Genetics", "Developmental Biology"], "users"=>["Timothy J. Mosca", "Thomas L. Schwarz"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0015223.t001", "stats"=>{"downloads"=>0, "page_views"=>2, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Frequency_of_Muscle_Patterning_Errors_in_importin_945_2_Mutant_Larvae_/485359", "title"=>"Frequency of Muscle Patterning Errors in <i>importin-α2</i> Mutant Larvae.", "pos_in_sequence"=>0, "defined_type"=>3, "published_date"=>"2010-12-06 01:29:19"}
  • {"files"=>["https://ndownloader.figshare.com/files/406145"], "description"=>"<div><p>Nuclear import is required for communication between the cytoplasm and the nucleus and to enact lasting changes in gene transcription following stimuli. Binding to an Importin-α molecule in the cytoplasm is often required to mediate nuclear entry of a signaling protein. As multiple isoforms of Importin-α exist, some may be responsible for the entry of distinct cargoes rather than general nuclear import. Indeed, in neuronal systems, Importin-α isoforms can mediate very specific processes such as axonal tiling and communication of an injury signal. To study nuclear import during development, we examined the expression and function of Importin-α2 in <em>Drosophila melanogaster</em>. We found that Importin-α2 was expressed in the nervous system where it was required for normal active zone density at the NMJ and axonal commissure formation in the central nervous system. Other aspects of synaptic morphology at the NMJ and the localization of other synaptic markers appeared normal in <em>importin-α2</em> mutants. Importin-α2 also functioned in development of the body wall musculature. Mutants in <em>importin-α2</em> exhibited errors in muscle patterning and organization that could be alleviated by restoring muscle expression of Importin-α2. Thus, Importin-α2 is needed for some processes in the development of both the nervous system and the larval musculature.</p> </div>", "links"=>[], "tags"=>["axon"], "article_id"=>140240, "categories"=>["Neuroscience", "Genetics", "Developmental Biology"], "users"=>["Timothy J. Mosca", "Thomas L. Schwarz"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0015223", "stats"=>{"downloads"=>0, "page_views"=>5, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Drosophila_Importin_2_Is_Involved_in_Synapse_Axon_and_Muscle_Development/140240", "title"=>"<em>Drosophila</em> Importin-α2 Is Involved in Synapse, Axon and Muscle Development", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-12-06 00:04:00"}
  • {"files"=>["https://ndownloader.figshare.com/files/814550"], "description"=>"<p>(A–E) Representative confocal projections of the larval NMJ at muscle 4 stained with antibodies to Synaptotagmin I, Cysteine String Protein (CSP), Fasciclin II, and HRP, as indicated, in wild-type (<i>y,w; FRT42D; +; +</i>) larvae. (F–J) Representative confocal projections of <i>importin-α2</i> mutant larvae (<i>y,w; imp-α2<sup>D14</sup>; +; +</i>) stained as in (A–E). As in wild-type larvae, synaptic vesicle proteins and Fas II properly localize at the NMJ in <i>importin-α2</i> mutants. Scale bar = 10 µm. (K–M) Quantification of the intensity of immunoreactivity for Syt I (K), CSP (L) and Fas II (M) in wild-type and <i>importin-α2</i> mutant larvae. No statistically significant differences were observed. In all cases, <i>p</i>>0.4, <i>n</i> = 6 animals, 12 NMJs and the error bars indicate S.E.M.</p>", "links"=>[], "tags"=>["synaptic", "markers"], "article_id"=>484920, "categories"=>["Neuroscience", "Genetics", "Developmental Biology"], "users"=>["Timothy J. Mosca", "Thomas L. Schwarz"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0015223.g004", "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Common_Synaptic_Markers_are_Normal_in_importin_945_2_Mutants_/484920", "title"=>"Common Synaptic Markers are Normal in <i>importin-α2</i> Mutants.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-12-06 01:22:00"}
  • {"files"=>["https://ndownloader.figshare.com/files/814860"], "description"=>"<p>(A) Representative confocal image of a wild-type (<i>y,w; FRT42D; +; +</i>) larval muscle field stained with Texas Red-conjugated phalloidin. Muscles are labeled according to Crossley <a href=\"http://www.plosone.org/article/info:doi/10.1371/journal.pone.0015223#pone.0015223-Crossley1\" target=\"_blank\">[47]</a>. This represents a single hemisegment between the midline (at left of image) and muscle 4. Muscle 8 from the next anterior hemisegment is also visible. (B) An <i>importin-α2</i> mutant (<i>y,w; imp-α2<sup>D14</sup>; +; +</i>) with incorrect positioning of muscles 6 and 7 (arrowheads). In wild-type larvae (A), these muscles are parallel but they cross in some <i>importin-α2</i> mutants. Also, an abnormally thin muscle 5 is evident in this example (arrow). (C) Representative example of an <i>importin-α2</i> mutant where muscle 5 is absent but other muscles are normal. (D) Representative example of an <i>importin-α2</i> mutant in which an abnormally branched fiber has formed (arrowhead). In this case, the branch derives from muscle 6. These four categories of defect were observed in <i>importin-α2</i> mutants. Scale bar = 50 µm.</p>", "links"=>[], "tags"=>["patterning", "defects"], "article_id"=>485230, "categories"=>["Neuroscience", "Genetics", "Developmental Biology"], "users"=>["Timothy J. Mosca", "Thomas L. Schwarz"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0015223.g006", "stats"=>{"downloads"=>0, "page_views"=>2, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Muscle_Patterning_Defects_in_importin_945_2_Mutants_/485230", "title"=>"Muscle Patterning Defects in <i>importin-α2</i> Mutants.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-12-06 01:27:10"}
  • {"files"=>["https://ndownloader.figshare.com/files/814184"], "description"=>"<p><b>Mutants.</b> (A–B) Representative confocal images of third-instar larval NMJs from wild-type (<i>y,w; FRT42D; +; +</i>) and <i>importin-α2</i> mutants (<i>y,w; imp-α2<sup>D14</sup>; +; +</i>) stained with antibodies to HRP. Scale bar = 10 µm. (C–E) Quantification of bouton number (C), bouton number normalized to muscle surface area (D) and muscle surface area (E) in <i>wild-type</i> and <i>importin-α2</i> larvae. No significant differences were detected. In all cases, <i>p</i>>0.2 and error bars represent S.E.M. For <i>wild-type</i>: <i>n</i> = 29 NMJs, 15 larvae and for <i>importin-α2</i>: <i>n</i> = 22 NMJs, 12 larvae.</p>", "links"=>[], "tags"=>["bouton"], "article_id"=>484568, "categories"=>["Neuroscience", "Genetics", "Developmental Biology"], "users"=>["Timothy J. Mosca", "Thomas L. Schwarz"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0015223.g002", "stats"=>{"downloads"=>0, "page_views"=>0, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Normal_Bouton_Number_in_importin_945_2_/484568", "title"=>"Normal Bouton Number in <i>importin-α2</i>", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-12-06 01:16:08"}
  • {"files"=>["https://ndownloader.figshare.com/files/814737"], "description"=>"<p>(A–B) Representative confocal projections of the ventral nerve cord midline in both wild-type (<i>y,w; FRT42D; +; +</i>) and <i>importin-α2</i> mutant (<i>y,w; imp-α2<sup>D14</sup>; +; +</i>) third-instar larvae stained with antibodies to Fasciclin III. In wild-type larvae, Fas III antibodies stained the central-most longitudinal tracts and commissural axons crossing the midline. While both the longitudinal tracts and commissures were visible in <i>importin-α2</i> mutants, there were frequent discontinuities in the staining of the commissures (red asterisks). Scale bar = 50 µm. (C) Quantification of larvae possessing no errors, single abnormal commissures or multiple abnormalities in wild-type controls, <i>importin-α2</i> mutant larvae, and <i>importin-α2</i> mutant larvae where neuronal expression has been restored (<i>y,w; imp-α2<sup>D14</sup>; elav-GAL4/UASt-Importin-α2; +</i>). For each genotype, <i>n</i>≥20 animals.</p>", "links"=>[], "tags"=>["commissure", "defects", "midline"], "article_id"=>485107, "categories"=>["Neuroscience", "Genetics", "Developmental Biology"], "users"=>["Timothy J. Mosca", "Thomas L. Schwarz"], "doi"=>"https://dx.doi.org/10.1371/journal.pone.0015223.g005", "stats"=>{"downloads"=>1, "page_views"=>2, "likes"=>0}, "figshare_url"=>"https://figshare.com/articles/_Axon_Commissure_Defects_at_the_Midline_of_importin_945_2_Mutants_/485107", "title"=>"Axon Commissure Defects at the Midline of <i>importin-α2</i> Mutants.", "pos_in_sequence"=>0, "defined_type"=>1, "published_date"=>"2010-12-06 01:25:07"}

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