Allele Dmel\Nrg^l4

General Information
Symbol	Dmel\Nrg^l4	Species	D. melanogaster
Name		FlyBase ID	FBal0013169
Feature type	allele	Associated gene	Dmel\Nrg
Also Known As	Nrg¹, Nrg¹⁴, l(1)RA35

Allele class	amorphic allele - genetic evidence, loss of function allele
Mutagen	X ray
Recent Updates
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FB2013_03
FB2013_02
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Nature of the Allele
Allele class	amorphic allele - genetic evidence (Hortsch and Goodman, 1993, ) loss of function allele (Hall and Bieber, 1997, Speicher et al., 1998)
Mutagen	X ray (Hall and Bieber, 1997)
Mutations Mapped to the Genome

Type Location Additional Notes References
Associated Sequence Data
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name

UniProtKB/Swiss-Prot
UniProtKB/TrEMBL

Progenitor genotype
Nature of the lesion	Statement Reference Inversion breakpoint within the Nrg transcription unit. (Hall and Bieber, 1997)
Caused by aberration	In(1)RA35 (Hall and Bieber, 1997, King et al., 1986, Bieber et al., 1989, Kania and Bellen, 1995)
Cytology
Phenotypic Data
Phenotypic Class
	courtship behavior defective \| female (with Nrg⁸⁴⁹) (Carhan et al., 2005) courtship behavior defective \| female (with Nrg⁸⁹²) (Carhan et al., 2005) courtship behavior defective \| female (with Nrg^ibx) (Carhan et al., 2005) lethal \| embryonic stage (Bieber et al., 1989) lethal \| embryonic stage \| recessive (Hall and Bieber, 1997) lethal \| heat sensitive (with Nrg^l10) (Carhan et al., 2005) lethal \| recessive (Speicher et al., 1998) neuroanatomy defective (Yamamoto, 2006, Schwabe et al., 2005)
Phenotype Manifest In
	abdominal lateral pentascolopidial chordotonal organ lch5 (Banerjee et al., 2006) abdominal segmental nerve (Hall and Bieber, 1997) aCC neuron (Hall and Bieber, 1997) axon & nerve (Banerjee et al., 2006) cell cortex & cortical cytoskeleton (Schwabe et al., 2005) dorsal multidendritic neuron ddaD & axon (Yamamoto, 2006) dorsal multidendritic neuron ddaD & dendritic tree (Yamamoto, 2006) dorsal multidendritic neuron ddaE & axon (Yamamoto, 2006) dorsal multidendritic neuron ddaE & axon \| somatic clone (Yamamoto, 2006) embryonic epidermis & pleated septate junction (Faivre-Sarrailh et al., 2004) embryonic salivary gland & epithelial cell (Genova and Fehon, 2003) lateral cord surface glia & nucleus (Schwabe et al., 2005) peripheral glial cell (Banerjee et al., 2006) peripheral nervous system (Hall and Bieber, 1997) scolopidium & abdominal lateral pentascolopidial chordotonal organ lch5 (Banerjee et al., 2006) septate junction (Genova and Fehon, 2003) septate junction & peripheral glial cell (Banerjee et al., 2006) ventral nerve cord (Hijazi et al., 2009)
Detailed Description
	Statement Reference Dextran dye can penetrate in the nerve cord after it has been injected into 22 hour mutant embryos, indicating a defect in the integrity of the nerve cord paracellular barrier. (Hijazi et al., 2009) In Nrg^l4 homozygotes lateral pentascolopidial chordotonal organs have a disorganised morphology and rounded, rather than fusiform scolopales. Unlike in wild-type embryos, lateral pentascolopidial chordotonal organs in these embryos fail to exclude a 10kDa dextran dye. Ultrastructural analyses of peripheral nerves in these mutants at late stages of embryogenesis show that, while glial membranes are present around these nerves, the associated septate junctions linking inner and outer glial sheath cells are missing. and the inner and outer glial membranes are abnormally far apart. In addition, axons are often missing from bundles and replaced by protrusions from the inner glial cells. (Banerjee et al., 2006) 29% of dorsal da neuron E (ddaE) and 38% of ddaD neurons show ectopic branches on their axons in Nrg^l4 embryos. The dendritic branches of ddaD cover a smaller field and have fewer terminals than wild-type ddaD neurons. Nrg^l4 ddaE clones form ectopic branches on their axons. (Yamamoto, 2006) Nrg[l10]/Nrg[l4] females are viable at 18[o]C but are inviable at 25[o]C. Nrg[l4]/Nrg[849] and Nrg[l4]/Nrg[892] females show reduced sexual receptivity compared to wild-type controls when mated to wild-type males. (Carhan et al., 2005) Fluorescent dye injected into the body cavity of Nrg^l4 embryos penetrates into the nerve cord after the stage at which the nerve cord is sealed in wild-type embryos, showing that formation of the blood-brain barrier if defective. The surface glial cell shape and cortical actin distribution is only mildly disrupted. (Schwabe et al., 2005) In the epidermis of about 80% of stage 15 Nrg^l4 mutant embryos, occasional clusters of septa are formed, often basal in their position. (Faivre-Sarrailh et al., 2004) Nrg^l4 mutants exhibit disruption of the paracellular barrier in the embryonic salivary gland. In Nrg^l4 homozygous mutants the adherens junction remains intact, with the regular spacing of plasma membranes maintained. However, the septae normally located between these membranes are reduced in number or absent. Nrg^l4/Nrg^l4 somatic clones generated in larvae (approx. 72 hours after egg laying) do not survive in adult tissues. (Genova and Fehon, 2003) Fas2 expressing axons appear normal in homozygous embryos. (Speicher et al., 1998) Homozygous embryos show a general disorganisation of the peripheral nervous system cell bodies. The aCC and SNb motoneurons often show a stalled phenotype, failing to extend normally, and SNb motoneurons sometimes show abnormal contacts with their targets. (Hall and Bieber, 1997)
External Data
Linkouts
Interactions
	Show the genetic interaction network for Enhancers & Suppressors
Phenotypic Class
Enhancer of
Statement Reference Nrg^l4/Nrg[+] is an enhancer of neuroanatomy defective \| larval stage phenotype of Nak^{dsRNA.Scer\UAS}, Scer\GAL4^elav-C155 (Yang et al., 2011)
NOT Suppressor of
Statement Reference Nrg^l4 is a non-suppressor of neuroanatomy defective \| embryonic stage phenotype of rap^ie28 (Silies and Klämbt, 2010)
Phenotype Manifest In
Enhancer of
Statement Reference Nrg^l4/Nrg[+] is an enhancer of dendrite \| larval stage phenotype of Nak^{dsRNA.Scer\UAS}, Scer\GAL4^elav-C155 (Yang et al., 2011) Nrg^l4/Nrg[+] is an enhancer of dendritic arborizing neuron \| larval stage phenotype of Nak^{dsRNA.Scer\UAS}, Scer\GAL4^elav-C155 (Yang et al., 2011) Nrg^l4 is an enhancer of axon \| embryonic stage phenotype of Nrt¹ (Speicher et al., 1998) Nrg^l4 is an enhancer of commissure phenotype of Nrt⁵ (Speicher et al., 1998) Nrg^l4 is an enhancer of dMP2 neuron phenotype of Nrt⁵ (Speicher et al., 1998) Nrg^l4 is an enhancer of longitudinal connective phenotype of Nrt⁵ (Speicher et al., 1998) Nrg^l4 is an enhancer of MP1 neuron phenotype of Nrt⁵ (Speicher et al., 1998) Nrg^l4 is an enhancer of SP1 neuron phenotype of Nrt⁵ (Speicher et al., 1998)
NOT Enhancer of
Statement Reference Nrg^l4, Scer\GAL4^GMR.PF/Scer\GAL4^GMR.PF is a non-enhancer of eye photoreceptor cell phenotype of Scer\GAL4^GMR.PF/Scer\GAL4^GMR.PF, ed^Scer\UAS.cBa (Islam et al., 2003)
Suppressor of
Statement Reference Nrg^l4, Scer\GAL4^GMR.PF/Scer\GAL4^GMR.PF is a suppressor of cone cell phenotype of Scer\GAL4^GMR.PF/Scer\GAL4^GMR.PF, ed^Scer\UAS.cBa (Islam et al., 2003)
NOT Suppressor of
Statement Reference Nrg^l4, Scer\GAL4^GMR.PF/Scer\GAL4^GMR.PF is a non-suppressor of eye photoreceptor cell phenotype of Scer\GAL4^GMR.PF/Scer\GAL4^GMR.PF, ed^Scer\UAS.cBa (Islam et al., 2003) Nrg^l4 is a non-suppressor of glial cell phenotype of rap^ie28 (Silies and Klämbt, 2010)
Other
Statement Reference Nrg^l4, Nrt^rP668 has axon \| embryonic stage phenotype (Speicher et al., 1998, Speicher et al., 1998)
Additional Comments
Genetic Interactions
Statement Reference Nrg[l4] dominantly enhances the shortening of dendritic length and the reduction in the number of dendritic endpoints of dorsal dendritic arborisation neurons which is seen in larvae expressing Nak[dsRNA.Scer\UAS] under the control of Scer\GAL4[elav-C155]. (Yang et al., 2011) The glial cell migration defects of rap[ie28] embryos are not suppressed by Nrg[l4]. (Silies and Klämbt, 2010) The addition of Nrg^l4/+ significantly suppresses the loss of cone cell phenotype seen in ed^Scer\UAS.cBa, Scer\GAL4^GMR.PF animals but has no effect on photoreceptor loss. (Islam et al., 2003) Nrg^l4 Nrt⁵ double mutant embryos have a severe CNS phenotype, with thinning or complete interruption of the longitudinal connectives, and fusion of the commissures. Defects in Fas2 expressing neurons similar to those seen in Nrt⁵ single mutants are seen, although with much higher expressivity and penetrance. Axons of the dMP2 and MP1 neurons either stall or delay their extension considerably in 29% of cases. The ventral unpaired medial (VUM) neurons show misguidance phenotypes in 15% of cases, and anomalies in the trajectory of the SP1 axon are observed rarely. The pCC and vMP2 axons grow normally in most hemisegments, and the aCC and U neurons are normal. These defects are rescued by Nrt^Scer\UAS.cSa expressed under the control of Scer\GAL4^Mz1277. Nrt^rP668 Nrg^l4 double mutant embryos have defects in Fas2 expressing axons. Nrt^R20F Nrg^l4 embryos have essentially normal Fas2 expressing axons. (Speicher et al., 1998)
Xenogenetic Interactions
Statement Reference
Complementation & Rescue Data
Fails to complement	Nrg⁸⁹² (Carhan et al., 2005) Nrg^ibx (Carhan et al., 2005) Nrg^l7 (Hall and Bieber, 1997) Nrg^l10 (Hall and Bieber, 1997)
Comments	Nrg[l4] fails to complement the reduced female receptivity phenotype of Nrg[ibx] and of Nrg[892]. Nrg[l4] fails to complement the reduced female receptivity phenotype of Nrg[892]. (Carhan et al., 2005)
Stocks ( 2 )
Bloomington	5708 Nrg^l4/FM7c
Kyoto	108384 Nrg^l4/FM7c
Notes on Origin
Discoverer	Lefevre.

External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 10 )
Reported As
Symbol Synonym	l(1)7Fa⁴ l(1)RA35 (Spradling et al., 1994, Hortsch and Goodman, 1993, Bieber et al., 1989, ) Nrg¹ (Faivre-Sarrailh et al., 2004) nrg¹ (Islam et al., 2003, Jenkins et al., 2001, Bouley et al., 2000, Speicher et al., 1998, Hall and Bieber, 1997, Yamamoto, 2006, Banerjee et al., 2006, Carhan et al., 2005) Nrg¹⁴ (Read et al., 2005, Genova and Fehon, 2003, Schwabe et al., 2005, Godenschwege et al., 2006) nrg¹⁴ (Stork et al., 2008, Hijazi et al., 2009, Moyer and Jacobs, 2008, Yang et al., 2011) Nrg^l4 (Popodi et al., 2010-, Mao and Freeman, 2009) nrg^l4 (Silies and Klämbt, 2010) RA35 (Eeken et al., 1985) unnamed (Kania and Bellen, 1995, King et al., 1986)
Name Synonym
Secondary FlyBase IDs

References ( 26 )

Generate a list of	All references relating to this allele ( 26 )

List References by type	Research paper ( 24 ) Supplementary material ( 1 ) Personal communication to FlyBase ( 1 )
Recent research papers ( 1 )
	Yang et al., 2011, Neuron 72(2): 285--299 Nak regulates localization of clathrin sites in higher-order dendrites to promote local dendrite growth. [FBrf0216477] Show all research papers ...

Allele Dmel\Nrgl4

Allele Dmel\Nrg^l4