Allele Dmel\ben¹

General Information
Symbol	Dmel\ben¹	Species	D. melanogaster
Name		FlyBase ID	FBal0001101
Feature type	allele	Associated gene	Dmel\ben

Map ( GBrowse )
Allele class	amorphic allele - genetic evidence
Mutagen	ethyl methanesulfonate
Recent Updates
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FB2012_01
FB2011_10
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Nature of the Allele
Allele class	amorphic allele - genetic evidence (Thomas and Wyman, 1984, Edgecomb et al., 1993)
Mutagen	ethyl methanesulfonate (Oh et al., 1994)
Mutations Mapped to the Genome

Type Location Additional Notes References point mutation X:13,892,482..13,892,482 na_change=C13892482T evidence=experimental pr_change=P97S\|ben-PA reported_na_change=C490T reported_pr_change=P97S (Muralidhar and Thomas, 1993)
Associated Sequence Data
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name

UniProtKB/Swiss-Prot
UniProtKB/TrEMBL

Progenitor genotype
Nature of the lesion	Statement Reference Nucleotide substitution: C490T. Amino acid replacement: P97S. (Muralidhar and Thomas, 1993)
Cytology
Phenotypic Data
Phenotypic Class
	behavior defective \| recessive (Thomas and Wyman, 1984) grooming behavior defective \| recessive (Phillis et al., 1993) jumping defective \| recessive (Thomas and Wyman, 1984, ) lethal \| embryonic stage \| non-rescuable maternal effect (with Df(1)KA10) (Merkle et al., 2009) lethal \| embryonic stage \| non-rescuable maternal effect \| recessive (Merkle et al., 2009) lethal \| partially \| recessive (Oh et al., 1994) lethal \| pupal stage \| recessive (Edgecomb et al., 1993) locomotor behavior defective (Edgecomb et al., 1993) locomotor behavior defective \| recessive (Oh et al., 1994) neuroanatomy defective (Muralidhar and Thomas, 1993) neuroanatomy defective \| recessive (Oh et al., 1994) uncoordinated \| recessive (Oh et al., 1994)
Phenotype Manifest In
	centrosome \| non-rescuable maternal effect (Merkle et al., 2009) centrosome \| non-rescuable maternal effect (with Df(1)KA10) (Merkle et al., 2009) coxal tergal remotor muscle (Edgecomb et al., 1993) giant fiber neuron (Muralidhar and Thomas, 1993, Oh et al., 1994) lamina anlage (Muralidhar and Thomas, 1993) lateral oblique dorsal muscle (Edgecomb et al., 1993) medulla anlage (Muralidhar and Thomas, 1993) mesothoracic extracoxal depressor muscle 66 (Oh et al., 1994, Edgecomb et al., 1993) optic cartridge (Oh et al., 1994) photoreceptor cell R1 (Oh et al., 1994) photoreceptor cell R2 (Oh et al., 1994) photoreceptor cell R3 (Oh et al., 1994) photoreceptor cell R4 (Oh et al., 1994) photoreceptor cell R5 (Oh et al., 1994) photoreceptor cell R6 (Oh et al., 1994) photoreceptor cell R7 (Oh et al., 1994) photoreceptor cell R8 (Oh et al., 1994) retina (Muralidhar and Thomas, 1993) rhabdomere R1 (Muralidhar and Thomas, 1993, Oh et al., 1994) rhabdomere R2 (Muralidhar and Thomas, 1993, Oh et al., 1994) rhabdomere R3 (Muralidhar and Thomas, 1993, Oh et al., 1994) rhabdomere R4 (Muralidhar and Thomas, 1993, Oh et al., 1994) rhabdomere R5 (Muralidhar and Thomas, 1993, Oh et al., 1994) rhabdomere R6 (Muralidhar and Thomas, 1993, Oh et al., 1994) rhabdomere R7 (Muralidhar and Thomas, 1993, Oh et al., 1994) rhabdomere R8 (Oh et al., 1994) spindle \| non-rescuable maternal effect (Merkle et al., 2009) spindle \| non-rescuable maternal effect (with Df(1)KA10) (Merkle et al., 2009) tergosternal muscle (Edgecomb et al., 1993)
Detailed Description
	Statement Reference Embryos derived from homozygous females mated to wild-type males fail to develop; 72% contain only one nucleus, 12% contain 2-8 nuclei ad the remaining 16% have more than 8 nuclei. Only 2% hatch. 72% of the embryos have one acentrosomal spindle. These single spindles appear to be mitotic rather than meiotic; their presence requires fertilisation, they are positioned deep within the egg interior where the first mitotic spindle normally resides, polar bodies are present (indicating completion of meiotic divisions) and centrosomes are occasionally seen near the spindle. 80% of the embryos have a majority of spindles that are barrel-shaped and/or lacking centrosomes. Misaligned chromosomes are often seen. Embryos derived from ben[1]/Df(1)KA10 females mated to wild-type males fail to develop; 40% contain only one nucleus, 43% contain 2-8 nuclei ad the remaining 17% have more than 8 nuclei. None of these embryos hatch. 50% of the embryos have a majority of spindles that are barrel-shaped and/or lacking centrosomes. (Merkle et al., 2009) Homozygous flies appear lethargic and uncoordinated, and do not show normal climbing behaviour. Adults are capable of flight, but will not initiate flight when dropped from a height. Viability is reduced, with approximately 60% of pupae failing to eclose successfully and dying during emergence. The giant fibre (GF) drives the tergotrochanteral muscle (TTM) at abnormally long latencies and the response fails completely at moderate frequencies in homozygous flies. Stimulation of the dorsal longitudinal muscle (DLM), and the three dorsoventral muscles (DVM I, II and III) by the GF appears normal. The terminal bend of the GF is abnormal. Gynandromorph analysis suggests that the GF phenotype is determined by the genotype of the head, which contains the cell body of the giant axon. Homozygous flies prefer visible light over UV light in a choice test, in contrast to wild-type flies. Photoreceptor R7 cell rhabdomeres appear deformed and displaced, and rhabdomeres of other photoreceptor cells may also show less severe abnormalities. The arrangement of the optic cartridges within the lamina is completely disrupted. Photoreceptor axons which reach the medulla make shallow disordered projections into it, and photoreceptor axon projections appear irregular and disordered after exiting the optic stalk. (Oh et al., 1994) TDT attachment sites vary. The muscle may be found in the wild type position, posterior to the intrascutal suture and displaced medially, anterior to the intrascutal suture or missing entirely. Muscle may also be reduced in size or split dorsally attaching to two separate sites on the scutum. Cytology of the TDT is also altered, muscles may have fibres that are swollen and stain abnormally, other fibres may have large, axially aligned holes. Attachment pattern of the DVMs is also altered. DVMs remain within their respective attachment regions or extend into the region normally occupied by an adjacent DVM. DVMs may be entirely missing. Many flies fail to eclose, dying during late pupal stages. Flies that do eclose show impaired motility. Flies can get stuck in their food, falling into the food causes their femurs to become swollen and bent. (Edgecomb et al., 1993) Giant fibre (GF) fails to extend laterally along the tergotrochanteral jump muscle motorneuron (TTMmn) and instead terminates at the midline where it synapses with the peripherally synapsing interneuron (PSI). This results in a lack of direct synaptic connection between the GF and TTMmn. (Muralidhar and Thomas, 1993) Ethanol is capable of evoking a walk response at concentrations lower than 10%. (Venard and Stocker, 1991)
External Data
Linkouts
Interactions
Phenotypic Class
Phenotype Manifest In
Additional Comments
Genetic Interactions
Statement Reference The giant fibre (GF) of Ubx^abx-1 Ubx^bx-3 Ubx^pbx-1/Df(3R)P2 mutants makes a midline tuft and extends a lateral process to T2 and T3 segment. In the presence of ben¹ the GF is capable of extending a process into the T3 segment but no lateral processes from the midline. R7 rhabdomeres are reduced in size and often misplaced, outer rhabdomeres are loosely packed maintaining position further from the centre of the ommatidia. The ordered arrangement of photoreceptor neuron axons entering the lamina is disrupted. Also the medula is abnormally rotated in relation to the lamina. (Muralidhar and Thomas, 1993)
Xenogenetic Interactions
Statement Reference
Complementation & Rescue Data
Rescued by	ben¹ is rescued by ben^+t6 (Oh et al., 1994) ben¹ is rescued by ben^+t8 (Muralidhar and Thomas, 1993) ben¹ is rescued by ben^+t10 (Muralidhar and Thomas, 1993) ben¹ is rescued by ben^+t18 (Oh et al., 1994) ben¹ is rescued by ben^+t35 (Muralidhar and Thomas, 1993)
Comments
Stocks ( 1 )
Bloomington	9565 w¹¹⁸ ben¹/C(1)A, y¹
Notes on Origin
Discoverer	Wyman and Thomas.

Comments
	Allelic series of TDT phenotype: ben^P2/ben^P2 < ben^P2/Df(1)HA92 < ben¹/ben^P2 < ben¹/ben¹ = ben¹/Df(1)HA92. (Edgecomb et al., 1993) The defect in the pathway responsible for the abnormal TTM muscle response is at the GF-TTM motor neuron junction. (Thomas and Wyman, 1984)
External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 2 )
Reported As
Symbol Synonym	ben¹ (Merkle et al., 2009) unnamed (Phillis et al., 1993)
Name Synonym
Secondary FlyBase IDs

References ( 8 )
Research paper	Merkle et al., 2009, Development 136(3): 449--459 no poles encodes a predicted E3 ubiquitin ligase required for early embryonic development of Drosophila. [FBrf0206570] Oh et al., 1994, J. Neurosci. 14(5 Pt. 2): 3166--3179 bendless, a Drosophila gene affecting neuronal connectivity, encodes a ubiquitin-conjugating enzyme homolog. [FBrf0074062] Edgecomb et al., 1993, J. Neurogenet. 8(4): 201--219 Bendless alters thoracic musculature in Drosophila. [FBrf0058914] Muralidhar and Thomas, 1993, Neuron 11(2): 253--266 The Drosophila bendless gene encodes a neural protein related to ubiquitin-conjugating enzymes. [FBrf0063736] Phillis et al., 1993, Genetics 133(3): 581--592 Isolation of mutations affecting neural circuitry required for grooming behavior in Drosophila melanogaster. [FBrf0058570] Venard and Stocker, 1991, J. Insect Behav. 4(6): 683--705 Behavioral and electroantennogram analysis of olfactory stimulation in lozenge: a Drosophila mutant lacking antennal basiconic sensilla (Diptera: Drosophilidae). [FBrf0074701] Thomas and Wyman, 1984, J. Neurosci. 4: 530--538 Mutations altering synaptic connectivity between identified neurons in Drosophila. [FBrf0041358]
Book	Lindsley and Zimm, 1992, The Genome of Drosophila melanogaster. The Genome of Drosophila melanogaster. [FBrf0066905]

Allele Dmel\ben1

Allele Dmel\ben¹