Allele Dmel\Camⁿ³³⁹

General Information
Symbol	Dmel\Camⁿ³³⁹	Species	D. melanogaster
Name		FlyBase ID	FBal0048074
Feature type	allele	Associated gene	Dmel\Cam
Also Known As	Cam^null

Allele class	amorphic allele - genetic evidence, loss of function allele
Mutagen	P-element activity
Recent Updates
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FB2013_03
FB2013_02
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Nature of the Allele
Allele class	amorphic allele - genetic evidence (Heiman et al., 1996) loss of function allele (Scott et al., 1997, Harrisingh et al., 2007)
Mutagen	P-element activity (Heiman et al., 1996)
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	Cam² (Heiman et al., 1996, Arredondo et al., 1998)
Nature of the lesion	Statement Reference Mobilisation of the P{lacW} insertion generating a deletion of exons 0 and 1 and the initiator codon. (Heiman et al., 1996)
Cytology
Phenotypic Data
Phenotypic Class
	auditory perception defective \| adult stage (with Cam⁵) (Senthilan et al., 2012) behavior defective (Arredondo et al., 1998) behavior defective, with Cam⁷, Cam^Scer\UAS.cWa, Scer\GAL4^how-24B (Wang et al., 2003) behavior defective (with Cam⁷), with Cam^Scer\UAS.cWa, Scer\GAL4^how-24B (Wang et al., 2003) behavior defective \| larval stage (Heiman et al., 1996) flight defective (Arredondo et al., 1998) hypoactive \| larval stage (Wang et al., 2003) hypoactive \| larval stage (with Cam⁷) (Wang et al., 2003) lethal (with Cam⁷) (Wang et al., 2002) lethal \| larval stage, with Cam^{B34Q.Scer\UAS}/Cam⁷, Scer\GAL4^how-24B (Wang et al., 2002) lethal \| larval stage (with Cam⁷), with Cam^{B34Q.Scer\UAS}, Scer\GAL4^how-24B (Wang et al., 2002) lethal \| larval stage (with Cam^{B34Q.Scer\UAS}), with Cam⁷, Scer\GAL4^how-24B (Wang et al., 2002) lethal \| larval stage \| recessive (Heiman et al., 1996) lethal \| pharate adult stage (with Cam⁷) (Wang et al., 2003) lethal \| recessive (Nelson et al., 1997, Scott et al., 1997) locomotor behavior defective (Arredondo et al., 1998) locomotor behavior defective \| larval stage (Wang et al., 2003) locomotor behavior defective \| larval stage (with Cam³⁵²) (Wang et al., 2003) locomotor behavior defective \| larval stage (with Cam⁰³⁹⁰⁹) (Wang et al., 2003) mating defective, with Cam⁷, Cam^Scer\UAS.cWa, Scer\GAL4^how-24B (Wang et al., 2003) mating defective (with Cam⁷), with Cam^Scer\UAS.cWa, Scer\GAL4^how-24B (Wang et al., 2003) neuroanatomy defective (with Cam^3c1) (Arredondo et al., 1998) neurophysiology defective (with Cam³⁵²) (Liu et al., 2008) uncoordinated (with Cam^3c1) (Arredondo et al., 1998) visual behavior defective \| recessive (Scott et al., 1997)
Phenotype Manifest In
	adult head (with Cam⁷) (Wang et al., 2003) adult head \| P-stage (with Cam⁷) (Wang et al., 2002) bouton & abdominal 2 ventral longitudinal muscle 2 (Arredondo et al., 1998) bouton & abdominal 3 ventral longitudinal muscle 2 (Arredondo et al., 1998) bouton & abdominal 4 ventral longitudinal muscle 2 (Arredondo et al., 1998) bouton & abdominal 5 ventral longitudinal muscle 2 (Arredondo et al., 1998) bouton & abdominal 6 ventral longitudinal muscle 2 (Arredondo et al., 1998) bouton & abdominal 7 ventral longitudinal muscle 2 (Arredondo et al., 1998) longitudinal muscle (with Cam⁷) (Wang et al., 2003) ommatidium (with Cam³⁵²) (Liu et al., 2008) pupa (with Cam⁷) (Wang et al., 2002, Wang et al., 2003) pupal cuticle (with Cam⁷) (Wang et al., 2003) wing vein \| ectopic (Nelson et al., 1997)
Detailed Description
	Statement Reference Sound-evoked compound action potentials in the antennal nerve are not significantly different from wild type in Cam[5]/Cam[n339] flies. However, nonlinear mechanical amplification is significantly increased compared to wild type. (Senthilan et al., 2012) Camⁿ³³⁹ heterozygotes exhibit little effect on the average period of free-running. (Harrisingh et al., 2007) Homozygous larvae are sluggish and show spontaneous backward locomotion. Cam⁷/Camⁿ³³⁹ larvae are morphologically normal and show normal forward locomotion with no spontaneous backward movement. However, they are sluggish with a body wall contraction (BWC) rate about one-third that of controls. Cam⁷/Camⁿ³³⁹ wandering third instar larvae become progressively incapable of relaxing at the end of each body-wall contraction and show increasing stiffness during locomotion. Third instar body wall muscles that have been treated with ether (to artificially relax them) all show longitudinal muscles that have a "bunched" appearance, with structural disorganisation and misaligned myofibrils. Cam⁷/Camⁿ³³⁹ animals form pupal cases with deep indentations at the larval segment boundaries. On average the mutant larvae decrease their body length by 50% during pupariation, compared to a decrease of one-third in wild-type larvae. Only the long axis is abnormally compressed in the mutant animals. Cam⁷/Camⁿ³³⁹ pupae never eclose, but most develop into pharate adults with head defects. Three types of heads are seen; normal heads with no obvious defects, malformed heads that are partially everted and "inside-out" heads where head eversion has completely failed and head development proceeds in a noneverted head sac buried in the thorax. There is a correlation between the pupal length:width ratio and the degree of head abnormality; the more severe head defects are associated with shorter pupal cases. Cam⁷/Camⁿ³³⁹ animals rescued to adulthood by expression of Cam^Scer\UAS.cWa under the control of Scer\GAL4^how-24B are behaviourally abnormal; they fail to climb after being gently knocked to the bottom of a vial, and they show poor flight and a reduced ability to right themselves after a brief vortexing. The initial stages of male courtship are normal, but the rescued males cannot bend their abdomens sufficiently to achieve penetration. The amount of backward movement seen in Cam⁰³⁹⁰⁹/Camⁿ³³⁹ larvae is increased by high light intensity. The amount of backward movement seen in Cam³⁵²/Camⁿ³³⁹ larvae is increased by high light intensity. (Wang et al., 2003) Cam⁷/Camⁿ³³⁹ pupal cases are shorter than wild type and have deeply indented rings at the larval segment boundaries. Many animals have head eversion defects and none eclose. Expression of both Cam^{B12Q.Scer\UAS} and Cam^{B34Q.Scer\UAS} under the control of Scer\GAL4^how-24B in Cam⁷/Camⁿ³³⁹ animals results in 100% larval lethality. (Wang et al., 2002) Cam^3c1/Camⁿ³³⁹ flies exhibit reduced locomotion, coordination and flight ability. A slight dominant effect can be detected for both Cam^3c1 and Camⁿ³³⁹. The recessive mutant effects are more substantial. In the larval neuromuscular junction voltage clamp preparation, working on muscle 6, ejc amplitude for Cam^3c1/Camⁿ³³⁹ is normal. Ejc amplitude is increased approximately three fold by quinidine at low external calcium concentrations (at 0.4mM external calcium quinidine has no effect. The mejc amplitude and frequency is unchanged, suggesting the increase in ejc reflects increased neurotransmitter release. Cam^3c1/Camⁿ³³⁹ larvae show structural synaptic abnormalities: the terminal arbor forms a thickened, or large, misshapen structure with few distinct boutons. This results in a reduced number of boutons and a nearly complete lack of terminal branching in pleural external longitudinal muscle 13. Muscle 13 shows the same ejc phenotype as ventral internal longitudinal muscle 6. No abnormalities in nerve terminals on muscles ventral internal longitudinal muscle 6, 7 or pleural external longitudinal muscle 12 have been observed. (Arredondo et al., 1998) Transheterozygous combinations with other Cam mutations produces an incompletely penetrant ectopic wing vein phenotype. (Nelson et al., 1997) Camⁿ³³⁹/Cam³⁵² animals do not eclose from the pupal case. Camⁿ³³⁹/Cam³⁵² photoreceptors have dramatic defects in deactivation kinetics, displaying greatly prolonged deactivation times. Similar deactivation kinetics are seen at different concentrations of extracellular calcium ions, in contrast to wild-type. A single photon produces a train of quantum bumps in Cam³⁵²/Camⁿ³³⁹ double mutant photoreceptors, in contrast to wild-type. (Scott et al., 1997) Maternal Cam supports Camⁿ³³⁹ individuals throughout embryogenesis but they die within 2 days of hatching as first instar larvae. During feeding larvae show a high frequency of head swinging compared to heterozygous siblings. Larvae exhibit a significant decrease in the number of locomotive contraction waves due to a lower frequency of contraction initiation and most locomotion in the larvae is spontaneous backward movement. This spontaneous avoidance behaviour does not originate from any obvious morphological defects in the neuromuscular apparatus so starvation could be a potential cause of the aberrant locomotion. (Heiman et al., 1996)
External Data
Linkouts
Interactions
	Show the genetic interaction network for Enhancers & Suppressors
Phenotypic Class
Suppressed by
Statement Reference Cam⁷/Camⁿ³³⁹ has lethal phenotype, suppressible \| partially by Ca-α1D^X7/Ca-alpha1D[+] (Wang et al., 2003) Cam⁷/Camⁿ³³⁹ has lethal phenotype, suppressible by Rya-r44F¹⁶/Rya-r44F[+] (Wang et al., 2003)
NOT suppressed by
Statement Reference Cam⁷/Camⁿ³³⁹ has lethal phenotype, non-suppressible by Ca-α1D^AR66/Ca-alpha1D[+] (Wang et al., 2003) Cam⁷/Camⁿ³³⁹ has lethal phenotype, non-suppressible by cn[+]/cn¹ (Wang et al., 2003) Cam⁷/Camⁿ³³⁹ has lethal phenotype, non-suppressible by Df(2R)H3E1/+ (Wang et al., 2003)
Other
Statement Reference Cam⁷/Camⁿ³³⁹, Rya-r44F¹⁶/Rya-r44F[+] has locomotor behavior defective phenotype (Wang et al., 2003) Cam⁷/Camⁿ³³⁹, Rya-r44F¹⁶ has locomotor behavior defective phenotype (Wang et al., 2003) Camⁿ³³⁹, l(2)C43^C43 has lethal phenotype (Nelson et al., 1997, Nelson et al., 1997)
Phenotype Manifest In
NOT Enhanced by
Statement Reference Cam⁷/Camⁿ³³⁹ has adult head phenotype, non-enhanceable by cn[+]/cn¹ (Wang et al., 2003) Cam⁷/Camⁿ³³⁹ has pupa phenotype, non-enhanceable by cn[+]/cn¹ (Wang et al., 2003)
Suppressed by
Statement Reference Cam⁷/Camⁿ³³⁹ has adult head phenotype, suppressible \| partially by Df(2R)H3E1/+ (Wang et al., 2003) Cam⁷/Camⁿ³³⁹ has adult head phenotype, suppressible by Rya-r44F¹⁶/Rya-r44F[+] (Wang et al., 2003) Cam⁷/Camⁿ³³⁹ has pupal cuticle phenotype, suppressible by Rya-r44F¹⁶/Rya-r44F[+] (Wang et al., 2003) Cam⁷/Camⁿ³³⁹ has pupa phenotype, suppressible \| partially by Ca-α1D^AR66/Ca-alpha1D[+] (Wang et al., 2003) Cam⁷/Camⁿ³³⁹ has pupa phenotype, suppressible \| partially by Ca-α1D^X7/Ca-alpha1D[+] (Wang et al., 2003) Cam⁷/Camⁿ³³⁹ has pupa phenotype, suppressible \| partially by Df(2R)H3E1/+ (Wang et al., 2003) Cam⁷/Camⁿ³³⁹ has pupa phenotype, suppressible by Rya-r44F¹⁶/Rya-r44F[+] (Wang et al., 2003)
NOT suppressed by
Statement Reference Cam⁷/Camⁿ³³⁹ has adult head phenotype, non-suppressible by cn[+]/cn¹ (Wang et al., 2003) Cam⁷/Camⁿ³³⁹ has pupa phenotype, non-suppressible by cn[+]/cn¹ (Wang et al., 2003)
Other
Statement Reference Cam⁷/Camⁿ³³⁹, Rya-r44F¹⁶/Rya-r44F[+] has wing phenotype (Wang et al., 2003) Cam⁷/Camⁿ³³⁹, Rya-r44F¹⁶ has wing phenotype (Wang et al., 2003) Ca-α1D^X7, Cam⁷/Camⁿ³³⁹ has wing phenotype (Wang et al., 2003) Ca-α1D^X7/Ca-alpha1D[+], Cam⁷/Camⁿ³³⁹ has wing phenotype (Wang et al., 2003)
Additional Comments
Genetic Interactions
Statement Reference Rya-r44F¹⁶/+ rescues the pupal defects and lethality of Cam⁷/Camⁿ³³⁹ animals; the pupae are smooth and indentation free in the anterior and only have mild ridges in the posterior, their length:width ratios are actually higher than wild type, more than 40% of the pupae eclose and those that do not eclose do not show head defects. Some of the eclosed animals do not show wing expansion and they all perform poorly in a climb test. Df(2R)H3E1/+ has a slight suppressing effect on the Cam⁷/Camⁿ³³⁹ phenotype; pupae show some increase in the pupal length:width ratio, but very few pupal cases have decreased indentations, and fewer head defects are seen in pupae, but they fail to eclose. Ca-α1D^X7/+ has a slight suppressing effect on the Cam⁷/Camⁿ³³⁹ phenotype; there is a small but significant increase in the pupal length:width ratio, but no obvious effect on pupal case indentations, and a small fraction of the animals eclose as weak, uncoordinated adults with no wing expansion. Ca-α1D^AR66/+ has a slight suppressing effect on the Cam⁷/Camⁿ³³⁹ phenotype; there is an increase in the pupal length:width ratio, but none of the pupae eclose. (Wang et al., 2003) In combination with l(2)C43^C43 mutants exhibit completely penetrant lethality, mutants survive to pupation. (Nelson et al., 1997) trp⁹; Cam³⁵²/Camⁿ³³⁹ double mutant photoreceptors do not have transient light responses. (Scott et al., 1997)
Xenogenetic Interactions
Statement Reference A heterozygous Camⁿ³³⁹ background in flies expressing Mmmm\PV^{Scer\UAS.T:Hsap\MYC} under the control of Scer\GAL4^P2.4.Pdf has no effect on the length of the free-running period, compared to controls, after 1-5 weeks in constant darkness. (Harrisingh et al., 2007)
Complementation & Rescue Data
Partially complements	Cam^3c1 (Nelson et al., 1997) Cam^4c1 (Nelson et al., 1997) Cam^6c4 (Nelson et al., 1997)
Fails to complement	Cam⁵ (Nelson et al., 1997) Cam⁷ (Nelson et al., 1997) Cam^8t (Nelson et al., 1997)
Rescued by	Cam^3c1/Camⁿ³³⁹ is rescued by Cam^hs.PA (Arredondo et al., 1998) Cam⁷/Camⁿ³³⁹ is rescued by Scer\GAL4^how-24B/Cam^Scer\UAS.cWa/Scer\GAL4^how-24B (Wang et al., 2002) Cam³⁵²/Camⁿ³³⁹ is rescued by Cam^hs.PDH (Scott et al., 1997) Camⁿ³³⁹ is rescued by Cam^{V91G.Scer\UAS}/Scer\GAL4^elav.PLu (Wang et al., 2003)
Partially rescued by	Cam⁷/Camⁿ³³⁹ is partially rescued by Scer\GAL4^how-24B/Cam^Scer\UAS.cWa/Scer\GAL4^how-24B (Wang et al., 2003) Cam⁷/Camⁿ³³⁹ is partially rescued by Scer\GAL4^how-24B/Scer\GAL4^how-24B/Cam^{B12Q.Scer\UAS} (Wang et al., 2002) Camⁿ³³⁹ is partially rescued by Cam^Scer\UAS.cWa/Scer\GAL4^elav.PLu (Wang et al., 2003)
Not rescued by	Cam⁷/Camⁿ³³⁹ is not rescued by Cam^{B34Q.Scer\UAS}/Scer\GAL4^how-24B/Scer\GAL4^how-24B (Wang et al., 2002) Cam⁷/Camⁿ³³⁹ is not rescued by Cam^{B1234Q.Scer\UAS}/Scer\GAL4^how-24B/Scer\GAL4^how-24B (Wang et al., 2002) Cam⁷/Camⁿ³³⁹ is not rescued by Cam^Scer\UAS.cWa/Scer\GAL4^elav.PLu (Wang et al., 2003) Cam⁷/Camⁿ³³⁹ is not rescued by Scer\GAL4^sca.PU/Cam^Scer\UAS.cWa (Wang et al., 2003) Camⁿ³³⁹ is not rescued by Cam^{B12Q.Scer\UAS}/Scer\GAL4^elav.PLu (Wang et al., 2003) Camⁿ³³⁹ is not rescued by Cam^{B34Q.Scer\UAS}/Scer\GAL4^elav.PLu (Wang et al., 2003) Camⁿ³³⁹ is not rescued by Cam^{B1234Q.Scer\UAS}/Scer\GAL4^elav.PLu (Wang et al., 2003) Camⁿ³³⁹ is not rescued by Scer\GAL4^how-24B/Cam^Scer\UAS.cWa/Scer\GAL4^how-24B (Wang et al., 2003) Camⁿ³³⁹ is not rescued by Scer\GAL4^sca.PU/Cam^Scer\UAS.cWa (Wang et al., 2003)
Comments	The lethality and pupariation defects seen in Cam⁷/Camⁿ³³⁹ animals are rescued by expression of Cam^Scer\UAS.cWa under the control of Scer\GAL4^how-24B; the pupal cases are morphologically wild type and have more normal length:width ratios, almost all of the rescued pupae eclose and no aberrant pharate head structures are seen. The lethality and pupariation defects seen in Cam⁷/Camⁿ³³⁹ animals are not rescued by expression of Cam^Scer\UAS.cWa under the control of Scer\GAL4^elav.PLu or Scer\GAL4^sca.PU. The sluggish larval locomotion of Cam⁷/Camⁿ³³⁹ animals is incompletely rescued by expression of Cam^Scer\UAS.cWa under the control of Scer\GAL4^how-24B or Scer\GAL4^elav.PLu. The spontaneous backward movement seen in homozygous Camⁿ³³⁹ larvae is not rescued by expression of Cam^Scer\UAS.cWa under the control of Scer\GAL4^how-24B or Scer\GAL4^sca.PU. The spontaneous backward movement seen in homozygous Camⁿ³³⁹ larvae is rescued by expression of Cam^Scer\UAS.cWa under the control of Scer\GAL4^elav.PLu, although the larvae are still sluggish. (Wang et al., 2003) The Camⁿ³³⁹/Cam³⁵² photoreceptor deactivation time phenotype can be partially rescued by Cam^hs.PDH, the degree of rescue depending on the number of heat shocks given. (Scott et al., 1997)
Stocks ( 6 )
Bloomington	6806 y¹ w^; Camⁿ³³⁹/CyO, y⁺ 6809 y¹ w^; Camⁿ³³⁹/CyO, y⁺; P{UAS-Cam.W}3 6805 y¹ w^; Camⁿ³³⁹/CyO, y⁺; P{UAS-Cam.B34Q}3 6808 y¹ w^; Camⁿ³³⁹/CyO, y⁺; P{UAS-Cam.B12Q}3 6807 y¹ w^; Camⁿ³³⁹/CyO, y⁺; P{UAS-Cam.V91G}3/+ 8472 y¹ w^; Camⁿ³³⁹/CyO, y⁺; P{UAS-Cam.B1234Q}3
Notes on Origin
Discoverer

External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 4 )
Reported As
Symbol Synonym	cam³³⁹ (Liu et al., 2008) camⁿ³³⁹ (Alloway and Dolph, 1999) Camⁿ³³⁹ (Christensen et al., 2008.4.15, Harrisingh et al., 2007, Senthilan et al., 2012) Cam^null (Arredondo et al., 1998, Nelson et al., 1997)
Name Synonym
Secondary FlyBase IDs

References ( 11 )
Research paper	Senthilan et al., 2012, Cell 150(5): 1042--1054 Drosophila auditory organ genes and genetic hearing defects. [FBrf0219321] Liu et al., 2008, Neuron 59(5): 778--789 Ca2+-dependent metarhodopsin inactivation mediated by calmodulin and NINAC myosin III. [FBrf0205960] Harrisingh et al., 2007, J. Neurosci. 27(46): 12489--12499 Intracellular Ca[2+] Regulates Free-Running Circadian Clock Oscillation In Vivo. [FBrf0204998] Wang et al., 2003, Genetics 165(3): 1255--1268 Drosophila calmodulin mutants with specific defects in the musculature or in the nervous system. [FBrf0167631] Wang et al., 2002, genesis 34(1-2): 86--90 Calmodulin UAS-constructs and the in vivo roles of calmodulin: analysis of a muscle-specific phenotype. [FBrf0152385] Alloway and Dolph, 1999, Proc. Natl. Acad. Sci. U.S.A. 96(11): 6072--6077 A role for the light-dependent phosphorylation of visual arrestin. [FBrf0108568] Arredondo et al., 1998, Genetics 150(1): 265--274 Increased transmitter release and aberrant synapse morphology in a Drosophila calmodulin mutant. [FBrf0104398] Nelson et al., 1997, Genetics 147(4): 1783--1798 Calmodulin point mutations affect Drosophila development and behavior. [FBrf0099767] Scott et al., 1997, Cell 91(3): 375--383 Calmodulin regulation of Drosophila light-activated channels and receptor function mediates termination of the light response in vivo. [FBrf0099505] Heiman et al., 1996, Proc. Natl. Acad. Sci. U.S.A. 93(6): 2420--2425 Spontaneous avoidance behavior in Drosophila null for calmodulin expression. [FBrf0086449]
Personal communication to FlyBase	Christensen et al., 2008.4.15, Isolation and characterization of Df(2R)BSC463. Isolation and characterization of Df(2R)BSC463. [FBrf0205026]

Allele Dmel\Camn339

Allele Dmel\Camⁿ³³⁹