Allele Dmel\chico¹

General Information
Symbol	Dmel\chico¹	Species	D. melanogaster
Name		FlyBase ID	FBal0031303
Feature type	allele	Associated gene	Dmel\chico
Also Known As	chico1, fs(2)4¹

Allele class	amorphic allele - molecular evidence, loss of function allele
Mutagen	P-element activity
Recent Updates
Description	What does this section display? This section contains items that were added to this record for each release. It currently only tracks new links between this FlyBase report and other FlyBase data classes (e.g. genes, references, stocks) or controlled vocabulary terms (e.g. GO, anatomy terms). What does this section not display? This section does not currently display links that were removed or gene model changes.
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FB2013_03
FB2013_02
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Nature of the Allele
Allele class	amorphic allele - molecular evidence (Naganos et al., 2012) loss of function allele (Shingleton et al., 2005)
Mutagen	P-element activity (Berg and Spradling, 1991, Riesgo-Escovar, 1998.3.10, Bohni et al., 1999)
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 P{ry11} is inserted 80bp downstream of the putative translation initiation site in the PH domain. (Bohni et al., 1999) P{ry11} is inserted into, and in the same orientation as, the chico transcription unit. (Riesgo-Escovar et al., 1996)
Caused by insertion	P{ry11}chico¹ (Berg and Spradling, 1991, Takeo et al., 2005, Bohni et al., 1999, Wessells et al., 2004, Riesgo-Escovar, 1998.3.10, Parrish et al., 2009)
Cytology
Phenotypic Data
Phenotypic Class
	decreased cell number (with chico^flp147E) (Junger et al., 2003) decreased cell size (with chico^flp147E) (Junger et al., 2003) developmental rate defective \| recessive (Bohni et al., 1999) female sterile (Clancy et al., 2001, Richard et al., 2005) female sterile \| recessive (Spradling, 1990.8.1, Clegg et al., 1993) learning defective (with chico^flp147E) (Naganos et al., 2012) learning defective \| recessive (Naganos et al., 2012) long lived (Clancy et al., 2001) long lived \| dominant (Wessells et al., 2004) long lived \| dominant \| female limited (Albrecht et al., 2011) male semi-sterile \| recessive (Spradling, 1990.8.1) neuroanatomy defective \| adult stage (Naganos et al., 2012) partially lethal - majority die (Naganos et al., 2012) partially lethal - majority die (with chico^flp147E) (Naganos et al., 2012) partially lethal - majority die \| recessive (Spradling, 1990.8.1) partially lethal - majority live \| recessive (Riesgo-Escovar et al., 1998) size defective (Shingleton et al., 2005) size defective \| larval stage (Parrish et al., 2009) small body (Hulbert et al., 2004, Wessells et al., 2004, Bohni et al., 1999, Richard et al., 2005) small body (with chico^flp147E) (Junger et al., 2003) small body \| adult stage (Wittwer et al., 2005, Naganos et al., 2012) small body \| recessive (Clancy et al., 2001) stress response defective (Vigne et al., 2009) viable (Clancy et al., 2001) visible (with chico^flp147E) (Junger et al., 2003) visible \| recessive (Hulbert et al., 2004, Bohni et al., 1999)
Phenotype Manifest In
	adult (with chico^flp147E) (Junger et al., 2003) adult brain (Naganos et al., 2012) adult gut (Biteau et al., 2010) adult heart (Wessells et al., 2004) adult mushroom body (Naganos et al., 2012) egg chamber (Drummond-Barbosa and Spradling, 2001) eye (Wittwer et al., 2005) eye (with chico^flp147E) (Junger et al., 2003) eye \| cell autonomous \| somatic clone (Bohni et al., 1999) follicle cell (Drummond-Barbosa and Spradling, 2001) genital arch \| male \| somatic clone (Shingleton et al., 2005) head capsule \| somatic clone (Bohni et al., 1999) Kenyon cell (Naganos et al., 2012) larva (Parrish et al., 2009) maxillary palpus \| male \| somatic clone (Shingleton et al., 2005) nurse cell (Richard et al., 2005) ommatidium (Wittwer et al., 2005) ommatidium (with chico^flp147E) (Junger et al., 2003) ovary (Richard et al., 2005) ovary (with chico^flp147E) (Werz et al., 2009) wing (Shingleton et al., 2005) wing (with chico^flp147E) (Junger et al., 2003)
Detailed Description
	Statement Reference Homozygous and chico[1]/chico[flp147E] adults show normal odour and electric shock reactivity. Homozygous and chico[1]/chico[flp147E] adults show significant impairment in 3 minute, 1 hour and 3 hour memory after olfactory aversive conditioning. Analysis using a short-duration training protocol indicates that the defect appears to be in the initial learning, since after normalising 3 minute memory in chico[1] and wild-type flies, subsequent memory decay is indistinguishable in the two genotypes. The brains of homozygous flies are significantly smaller than normal, with a significant reduction in the number of Kenyon cells and in the size of the mushroom bodies. (Naganos et al., 2012) Heterozygous females show an extension of median lifespan compared to wild-type controls. These females show accelerated accumulation of cytosolic hydrogen peroxide in the gut compared to wild-type controls. Heterozygous males do not show an extension of median lifespan compared to wild-type controls. These males show a delayed accumulation of cytosolic hydrogen peroxide in the gut compared to wild-type controls. (Albrecht et al., 2011) The frequency of proliferating cells in the intestine of aging chico[1] homozygotes is significantly lower than that of age-matched controls. (Biteau et al., 2010) Heterozygous flies show increased resistance to dry starvation compared to control flies. (Hull-Thompson et al., 2009) The length of the dorsal hemisegments along the antero-posterior axis in chico[1] mutant larvae is only 10% of that of wild-types. (Parrish et al., 2009) In a fed state, chico[1] flies are anoxia tolerant (heterozygotes are not). (Vigne et al., 2009) chico[1]/chico[flp147E] flies have elevated levels of lipids compared to control flies. (Werz et al., 2009) Heterozygous ovaries transplanted into homozygous females usually show a robust vitellogenic response, with vitellogenic oocytes spanning the entire range of stage 8 to mature stage 14 eggs being seen in the transplanted ovaries. In all cases, the oocytes of the homozygous host female terminate development at the previtellogenic stage 7. Homozygous ovaries transplanted into heterozygous or wild-type females fail to develop oocytes beyond stage 7, even though the host ovaries become fully vitellogenic in each recipient female. Methoprene does not stimulate vitellogenesis or egg production in homozygous females. On day 1 post-eclosion, homozygous females contain an average of 11.2 nurse cells per egg chamber, and on day 4 they have an average of 14.8 nurse cells per egg chamber (a statistically significant difference). (Richard et al., 2005) chico¹ mutants show a reduction in wing size compared to wild-type controls. However, this phenotype is not temperature sensitive as chico¹ flies raised at 25^oC show a similar decrease in wing area compared to chico¹ flies raised at 18^oC to the decrease in wing area observed for control flies at these two temperatures. Males that carry chico¹ Minute clones throughout one side of the genitals and are heterozygous for chico¹ on the other side show smaller genital arches on the homozygous clone side than on the heterozygous side. The difference in size is 16%. Maxillary palps consisting of chico¹ homozygous clones are 45% smaller than paired palps on the same male. (Shingleton et al., 2005) chico¹ homozygous adults are abnormally small and have reduced numbers of ommatidia per eye. (Wittwer et al., 2005) chico¹ homozygous mutants are small, weighing on average 44% of the mass of wild-types. Mean oxygen consumption is not significantly higher than in wild-type flies. Mass-specific resting heat production is the same in chico¹ mutants as wild-type flies. Dietary-restriction of chico¹ mutants has no effect on resting heat production. While the body mass of wild-type flies fed on an energy restricted medium is reduced, chico¹ mutants show no decrease in body mass. (Hulbert et al., 2004) chico¹ flies are small and live ~50% longer than wild-type flies. These flies have low rates of pacing-induced heart failure and arrest, and their heart beat frequency does not increase with age, compared to wild-type flies. chico¹/+ flies have lifespans that are longer than wild-type flies, but shorter than chico¹ homozygotes. Like wild-type flies, these chico¹/+ flies experience stress-induced heart failure and their heart beat frequency decreases with age. There is no difference between male or female flies for any of the heart functions tested in homozygous or heterozygous chico¹ flies. (Wessells et al., 2004) chico¹/chico^flp147E flies show lower body weight, fewer ommatidia, smaller wings, fewer cells per wing and smaller wing cell area than wild type. (Junger et al., 2003) Photoreceptor cell projection patterns are indistinguishable from wild type in homozygous larvae and in mosaic larvae in which homozygous clones have been induced in the eye. The pattern of R7 and R8 projections in the medulla is similar to wild type in mosaic adults in which homozygous clones have been induced in the eye. Heterozygotes have normal photoreceptor cell projections in the medulla. (Song et al., 2003) The relation between life-span and food concentration is right-shifted in chico¹ females compared to controls. Control and chico¹ females show similar peak life-spans under dietary restriction, but chico¹ females show a peak in mean lifespan at a food concentration 0.8 times that of normal, while control females show a peak in mean lifespan at a food concentration 0.65 times that of normal. (Clancy et al., 2002) chico¹ adults are resistant to starvation conditions compared to wild type. (Oldham et al., 2002) Homozygous adults are dwarf in size. Homozygous females show an increase of median and maximum life-span of up to 48 and 41% respectively compared to controls. Homozygous males are slightly short-lived. Heterozygous females and males show an increase in median life-span of up to 36 and 13% respectively. Heterozygous females have reduced fecundity and homozygotes are almost sterile. Mutant flies show no resistance to heat stress at 37^oC. Slight resistance to oxidative stress (methyl viologen) is seen in heterozygotes but not in homozygotes. Heterozygotes and homozygotes show some resistance to starvation. (Clancy et al., 2001) The chico¹ mutation partially impaired the ability of follicle cells to proliferate faster in the presence of abundant nutrients. Egg chambers do not develop beyond vitellogenic stages in homozygotes, even in the presence of abundant food. (Drummond-Barbosa and Spradling, 2001) chico¹ flies do not show unexpanded wings, are not dark and do not show male sterility. (Riesgo-Escovar, 2001.2.7) Homozygous clones are not visible in the adult eye because they are out-competed by heterozygous cells. (Wittwer et al., 2001) Homozygosity for chico¹ causes semilethality and an overall delay in development. Flies eclose 2-3 days after their heterozygous siblings. In non-crowded culture conditions, homozygous mutant mothers can produce few viable progeny lacking both maternal and zygotic chico function. Homozygotes have a drastic weight reduction (65% in females, 55% in males) compared to wild-type. Body size reduction is observed at all developmental stages but does not alter the overall proportions of the flies. Homozygous clones in the eye produce morphologically normal ommatidia, except that they are more than 50% smaller than wild-type. Otherwise normal ommatidia with both smaller mutant and larger wild-type photoreceptors are observed at the edges of clones, indicating that chico acts cell autonomously. Autonomy of cell size control is also observed in mutant clones in the wing. Selective removal of chico¹ function in the eye imaginal disc cells generates flies with a strongly reduced head capsule and reduced eyes, whereas the proboscis and the rest of the body are of wild type size. Homozygous mutant clones in the adult eye are rarer, more variable in size and on average smaller than their wild-type sister clones. Clones are more frequently observed in the anterior half of the eye around the equator. In the eye disc, clones are also more variable in size and on average smaller than their wild-type sister clones. TUNEL analysis shows no increase in apoptotic cells in homozygous clones. Clones produced in a Minute background do not reveal enhanced apoptosis compared to wild type clones, or any increase in morphological signs of programmed cell death, in either the imaginal discs or the adult eye. No significant difference is seen in the apoptotic sub-G1 fraction of homozygous mutant cells compared to heterozygous cells. (Bohni et al., 1999) Mutant flies develop slower than sibling controls and have a reduced body size. chico mutant cells differentiate normally but are significantly reduced in size. (Riesgo-Escovar et al., 1998) male fertility poor female-sterile
External Data
Linkouts
Interactions
	Show the genetic interaction network for Enhancers & Suppressors
Phenotypic Class
Enhanced by
Statement Reference chico^flp147E/chico¹ has decreased cell size phenotype, enhanceable by foxo²⁵/foxo²¹ (Junger et al., 2003)
Suppressed by
Statement Reference chico¹, foxo^Δ94 has lethal phenotype, suppressible by foxo^{Scer\UAS.P\T.cSa}/Scer\GAL4^da.G32 (Slack et al., 2011) chico¹ has small body \| adult stage phenotype, suppressible \| partially by B4^susi-1 (Wittwer et al., 2005) chico^flp147E/chico¹ has decreased cell number phenotype, suppressible by foxo²¹/foxo[+] (Junger et al., 2003) chico^flp147E/chico¹ has decreased cell number phenotype, suppressible by foxo²⁵/foxo²¹ (Junger et al., 2003) chico^flp147E/chico¹ has small body phenotype, suppressible \| partially by foxo²⁵/foxo²¹ (Junger et al., 2003) chico^flp147E/chico¹ has visible phenotype, suppressible by foxo²¹/foxo[+] (Junger et al., 2003) chico^flp147E/chico¹ has visible phenotype, suppressible by foxo²⁵/foxo²¹ (Junger et al., 2003)
NOT suppressed by
Statement Reference chico^flp147E/chico¹ has decreased cell size phenotype, non-suppressible by foxo²¹/foxo[+] (Junger et al., 2003)
NOT Enhancer of
Statement Reference chico¹/chico[+] is a non-enhancer of size defective \| adult stage phenotype of sl² (Murillo-Maldonado et al., 2011) chico¹ is a non-enhancer of visible phenotype of os^GMR.PB (Bach et al., 2003)
Suppressor of
Statement Reference chico¹/chico[+] is a suppressor \| partially of increased cell number \| adult stage phenotype of sl² (Murillo-Maldonado et al., 2011)
NOT Suppressor of
Statement Reference chico¹/chico[+] is a non-suppressor of large body phenotype of Scer\GAL4^29BD, dally^{sec.Scer\UAS.T:Hsap\MYC} (Takeo et al., 2005) chico¹ is a non-suppressor of lethal \| pupal stage phenotype of Csk^j1D8/Csk^S030003 (Read et al., 2004) chico¹ is a non-suppressor of visible phenotype of os^GMR.PB (Bach et al., 2003)
Other
Statement Reference chico¹, foxo^Δ94 has lethal phenotype (Slack et al., 2011) Df(2L)Exel6026/+, Lnk^6S2/Lnk^4Q3, chico¹ has viable phenotype (Werz et al., 2009) Lnk^6S2/Lnk^4Q3, chico^flp147E/chico¹ has lethal phenotype (Werz et al., 2009)
Phenotype Manifest In
Enhanced by
Statement Reference chico^flp147E/chico¹ has ommatidium phenotype, enhanceable by foxo²⁵/foxo²¹ (Junger et al., 2003)
Suppressed by
Statement Reference chico¹ has ommatidium phenotype, suppressible by B4⁰⁵³³⁷ (Wittwer et al., 2005) chico¹ has ommatidium phenotype, suppressible by B4^susi-1 (Wittwer et al., 2005) chico^flp147E/chico¹ has ommatidium phenotype, suppressible by foxo²¹/foxo[+] (Junger et al., 2003) chico^flp147E/chico¹ has ommatidium phenotype, suppressible by foxo²⁵/foxo²¹ (Junger et al., 2003) chico^flp147E/chico¹ has wing phenotype, suppressible by foxo²¹/foxo[+] (Junger et al., 2003) chico^flp147E/chico¹ has wing phenotype, suppressible by foxo²⁵/foxo²¹ (Junger et al., 2003)
NOT suppressed by
Statement Reference chico^flp147E/chico¹ has ommatidium phenotype, non-suppressible by foxo²¹/foxo[+] (Junger et al., 2003)
Enhancer of
Statement Reference chico¹/chico[+] is an enhancer of eye disc phenotype of Pten^Scer\UAS.cHa, Scer\GAL4^ey.PH (Huang et al., 1999) chico¹/chico[+] is an enhancer of eye phenotype of Pten^Scer\UAS.cHa, Scer\GAL4^ey.PH (Huang et al., 1999) chico¹/chico[+] is an enhancer of ommatidium phenotype of Pten^Scer\UAS.cHa, Scer\GAL4^ey.PH (Huang et al., 1999)
NOT Enhancer of
Statement Reference chico¹/chico[+] is a non-enhancer of wing blade phenotype of sl² (Murillo-Maldonado et al., 2011) chico¹/chico[+] is a non-enhancer of wing vein \| ectopic phenotype of sl² (Murillo-Maldonado et al., 2011) chico¹ is a non-enhancer of eye phenotype of os^GMR.PB (Bach et al., 2003)
Suppressor of
Statement Reference chico¹/chico[+] is a suppressor \| partially of photoreceptor cell R7 \| adult stage phenotype of sl² (Murillo-Maldonado et al., 2011)
NOT Suppressor of
Statement Reference chico¹/chico[+] is a non-suppressor of pupa phenotype of Scer\GAL4^29BD, dally^{sec.Scer\UAS.T:Hsap\MYC} (Takeo et al., 2005) chico¹/chico[+] is a non-suppressor of wing phenotype of Scer\GAL4^29BD, dally^{sec.Scer\UAS.T:Hsap\MYC} (Takeo et al., 2005) chico¹ is a non-suppressor of eye phenotype of os^GMR.PB (Bach et al., 2003)
Additional Comments
Genetic Interactions
Statement Reference One copy of chico[1] does not enhance the reduction in wing blade area seen in homozygous sl[2] males. One copy of chico[1] does not enhance the ectopic wing vein phenotype seen in sl[2] homozygotes. One copy of chico[1] partially suppresses the percentage of sl[2] mutant ommatidia that contain extra R7 photoreceptors. (Murillo-Maldonado et al., 2011) Expression of NLaz[Scer\UAS.cHTa] under the control of Scer\GAL4[ppl.PP] can not further increase the starvation resistance of chico[1]/+ flies. (Hull-Thompson et al., 2009) Dg[323]/chico[1] double heterozygous mutant third instar larval eye discs do not show disrupted axon targeting from photoreceptor neurons to the brain optic lobes. (Shcherbata et al., 2007) chico¹/+; dally^{sec.Scer\UAS.T:Hsap\MYC}/Scer\GAL4^29BD pupae show the same large pupae and disordered wing patterning phenotype as wild-type animals expressing dally^{sec.Scer\UAS.T:Hsap\MYC} under the control of Scer\GAL4^29BD. (Takeo et al., 2005) The small body size and reduced numbers of ommatidia seen in chico¹ homozygous adults are partially suppressed by B4^susi-1/B4^susi-1. The eye phenotype is also partially suppressed by B4⁰⁵³³⁷/B4⁰⁵³³⁷ (Wittwer et al., 2005) Homozygous chico;foxo double mutants have more, and even slightly smaller, cells than homozygous chico single mutants. (Junger et al., 2003) No obvious defects in photoreceptor axon projections in the medulla are seen in chico¹/+ ; InR^ex15/+ double heterozygous adults. (Song et al., 2003) ovo^D1 chico¹ double heterozygotes live as long as chico¹/+ single heterozygotes and significantly longer than ovo^D1/+ single heterozygotes. (Clancy et al., 2001)
Xenogenetic Interactions
Statement Reference
Complementation & Rescue Data
Rescued by	chico¹ is rescued by chico^+t8 (Bohni et al., 1999, Clancy et al., 2001, Oldham et al., 2002) chico¹ is rescued by chico^Drkmut (Oldham et al., 2002) chico¹ is rescued by chico^PTBmut (Oldham et al., 2002) chico¹ is rescued by chico^Scer\UAS.cNa/Scer\GAL4^elav.PLu (Naganos et al., 2012) chico¹ is rescued by Scer\GAL4^c309/chico^Scer\UAS.cNa (Naganos et al., 2012)
Partially rescued by	chico¹ is partially rescued by Scer\GAL4^ey-OK107/chico^Scer\UAS.cNa (Naganos et al., 2012)
Not rescued by	chico¹ is not rescued by chico^PHmut (Oldham et al., 2002) chico¹ is not rescued by chico^PI3Kmut (Oldham et al., 2002) chico¹ is not rescued by chico^Scer\UAS.cNa/Scer\GAL4^GH146 (Naganos et al., 2012) chico¹ is not rescued by Scer\GAL4^{elav.Switch.PO}/chico^Scer\UAS.cNa (Naganos et al., 2012)
Comments	Expression of chico[Scer\UAS.cNa] under the control of Scer\GAL4[elav.PLu] rescues the defects in 1 hour memory seen after olfactory aversive conditioning in chico[1] adults. Expression of chico[Scer\UAS.cNa] limited to the adult stage (driven by Scer\GAL4[elav.Switch.PO] in the presence of RU486) does not rescue the defects in 1 hour memory seen after olfactory aversive conditioning in chico[1] adults. Expression of chico[Scer\UAS.cNa] under the control of Scer\GAL4[ey-OK107] rescues the number of Kenyon cells and partially rescues reduced mushroom body size in chico[1] adults. Olfactory learning is rescued in these flies. Expression of chico[Scer\UAS.cNa] under the control of Scer\GAL4[c309], but not under the control of Scer\GAL4[GH146], rescues olfactory learning in chico[1] adults. (Naganos et al., 2012) chico^+t8 significantly reduces the extended life-span seen in chico¹ heterozygous males and females. (Clancy et al., 2001)
Stocks ( 1 )
Bloomington	10738 cn¹ P{ry11}chico¹/CyO; ry⁵⁰⁶
Notes on Origin
Discoverer	A. Spradling.

Comments
	Complements: flp^hd. (Riesgo-Escovar, 2001.2.7) Designation as "flp^ry4" was a symbol misassignment. (Spradling, 1990.8.1)
External Crossreferences & Linkouts
Other Crossreferences

Linkouts

Synonyms & Secondary IDs ( 10 )
Reported As
Symbol Synonym	chic¹ chico1 (Wagaman et al., 2004, Bartke et al., 2002, Vigne et al., 2009) chico-1 (Tu et al., 2002) chico¹ (Wagaman and Rand, 2007, Christensen and Cook, 2006.8.30, Shingleton et al., 2005, Wang et al., 2011, Albrecht et al., 2011, Wong et al., 2009, Hull-Thompson et al., 2009, Richard et al., 2005, Werz et al., 2009, Shcherbata et al., 2007, Wang et al., 2008, Naganos et al., 2012, Biteau et al., 2010, Murillo-Maldonado et al., 2011, Slack et al., 2011, Chakrabarti et al., 2012) flp^ry4 (Spradling, 1990.8.1) fs(2)4 (Riesgo-Escovar, 2001.2.7, ) fs(2)4¹ (Goberdhan et al., 1999, Riesgo-Escovar et al., 1996) fs(2)ry4 fs(2)ry⁴ (Clegg et al., 1993) unnamed (Riesgo-Escovar et al., 1998, Berg and Spradling, 1991)
Name Synonym
Secondary FlyBase IDs
	FBal0034657 FBal0086688
References ( 55 )

Generate a list of	All references relating to this allele ( 55 )

List References by type	Research paper ( 40 ) Review ( 2 ) Personal communication to FlyBase ( 6 ) Abstract ( 7 )
Recent research papers ( 6 )
	Chakrabarti et al., 2012, Cell Host Microbe 12(1): 60--70 Infection-induced host translational blockage inhibits immune responses and epithelial renewal in the Drosophila gut. [FBrf0218984] Naganos et al., 2012, Neurosci. Res. 73(1): 49--55 Mutations in the Drosophila insulin receptor substrate, CHICO, impair olfactory associative learning. [FBrf0217997] Albrecht et al., 2011, Cell Metab. 14(6): 819--829 In Vivo Mapping of Hydrogen Peroxide and Oxidized Glutathione Reveals Chemical and Regional Specificity of Redox Homeostasis. [FBrf0216929] Murillo-Maldonado et al., 2011, PLoS ONE 6(11): e28067 Insulin Receptor-Mediated Signaling via Phospholipase C-γ Regulates Growth and Differentiation in Drosophila. [FBrf0216849] Slack et al., 2011, Aging Cell 10(5): 735--748 dFOXO-independent effects of reduced insulin-like signaling in Drosophila. [FBrf0215229] Wang et al., 2011, Cell 145(4): 596--606 A hormone-dependent module regulating energy balance. [FBrf0213699] Show all research papers ...
Recent reviews (0)
	All reviews listed in FlyBase were published before 2011 Show reviews ...

Allele Dmel\chico1

Allele Dmel\chico¹