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Gene Dmel\ph-p

General Information
Symbol	Dmel\ph-p	Species	D. melanogaster
Name	polyhomeotic proximal	Annotation symbol	CG18412
Feature type	protein_coding_gene	FlyBase ID	FBgn0004861
Created / Updated	2003-12-01/2003-12-01
Genomic Location
Chromosome (arm)	X	Recombination map	1-0.5
Cytogenetic map	2D2-2D3	Sequence location	X:2,020,237..2,030,578 [-]
Map ( GBrowse )	Decorated FastA GenBankGFF Gene region Extended Gene region CDSIntronsExonsTranslationsTranscripts3' UTR5' UTR
Summary Information
Automatically generated summary See sections below for more information	The gene polyhomeotic proximal is referred to in FlyBase by the symbol ph-p (CG18412, FBgn0004861). It has the cytological map location 2D2-2D3. Its sequence location is X:2020237..2030578. Its molecular function is described as: DNA binding; chromatin binding; chaperone binding; zinc ion binding. It is involved in the biological processes: gene silencing; germarium-derived female germ-line cyst encapsulation; ovarian follicle cell stalk formation; chromatin silencing; syncytial blastoderm mitotic cell cycle; response to ecdysone; central nervous system neuron development. 42 alleles are reported. The phenotypes of these alleles are annotated with 30 unique terms, many of which group under: adult segment; organ system; adult brain; germarium; nervous system; peripheral nervous system; larval abdominal segment; metatarsus; adult mesothoracic segment; follicle cell. It has one annotated transcript and one annotated polypeptide.
External Summaries
Phenotypic Description from the Red Book (Lindsley & Zimm 1992)
Gene/Allele symbols may differ from current usage	ph: polyhomeotic (P. Santamaria) A member of the Polycomb group of genes; seems to be the only one that is strongly required both maternally and zygotically for normal embryonic development. Two mutagenic events are necessary to produce null mutations, suggesting that the locus is complex, and in fact molecular determinations indicate that the locus comprises a direct repeat. Single-event mutations are viable as males and homozygous females; such mutations produce transformations similar to those of known dominant gain-of-function mutants in the ANTC and BXC, i.e., transformation of wings to halteres, second and third legs to first legs, and anterior abdominal segments to more posterior segments (mutants may also show loss of the humerus). Two-event lethal mutations die in mid embryogenesis and completely lack ventral and abdominal epidermal derivatives; they show transformations of most of the segments toward the eighth abdominal segment (Dura et al., 1988). There is an alteration in the pattern of axon pathways in the CNS. The axons fail to form commissures or connectives but instead form large bundles in the middle of each hemi-ganglion (Smouse et al., 1988). Trans heterozygotes between viable and lethal alleles or between viable alleles and a deficiency for ph die in late embryogenesis and exhibit posteriorly directed transformations; i.e. viable alleles are haplo insufficient. Dura et al. (1987) postulate that mutations can occur in either or both of the repeated sequences such that -+/-+ = +-/+- = -+/+- = viable mutant constitutions (--/++ not stated to be mutant); -+/-- = +-/-- = late embryonic lethals; and --/-- = mid-embryonic-stage lethals. The ph+ product is required autonomously in imaginal cells. A total lack of ph+ function prevents viability of the cuticular derivatives of these cells, but amorphic ph clones induced in late third instar survive. ph has a strong maternal effect on segmental identity and epidermal development that cannot be rescued by a single paternally supplied dose of ph+ in the zygote (Dura et al., 1988). The expression of ph is inversely related to dosage of the ANTC and the BXC; the gene interacts with Pc, Pcl, and esc (Dura et al., 1985). At the shortened germ band stage (but not at the blastoderm stage), ph seems to be involved in the regulation, not only of the homeotic genes Scr and Ubx, but also in the regulation of the segmentation genes ftz, eve, and en (Dura and Ingham, 1988).
Detailed Mapping Data
FlyBase Computed Cytological Location
Cytogenetic map Evidence for location 2D2-2D3   Limits computationally determined from genome sequence between P{EP}ActnEP1193&P{EP}CG4322EP1631 and P{EP}EP1460&P{EP}CG2924EP1596
Experimentally Determined Cytological Location
Cytogenetic map Notes References
Experimentally Determined Recombination Data
Location	1-0.5
Left of (cM)
Right of (cM)
Notes
Molecular Map Data
Gene Order (in direction of increasing cytology) References Gene Order (overall orientation not stated) References Overall orientation not stated: ph-p+ ph-d+ (Deatrick et al., 1991)
Gene Model & Products
Please see the GBrowse view of Dmel\ph-p for information on other features
Comments on Gene Model
	DGC clone GH08934 appears problematic: incomplete CDS
Transcript Data
Annotated Transcripts
Name FlyBase ID RefSeq ID Length (nt) Associated CDS (aa) ph-p-RA FBtr0070432  NM_057523   5869   1589
Additional Transcript Data & Comments
Reported size (kB)	6.4, 6.1 (northern blot) (Hodgson et al., 1997)
Comments	Transcripts of 6.4kb and 6.1kb are observed in the region (FBrf457872) but it\\'s not clear if they are transcribed from ph-p or ph-d. (Deatrick et al., 1991)
External Data
Crossreferences
Polypeptide Data
Annotated Polypeptides
Name FlyBase ID Predicted MW (kD) Length (aa) Theoretical pI RefSeq ID GenBank protein ph-p-PA   FBpp0070416   167.3   1589   9.55     NP_476871   AAF45727
Additional Polypeptide Data & Comments
Reported size (kD)	1589 (aa); 170 (kD observed); 169 (kD predicted) (DeCamillis et al., 1992) 1589 (aa); 170 (kD observed) (Hodgson et al., 1997)
Comments	Antibodies specific to ph-p protein were raised. (Hodgson et al., 1997) ph-p protein binds to about 80 sites along the polytene chromosomes that are distributed throughout the genome. The bound sites largely overlap those bound by Pc protein. ph-p binds to insertion sites of constructs containing sequences from the bxd region of Ubx showing that the binding is DNA-sequence dependent. (DeCamillis et al., 1992) The sequences of ph-p and ph-d may be aligned along a stretch of 1442 amino acids along which they show great similarity. (Deatrick et al., 1991)
External Data
Linkouts	PANTHER - Protein classification by function, families, and pathways • FBgn0004861
Crossreferences	InterPro domains - A database of protein families, domains, and functional sites • Sterile alpha motif SAM (IPR001660) Sterile alpha motif homology (IPR010993) Sterile alpha motif homology 2 (IPR011510) Zinc finger, FCS-type (IPR012313) Sterile alpha motif-type (IPR013761) PDB - Protein Data Bank. An information portal to biological macromolecular structures • 1KW4
Sequences Consistent with the Gene Model
DDBJ / EMBL / GenBank	DNA sequence Protein sequence Name AA951210   AE003423 AAF45727   AI109682   AI388747   AW940753   BP547191   BP552800   BP552801   BT016124 AAV37009   CG18412   CK131164   CO296405   CO300447   CO300650   CO305401   CO312983   CZ472451   EC065113   EC073921   EC074400   EC076514   EC200696   EC201718   EC204075   EC223140   EC225806   EC241618   EC247320   M64750   X63672 CAA45211
UniProtKB/Swiss-Prot	P39769
UniProtKB/TrEMBL	Q5U0W8
Maps to	87B1 (FlyBase, 1992-)
Does NOT map to
Identified with	GH19743 (Benos et al., 2001) GH08934 LD01444 (BDGP Project Members, 2000-)
Mapped Features & Mutations
Please see GBrowse or insertion reports for information on insertions of transgenic constructs and features not listed here Type Symbol & Location Additional Notes References deletion ph-p[2] X:2,022,856..2,024,102 comment=ph-p[2] is a 10kb deletion that removes the 3' end of ph-p and the 5' end of ph-d. The breakpoint maps within a 1247bp region that is identicle between the two genes but cannot be further localized. evidence=experimental (Saget and Randsholt, 1994) deletion ph-p[4] X:2,022,856..2,024,102 comment=ph-p[4] is a 10kb deletion that removes the 3' end of ph-p and the 5' end of ph-d. The breakpoint maps within a 1247bp region that is identicle between the two genes but cannot be further localized. evidence=experimental (Saget and Randsholt, 1994) protein binding site ph-p-protein_bind-16 X:2,034,644..2,034,670 bound_moiety=en-XP comment=Bound by en protein in DNaseI footprint analyses evidence=experimental (Serrano et al., 1995, Serrano and Maschat, 1998, Bergman et al., 2004) protein binding site ph-p-protein_bind-15 X:2,034,624..2,034,634 bound_moiety=en-XP comment=Bound by en protein in DNaseI footprint analyses evidence=experimental (Serrano and Maschat, 1998, Serrano et al., 1995, Bergman et al., 2004) protein binding site ph-p-protein_bind-14 X:2,034,521..2,034,542 bound_moiety=en-XP comment=Bound by en protein in DNaseI footprint analyses evidence=experimental (Serrano and Maschat, 1998, Serrano et al., 1995, Bergman et al., 2004) protein binding site ph-p-protein_bind-13 X:2,034,459..2,034,518 bound_moiety=en-XP comment=Bound by en protein in DNaseI footprint analyses evidence=experimental (Serrano et al., 1995, Serrano and Maschat, 1998, Bergman et al., 2004) protein binding site ph-p-protein_bind-12 X:2,034,350..2,034,411 bound_moiety=en-XP comment=Bound by en protein in DNaseI footprint analyses evidence=experimental (Serrano et al., 1995, Serrano and Maschat, 1998, Bergman et al., 2004) protein binding site ph-p-protein_bind-11 X:2,034,260..2,034,316 bound_moiety=en-XP comment=Bound by en protein in DNaseI footprint analyses evidence=experimental (Serrano et al., 1995, Bergman et al., 2004, Serrano and Maschat, 1998) protein binding site ph-p-protein_bind-9 X:2,019,566..2,019,602 bound_moiety=en-XP evidence=experimental comment=Bound by en protein in DNaseI footprint analyses (Bergman et al., 2004, Serrano et al., 1995, Serrano and Maschat, 1998) protein binding site ph-p-protein_bind-8 X:2,019,526..2,019,546 comment=Bound by en protein in DNaseI footprint analyses evidence=experimental bound_moiety=en-XP (Serrano et al., 1995, Serrano and Maschat, 1998, Bergman et al., 2004) protein binding site ph-p-protein_bind-7 X:2,019,414..2,019,499 bound_moiety=en-XP evidence=experimental comment=Bound by en protein in DNaseI footprint analyses (Serrano and Maschat, 1998, Bergman et al., 2004, Serrano et al., 1995) protein binding site ph-p-protein_bind-6 X:2,019,388..2,019,400 bound_moiety=en-XP evidence=experimental comment=Bound by en protein in DNaseI footprint analyses (Serrano and Maschat, 1998, Bergman et al., 2004, Serrano et al., 1995) protein binding site ph-p-protein_bind-5 X:2,019,327..2,019,372 bound_moiety=en-XP comment=Bound by en protein in DNaseI footprint analyses evidence=experimental (Bergman et al., 2004, Serrano and Maschat, 1998, Serrano et al., 1995) protein binding site ph-p-protein_bind-3 X:2,018,280..2,018,309 evidence=experimental bound_moiety=en-XP comment=Bound by en protein in DNaseI footprint analyses (Bergman et al., 2004, Serrano et al., 1995, Serrano and Maschat, 1998) protein binding site ph-p-protein_bind-2 X:2,018,164..2,018,255 evidence=experimental bound_moiety=en-XP comment=Bound by en protein in DNaseI footprint analyses (Serrano et al., 1995, Serrano and Maschat, 1998, Bergman et al., 2004) protein binding site ph-p-protein_bind-17 X:2,034,240..2,034,682 bound_moiety=en-XP comment=P1 fragment evidence=experimental linked_to=Sau3A-BamHI_rfrag (Serrano et al., 1995) protein binding site ph-p-protein_bind-19 X:2,019,323..2,019,619 bound_moiety=en-XP comment=D1 fragment evidence=experimental linked_to=SalI-Sau3A_rfrag (Serrano et al., 1995) protein binding site ph-p-protein_bind-25 X:2,018,146..2,018,483 bound_moiety=en-XP comment=D2 fragment evidence=experimental linked_to=DdeI-SalI_rfrag (Serrano et al., 1995) regulatory region ph-p-misc_feature-1 X:2,031,475..2,035,436 comment=Regulatory fragment associated with variegation and transvection effects on an adjacent miniwhite gene when introduced into the genome on a P element vector. evidence=experimental linked_to=EcoRI-KpnI_rfrag (Fauvarque and Dura, 1993) regulatory region ph-p-misc_feature-9 X:2,030,529..2,033,446 comment=regulatory fragment associated with variegation and transvection effects on an adjacent miniwhite gene when introduced into the genome on a P element vector evidence=experimental linked_to=XbaI-PstI_rfrag (Fauvarque and Dura, 1993)
External Data
Linkouts	DEDB - Drosophila exon database: splicing graphs • 254
Crossreferences
Expression Data
FlyBase-Curated Data
Transcript and Protein data	Please see the FlyBase Gene Expression Report for details of gene expression from the literature.
Summary of Transcript Expression
Stage Tissue/Position Reference  oogenesis stage,adult stage | stage S1-S10  follicle cell (de Camillis and Brock, 1994)  embryonic stage-adult stage   (Hodgson et al., 1997)  embryonic stage | cellular blastoderm   (Fauvarque et al., 1995)  embryonic stage  ectoderm (Fauvarque et al., 1995)  larval stage | third instar  ventral thoracic disc (de Camillis and Brock, 1994)  embryonic stage | stage >=12  embryonic central nervous system (de Camillis and Brock, 1994)  larval stage | third instar  dorsal metathoracic disc (de Camillis and Brock, 1994)  embryonic stage   (de Camillis and Brock, 1994)  oogenesis stage,adult stage  germarium (de Camillis and Brock, 1994)  embryonic stage-adult stage   (Hodgson et al., 1997)  oogenesis stage,adult stage | stage S1-S10  nurse cell (de Camillis and Brock, 1994)  larval stage | third instar  dorsal mesothoracic disc (de Camillis and Brock, 1994)  embryonic stage | blastoderm   (de Camillis and Brock, 1994)  embryonic stage | cellular blastoderm   (Fauvarque et al., 1995)  larval stage | third instar  eye-antennal disc (de Camillis and Brock, 1994)  embryonic stage | stage 8-11  parasegment | restricted (de Camillis and Brock, 1994)  embryonic stage | stage 8-11  germ band (de Camillis and Brock, 1994)  embryonic stage  embryonic central nervous system (Fauvarque et al., 1995)
Marker for
Subcellular Localization
CV Term
Summary of Polypeptide Expression
Stage Tissue/Position Reference  larval stage | third instar  eye-antennal disc (de Camillis and Brock, 1994)  larval stage | third instar  dorsal mesothoracic disc (de Camillis and Brock, 1994)  embryonic stage | late  sensillum campaniformium (de Camillis and Brock, 1994)  embryonic stage | late  embryonic peripheral nervous system (de Camillis and Brock, 1994)  larval stage | third instar  dorsal metathoracic disc (de Camillis and Brock, 1994)  larval stage | third instar  ventral thoracic disc (de Camillis and Brock, 1994)  embryonic stage  cell body (de Camillis and Brock, 1994)  embryonic stage   (de Camillis and Brock, 1994)  embryonic stage  embryonic central nervous system (de Camillis and Brock, 1994)
Marker for
Subcellular Localization
CV Term	nucleus (de Camillis and Brock, 1994)
External Data & Images
Linkouts	FLIGHT - Cell culture data for RNAi and other high-throughput technologies • FBgn0004861 GEO (NCBI) - Gene expression data: microarray and other high-throughput technologies • FBgn0004861
Alleles & Phenotypes
Summary of Allele Phenotypes
Lethality Allele viable ph-p409 ph-p410 ph-p5PS ph-p7 ph-plac lethal | recessive | conditional - heat sensitive ph-p2 lethal, with Scer\GAL4hs.PB ph-pScer\UAS.cNa lethal, with Scer\GAL4ptc-559.1 ph-pScer\UAS.cNa lethal, with Scer\GAL4c355 ph-pGS1032 lethal, with Scer\GAL4da.G32 ph-pScer\UAS.cNa lethal, with Scer\GAL4e22c ph-pScer\UAS.cNa lethal, with Scer\GAL429BD ph-pGS1032 semi-lethal, with Scer\GAL4dpp.blk1 ph-pGS1032 semi-lethal, with Scer\GAL4hs.2sev ph-pGS1032 Other Phenotypes Allele wild-type ph-pP1 visible ph-p410 ph-p5PS visible | recessive ph-p410 visible | dominant | homeotic ph-p410 visible | homeotic ph-p40 ph-p410 visible, with Scer\GAL4hs.2sev ph-pGS1032 eye color defective ph-p600 ph-p602 suppressor of variegation | dominant, with P{wA}4-4 ph-p410 suppressor of variegation | dominant, with P{hsp26-pt-T}39C-5 ph-p410 suppressor of variegation | dominant, with P{hsp26-pt-T}39C-27 ph-p410 suppressor of variegation | dominant, with P{hsp26-pt-T}39C-31 ph-p410 mitotic cell cycle defective ph-plac mitotic cell cycle defective | embryonic stage ph-p409 ph-p504 neuroanatomy defective | somatic clone ph-p602 Phenotype manifest in Allele mesothoracic leg ph-p2 ph-p4 ph-p410 metathoracic leg ph-p2 ph-p4 ph-p410 wing | posterior compartment ph-p2 ph-p4 ph-p410 abdominal segment 1 ph-p2 ph-p4 abdominal segment 4 ph-p2 ph-p4 ph-p410 postpronotum ph-p2 ph-p4 sex comb ph-p2 ph-p409 ph-p410 ph-p5PS metathoracic leg & sex comb tooth | ectopic ph-p40 mesothoracic leg & sex comb tooth | ectopic ph-p40 adult abdominal segment 4 ph-p40 adult abdominal segment 5 | ectopic ph-p40 wing ph-p40 haltere | ectopic ph-p40 ovary, with Scer\GAL4da.G32 ph-p4061 interfollicle cell, with Scer\GAL4da.G32 ph-p4061 egg chamber, with Scer\GAL4da.G32 ph-p4061 germarium region 2b, with Scer\GAL4da.G32 ph-p4061 germarium region 3, with Scer\GAL4da.G32 ph-p4061 germline cyst, with Scer\GAL4da.G32 ph-p4061 wing, with Scer\GAL4da.G32 ph-p4061 haltere | ectopic, with Scer\GAL4da.G32 ph-p4061 anaphase & embryo ph-p409 ph-p504 telophase & embryo ph-p409 ph-p504 nucleus & embryo ph-p409 ph-p504 sex comb (with Df(3R)ry27) ph-p409 sex comb (with Df(3R)ry619) ph-p409 sex comb (with Tp(3;3)MRS) ph-p409 sex comb | ectopic ph-p410 larval abdominal segment 4 ph-p410 larval abdominal segment 5 | ectopic ph-p410 abdominal segment 6 ph-p410 prothoracic leg | ectopic ph-p410 sex comb tooth ph-p410 mesothoracic leg & sex comb | ectopic ph-p410 genitalia ph-p5PS eye ph-p600 ph-p602 optic lobe neuron | somatic clone ph-p602 Kenyon cell | somatic clone ph-p602 antennal lobe | somatic clone ph-p602 dorsal cluster neuron | somatic clone ph-p602 ellipsoid body | somatic clone ph-p602 eye, with Scer\GAL4hs.2sev ph-pGS1032 egg chamber | somatic clone, with Scer\GAL4Act5C.PP ph-pScer\UAS.cNa interfollicle cell | somatic clone, with Scer\GAL4Act5C.PP ph-pScer\UAS.cNa germarium | somatic clone, with Scer\GAL4Act5C.PP ph-pScer\UAS.cNa germarium region 2b | somatic clone, with Scer\GAL4Act5C.PP ph-pScer\UAS.cNa germarium region 3 | somatic clone, with Scer\GAL4Act5C.PP ph-pScer\UAS.cNa egg chamber ph-plac germarium ph-plac germarium region 3 ph-plac interfollicle cell ph-plac interfollicle cell | ectopic ph-plac polar follicle cell | ectopic ph-plac
Classical Alleles ( 32 )
For All Classical Alleles Show
Allele of ph-p Class Mutagen Stocks Known lesion ph-p+11 Δ2-3 0 Yes ph-p+12 Δ2-3 0 Yes ph-p+14 Δ2-3 0 Yes ph-p+18 Δ2-3 0 Yes ph-p+22 Δ2-3 0 Yes ph-p+27 Δ2-3 0 Yes ph-p10a X ray 0 Yes ph-p15 1 -- ph-p2 PM hybrid dysgenesis 0 Yes ph-p3 PM hybrid dysgenesis 0 Yes ph-p4061 P-element activity 0 Yes ph-p409 loss of function, hypomorph X ray 0 Yes ph-p40 0 -- ph-p410 X ray 2 Yes ph-p4 PM hybrid dysgenesis 0 Yes ph-p504 loss of function 1 -- ph-p505 loss of function 0 -- ph-p5PS hypomorph 0 -- ph-p600 X ray 0 Yes ph-p602 X ray 2 Yes ph-p603 X ray 0 Yes ph-p604 X ray 0 Yes ph-p609 X ray 0 Yes ph-p7 PM hybrid dysgenesis 0 Yes ph-pFB spontaneous 0 Yes ph-pGS1032 Δ2-3 1 Yes ph-pP1 P-element activity 0 Yes ph-pd01798 P-element activity 1 -- ph-plac+2 Δ2-3 0 Yes ph-plac+3 Δ2-3 1 Yes ph-plac hypomorph P-element activity 1 Yes ph-prIIA 0 --
Alleles Carried on Transgenic Constructs ( 10 )
For All Alleles Carried on Transgenic Constructs Show
Allele of ph-p Class Mutagen Stocks Known lesion ph-pGD17398 in vitro construct | RNAi in vitro construct | regulatory fusion 2 Yes ph-pGD4480 <