OCA2 contains regions for the numerous eye colors, but one SNP is a strong predictor for brown/blue eyes. Heterochromia, although not viewed as a severe disorder, affects many individuals. 1998; Flanagan et al. (1997), suggesting that these sequences are indeed associated with iris pigmentation as suggested by these authors, although we note that the associations described by these authors were with blue irises and at the level of the SNP, while those that we observed were with green irises and apparent only at the level of the haplotypes and diplotypes. Nat Genet. SNP discovery: We obtained candidate SNPs from the National Center for Biotechnology Information (NCBI) Single Nucleotide Polymorphism Database (dbSNP), which generally provided more candidate SNPs than were possible to genotype. The second parent has a non-mutated HERC2 allele but does not have the coding for brown eyes in the OCA2 gene. If you have no pigment you have either blue or gray eyes. Although this could indicate that the SNPs are in LD with other phenotypically active loci, it may also be a reflection that variability in message transcription and/or turnover may explain part of the variability observed in human iris colors. Incomplete dominance shows in individuals with lighter shades of brown and hazel. Eye color is determined by the pattern of brown and red pigment, collagen fibers, and the topography of the iris. Refer to TABLE II. The reason many white, non-Hispanic babies are born with blue eyes is that they don't have the full amount of melanin present in their irises at birth. The quantity and quality of melanin in the cytoplasm determines the observed color of the eye. Statistical methods: To test the departures from independence in allelic state within and between loci, we used the exact test, described in Zaykin et al. 1991; Boissy et al. In the most elementary form, the inheritance of eye color is classified as a Mendelian trait.1 On the basis of the observation of more than two phenotypes, eye color has a more complex pattern of inheritance. In the meantime, to ensure continued support, we are displaying the site without styles Indeed, one of those for which the evidence of lack of HWE was the strongest was validated as a legitimate SNP through direct DNA sequencing (data not shown). We identified numerous SNPs, haplotypes, and diplotypes (diploid pairs of haplotypes) within the OCA2, MYO5A, TYRP1, AIM, DCT, and TYR genes and the CYP1A2-15q22-ter, CYP1B1-2p21, CYP2C8-10q23, CYP2C9-10q24, and MAOA-Xp11.4 regions as significantly associated with iris colors. To identify SNP loci associated with variable human pigmentation, we genotyped for 754 SNPs: 335 SNPs within pigmentation genes (AP3B1, ASIP, DCT, MC1R, OCA2, SILV, TYR, TYRP1, MYO5A, POMC, AIM, AP3D1, and RAB; Table 1), and 419 other SNPs distributed throughout the genome. Sturm, R., Duffy, D., Zhao, Z., Leite, F., Stark, M., Hayward, N. et al. https://doi.org/10.1038/jhg.2010.126, DOI: https://doi.org/10.1038/jhg.2010.126. In Drosophila, iris pigmentation defects have been ascribed to mutations in >85 loci contributing to a variety of cellular processes in melanocytes (Ooi et al. 1997; Smith et al. In the population sample, we were also able to examine the correlation between genotype at the W locus and iris color . During pigment distribution in an infant, a diet low in cysteine or methionine, which it is synthesized from, would likely affect the color of the child's eyes until the amino acid is supplemented. Rebbeck et al. What colour are your eyes? Teaching the genetics of eye colour - Nature Melanocortin 1 receptor (MC1R) instructs a melanocyte to switch production between eumelanin and pheomelanin.3, 4, 5 Therefore, these two proteins affect the quality and quantity of the melanin in the cell. This same phenomenon is the reason why the pupil appears black. To an investigator interested in elucidating a biological mechanism, association due to population structure might not seem to be very satisfying, but when classification is the goal rather than the elucidation of a biological mechanism, it would seem to matter little why a marker is associated with a trait. We also thank Robert White for his help with sample collection. This also explains why deletions within HERC2 would cause a decrease in melanin without interacting with the P protein itself. .. Kwon H Y, Bultman S J, Loffler C, Chen W-J, Furdon P J et al. As different genes may be transcribed in various cells, certain cells will produce more pigment or a different type of pigment than other adjacent cells. TYR, the enzyme responsible for pigment throughout the body, uses tyrosine to begin the chemical pathway. To form eumelanin, dopachrome tautomerase, TYR, and TYR-related protein 1 complete the chemical pathway from dopaquinone.3, Although the aforementioned proteins are responsible for the production of melanin, once it has been produced in the melanosomes, other proteins are responsible for melanin maturation. For example, dissection of the oculocutaneous albinism (OCA) trait in humans has shown that many pigmentation defects are due to lesions in the TYR gene, resulting in their designation as TYR-negative OCAs (Oetting and King 1991, 1992, 1993, 1999; see albinism database at http://www.cbc.umn.edu/tad/). We focused on human pigmentation and xenobiotic metabolism genes, selected on the basis of their gene identities, not their chromosomal position. Our results show that a surprisingly large number of polymorphisms in a large number of genes are associated with iris colors, suggesting that the genetics of iris color pigmentation are quite complex. The mouse pink-eyed dilution gene: association with human Prader-Willi and Angelman Syndromes. For those remaining, only a single round of PCR was performed. Eumelanin (brown pigment) is a light-absorbing polymer synthesized in specialized melanocyte lysosomes called melanosomes. The iris consists of five cell layers, the anterior border layer, stroma, the sphincter and dilator muscles fibers, and the posterior pigment epithelium ( Figure 1 ), of which the most important for the appearance of eye colour are the anterior layer and its underlying stroma ( Eagle, 1988; Imesch et al., 1996; Wilkerson et al., 1996 ). a) Give the genotype of an individual, who is homozygous dominant for Brown eye color, where "B" is the letter used to distinguish this trait. Attached earlobes. id List the possible genotypes of a blue eyed, dimple chinned individual. Anyone you share the following link with will be able to read this content: Sorry, a shareable link is not currently available for this article. The disorder is characterized by different-colored irises or different colors within the iris. Liu, F., Wollstein, A., Hysi, P. G., Ankra-Badu, G. A., Spector, T. D., Park, D. et al. One SNP has been studied to show a large significance for eye color. 1997; Box et al. 1991; Chintamaneni et al. In this case, pleiotropic effects change eye color. Genotype-phenotype correlations have been reported with specific mutations possibly associated with certain angle abnormalities. As one might expect from the proximity of these two regions, CYP2C8-CYP2C9 marker pairs were found to be in tight LD with one another (P < 0.001 for each possible pair). Chromosome 15 contains HERC1 and HERC2. Gene 277, 4962 (2001). .. Schioth H B, Phillips S R, Rudzish R, Birch-Machin M A, Wikberg J E et al. For some genes, the number of SNPs in the database was low and/or some of the SNPs were strongly associated with iris colors, warranting a deeper investigation. In addition, we independently isolated the red hair/blue iris SNP alleles described by Valverde et al. Multiple SNPs were identified on chromosome 10q; the CYP2C8-10p23 region had 1 marginally associated SNP, and the neighboring region, CYP2C9-10p24, also had one. A dominant allele of this gene (P) causes pigment to be deposited in the front of the iris, thus masking the blue to various degrees. Further, certain of our results support the previous literature. However, it is yet to be completely understood. Each chromosome contains thousands of individual genes. 2003) and it is possible that alleles for these SNPs are associated with elements of population structure that correlate with iris colors. Knoll, J. H. M., Nicholls, R. D., Magenis, R. E., Glatt, K., Graham, Jr J. M., Kaplan, L. et al. In studies with HERC2 functions, deletions caused hypopigmentation even though the protein has nothing to do with pigmentation. Pedigree studies in the mid-1970s suggested that iris color variation is a function of two loci: a single locus responsible for depigmentation of the iris, not affecting skin or hair, and another pleiotropic gene for reduction of pigment in all tissues (Brues 1975). Question: In albinism (a recessive disorder), the formation of melanin, a dark skin pigment, are blocked so that albinos have extremely light skin and hair. Diplotypes for these genes explain 15% of iris color variation. Genotyping was performed for individual DNA specimens using a single base primer extension protocol and an SNPstream 25K/ultra-high throughput (UHT) instrument (Beckman Coulter, Fullerton, CA, and Orchid Biosystems, Princeton, NJ). Human Iris Color. Chromosome 15q harbored the majority (14/27) of the SNPs that were marginally associated with iris colors, and all but one of these 14 were found in two different genes: OCA2 and MYO5A (Table 2). The SNP, rs12913832, causes a phenotype change from brown to blue eyes, respectively. OCA2 associations were by far the most significant of any gene or region we tested, while MYO5A SNPs were only weakly associated (but haplotypes and diplotypes more strongly). Google Scholar. These two seemingly unrelated genes have a major effect on eye color in humans. Although such an error is tolerable for identifying sequences marginally associated with iris colors, the use of the sequences described herein for iris color classification would therefore likely require digitally quantified iris colors (which we have begun to accumulate and will present elsewhere). Melanopsin signalling in mammalian iris and retina | Nature Specify the following genotypes using A and a to indicate dominant and recessive alleles, respectively. We did not confine this higher-order analysis to those genes with marginal SNP associations, but we grouped all of the high-frequency SNPs tested for each gene. We selected those for which at least two instances of PHRED identified variants that scored 24, and each of these SNPs discovered through resequencing were used for genotyping. This page titled 8.2: Human Traits Determined by Single Genes is shared under a CC BY 4.0 license and was authored, remixed, and/or curated by Ellen Genovesi, Laura Blinderman, & Patrick Natale via source content that was edited to the style and standards of the LibreTexts platform; a detailed edit history is available upon request. Sturm, R. A., Teasdale, R. D. & Box, N. F. Human pigmentation genes: identification, structure and consequences of polymorphic variation. No significant SNP associations within the pigmentation genes SILV, MC1R, ASIP, POMC, RAB, or TYR were found, although TYR had one SNP with a P = 0.06. The rest were found in or around xenobiotic metabolism genes, which we have previously shown exhibit dramatic sequence variation, Candidate genes tested for sequence associations with human iris pigmentation. Red and violet eyes come from a lack of pigment. For example, the OCA2 has 200 known candidate SNPs in NCBI's dbSNP, and it is possible that this gene has more to teach us about variable human iris pigmentation than what we have learned from the work presented herein. 1997). Linkage studies have implicated certain pigmentation genes as specifically relevant for pigmentation phenotypes, and most of the pigmentation gene SNPs that we identified clustered to certain genes such as OCA2, MYO5A, TYRP1, and AIM. Sequences Associated With Human Iris Pigmentation For these subjects, we obtained digital photographs of the right iris, where subjects peered into a box at one end at the camera at the other end to standardize lighting conditions and distance and from which a judge assigned the sample to a color group. Using a chi-square test, determine whether those numbers are consistent with . 2. Following your lab manual and your tutor's instructions fill out this table: Trait Phenotype Possible genotypes Class frequency Pigmented iris Pigment No pigment PP Pp pp Pigmented iris = 79% No pigment= 21% Tongue rolling Yes no RR Rr rr 78% can 22% cannot Bent little finger Yes No BB Bb bb 20% can 80% cannot Widow's peak Yes No WW Ww ww . Pigmented iris A person with the B allele has brown eyes. Now, that color depends on the kind and density of melanin a person is born with. To correct for multiple tests, we used the empirical Bayes adjustments for multiple results method described by Steenland et al. For each gene, we inferred haplotypes and used contingency analyses to determine which haplotypes were statistically associated with iris colors. 1997, 2001; Akey et al. Annals of Hum Genet 73, 160170 (2009). 1999; Flanagan et al. IRIS pigmentation is a complex genetic trait that has long interested geneticists, anthropologists, and the public at large. Principles of Biology Lab 5 Worksheet-2.docx .. Hamabe J, Fukushima Y, Harada N, Abe K, Matsuo N et al. Problems with just HERC2 lead to nerve tissue malfunctioning, small size and semi-sterility or sterility. 1995; Koppula et al. In fact, study of the transmission genetics for pigmentation traits in humans and various model systems suggests that variable pigmentation is a function of multiple heritable factors whose interactions appear to be quite complex (Brauer and Chopra 1978; Bito et al. b) Give the genotype of an individual who is homozygous recessive for brown eye color. Forensic Sci Int: Genet. The Genetics of Eye Color - HudsonAlpha Institute for Biotechnology Although TYR does not code for color, a nonfunctioning TYR masks any other gene responsible for pigmentation. At the cellular level, variable iris color in healthy humans is the result of the differential deposition of melanin pigment granules within a fixed number of stromal melanocytes in the iris (Imesch et al. Eumelanin (brown pigment) is a light-absorbing polymer synthesized in specialized melanocyte lysosomes called melanosomes. P_ Pigmented Iris (Additional genes give specific color, e.g. Fig. More than likely, their offspring would have blue eyes, but a 25% chance stands that offspring would have brown eyes. Use the Previous and Next buttons to navigate the slides or the slide controller buttons at the end to navigate through each slide. For example, forensics investigators construct physical profiles using surprisingly unscientific means; only in rare cases are eye-witness accounts available, and in certain circumstances these accounts are subjective and unreliable. E-mail: Search for other works by this author on: The human homolog of the mouse brown gene maps to the short arm of chromosome 9 and extends the known region of homology with mouse chromosome 4, Interaction between the melanocortin-1 receptor and P genes contributes to inter-individual variation in skin pigmentation phenotypes in a Tibetan population, Discrete visual defects in pearl mutant mice, Human tyrosinase gene, mapped to chromosome 11 (q14-q21), defines second region of homology with mouse chromosome 7, Iris color changes past early childhood. Frequency of the minor allele and the major and minor allele nucleotide. Two major genes on chromosome 15 affect the quantity and quality of the melanin produced by melanogenesis. 2001; Sturm et al. 2001), there appears to be only a minor dominance component for mammalian iris color determination (Brauer and Chopra 1978), and minimal correlation exists among skin, hair, and iris color within or between individuals of a given population. These analyses resulted in the identification of 61 SNPs in 16 genes/chromosomal regions associated with iris colors on one level or another; details for each and whether the SNP is marginally associated or associated within the context of the haplotype and/or diplotype are shown in Table 2. PubMedGoogle Scholar. The mammalian iris has three main tissue layers, all pigmented with melanin: an anterior fibrovascular stroma; a middle smooth-muscle layer consisting of the circumferential sphincter muscle at . The "P" allele produces the pigment which gives you eye color. Multiple SNPs were also identified on chromosome 2; the C/C genotype for the POMC SNP located at 2p23 was associated with blue iris color (Table 3) and a CYP1B1-2p21-region SNP was also marginally associated at the level of iris shade (Table 2), as well as within the context of a 2-SNP haplotype (Table 3). We developed a program (T. Frudakis, M. Thomas, Z. Gaskin, K. Venkateswarlu, K. Suresh Chandra, S. Ginjupalli, S. Gunturi, S. Natrajan, V. K. Ponnuswamy and K. N. Ponnuswamy, unpublished results) to design resequencing primers in a manner respectful of homologous sequences in the genome, to ensure that we did not coamplify pseudogenes or amplify from within repeats. Of course, identifying markers in LD with phenotypically active loci (or the phenotypically active loci themselves) would provide for more accurate classification (as well as for a better understanding of biological mechanism), but the hunt for these elusive loci in heterogeneous populations is still impractical because LD extends only for a few kilobases and the economics of genome-wide scans in heterogeneous samples with full LD coverage are out of reach for most labs. The strongest associations were observed for genes with SNPs that were marginally associated (Table 2) and most of the genes with marginal SNP associations had haplotypes and diplotypes (sometimes referred to as multilocus gene-wise genotypes or diploid pairs of haplotypes) positively (agonist) or negatively (antagonist) associated with at least one iris color (Table 3). An individual with this disorder produces little or no pigment in their ocular melanocytes. When a T is replaced with a C in rs12913832 of intron 86, OCA2 transcription is depressed, resulting in a blue-eyed individual. MYO5A alleles were not found to be in LD with those of OCA2, suggesting that these results were independently obtained and that Eiberg and Mohr's results may have been a reflection of the activity of two separate genes. Pigmented irises. Genetics 165, 20712083 (2003). Overall, the diversity of haplotypes associated with brown irises was similar to that of haplotypes associated with blue irises. Last, we thank the reviewers of this manuscript who suggested a number of important improvements. It is around 12 . The colored area at the front of the eye is called the iris. Tyrosinase (TYR), the enzyme responsible for pigment production in the body, starts the synthesis of both types of melanin by catalyzing a reaction between tyrosine and dopa, forming dopaquinone. The "P" allele produces the pigment which gives you eye color. Your Blue Eyes Aren't Really Blue - American Academy of Ophthalmology An ASIP polymorphism is reported to be associated with both brown iris and hair color (Kanetsky et al. When a pigment is deposited in the front layer of the iris, this masks the blue layer to varying degrees. Peripheral scalloping of the posterior pigmented iris layer. Indeed, some, but not all, of our nonpigment gene SNPs are found in regions within the vicinity of pigmentation genes; CYP2C8 and CYP2C9 are located on chromosome 10 near the HPS1 and HPS2 pigmentation genes (which we did not test directly), CYP1A2 is located at 15q22ter on the same arm as OCA2 and MYO5A, CYP1B1 is located at 2p21 in the vicinity of the POMC gene at 2p23, and MAOA is located on the same arm of chromosome X (Xp11.411.3) as the OA1 pigmentation gene (which we also did not test directly). Human Genetics: Simple inheritance - Antranik Sequences associated with human iris pigmentation. .. Krude H, Biebermann H, Luck W, Horn R, Brabant G et al. 1, 105110 (2007). Similar to membrane-associated transporter protein, it transports melanosomes, but additionally, it controls their pH.3, 13 Therefore, the P protein encoded by OCA2 affects the amount and quality of melanin that deposits in melanocytes. The density of granules appears to reach genetically determined levels by early childhood and usually remains constant throughout later life, although a small minority of individuals exhibit changes in color during later stages of life (Bito et al. Albinism - EyeWiki Petunias with genotype R1R1 are red flowered, R1R2 are pink flowered and R2R2 are white flowered. Half of the associated SNPs were located on chromosome 15, which corresponds with results that others have previously obtained from linkage analysis. Many more genes affect brown and blue more than the other eye colors. Predicting phenotype from genotype: normal pigmentation. Clearly work remains to be done, objectifying the collection of iris colors from subjects, enhancing the sample size so that epistatic interactions can be explored, possibly screening other regions of the genome not screened here, and modeling the sequences that we have described to enable classification of iris colors from DNA. Green eyes require more pigment than blue and not much less than brown, and because the shades of hazel (brown with blue or green) are more versatile, hazel is still more popular than green. .. Steenland K, Bray I, Greenland S, Boffetta P. Strobel M C, Seperack P K, Copeland N G, Jenkins N A. Valverde P, Healy E, Jackson I, Rees J L, Thody A J. Wilson S M, Yip R, Swing D A, OSullivan T N, Zhang Y et al. Specimens for genotyping were of self-reported European descent, of different age, sex, hair, iris, and skin shades and they were collected using informed consent guidelines under Investigational Review Board guidance. Dopachrome tautomerase (13q32) and TYR-related protein 1 (9p23) will continue the pathway to form eumelanin.15 Therefore, if any of these proteins are not coded for correctly, the eumelanin production may be hindered, producing lighter eyes. The SNPs between the 2p21 and 2p23 regions were also in LD (P < 0.01). 2002). All visible light is absorbed by the retina. Use two alleles per trait for the genotype. Abbott C, Jackson I J, Carritt B, Povey S. Akey J M, Wang H, Xiong M, Wu H, Liu W et al. One method of grouping colors is light = blue + green and dark = hazel + brown, and this grouping would seem to more clearly distinguish individuals with respect to the detectible level of eumelanin (brown pigment). In the presence of cysteine, the reaction will proceed to form pheomelanin. Angelman syndrome: three molecular classes identified with chromosome 15q11q13-specific DNA markers. Cassidy, S. B. For R2 computation, we used the following function: Adj-R2 = 1 [n/(n p)](1 R2), where n is the model degrees of freedom and n p is the error degrees of freedom. Allele Variations in OCA2 gene (pink-eyed-dilution locus) are associated with genetic susceptibility to melanoma. It was unclear from the outset whether we would have better success considering iris color in terms of four colors (blue, green, hazel, and brown) or in terms of groups of colors. .. Rebbeck T R, Kanetsky P A, Walker A H, Holmes R, Halpern A C et al. Blue Iris (non-pigmented) MG-3: Jeremy has attached earlobes and pigmented irises. Cell Mol Life Sci 62, 18261838 (2005). In the case of TYR, melanin production will halt entirely, resulting in albinism in the entire body. The chromosomal distribution of the SNPs that were significantly associated in a marginal sense was found to be independent of the distribution of SNPs actually surveyed, indicating that the associations were not merely a function of SNP sampling and the same was true for the distribution of all the SNPs shown in Table 2 (data not shown). Comparing the results of the two methods of classification, 86 of the classifications matched. Correspondence to Further studies of this region and its sequence revealed that a change in one nucleotide, single-nucleotide polymorphism (SNP), regulates the binding site for the transcription of the OCA2 gene, altering its expression.9 The base changes from a thymine to a cytosine. Although corrections for multiple testing left most of the SNP-level associations intact, a number of the associations we found did not pass the multiple-testing examination, but nonetheless we present them here to avoid possible type II error; the sequences may be weakly associated with iris colors and possibly relevant within a multiple-gene model for classification (i.e., epistasis). 1997), and other genes (reviewed by Sturm et al. .. Kwon B S, Chintamaneni C, Kozak C A, Copeland N G, Gilbert D J et al. In the traditional view of eye color, it was taught that two blue-eyed individuals could only produce blue-eyed offspring. PubMed Central On the basis of population studies, scientists speculate that the blue-eyed mutation originated in peoples of Northern Europe (Scandinavian countries). Linkage disequilibrium (LD) for pairs of SNPs within a gene was determined using the Zaykin exact test and a cutoff value of |D| 0.05 (P value < 0.05; Zaykin et al. To test this, we performed a corrected ANOVA analysis for our data on each of these three levels. Genetic traits - University of Northern Iowa A battery of genetic tests, of which one for the inference of iris color could be a part, could enable the construction of a more objective and science-based (partial) physical profile from crime-scene DNA, and an investigator using these tests would be less interested in the biological mechanism of the phenotype than in an ability to make an accurate inference of trait value. The front layer of the iris (called the stroma) can make eyes appear brown, blue or green. Producing multicolored irises, heterochromia stems from mutations in certain cells of the iris. E_ Free earlobes. Solved P>p Trait Genotype Phenotypic Effect Relationship P. | Chegg.com We will explore some of these single gene traits in the laboratory. Genotypes for these 754 candidate SNPs were scored for 851 European-derived individuals of self-reported iris colors (292 blue, 100 green, 186 hazel, and 273 brown). Eye colors are green, hazel, brown or black. homework 5 ans. 11. In mice and humans where the P protein is nonfunctional, albinism occurs, indicating its crucial role in pigmentation.13, 14 The gene located 11.7kb from HERC2 requires 345kb, but it requires only 24 exons to produce a 110kDa protein with 838 residues. Most of the SNPs that we identified were on chromosome 15, which Eiberg and Mohr (1996) described from linkage analyses as the primary chromosome for the determination of brownness. As suggested by these authors, the candidate gene within the interval containing this locus (BEY2) is most likely the OCA2 gene, although the MYO5A gene is also present within this interval and, as shown here, associated with iris colors. Iris phenotypes and pigment dispersion caused by genes influencing Genotype. P is for pigment and dimpled chins (D) are dominant over undimpled chins. Supplement Series 1, 544546 (2008). Asterisks represent P values that remained significant after the correction for multiple tests and P values in italic are those that were statistically significant (P 0.05). Google Scholar. Human Earlobe attachment. To determine the extent to which extant iris color variation could be explained by various models, we calculated R2 values for SNPs, haplotypes, and multilocus genotype data by first assigning the phenotypic value for blue eye color as 1, green eye color as 2, hazel eye color as 3, and brown eye color as 4. Twin Res 7, 197210 (2004). Article Pigment Cell Res 14, 8693 (2001). Before the revelation of the effect of HERC2, rs1800407 in exon nine was thought to be the main factor for eye color. This gene is often referred to as the red-headed gene because of its prevalent expression in people with red hair and green eyes.4 Dopachrome tautomerase also contains regions for hazel and green eyes.5 Regions for brown eyes dominate the effects of these genes, though. However, the results presented herein constitute a good first step toward solving what our results confirm is a very complex genetics problem. (2000) with adjusted residuals to compensate for this risk. CAS Lack of HWE is usually an indication of a poorly designed genotyping assay, but none of the remaining 7 SNPs exhibited genotyping patterns that we have previously associated with such problems (such as the complete absence of an expected genotype class or all genotypes registering as heterozygotes). Blue is confined mostly to people who originated from Europe.11 Green eyes permeate the lowest amount of the population (excluding the disorders), probably due to the lack of coding for it within the genome.
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