{"id":2752,"date":"2016-06-13T17:05:12","date_gmt":"2016-06-13T17:05:12","guid":{"rendered":"https:\/\/courses.lumenlearning.com\/biologyxwaymakerxmaster\/?post_type=chapter&#038;p=2752"},"modified":"2024-04-29T16:32:51","modified_gmt":"2024-04-29T16:32:51","slug":"reading-multiple-alleles","status":"publish","type":"chapter","link":"https:\/\/courses.lumenlearning.com\/wm-biology1\/chapter\/reading-multiple-alleles\/","title":{"raw":"Multiple Alleles","rendered":"Multiple Alleles"},"content":{"raw":"<div class=\"textbox learning-objectives\">\r\n<h3>Learning Outcomes<\/h3>\r\n<ul>\r\n \t<li>Explain how mutli-allele inheritance will impact a trait within in a population<\/li>\r\n<\/ul>\r\n<\/div>\r\nMendel implied that only two alleles, one dominant and one recessive, could exist for a given gene. We now know that this is an oversimplification. Although individual humans (and all diploid organisms) can only have two alleles for a given gene, multiple alleles may exist at the population level such that many combinations of two alleles are observed. Note that when many alleles exist for the same gene, the convention is to denote the most common phenotype or genotype among wild animals as the\u00a0<strong>wild type<\/strong> (often abbreviated \"+\"); this is considered the standard or norm. All other phenotypes or genotypes are considered <strong>variants<\/strong> of this standard, meaning that they deviate from the wild type. The variant may be recessive or dominant to the wild-type allele.\r\n\r\nAn example of multiple alleles is coat color in rabbits (Figure 1). Here, four alleles exist for the\u00a0<em>c<\/em> gene. The wild-type version, <em>C<sup data-redactor-tag=\"sup\">+<\/sup>C<sup>+<\/sup><\/em>, is expressed as brown fur. The chinchilla phenotype, <em>c<i data-redactor-tag=\"i\"><sup>ch<\/sup><\/i>c<i><sup data-redactor-tag=\"sup\">ch<\/sup><\/i><\/em>, is expressed as black-tipped white fur. The Himalayan phenotype, <em>c<i data-redactor-tag=\"i\"><sup>h<\/sup><\/i>c<i><sup data-redactor-tag=\"sup\">h<\/sup><\/i><\/em>, has black fur on the extremities and white fur elsewhere. Finally, the albino, or \"colorless\" phenotype, <em>cc<\/em>, is expressed as white fur. In cases of multiple alleles, dominance hierarchies can exist. In this case, the wild-type allele is dominant over all the others, chinchilla is incompletely dominant over Himalayan and albino, and Himalayan is dominant over albino. This hierarchy, or allelic series, was revealed by observing the phenotypes of each possible heterozygote offspring.\r\n\r\n[caption id=\"attachment_1412\" align=\"aligncenter\" width=\"800\"]<img class=\"size-full wp-image-1412\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/110\/2016\/05\/02182457\/Figure_12_02_05.jpg\" alt=\"This illustration shows the four different variants for coat color in rabbits at the c allele. The genotype CC produces the wild type phenotype, which is brown. The genotype c^{ch}c^{ch} produces the chinchilla phenotype, which is black-tipped white fur. The genotype c^{h}c^{h} produces the Himalayan phenotype, which is white on the body and black on the extremities. The genotype cc produces the recessive phenotype, which is white\" width=\"800\" height=\"574\" \/> Figure 1. Four different alleles exist for the rabbit coat color (<i>C<\/i>) gene.[\/caption]\r\n\r\n[caption id=\"attachment_3251\" align=\"alignright\" width=\"350\"]<img class=\"wp-image-3251\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/110\/2016\/06\/27185610\/Figure_12_02_06-e1467053834271.jpg\" alt=\"This photo shows Drosophila that has normal antennae on its head, and a mutant that has legs on its head.\" width=\"350\" height=\"315\" \/> Figure 2. As seen in comparing the wild-type <i>Drosophila<\/i> (left) and the <i>Antennapedia<\/i> mutant (right), the Antennapedia mutant has legs on its head in place of antennae.[\/caption]\r\n\r\nThe complete dominance of a wild-type phenotype over all other mutants often occurs as an effect of \"dosage\" of a specific gene product, such that the wild-type allele supplies the correct amount of gene product whereas the mutant alleles cannot. For the allelic series in rabbits, the wild-type allele may supply a given dosage of fur pigment, whereas the mutants supply a lesser dosage or none at all. Interestingly, the Himalayan phenotype is the result of an allele that produces a temperature-sensitive gene product that only produces pigment in the cooler extremities of the rabbit's body.\r\n\r\nAlternatively, one mutant allele can be dominant over all other phenotypes, including the wild type. This may occur when the mutant allele somehow interferes with the genetic message so that even a heterozygote with one wild-type allele copy expresses the mutant phenotype. One way in which the mutant allele can interfere is by enhancing the function of the wild-type gene product or changing its distribution in the body.\r\n\r\nOne example of this is the <em>Antennapedia<\/em> mutation in <em>Drosophila<\/em> (Figure 2). In this case, the mutant allele expands the distribution of the gene product, and as a result, the <em>Antennapedia<\/em> heterozygote develops legs on its head where its antennae should be.\r\n<div class=\"key-takeaways textbox\">\r\n<h3>Multiple Alleles Confer Drug Resistance in the Malaria Parasite<\/h3>\r\nMalaria is a parasitic disease in humans that is transmitted by infected female mosquitoes, including\u00a0<em>Anopheles gambiae<\/em> (Figure 3a), and is characterized by cyclic high fevers, chills, flu-like symptoms, and severe anemia. <em>Plasmodium falciparum<\/em> and <em>P. vivax<\/em> are the most common causative agents of malaria, and <em>P. falciparum<\/em> is the most deadly (Figure 3b)<em>.<\/em> When promptly and correctly treated, <em>P. falciparum<\/em>malaria has a mortality rate of 0.1 percent. However, in some parts of the world, the parasite has evolved resistance to commonly used malaria treatments, so the most effective malarial treatments can vary by geographic region.\r\n\r\n[caption id=\"attachment_1414\" align=\"aligncenter\" width=\"969\"]<img class=\"size-full wp-image-1414\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/110\/2016\/05\/02182928\/Figure_12_02_07.jpg\" alt=\"Photo a shows the Anopheles gambiae mosquito, which carries malaria. Photo b shows a micrograph of sickle-shaped Plasmodium falciparum, the parasite that causes malaria. The Plasmodium is about 0.75 microns across.\" width=\"969\" height=\"489\" \/> Figure 3. The (a) Anopheles gambiae, or African malaria mosquito, acts as a vector in the transmission to humans of the malaria-causing parasite (b) Plasmodium falciparum, here visualized using false-color transmission electron microscopy. (credit a: James D. Gathany; credit b: Ute Frevert; false color by Margaret Shear; scale-bar data from Matt Russell)[\/caption]\r\n\r\nIn Southeast Asia, Africa, and South America,\u00a0<em>P. falciparum<\/em> has developed resistance to the anti-malarial drugs chloroquine, mefloquine, and sulfadoxine-pyrimethamine. <em>P. falciparum<\/em>, which is haploid during the life stage in which it is infectious to humans, has evolved multiple drug-resistant mutant alleles of the\u00a0<em>dhps<\/em> gene. Varying degrees of sulfadoxine resistance are associated with each of these alleles. Being haploid, <em>P. falciparum<\/em> needs only one drug-resistant allele to express this trait.\r\n\r\nIn Southeast Asia, different sulfadoxine-resistant alleles of the\u00a0<em>dhps<\/em> gene are localized to different geographic regions. This is a common evolutionary phenomenon that occurs because drug-resistant mutants arise in a population and interbreed with other <em>P. falciparum<\/em> isolates in close proximity. Sulfadoxine-resistant parasites cause considerable human hardship in regions where this drug is widely used as an over-the-counter malaria remedy. As is common with pathogens that multiply to large numbers within an infection cycle, <em>P. falciparum<\/em> evolves relatively rapidly (over a decade or so) in response to the selective pressure of commonly used anti-malarial drugs. For this reason, scientists must constantly work to develop new drugs or drug combinations to combat the worldwide malaria burden.[footnote]Sumiti Vinayak, et al., \"Origin and Evolution of Sulfadoxine Resistant\u00a0<em>Plasmodium falciparum<\/em>,\" <em>Public Library of Science Pathogens<\/em> 6, no. 3 (2010): e1000830, doi:10.1371\/journal.ppat.1000830.[\/footnote]\r\n\r\n<\/div>\r\n<h2>Multiple Alleles (ABO Blood Types) and Punnett Squares<\/h2>\r\nhttps:\/\/youtu.be\/9O5JQqlngFY\r\n<div class=\"textbox tryit\">\r\n<h3>Try It<\/h3>\r\nhttps:\/\/assess.lumenlearning.com\/practice\/d7712e83-6e4b-4a8e-ba95-516fa0d0e39b\r\n<\/div>","rendered":"<div class=\"textbox learning-objectives\">\n<h3>Learning Outcomes<\/h3>\n<ul>\n<li>Explain how mutli-allele inheritance will impact a trait within in a population<\/li>\n<\/ul>\n<\/div>\n<p>Mendel implied that only two alleles, one dominant and one recessive, could exist for a given gene. We now know that this is an oversimplification. Although individual humans (and all diploid organisms) can only have two alleles for a given gene, multiple alleles may exist at the population level such that many combinations of two alleles are observed. Note that when many alleles exist for the same gene, the convention is to denote the most common phenotype or genotype among wild animals as the\u00a0<strong>wild type<\/strong> (often abbreviated &#8220;+&#8221;); this is considered the standard or norm. All other phenotypes or genotypes are considered <strong>variants<\/strong> of this standard, meaning that they deviate from the wild type. The variant may be recessive or dominant to the wild-type allele.<\/p>\n<p>An example of multiple alleles is coat color in rabbits (Figure 1). Here, four alleles exist for the\u00a0<em>c<\/em> gene. The wild-type version, <em>C<sup data-redactor-tag=\"sup\">+<\/sup>C<sup>+<\/sup><\/em>, is expressed as brown fur. The chinchilla phenotype, <em>c<i data-redactor-tag=\"i\"><sup>ch<\/sup><\/i>c<i><sup data-redactor-tag=\"sup\">ch<\/sup><\/i><\/em>, is expressed as black-tipped white fur. The Himalayan phenotype, <em>c<i data-redactor-tag=\"i\"><sup>h<\/sup><\/i>c<i><sup data-redactor-tag=\"sup\">h<\/sup><\/i><\/em>, has black fur on the extremities and white fur elsewhere. Finally, the albino, or &#8220;colorless&#8221; phenotype, <em>cc<\/em>, is expressed as white fur. In cases of multiple alleles, dominance hierarchies can exist. In this case, the wild-type allele is dominant over all the others, chinchilla is incompletely dominant over Himalayan and albino, and Himalayan is dominant over albino. This hierarchy, or allelic series, was revealed by observing the phenotypes of each possible heterozygote offspring.<\/p>\n<div id=\"attachment_1412\" style=\"width: 810px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" aria-describedby=\"caption-attachment-1412\" class=\"size-full wp-image-1412\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/110\/2016\/05\/02182457\/Figure_12_02_05.jpg\" alt=\"This illustration shows the four different variants for coat color in rabbits at the c allele. The genotype CC produces the wild type phenotype, which is brown. The genotype c^{ch}c^{ch} produces the chinchilla phenotype, which is black-tipped white fur. The genotype c^{h}c^{h} produces the Himalayan phenotype, which is white on the body and black on the extremities. The genotype cc produces the recessive phenotype, which is white\" width=\"800\" height=\"574\" \/><\/p>\n<p id=\"caption-attachment-1412\" class=\"wp-caption-text\">Figure 1. Four different alleles exist for the rabbit coat color (<i>C<\/i>) gene.<\/p>\n<\/div>\n<div id=\"attachment_3251\" style=\"width: 360px\" class=\"wp-caption alignright\"><img loading=\"lazy\" decoding=\"async\" aria-describedby=\"caption-attachment-3251\" class=\"wp-image-3251\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/110\/2016\/06\/27185610\/Figure_12_02_06-e1467053834271.jpg\" alt=\"This photo shows Drosophila that has normal antennae on its head, and a mutant that has legs on its head.\" width=\"350\" height=\"315\" \/><\/p>\n<p id=\"caption-attachment-3251\" class=\"wp-caption-text\">Figure 2. As seen in comparing the wild-type <i>Drosophila<\/i> (left) and the <i>Antennapedia<\/i> mutant (right), the Antennapedia mutant has legs on its head in place of antennae.<\/p>\n<\/div>\n<p>The complete dominance of a wild-type phenotype over all other mutants often occurs as an effect of &#8220;dosage&#8221; of a specific gene product, such that the wild-type allele supplies the correct amount of gene product whereas the mutant alleles cannot. For the allelic series in rabbits, the wild-type allele may supply a given dosage of fur pigment, whereas the mutants supply a lesser dosage or none at all. Interestingly, the Himalayan phenotype is the result of an allele that produces a temperature-sensitive gene product that only produces pigment in the cooler extremities of the rabbit&#8217;s body.<\/p>\n<p>Alternatively, one mutant allele can be dominant over all other phenotypes, including the wild type. This may occur when the mutant allele somehow interferes with the genetic message so that even a heterozygote with one wild-type allele copy expresses the mutant phenotype. One way in which the mutant allele can interfere is by enhancing the function of the wild-type gene product or changing its distribution in the body.<\/p>\n<p>One example of this is the <em>Antennapedia<\/em> mutation in <em>Drosophila<\/em> (Figure 2). In this case, the mutant allele expands the distribution of the gene product, and as a result, the <em>Antennapedia<\/em> heterozygote develops legs on its head where its antennae should be.<\/p>\n<div class=\"key-takeaways textbox\">\n<h3>Multiple Alleles Confer Drug Resistance in the Malaria Parasite<\/h3>\n<p>Malaria is a parasitic disease in humans that is transmitted by infected female mosquitoes, including\u00a0<em>Anopheles gambiae<\/em> (Figure 3a), and is characterized by cyclic high fevers, chills, flu-like symptoms, and severe anemia. <em>Plasmodium falciparum<\/em> and <em>P. vivax<\/em> are the most common causative agents of malaria, and <em>P. falciparum<\/em> is the most deadly (Figure 3b)<em>.<\/em> When promptly and correctly treated, <em>P. falciparum<\/em>malaria has a mortality rate of 0.1 percent. However, in some parts of the world, the parasite has evolved resistance to commonly used malaria treatments, so the most effective malarial treatments can vary by geographic region.<\/p>\n<div id=\"attachment_1414\" style=\"width: 979px\" class=\"wp-caption aligncenter\"><img loading=\"lazy\" decoding=\"async\" aria-describedby=\"caption-attachment-1414\" class=\"size-full wp-image-1414\" src=\"https:\/\/s3-us-west-2.amazonaws.com\/courses-images\/wp-content\/uploads\/sites\/110\/2016\/05\/02182928\/Figure_12_02_07.jpg\" alt=\"Photo a shows the Anopheles gambiae mosquito, which carries malaria. Photo b shows a micrograph of sickle-shaped Plasmodium falciparum, the parasite that causes malaria. The Plasmodium is about 0.75 microns across.\" width=\"969\" height=\"489\" \/><\/p>\n<p id=\"caption-attachment-1414\" class=\"wp-caption-text\">Figure 3. The (a) Anopheles gambiae, or African malaria mosquito, acts as a vector in the transmission to humans of the malaria-causing parasite (b) Plasmodium falciparum, here visualized using false-color transmission electron microscopy. (credit a: James D. Gathany; credit b: Ute Frevert; false color by Margaret Shear; scale-bar data from Matt Russell)<\/p>\n<\/div>\n<p>In Southeast Asia, Africa, and South America,\u00a0<em>P. falciparum<\/em> has developed resistance to the anti-malarial drugs chloroquine, mefloquine, and sulfadoxine-pyrimethamine. <em>P. falciparum<\/em>, which is haploid during the life stage in which it is infectious to humans, has evolved multiple drug-resistant mutant alleles of the\u00a0<em>dhps<\/em> gene. Varying degrees of sulfadoxine resistance are associated with each of these alleles. Being haploid, <em>P. falciparum<\/em> needs only one drug-resistant allele to express this trait.<\/p>\n<p>In Southeast Asia, different sulfadoxine-resistant alleles of the\u00a0<em>dhps<\/em> gene are localized to different geographic regions. This is a common evolutionary phenomenon that occurs because drug-resistant mutants arise in a population and interbreed with other <em>P. falciparum<\/em> isolates in close proximity. Sulfadoxine-resistant parasites cause considerable human hardship in regions where this drug is widely used as an over-the-counter malaria remedy. As is common with pathogens that multiply to large numbers within an infection cycle, <em>P. falciparum<\/em> evolves relatively rapidly (over a decade or so) in response to the selective pressure of commonly used anti-malarial drugs. For this reason, scientists must constantly work to develop new drugs or drug combinations to combat the worldwide malaria burden.<a class=\"footnote\" title=\"Sumiti Vinayak, et al., &quot;Origin and Evolution of Sulfadoxine Resistant\u00a0Plasmodium falciparum,&quot; Public Library of Science Pathogens 6, no. 3 (2010): e1000830, doi:10.1371\/journal.ppat.1000830.\" id=\"return-footnote-2752-1\" href=\"#footnote-2752-1\" aria-label=\"Footnote 1\"><sup class=\"footnote\">[1]<\/sup><\/a><\/p>\n<\/div>\n<h2>Multiple Alleles (ABO Blood Types) and Punnett Squares<\/h2>\n<p><iframe loading=\"lazy\" id=\"oembed-1\" title=\"Multiple Alleles (ABO Blood Types) and Punnett Squares\" width=\"500\" height=\"281\" src=\"https:\/\/www.youtube.com\/embed\/9O5JQqlngFY?feature=oembed&#38;rel=0\" frameborder=\"0\" allowfullscreen=\"allowfullscreen\"><\/iframe><\/p>\n<div class=\"textbox tryit\">\n<h3>Try It<\/h3>\n<p>\t<iframe id=\"assessment_practice_d7712e83-6e4b-4a8e-ba95-516fa0d0e39b\" class=\"resizable\" src=\"https:\/\/assess.lumenlearning.com\/practice\/d7712e83-6e4b-4a8e-ba95-516fa0d0e39b?iframe_resize_id=assessment_practice_id_d7712e83-6e4b-4a8e-ba95-516fa0d0e39b\" frameborder=\"0\" style=\"border:none;width:100%;height:100%;min-height:300px;\"><br \/>\n\t<\/iframe>\n<\/div>\n\n\t\t\t <section class=\"citations-section\" role=\"contentinfo\">\n\t\t\t <h3>Candela Citations<\/h3>\n\t\t\t\t\t <div>\n\t\t\t\t\t\t <div id=\"citation-list-2752\">\n\t\t\t\t\t\t\t <div class=\"licensing\"><div class=\"license-attribution-dropdown-subheading\">CC licensed content, Shared previously<\/div><ul class=\"citation-list\"><li>Biology 2e. <strong>Provided by<\/strong>: OpenStax. <strong>Located at<\/strong>: <a target=\"_blank\" href=\"http:\/\/cnx.org\/contents\/185cbf87-c72e-48f5-b51e-f14f21b5eabd@10.8\">http:\/\/cnx.org\/contents\/185cbf87-c72e-48f5-b51e-f14f21b5eabd@10.8<\/a>. <strong>License<\/strong>: <em><a target=\"_blank\" rel=\"license\" href=\"https:\/\/creativecommons.org\/licenses\/by\/4.0\/\">CC BY: Attribution<\/a><\/em>. <strong>License Terms<\/strong>: Access for free at https:\/\/openstax.org\/books\/biology-2e\/pages\/1-introduction<\/li><\/ul><div class=\"license-attribution-dropdown-subheading\">All rights reserved content<\/div><ul class=\"citation-list\"><li>Multiple Alleles (ABO Blood Types) and Punnett Squares. <strong>Authored by<\/strong>: Amoeba Sisters. <strong>Located at<\/strong>: <a target=\"_blank\" href=\"https:\/\/youtu.be\/9O5JQqlngFY\">https:\/\/youtu.be\/9O5JQqlngFY<\/a>. <strong>License<\/strong>: <em>All Rights Reserved<\/em>. <strong>License Terms<\/strong>: Standard YouTube License<\/li><\/ul><\/div>\n\t\t\t\t\t\t <\/div>\n\t\t\t\t\t <\/div>\n\t\t\t <\/section><hr class=\"before-footnotes clear\" \/><div class=\"footnotes\"><ol><li id=\"footnote-2752-1\">Sumiti Vinayak, et al., \"Origin and Evolution of Sulfadoxine Resistant\u00a0<em>Plasmodium falciparum<\/em>,\" <em>Public Library of Science Pathogens<\/em> 6, no. 3 (2010): e1000830, doi:10.1371\/journal.ppat.1000830. <a href=\"#return-footnote-2752-1\" class=\"return-footnote\" aria-label=\"Return to footnote 1\">&crarr;<\/a><\/li><\/ol><\/div>","protected":false},"author":17,"menu_order":10,"template":"","meta":{"_candela_citation":"[{\"type\":\"cc\",\"description\":\"Biology 2e\",\"author\":\"\",\"organization\":\"OpenStax\",\"url\":\"http:\/\/cnx.org\/contents\/185cbf87-c72e-48f5-b51e-f14f21b5eabd@10.8\",\"project\":\"\",\"license\":\"cc-by\",\"license_terms\":\"Access for free at https:\/\/openstax.org\/books\/biology-2e\/pages\/1-introduction\"},{\"type\":\"copyrighted_video\",\"description\":\"Multiple Alleles (ABO Blood Types) and Punnett Squares\",\"author\":\"Amoeba Sisters\",\"organization\":\"\",\"url\":\"https:\/\/youtu.be\/9O5JQqlngFY\",\"project\":\"\",\"license\":\"arr\",\"license_terms\":\"Standard YouTube License\"}]","CANDELA_OUTCOMES_GUID":"f89c49c8-b9d0-4a0a-bafa-6d50770a79bd, 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