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66-513: A complete deletion of this complex results in the posterior segments of the Drosophila embryo becoming copies of the second thoracic segment. The first homeotic mutation to be isolated was the bithorax (bx) mutant in 1915 by Calvin Bridges, which was later worked on and better characterized as a gene cluster by Edward B. Lewis . Lewis found that BX-C contained genes that code for proteins controlling

132-438: A hypoxia treatment experience decreased thorax length, while hyperoxia produces smaller flight muscles, suggesting negative developmental effects of extreme oxygen levels. Circadian rhythms are also subject to developmental plasticity. Light conditions during development affect daily activity patterns in Drosophila melanogaster , where flies raised under constant dark or light are less active as adults than those raised under

198-591: A BA in Biostatistics from the University of Minnesota in 1939, where he worked on Drosophila melanogaster in the lab of C.P. Oliver . In 1942 Lewis received a PhD from California Institute of Technology (Caltech), working under the guidance of Alfred Sturtevant . In 1939, Edward B. Lewis arrived at Caltech and finished his PhD within three years. Lewis enrolled in the U.S. Army Air Corps training program in meteorology in 1942 and later received his master's degree in

264-833: A Pittsburgh physicist, argued that the linearity thesis was confirmed by the research of Alice Stewart . (See also John Gofman ) On November 20, 2001, Lewis was interviewed by Elliot Meyerowitz in the Kerckhoff Library at the California Institute of Technology, Pasadena, California. This interview was released on DVD in 2004 as "Conversations in Genetics: Volume 1, No. 3 – Edward B. Lewis; An Oral History of Our Intellectual Heritage in Genetics" 67 min; Producer Rochelle Easton Esposito; The Genetics Society of America. Drosophila melanogaster Drosophila melanogaster

330-419: A cluster of spiky hairs (claspers) surrounding the reproducing parts used to attach to the female during mating. Extensive images are found at FlyBase . Drosophila melanogaster can be distinguished from related species by the following combination of features: gena ~1/10 diameter of eye at greatest vertical height; wing hyaline and with costal index 2.4; male protarsus with a single row of ~12 setae forming

396-498: A complete BX-C deletion, suggesting that the three transcription units are the only functional protein-coding units in the complex. This was confirmed when the entire complex was eventually sequenced in 1995. Only about 1.4% of the BX-C sequence codes for proteins. Deletion of the entire complex results in death of the embryo late in development. It also shows changes in the segmentation of the embryonic cuticle, with all abdominal segments and

462-448: A few hundred, very long (1.76 mm) sperm cells in seminal fluid to the female. Females store the sperm in a tubular receptacle and in two mushroom-shaped spermathecae ; sperm from multiple matings compete for fertilization. A last male precedence is believed to exist; the last male to mate with a female sires about 80% of her offspring. This precedence was found to occur through both displacement and incapacitation. The displacement

528-560: A fully formed adult fruit fly. Males perform a sequence of five behavioral patterns to court females. First, males orient themselves while playing a courtship song by horizontally extending and vibrating their wings. Soon after, the male positions himself at the rear of the female's abdomen in a low posture to tap and lick the female genitalia. Finally, the male curls his abdomen and attempts copulation. Females can reject males by moving away, kicking, and extruding their ovipositor. Copulation lasts around 15–20 minutes, during which males transfer

594-578: A genetic model for several human diseases including the neurodegenerative disorders Parkinson's , Huntington's , spinocerebellar ataxia and Alzheimer's disease . The fly is also being used to study mechanisms underlying aging and oxidative stress , immunity , diabetes , and cancer , as well as drug abuse . The life cycle of this insect has four stages: fertilized egg, larva, pupa, and adult. Embryogenesis in Drosophila has been extensively studied, as its small size, short generation time, and large brood size makes it ideal for genetic studies. It

660-529: A greater mating efficiency for experienced males over naïve males. This modification also appears to have obvious evolutionary advantages, as increased mating efficiency is extremely important in the eyes of natural selection . Both male and female D. melanogaster flies act polygamously (having multiple sexual partners at the same time). In both males and females, polygamy results in a decrease in evening activity compared to virgin flies, more so in males than females. Evening activity consists of those in which

726-773: A laboratory known as the Fly Room. The Fly Room was cramped with eight desks, each occupied by students and their experiments. They started off experiments using milk bottles to rear the fruit flies and handheld lenses for observing their traits. The lenses were later replaced by microscopes, which enhanced their observations. Morgan and his students eventually elucidated many basic principles of heredity, including sex-linked inheritance, epistasis , multiple alleles, and gene mapping . D. melanogaster had historically been used in laboratories to study genetics and patterns of inheritance. However, D. melanogaster also has importance in environmental mutagenesis research, allowing researchers to study

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792-460: A presentation to a Congressional committee on atomic energy in 1957. At the scientific level of the debate, the crucial question was whether the "threshold theory" was valid or whether, as Lewis insisted, the effects of radioactivity were " linear with no threshold ", where every exposure to radiation had a long-term cumulative effect. The issue of linearity versus threshold re-entered the debate on nuclear fallout in 1962, when Ernest Sternglass ,

858-443: A sex comb; male epandrial posterior lobe small and nearly triangular; female abdominal tergite 6 with dark band running to its ventral margin; female oviscapt small, pale, without dorsodistal depression and with 12-13 peg-like outer ovisensilla. Drosophila melanogaster flies can sense air currents with the hairs on their backs. Their eyes are sensitive to slight differences in light intensity and will instinctively fly away when

924-510: A shadow or other movement is detected. Under optimal growth conditions at 25 °C (77 °F), the D. melanogaster lifespan is about 50 days from egg to death. The developmental period for D. melanogaster varies with temperature, as with many ectothermic species. The shortest development time (egg to adult), seven days, is achieved at 28 °C (82 °F). Development times increase at higher temperatures (11 days at 30 °C or 86 °F) due to heat stress. Under ideal conditions,

990-461: A visual and physical, partial unilateral paralysis, which limited her mobility. Lewis maintained a constant exercise routine in his day, as he started his mornings with breakfast and exercise, until he suffered from cancer. Lewis would normally have lunch with his faculty members at the Anthenaeum, then take a nap and return to the laboratory in the evening. The most constant part of his daily routine

1056-447: Is a holometabolous insect, so it undergoes a full metamorphosis. Their life cycle is broken down into four stages: embryo, larva, pupa, adult. The eggs, which are about 0.5 mm long, hatch after 12–15 hours (at 25 °C or 77 °F). The resulting larvae grow for about four days (at 25 °C) while molting twice (into second- and third-instar larvae), at about 24 and 48 hours after hatching. During this time, they feed on

1122-439: Is a species of fly (an insect of the order Diptera) in the family Drosophilidae . The species is often referred to as the fruit fly or lesser fruit fly , or less commonly the " vinegar fly", " pomace fly", or " banana fly". In the wild, D. melanogaster are attracted to rotting fruit and fermenting beverages, and are often found in orchards, kitchens and pubs. Starting with Charles W. Woodworth 's 1901 proposal of

1188-409: Is also unique among model organisms in that cleavage occurs in a syncytium . During oogenesis, cytoplasmic bridges called "ring canals" connect the forming oocyte to nurse cells. Nutrients and developmental control molecules move from the nurse cells into the oocyte. In the figure to the left, the forming oocyte can be seen to be covered by follicular support cells. After fertilization of the oocyte,

1254-577: Is attributed to sperm handling by the female fly as multiple matings are conducted and is most significant during the first 1–2 days after copulation. Displacement from the seminal receptacle is more significant than displacement from the spermathecae. Incapacitation of first male sperm by second male sperm becomes significant 2–7 days after copulation. The seminal fluid of the second male is believed to be responsible for this incapacitation mechanism (without removal of first male sperm) which takes effect before fertilization occurs. The delay in effectiveness of

1320-443: Is followed by the name of the gene affected and a description of its phenotype. (Note: Recessive alleles are in lower case, while dominant alleles are capitalised.) Drosophila genes are traditionally named after the phenotype they cause when mutated. For example, the absence of a particular gene in Drosophila will result in a mutant embryo that does not develop a heart. Scientists have thus called this gene tinman , named after

1386-450: Is induced by the DNA damaging agents ultraviolet light and mitomycin C . Females become receptive to courting males about 8–12 hours after emergence. Specific neuron groups in females have been found to affect copulation behavior and mate choice. One such group in the abdominal nerve cord allows the female fly to pause her body movements to copulate. Activation of these neurons induces

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1452-474: Is located on the right arm of chromosome 3 alongside the other homeotic gene complex, the Antennapedia (Antp) complex . It is over 300 kb long and contains nine distinct cis-regulatory domains. The three transcription units, Ubx, abd-A, and Abd-B , are spread out evenly on the complex. A mutant embryo with a triple knock-out of Ubx, abd-A, and Abd-B has a phenotype that is identical to that of an embryo with

1518-422: Is one of the most widely used and genetically best-known of all eukaryotic organisms. All organisms use common genetic systems; therefore, comprehending processes such as transcription and replication in fruit flies helps in understanding these processes in other eukaryotes, including humans . Thomas Hunt Morgan began using fruit flies in experimental studies of heredity at Columbia University in 1910 in

1584-409: Is relatively very small and therefore often ignored, aside from its important eyeless gene. The D. melanogaster sequenced genome of 139.5 million base pairs has been annotated and contains around 15,682 genes according to Ensemble release 73. More than 60% of the genome appears to be functional non-protein-coding DNA involved in gene expression control. Determination of sex in Drosophila occurs by

1650-485: Is sent to a brain region that is a homolog of the hypothalamus and the hypothalamus then controls sexual behavior and desire. Gonadotropic hormones in Drosophila maintain homeostasis and govern reproductive output via a cyclic interrelationship, not unlike the mammalian estrous cycle . Sex peptide perturbs this homeostasis and dramatically shifts the endocrine state of the female by inciting juvenile hormone synthesis in

1716-660: Is served. Flies belonging to the family Tephritidae are also called "fruit flies". This can cause confusion, especially in the Mediterranean, Australia , and South Africa , where the Mediterranean fruit fly Ceratitis capitata is an economic pest. The term " Drosophila ", meaning "dew-loving", is a modern scientific Latin adaptation from Greek words δρόσος , drósos , " dew ", and φιλία , philía , "lover". The term " melanogaster " meaning "black-belly", comes from Ancient Greek μέλας , mélas , “black”, and γᾰστήρ , gastḗr , "belly". Unlike humans ,

1782-420: Is typically used in research owing to its rapid life cycle, relatively simple genetics with only four pairs of chromosomes , and large number of offspring per generation. It was originally an African species, with all non-African lineages having a common origin. Its geographic range includes all continents, including islands. D. melanogaster is a common pest in homes, restaurants, and other places where food

1848-400: The D. melanogaster homolog of WRN also cause increased physiologic signs of aging, such as shorter lifespan, higher tumor incidence, muscle degeneration, reduced climbing ability, altered behavior and reduced locomotor activity. Meiotic recombination in D. melanogaster appears to be employed in repairing damage in germ-line DNA as indicated by the findings that meiotic recombination

1914-659: The Oz character of the same name . Likewise changes in the Shavenbaby gene cause the loss of dorsal cuticular hairs in Drosophila sechellia larvae. This system of nomenclature results in a wider range of gene names than in other organisms. The genome of D. melanogaster (sequenced in 2000, and curated at the FlyBase database ) contains four pairs of chromosomes – an X/Y pair, and three autosomes labeled 2, 3, and 4. The fourth chromosome

1980-514: The X:A ratio of X chromosomes to autosomes, not because of the presence of a Y chromosome as in human sex determination. Although the Y chromosome is entirely heterochromatic , it contains at least 16 genes, many of which are thought to have male-related functions. There are three transferrin orthologs, all of which are dramatically divergent from those known in chordate models. A June 2001 study by National Human Genome Research Institute comparing

2046-409: The mesoderm , and posterior and anterior invagination of endoderm (gut), as well as extensive body segmentation until finally hatching from the surrounding cuticle into a first-instar larva. During larval development, tissues known as imaginal discs grow inside the larva. Imaginal discs develop to form most structures of the adult body, such as the head, legs, wings, thorax, and genitalia. Cells of

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2112-429: The microorganisms that decompose the fruit, as well as on the sugar of the fruit itself. The mother puts feces on the egg sacs to establish the same microbial composition in the larvae's guts that has worked positively for herself. Then the larvae encapsulate in the puparium and undergo a four-day-long metamorphosis (at 25 °C), after which the adults eclose (emerge). Drosophila melanogaster, commonly known as

2178-484: The abdomen. The black portions of the abdomen are the inspiration for the species name ( melanogaster = "black-bellied" ). The brick-red color of the eyes of the wild type fly are due to two pigments: xanthommatin, which is brown and is derived from tryptophan , and drosopterins, which are red and are derived from guanosine triphosphate . They exhibit sexual dimorphism ; females are about 2.5 mm (0.10 in) long; males are slightly smaller. Furthermore, males have

2244-475: The area a year later. As he left for military service in 1943, he was told by the university president Robert A. Millikan that he had a position as an instructor at Caltech when he returned. He served working mostly as a weather forecaster in Hawaii and Okinawa for four years. Lewis returned in 1946 and took his position at Caltech where his duties included helping in the laboratory for an introductory genetics course. He

2310-487: The chromosome in the same order as the parasegments that they act in.  Loss-of-function mutations of one sub-domain will result in a target parasegment becoming a copy of the parasegment that is immediately anterior to it. Edward B. Lewis Edward Butts Lewis (May 20, 1918 – July 21, 2004) was an American geneticist , a corecipient of the 1995 Nobel Prize in Physiology or Medicine . He helped to found

2376-906: The copulation of others). This behavior is sensitive to environmental conditions, and females copulate less in bad weather conditions. D. melanogaster males exhibit a strong reproductive learning curve. That is, with sexual experience, these flies tend to modify their future mating behavior in multiple ways. These changes include increased selectivity for courting only intraspecifically, as well as decreased courtship times. Sexually naïve D. melanogaster males are known to spend significant time courting interspecifically, such as with D. simulans flies. Naïve D. melanogaster will also attempt to court females that are not yet sexually mature, and other males. D. melanogaster males show little to no preference for D. melanogaster females over females of other species or even other male flies. However, after D. simulans or other flies incapable of copulation have rejected

2442-421: The corpus allatum. D. melanogaster is often used for life extension studies, such as to identify genes purported to increase lifespan when mutated . D. melanogaster is also used in studies of aging . Werner syndrome is a condition in humans characterized by accelerated aging. It is caused by mutations in the gene WRN that encodes a protein with essential roles in repair of DNA damage. Mutations in

2508-454: The development time at 25 °C (77 °F) is 8 + 1 ⁄ 2 days, at 18 °C (64 °F) it takes 19 days and at 12 °C (54 °F) it takes over 50 days. Under crowded conditions, development time increases, while the emerging flies are smaller. Females lay some 400 eggs (embryos), about five at a time, into rotting fruit or other suitable material such as decaying mushrooms and sap fluxes . Drosophila melanogaster

2574-405: The early embryo (or syncytial embryo ) undergoes rapid DNA replication and 13 nuclear divisions until about 5000 to 6000 nuclei accumulate in the unseparated cytoplasm of the embryo. By the end of the eighth division, most nuclei have migrated to the surface, surrounding the yolk sac (leaving behind only a few nuclei, which will become the yolk nuclei). After the 10th division, the pole cells form at

2640-448: The effects of radiation from X-rays , nuclear fallout and other sources as possible causes of cancer. He reviewed medical records from survivors of the atomic bombings of Hiroshima and Nagasaki , as well as radiologists and patients exposed to X-rays. Lewis concluded that "health risks from radiation had been underestimated". Lewis published articles in Science and other journals and made

2706-406: The effects of specific environmental mutagens. There are many reasons the fruit fly is a popular choice as a model organism: Genetic markers are commonly used in Drosophila research, for example within balancer chromosomes or P-element inserts, and most phenotypes are easily identifiable either with the naked eye or under a microscope. In the list of a few common markers below, the allele symbol

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2772-451: The female fruit fly and are present in the early stages of embryo development. They determine the embryo's main features and early development. For example, the gene called Bicoid regulates the formation of the embryo's anterior end, and its absence leads to an embryo lacking a head. Zygotic genes: These genes are activated in later stages of embryo development when the fruit fly embryo begins to produce its own genetic products. For example,

2838-494: The female to cease movement and orient herself towards the male to allow for mounting. If the group is inactivated, the female remains in motion and does not copulate. Various chemical signals such as male pheromones often are able to activate the group. Also, females exhibit mate choice copying . When virgin females are shown other females copulating with a certain type of male, they tend to copulate more with this type of male afterwards than naïve females (which have not observed

2904-455: The field of evolutionary developmental biology . Lewis was born in Wilkes-Barre, Pennsylvania , the second son of Laura Mary Lewis (née Histed) and Edward Butts Lewis, a watchmaker-jeweler. His full name was supposed to be Edward Butts Lewis Jr., but his birth certificate was incorrectly filled out with "B." as his middle name. Lewis graduated from E. L. Meyers High School . He received

2970-492: The flies participate other than mating and finding partners, such as finding food. The reproductive success of males and females varies, because a female only needs to mate once to reach maximum fertility. Mating with multiple partners provides no advantage over mating with one partner, so females exhibit no difference in evening activity between polygamous and monogamous individuals. For males, however, mating with multiple partners increases their reproductive success by increasing

3036-413: The fruit fly and human genome estimated that about 60% of genes are conserved between the two species. About 75% of known human disease genes have a recognizable match in the genome of fruit flies, and 50% of fly protein sequences have mammalian homologs . An online database called Homophila is available to search for human disease gene homologues in flies and vice versa. Drosophila is being used as

3102-485: The fruit fly, has been a significant model organism in embryonic development research. Many of its genes that regulate embryonic development and their mechanisms of action have been crucial in understanding the fundamental principles of embryonic development regulation in many multicellular organisms, including humans. Here are some important genes regulating embryonic development in Drosophila melanogaster and their modes of action: Maternal genes: These genes are encoded in

3168-529: The genetic diversity of their offspring. This benefit of genetic diversity is an evolutionary advantage because it increases the chance that some of the offspring will have traits that increase their fitness in their environment. The difference in evening activity between polygamous and monogamous male flies can be explained with courtship. For polygamous flies, their reproductive success increases by having offspring with multiple partners, and therefore they spend more time and energy on courting multiple females. On

3234-462: The gradient. For example, the Hedgehog morphogen regulates the differentiation of segments and segment identity in the fruit fly embryo. These genes and their modes of action form a complex regulatory network that guides the embryonic development of Drosophila melanogaster. They influence cell differentiation, segment formation, and axial patterning in the embryo, ultimately leading to the development of

3300-412: The groundwork for our current understanding of the universal, evolutionarily conserved strategies controlling animal development. He is credited with development of the complementation test . His key publications in the fields of genetics , developmental biology , radiation and cancer are presented in the book Genes, Development and Cancer , which was released in 2004. During the 1950s, Lewis studied

3366-482: The hunchback gene regulates the formation of segments in the embryo. Homeotic genes: This gene family regulates segmentation and axial patterning in development. They act as regulatory factors that determine cell fate in embryonic development. For example, the gene called Antennapedia regulates the formation of anterior limbs in the embryo. Morphogens: These are molecules that form gradients in embryonic development and regulate cell fate depending on their position in

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3432-768: The imaginal disks are set aside during embryogenesis and continue to grow and divide during the larval stages—unlike most other cells of the larva, which have differentiated to perform specialized functions and grow without further cell division. At metamorphosis, the larva forms a pupa , inside which the larval tissues are reabsorbed and the imaginal tissues undergo extensive morphogenetic movements to form adult structures. Biotic and abiotic factors experienced during development will affect developmental resource allocation leading to phenotypic variation , also referred to as developmental plasticity. As in all insects, environmental factors can influence several aspects of development in Drosophila melanogaster . Fruit flies reared under

3498-486: The incapacitation mechanism is believed to be a protective mechanism that prevents a male fly from incapacitating his own sperm should he mate with the same female fly repetitively. Sensory neurons in the uterus of female D. melanogaster respond to a male protein, sex peptide , which is found in semen. This protein makes the female reluctant to copulate for about 10 days after insemination . The signal pathway leading to this change in behavior has been determined. The signal

3564-509: The level of units called parasegments. The parasegments are slightly misaligned with the anatomical segments, each comprising the posterior part of one segment and the anterior part of the adjacent segment. The parasegment-specific expression patterns of the BX-C genes are controlled by large and complex cis-regulatory regions .  9 functionally distinct cis-regulatory sub-domains have been identified: abx/bx, bxd/pbx, iab-2, iab-3, iab-4, iab-5, iab-6, iab-7, and iab-8,9. They are arranged along

3630-502: The levels of thoracic and abdominal development, and postulated that at least four of these genes were regulated by cis-regulatory elements, as well as a separate locus (Polycomb) that acted as a repressor . However, subsequent research has found that the complex only contains three genes that are differentially expressed in each parasegment. Lewis’ work on BX-C earned him the Nobel Prize in Physiology or Medicine in 1995. The gene complex

3696-582: The males' advances, D. melanogaster males are much less likely to spend time courting nonspecifically in the future. This apparent learned behavior modification seems to be evolutionarily significant, as it allows the males to avoid investing energy into futile sexual encounters. In addition, males with previous sexual experience modify their courtship dance when attempting to mate with new females—the experienced males spend less time courting, so have lower mating latencies, meaning that they are able to reproduce more quickly. This decreased mating latency leads to

3762-638: The nuclei that have made a mistake detach from their centrosomes and fall into the centre of the embryo (yolk sac), which will not form part of the fly. The gene network (transcriptional and protein interactions) governing the early development of the fruit fly embryo is one of the best understood gene networks to date, especially the patterning along the anteroposterior (AP) and dorsoventral (DV) axes (See under morphogenesis ). The embryo undergoes well-characterized morphogenetic movements during gastrulation and early development, including germ-band extension , formation of several furrows, ventral invagination of

3828-455: The other hand, monogamous flies only court one female, and expend less energy doing so. While it requires more energy for male flies to court multiple females, the overall reproductive benefits it produces has kept polygamy as the preferred sexual choice. The mechanism that affects courtship behavior in Drosophila is controlled by the oscillator neurons DN1s and LNDs. Oscillation of the DN1 neurons

3894-479: The posterior end of the embryo, segregating the germ line from the syncytium. Finally, after the 13th division, cell membranes slowly invaginate, dividing the syncytium into individual somatic cells. Once this process is completed, gastrulation starts. Nuclear division in the early Drosophila embryo happens so quickly, no proper checkpoints exist, so mistakes may be made in division of the DNA. To get around this problem,

3960-404: The sex and physical appearance of fruit flies is not influenced by hormones . The appearance and sex of fruit flies is determined only by genetic information. Female fruit flies are substantially larger than male fruit flies, with females having bodies that are up to 30% larger than an adult male. Wild type fruit flies are yellow-brown, with brick-red eyes and transverse black rings across

4026-416: The third thoracic segment resembling the second thoracic segment. Individual mutations within the complex also result in segmental changes, though not as dramatic as the entire deletion of the complex. BX-C controls the differential patterning of the posterior thoracic segment (T3) and all 8 of the abdominal segments (A1-A8) of Drosophila . During embryonic development, regulation of BX-C expression occurs on

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4092-410: The use of this species as a model organism , D. melanogaster continues to be widely used for biological research in genetics , physiology , microbial pathogenesis , and life history evolution . D. melanogaster was the first animal to be launched into space in 1947. As of 2017, six Nobel Prizes have been awarded to drosophilists for their work using the insect. Drosophila melanogaster

4158-474: Was a humble man who did not always receive attention for his works. Over time, his work with Drosophila became more appreciated and he began to attract more attention. His Nobel Prize–winning studies with Drosophila , (including the discovery of the Drosophila Bithorax complex of homeotic genes , and elucidation of its function), founded the field of evolutionary developmental biology and laid

4224-416: Was found to be effected by sociosexual interactions , and is connected to mating-related decrease of evening activity. D. melanogaster remains one of the most studied organisms in biological research, particularly in genetics and developmental biology. It is also employed in studies of environmental mutagenesis. D. melanogaster was among the first organisms used for genetic analysis , and today it

4290-576: Was promoted in 1956 to a professor and became the Thomas Hunt Morgan Professor of Biology in 1966. In 1946, Edward B. Lewis met Pamela Harrah (1925–2018). She was an accomplished artist, but also shared Lewis' interests in animal life. She had gone to Stanford and studied genetics and later discovered the mutant Polycomb, that now is important in the understanding of gene regulation. They married and had three sons named Glenn, Hugh, and Keith. Pam developed an infection that caused her to have

4356-446: Was that he would do most of his work at night. He also enjoyed playing the flute and would allow himself to have time to play the flute at night. In addition, Lewis appreciated other aspects of life even though they interrupted his typical schedule. Some other things he did with his time include: jogging, swimming, playing on the beach, playing chamber music with friends, going to see movies, and attending opera performances. Edward B. Lewis

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