C Albicans

Word Count: 2965 |

The related yeasts Saccharyomyces cervisiae and Candida albicans can grow by producing a bud which is identical to the mother cell. What other forms of growth and development do these species undergo? Is there a connection between different growth forms and infection of humans by Candida?

C. albicans and S. cervisiae are both able to grow as pseudohyphae and yeast. In C. albicans there is a third growth form, the hyphal growth form.

In the presence of varying concentrations of nutrients, S. Cervisiae cells may adopt a number of different fates. In the presence of favourable environmental conditions, i.e. abundant nitrogen supply and abundant fermentable carbon source, S. Cervisiae will adopt the yeast form, characterized by asymmetric budding, and proliferate. If nutrients are limited but not exhausted, e.g. a limiting nitrogen source and an abundant fermentable carbon source, S. Cervisiae will switch from the yeast form to a filamentous form. This is known as pseudohyphal differentiation, and growth in this form produces pseudohyphae. When nutrients are exhausted, i.e. limiting nitrogen source, limiting fermentable carbon source, S. Cervisiae will undergo sporulation producing spores. (1, lecture notes).

C. albicans cells assume different growth forms and morhologies depending on environmental conditions. In cultures grown at low temperature and/or pH, e.g below 30 C or pH 4.0 the yeast form is prevalent. Hyphae develop from yeast cells in a response to a number of growth conditions including; temperatures above 34 C in the presence of Serum, in Lees medium at 37 C, in cultures at 37 C and neutral pH, and in the presence of N-acetylglucosamine. Given any of these environmental conditions the majority of growth is hyphal. Cultures grown in intermediate temperatures and pH are seen to contain pseudohyphae. Pseudohyphae can be reliably induced in cultures at pH 6.0 and 35 C, in Nitrogen limited growth on a solid medium or in the presence of high concentrations of phosphate. It is worth noting that pseudohyphal cultures in C. albicans always contain some yeast and/or hyphae and the proportions at which the various morphological forms are present depends on the environmental conditions. There are also growth conditions which have not been well characterised which induce filamentous growth, these include; engulfment by macrophages, growth in mouse kidneys and Iron deprivation (5, 3). In early study’s on C. albicans there was often little attempt to distinguish between hyphae and pseudohyphae with both forms being placed under the umbrella of filamentous growth. This is misleading as the two forms, despite superficial similarities, are morphologically distinct. In fact, despite their filamentous form, pseudohyphae may be more accurately described as yeast cells altered by polarized growth that don’t completely separate after cytokenesis (2, 3).

S. Cervisiae and C. albicans, in common with all yeasts, are able to grow by budding which produces a daughter cell identical to the mother. This type of growth is termed the yeast mode (2). The yeasts of the two species are similar in terms of shape, size and the order of cell cycle progression, although there are significant differences on a molecular level (5). Different stages in the yeast cell cycle are defined by the movements of actins and septins which make up the cytoskeleton and, along with the polarisome, regulate polarised growth. They also dictate the site of bud emergence, mitosis and cytokenisis. In the the first growth phase, G1, actin patches congregate at one pole of the cell and a septin ring appears in the same location, this is known as the pre-bud site. It is at this point that the new bud will envaginate. The location of this site is determined by the previous site of cytokenesis as well as cell type and environmental factors such as temperature (5, 6, 7). This leads to a regulatory point of the cell cycle known as START, located at the transition between G1 and S phase, at which point the cell is committed to reproduce. Passage through START is nessecary for the initiation of bud formation, DNA replication and spindle pole body duplication. After the passage through START a new bud emerges. The septin ring remains at the budding site, defining the junction between mother and bud known as the mother-bud neck. The actin patches and the polarisome remain at the tip of the cell resulting in polarised growth. As the cell cycle continues into the second growth phase, G2, growth in the bud switches from being polarised to isotropic (5). This switch occurs when the cell is about two thirds of its final size and results in even growth across the surface of the bud. This is reflected in the movement of the actin patches from localisation at the tip of the bud to even distribution throughout (3). At the same time, the nucleus, aided by microtubules, moves to the mother-bud neck in preparation for mitosis. In budding yeast cells, mitosis occurs across the mother-bud neck at the septin ring. The spindle pole bodies separate, dividing the nucleus, and cytokenesis takes place. This pulls one chromosomal complement into the mother cell and one into the daughter cell and separates the cytoplasm between mother and bud. After the completion of mitosis the septin ring splits into two, a true septum is formed, and mother and daughter cells separate. At the point of separation the daughter cell is slightly smaller than the mother cell. This means that the daughter cell begins the next cell cycle at a slightly later point resulting in an asymmetric growth pattern. Proliferation in the yeast mode produces colonys which are smooth and round (2, 3, 5, 6, 7).

A second mode common to both yeasts is pseudohyphal growth. When S. Cervisiae or C. albicans cells undergo pseudohyphal differentiation, and switch from producing single yeast cells, to producing pseudohyphae, a number of changes occur. Cells become elongated in relation to yeast cells, remain physically attached, and divide in a uni-polar budding pattern (2). These changes correspond to changes in the cell cycle which is in turn regulated by signalling pathways responding to environmental stimulus. The pseudohyphal cell cycle is extremely similar to the cell cycle in budding yeast and as such there is little need for a full description here. There are however, two main differences. Firstly, growth in the daughter cell is more polarised in pseudohyphae than in yeast, and cells remain in the second growth phase G2, for longer, this results in an elongated cell shape (5). Secondly, the mother and daughter cell do not fully separate after cytokenesis. The end result of the first cell cycle, starting with an unbudded yeast cell, is two attached but fully differentiated cells: a mother cell, which retains the same form as when the cycle started, and a daughter cell which is elongated. Following the initial switch of form, pseudohyphae continue to divide in the same manner, producing elongated daughter cells which generally remain attached to the mother cell at the site of the nucleur division. Unlike yeast which bud in a bi-polar pattern, budding in pseudohyphae is unipolar. This means that growth is apical, resulting in long filaments, constricted at the septum, which are able to invade a growth medium. The budding cycle is more synchronised in pseudohyphae than in yeast. This is a result of the longer period spent in G2 which means that mother and daughter cells reach START at roughly the same time (5). In S. cervisiae, pseudohyphae are induced by the presence of limiting nutrients. Given this, it has been postulated that the role of pseudohyphae in S. Cervisiae is to forage for nutrients in poor environmental conditions enabling this non-motile species to increase its chances of survival (1) In C. albicans, pseudohyphae have often been seen as an intermediate state between the yeast and hyphal forms with no unique function. This may seem intuitive as they occur in conditions intermeidiate to those which induce yeast or hyphae, and this remains a possibility. However, given that pseudohyphae are morphologically distinct to both yeast and true hyphae, it is equally likely that pseudohyphae may have unique biological properties and play a distinct role in the life cycle of C. albicans, perhaps in infection (2, 3). Consistent with this pseudohyphae rarely produce hyphae and visa versa. In both S. cervisiae and C. albicans pseudohyphae form invasive colonys which are fibrous and rough (5).

C. albicans has a third distinct growth mode: hyphal growth. In terms of cell cycle and morphology, hyphae in C. albicans are extremely different to both yeast and pseudohyphae. In fact, they bear more similarity to the hyphae of filamentous fungi. Hyphal growth results in narrow multicellular filaments know as germ tubes with parallel walls seperated by septai without constriction.( ) Rather than appearing at the mother-bud neck, the septum is formed in the elongating germ tube and mitosis occurs at this point. These differences reflect differences in cell cycle as well as the presense of the spitzenkorper, an organelle specific to hyphae, which along with the majority of cortical actin patches, remains at the tip of the growing germ tube. The spitzenkorper along with the polarisome is resposiblele for the continuously polarised growth seen in hyphae. In the hyphal cell cycle, germ tubes, unlike buds, appear before START during G1 and grow continuously throughout the cell cycle, even after cytokenesis has taken place. In mitosis, the nucleus migrates from the mother cell into the germ tube and undergos nucleur division there. One nucleus then migrates back into the mother cell while the other moves towards the tip of the growing germ tube. The movement of the nuclei is orchestrated by mictrotubules. Another distinguishing feature of hyphae is continuous linear growth. Linear growth and hyphal branching are regulated by cytoplasm and vacuole inheritance. In the first cell cycle a large vacoule appears in the mother cell. After cytokenesis the germ tube inherits mainly cytoplasm while the mother cell inherits mainly vacuole. This causes the mother cell to remain in the G1 phase for several generations, accumulating cytoplasmic mass until a certain threshold of cytoplasm is reached in relation to the vacuole. At this point the mother cell re-enters the cell cylce, elaborating another germ tube. While the mother cell remains in G1, the germ tube continues to grow and divide. This pattern is consistent in all subsequent cell divisions with the apical cell inheriting the majorityy of cytoplasm and the sub-apical cell inheriting the majority of vacuole. Sub-apical cells are only able to produce a new branch when a certain ratio of vacuole to cytoplasm is reached. As a result hyphal growth is linear and hyphae are less branched than pseudohyphae with a less regular branching pattern. (2, ,3 ,5).

C. albicans and S. cervisae also have other distinct morpholgical forms which relate to specific functions and environmental conditions. The haploid cells of S. Cervisiae are able to grow invasively if left on a rich growth medium for long periods of time. The main aim of S. Cervisiae haploid cells is to locate sexual partners and reproduce. Low concentrations of mating pheremones greatly stimulate haploid invasive growth and yeast cells are known to respond to gradients of mating pheremones. These facts suggest that haploid invasive growth in S. Cervisiae may be a mechanism by which haploid cells are able to find mating partners (1). C. albicans has several other distinct morphologys including chlamydospores and opaque cells. Chlamydospores are round projections, formed at the end of suspendor cells, with large thick walls and a high lipid and carbohydrate content. They may form in environments low in oxygen, light, temparature or nutrients.. Although the function of chlamydospores is unknown, there formation is associated with the pathogenic form of C. albicans and it is therefore thought that they may play some role in the pathogenic lifestyle (2). The opaque form is the best studyed example of ‘phenotype switching’ in C albicans and reprasents the state in which C. albicans, which was until recently considered to be an asexual orgainsm, is able to reproduce sexualy. Opaque cells are elongated and assymetrical with surface pimples. This compares with the usual form of C. albicans cell, White, which are relatively round and smooth. The white-opaque switch is refected in a change in the appearance of colonies from round and white to flat and grey (8).

Changes in the morpholgy of cells reflect an underlying change in cell cycle which is in turn regulated by signal transduction pathways responding to extrenal signals. There are two main signal transduction pathways which control morhogenisis and growth in C. albicans and S. Cervisiae. These are the MAP Kinase or mitogen-activated protein kinase cascades, and the nutrient-sensing cAMP or cyclic AMP pathway. The target of cAMP is the cAMP dependant protein PKA; protein Kinase A. These pathways consist of complex signalling cascades involvong posistive and negative regulation between receptors, proteins and transcription factors, ultimately regulating the expression of genes. These pathways, under various conditions, are able to give rise to different developmental fates (1). Although there are fundamental differences between the pathways in C.albicans and S.cervisae the basic features are conserved. MAP Kinase cascades contain three protein kinases which act in a series: a MAP kinase kinase kinase, a MAP kinase kinase, and a MAP kinase, with each activating the next protein in the cascade by phosphorilation. (9). The cAMP-PKA pathway involves cAMP which is produced in response to external factors, targeting PKA. PKA’s are made up of a regulatory sub-unit and catalytic sub-units. These pathways are not isolated and it has been shown that there is cross talk between them. In S.cervisiae the G protein Ras2 activates both the MAP Kinase and the cAMP-PKA pathways and both pathways together regulate the promoter of the FLO11 gene which is nessecary for pseudohyphal growth (2).

The roles played by signalling pathways can be investigated by studying mutant strains for genes involved in the pathways or disabling genes directly. By examining the phenotype of a mutant strain it is possible to deduce the role played by the gene in which it is deficient. One example of this can be seen in the transcription factor Flo8, which is part of the cAMP signalling pathway and plays a role in pseudohyphal differentiation in S. cervisae. The S. cervisiae strain S288C has a naturally occurring Flo8 mutation which means it is unbale to produce Flo8. This strain is unable to produce pseudohyphae (4).

The MAP kinase and cAMP-PKA pathways play a major role in regulating yeast, psedohyphae and the transition between them in S. cervisiae, and yeast, pseudohyphae, hyphae and the transition between them in C. albicans.

It has long been assumed that, rather than one single growth form being resposible for the pathogenic properties of C, albicans, it is the ability to switch morphologies, specifically between the yeast and hyphal modes of proliferation, that is required for pathogenesis (10) Although there is still no direct, unambiguous evidence for this, there is a wealth of experimental data which implies that this may be the case. Mutants for Efg1p and Cph1g which are locked into the yeast form are avirulent. Mutants for Tup1p and Nrg1p which are locked into the hyphal form are also avirulent.

1. Klaus, B. et al. (2000) Signal transduction Cascades Regulating Fungal Development and Virulence. Microbiology and Molecular Biology Reviews. Vol 64. 4:746-785.
2. Whiteway, M. Bachewich, C. (2007). Morphogenisis in Candida albicans. Annula review of Microbiology. 2007. 61:529-53.
3. Sudbery, P et al. (2004). The distinct morphogenic states of Candida albicans. Trends in Microbiology. From www.sciencedirect.com.
4. Cao, F et al. (2005). The Flo8 Transcription Factor is Essential for Hyphal Development and Virulence in Candida albicans. Molecular Biology of the Cell. Vol 17. 1:295-307.
5. Berman, J. (2006). Morphogenisis and cell cycle progression in Candida albicans. Current opinions in microbiology. 9:596-601.
6. Casamayor, A. Snyder, M. (2002). Bud site selction and cell polarity in buddin yeast. Current opinions in microbiology. 5:179-186.
7. Lew, D. Reed , S. (1995). Cell cycle control of morphogenesis in budding yeast. Current opinions in Genetics and Development. 5:17-23
8. Perez-Martin, J te al. (1999) Phenotype switching in Candida albicans is controlled by a SIR2 gene. The embo journal. Vol.18. 9:2580-2592.
9. Gustin, M et al. (1998) MAP Kinase pathways in the yeast Saccharyomyces cervisae. Microbiology and molecular biology reviews. Vol 62. 4:1264-1300
10. Saville, P et al. (2003). Engineered control of cell morphology In vivo reveals distinct roles for yeast and filamentous forms of Candida albicans during Infection. Vol 2. 5:1053-1060.
11. Roony, P. (2002) Limking fungal morhogenesis with virulence. Cellular microbiology. 4, 3:127-137.
12. Gow, N et al. (2002) Fungal morphogenesis and host invasion. Current opinion in microbiology. 5:366-371
13. Calderone. A, Fonzi, W. (2001).Virulence factors of Candida albicans.

You May Also Find These Documents Helpful

Allegory Of American Pie By Don Mc Lean

Ask anyone what was the defining moment in the rock history of the 1960s was and all you will get is a one word answer: Woodstock. The three day rock festival that defined an era was only one of many music festivals of the '60s. But Woodstock has come to symbolize, "an era of peaceful, free- loving, drug- taking hippie youth, carefree before harsher realities hit..." (Layman 40). The Woodstock festival ended a century filled with many metamorphoses of rock'n'roll, from the era of pop music to the rebirth of folk music to the invention of acid rock. But some cynics say that rock'n'roll died with the death of Buddy Holly before the 60s even began. One such person is Don McLean. The poet behind the haunting epic song about the death of 'danceable' music, McLean wrote the ever popular song, "American Pie" (appendix 1). The most important song in rock'n'roll history, "American Pie", is the song about the demise of rock'n'roll after Buddy Holly's death and the heathenism of rock that resulted. Although McLean himself won't reveal any symbolism in his songs, "American Pie" is one of the most analyzed pieces of literature in modern society. Although not all of its secrets have been revealed, many "scholars" of the sixties will agree that the mystery of this song is one of the reasons it has become so successful- everyone wants to know the meanings of its allegories. Proof of "American Pie's" truth lies in the allegory of the song. Many People enjoy the song but have no idea what it means- Who is the Jester? What is the levee? When the deeper story is found, the importance of the song is unearthed. "American Pie" is not only a song, it is an epic poem about the course of rock'n'roll...

Carl Orffs Philosophies In Music Education

While Carl Orff is a very seminal composer of the 20th century, his greatest success and influence has been in the field of Music Education. Born on July 10th in Munich, Germany in 1895, Orff refused to speak about his past almost as if he were ashamed of it. What we do know, however, is that Orff came from a Bavarian family who was very active in the German military. His father's regiment band would often play through some of the young Orff's first attempts at composing. Although Orff was adamant about the secrecy of his past, Moser's Musik Lexicon says that he studied in the Munich Academy of Music until 1914. Orff then served in the military in the first world war. After the war, he held various positions in the Mannheim and Darmstadt opera houses then returned home to Munich to further study music. In 1925, and for the rest of his life, Orff was the head of a department and co-founder of the Guenther School for gymnastics, music, and dance in Munich where he worked with musical beginners. This is where he developed his Music Education theories. In 1937, Orff's Carmina Burana premiered in Frankfurt, Germany. Needless to say, it was a great success. With the success of Carmina Burana, Orff orphaned all of his previous works except for Catulli Carmina and the En trata which were rewritten to be acceptable by Orff. One of Orff's most admired composers was Monteverdi. In fact, much of Orff's work was based on ancient material. Orff said: I am often asked why I nearly always select old material, fairy tales and legends for my stage works. I do not look upon them as old, but rather as valid material. The time element disappears, and only the spiritual power remains. My...

Johann Sebastian Bach Biography

Throughout the history of music, many great composers, theorists, and instrumentalists have left indelible marks and influences that people today look back on to admire and aspire to. No exception to this idiom is Johann Sebastian Bach, whose impact on music was unforgettable to say the least. People today look back to his writings and works to both learn and admire. He truly can be considered a music history great. Bach, who came from a family of over 53 musicians, was nothing short of a virtuosic instrumentalist as well as a masterful composer. Born in Eisenach, Germany, on March 21, 1685, he was the son of a masterful violinist, Johann Ambrosius Bach, who taught his son the basic skills for string playing. Along with this string playing, Bach began to play the organ which is the instrument he would later on be noted for in history. His instruction on the organ came from the player at Eisenach's most important church. He instructed the young boy rather rigorously until his skills surpassed anyone?s expectations for someone of such a young age. Bach suffered early trauma when his parents died in 1695. He went to go live with his older brother, Johann Christoph, who also was a professional organist at Ohrdruf. He continued his younger brother's education on that instrument, as well as introducing him to the harpsichord. The rigorous training on these instruments combined with Bach?s masterful skill paid off for him at an early age. After several years of studying with his older brother, he received a scholarship to study in Luneberg, Germany, which is located on the northern tip of the country. As a result, he left his brother?s tutelage and went to go and study there. The teenage years brought Bach to several parts of Germany where he...

Michelangelo

Michelangelo was pessimistic in his poetry and an optimist in his artwork. Michelangelo?s artwork consisted of paintings and sculptures that showed humanity in it?s natural state. Michelangelo?s poetry was pessimistic in his response to Strazzi even though he was complementing him. Michelangelo?s sculpture brought out his optimism. Michelangelo was optimistic in completing The Tomb of Pope Julius II and persevered through it?s many revisions trying to complete his vision. Sculpture was Michelangelo?s main goal and the love of his life. Since his art portrayed both optimism and pessimism, Michelangelo was in touch with his positive and negative sides, showing that he had a great and stable personality. Michelangelo?s artwork consisted of paintings and sculptures that showed humanity in it?s natural state. Michelangelo Buonarroti was called to Rome in 1505 by Pope Julius II to create for him a monumental tomb. We have no clear sense of what the tomb was to look like, since over the years it went through at least five conceptual revisions. The tomb was to have three levels; the bottom level was to have sculpted figures representing Victory and bond slaves. The second level was to have statues of Moses and Saint Paul as well as symbolic figures of the active and contemplative life- representative of the human striving for, and reception of, knowledge. The third level, it is assumed, was to have an effigy of the deceased pope. The tomb of Pope Julius II was never finished. What was finished of the tomb represents a twenty-year span of frustrating delays and revised schemes. Michelangelo had hardly begun work on the pope?s tomb when Julius commanded him to fresco the ceiling of the Sistine Chapel to complete the work done in the previous century under Sixtus IV. The overall organization consists of four large triangles at...

Oscar Wilde

Oscar Fingal O'Flahertie Wills Wilde was born in Dublin Ireland on October 16, 1854. He is one of the most talented and most controversial writers of his time. He was well known for his wit, flamboyance, and creative genius and with his little dramatic training showing his natural talent for stage and theatre. He is termed a martyr by some and may be the first true self-publicist and was known for his style of dress and odd behavior. Wilde, 1882 His Father, William Wilde, was a highly accredited doctor and his mother, Jane Francesca Elgee, was a writer of revolutionary poems. Oscar had a brother William Charles Kingsbury along with his father's three illegitimate children, Henry, Emily, and Mary. His sister, Isola Emily Francesca died in 1867 at only ten years of age from a sudden fever, greatly affecting Oscar and his family. He kept a lock of her hair in an envelope and later wrote the poem 'Requiescat' in her memory. Oscar and his brother William both attended the Protora Royal School at Enniskillen. He had little in common with the other children. He disliked games and took more interest in flowers and sunsets. He was extremely passionate about anything that had to do with ancient Greece and with Classics. Wilde during school years In 1871, he was awarded a Royal School Scholarship to Trinity College in Dublin and received many awards and earned the highest honor the college offered to an undergraduate, the Foundation Scholarship. In 1874, he also won the College's Berkley Gold Medal for Greek and was awarded a Demyship to Magdalen College, Oxford. After graduating from Oxford, Oscar moved to London with his friend Frank Miles, a well-known portrait painter of the time. In 1878 his poem Ravenna was published, for which he won the...

The History Of Greek Theater

Theater and drama in Ancient Greece took form in about 5th century BCE, with the Sopocles, the great writer of tragedy. In his plays and those of the same genre, heroes and the ideals of life were depicted and glorified. It was believed that man should live for honor and fame, his action was courageous and glorious and his life would climax in a great and noble death. Originally, the hero's recognition was created by selfish behaviors and little thought of service to others. As the Greeks grew toward city-states and colonization, it became the destiny and ambition of the hero to gain honor by serving his city. The second major characteristic of the early Greek world was the supernatural. The two worlds were not separate, as the gods lived in the same world as the men, and they interfered in the men's lives as they chose to. It was the gods who sent suffering and evil to men. In the plays of Sophocles, the gods brought about the hero's downfall because of a tragic flaw in the character of the hero. In Greek tragedy, suffering brought knowledge of worldly matters and of the individual. Aristotle attempted to explain how an audience could observe tragic events and still have a pleasurable experience. Aristotle, by searching the works of writers of Greek tragedy, Aeschulus, Euripides and Sophocles (whose Oedipus Rex he considered the finest of all Greek tragedies), arrived at his definition of tragedy. This explanation has a profound influence for more than twenty centuries on those writing tragedies, most significantly Shakespeare. Aristotle's analysis of tragedy began with a description of the effect such a work had on the audience as a "catharsis" or purging of the emotions. He decided that catharsis was the purging of two specific emotions, pity and...

Scholarship Essay About Goals

Ever since I was a young kid I have always been interested with aircraft. I was so curious of how airplane's fly. I remember taking my toys apart to see how it works. As a kid I wanted to go to the airport to watch the airplanes land and fly and pondered how this happens. Other kids wanted to go to the amusement places. As I grew older I became more and more interested in aircraft and the technology behind it. I always involved myself with aviation early on. I read books and magazines on aviation, took museum tours, built model airplanes. When I was younger my father would take me to aircraft repair facilities where I would watch in great fascination. In my teens, went up to the military bases and befriended many soldiers involved with aircraft and asked them numerous questions. I got to meet many aeronautics engineers and borrowed their old textbooks and read them till the wee hours of the morning. As technology improved with information superhighway, I logged on the web. Stayed up for hours and hours searching through web pages and web pages of information about aircraft and technology. I started my elementary school in the Philippines, then we moved to U.S. and continued my high school education and graduated. Enrolled at the CCSF to pursue my college education and now I am in the 2nd year in CCSF taking aeronautics. My goal now is to obtain my AS degree from the City College of San Francisco (CCSF) so I can transfer to a University and get a Bachelors degree and to continue for my Masters degree in Aeronautics Engineering. I will strive hard to reach the peak level of my career which is a Professor and hopefully to be an aeronautic professor so...

Circus Circus Enterprises Case Studies

Executive Summary: Circus Circus Enterprises is a leader and will continue to be in the gaming industry. In recent years, they have seen a decline in profit and revenue; management tends to blame the decrease on continuing disruptions from remodeling, expansion, and increased competition. Consequently, Circus has reported decreases in its net income for 1997 and 1998 and management believes this trend will continue as competition heightens. Currently the company is involved in several joint ventures, its brand of casino entertainment has traditionally catered to the low rollers and family vacationers through its theme park. Circus should continue to expand its existing operations into new market segments. This shift will allow them to attract the up scale gambler. Overview Circus Circus Enterprises, Inc founded in 1974 is in the business of entertainment, with its core strength in casino gambling. The company?s asset base, operating cash flow, profit margin, multiple markets and customers, rank it as one of the gaming industry leaders. Partners William G. Bennett an aggressive cost cutter and William N. Pennington purchased Circus Circus in 1974 as a small and unprofitable casino. It went public in 1983, from 1993 to 1997; the average return on capital invested was 16.5%. Circus Circus operates several properties in Las Vegas, Reno, Laughlin, and one in Mississippi, as well as 50% ownership in three other casinos and a theme park. On January 31,1998 Circus reported net income of 89.9 million and revenues of 1.35 billion, this is a down from 100 million on 1.3 billion in 1997. Management sees this decline in revenue due to the rapid and extensive expansion and the increased competition that Circus is facing. Well established in the casino gaming industry the corporation has its focus in the entertainment business and has particularly a popular theme resort concept....

Effect Of Civil War On American Economy

The Economies of the North and South, 1861-1865 In 1861, a great war in American history began. It was a civil war between the north and south that was by no means civil. This war would have great repercussions upon the economy of this country and the states within it. The American Civil War began with secession, creating a divided union of sorts, and sparked an incredibly cataclysmic four years. Although the actual war began with secession, this was not the only driving force. The economy of the Southern states, the Confederacy, greatly if not entirely depended on the institution of slavery. The Confederacy was heavily reliant on agriculture, and they used the profits made from the sale of such raw materials to purchase finished goods to use and enjoy. Their major export was cotton, which thrived on the warm river deltas and could easily be shipped to major ocean ports from towns on the Mississippi and numerous river cities. Slavery was a key part of this, as slaves were the ones who harvested and planted the cotton. Being such an enormous unpaid work force, the profits made were extraordinarily high and the price for the unfinished goods drastically low in comparison; especially since he invention of the cotton gin in 1793 which made the work all that much easier and quicker. In contrast, the economical structure of the Northern states, the Union, was vastly dependent on industry. Slavery did not exist in most of the Union, as there was no demand for it due to the type of industrial development taking place. As the Union had a paid work force, the profits made were lower and the cost of the finished manufactured item higher. In turn, the Union used the profits and purchased raw materials to use. This cycle...

Evaluation Of The Effectiveness Of Trade Embargoes

Although I am a strong critic of the use and effectiveness of economic sanctions, such as trade embargoes, for the sake of this assignment, I will present both their theoretical advantages and their disadvantages based upon my research. Trade embargoes and blockades have traditionally been used to entice nations to alter their behavior or to punish them for certain behavior. The intentions behind these policies are generally noble, at least on the surface. However, these policies can have side effects. For example, FDR's blockade of raw materials against the Japanese in Manchuria in the 1930s arguably led to the bombing of Pearl Harbor, which resulted in U.S. involvement in World War II. The decades-long embargo against Cuba not only did not lead to the topple of the communist regime there, but may have strengthened Castro's hold on the island and has created animosity toward the United States in Latin America and much suffering by the people of Cuba. Various studies have concluded that embargoes and other economic sanctions generally have not been effective from a utilitarian or policy perspective, yet these policies continue. Evaluation of the effectiveness of Trade Embargoes Strengths Trade embargoes and other sanctions can give the sender government the appearance of taking strong measures in response to a given situation without resorting to violence. Sanctions can be imposed in conjunction with other measures to achieve conflict prevention and mitigation goals. Sanctions may be ineffective: goals may be too elusive, the means too gentle, or cooperation from other countries insufficient. It is usually difficult to determine whether embargoes were an effective deterrent against future misdeeds: embargoes may contribute to a successful outcome, but can rarely achieve ambitious objectives alone. Some regimes are highly resistant to external pressures to reform. At the same time, trade sanctions may narrow the...