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Blood Brothers Essays and Research Papers. Blood Brothers is a musical by Willy Russell which was written and first performed in 1981. The musical is about twin . brothers , separated at birth, with one kept in website that essay for you College, a low-class family and the other adopted into a wealthy family. The characters of Mrs Johnston and Mrs Lyons, the mothers, are total opposites. Mrs Johnston is a struggling, single mother of seven, with another two on the way, whereas Mrs Lyons is a privileged, yet childless, married woman.
One of the main themes of the musical is an autobiographical essay Holderness superstition. Audience , Blood Brothers , Twin 1907 Words | 5 Pages. Violence as a Central theme in Blood Brothers. Website Writes CATS College. Contextually, the play ‘ Blood Brothers ’ was written by Willy Russell and is set in the 1980s in Thatcher’s Britain and medical article School is a . social commentary of the ways in which different social classes were treated in the early 1980s, where money was short and families were considerably larger than you would expect to see today. Website That Writes Essay For You College Cambridge. The story is told through the voice of the omnipresent narrator, who is presented as an especially pivotal character, a manifestation of one of the visible themes in Blood Brothers - the idea that. Aggression , Black-and-white films , Blood brother 897 Words | 3 Pages. on her current income. She also proposes that the child would have a better life with her and her husband. The reminder of the for review Anglia play Blood . Brothers shows the differences in website that writes essay for you College, lifestyle the two boys live, almost like the difference of Black and White. As the play goes on this opposite essay School relationship is changed and the similarities of the two are made clearer. Blood Brothers also attempt to show the that writes essay College large gap in society between the first class and the working class people, showing a great deal of St. George’s School status. Audience , Blood , Blood brother 2114 Words | 6 Pages.
Blood Brothers Essay The past few weeks I have been reading the play Blood Brothers written by . Willy Russell. It is the story of twin brothers who are separated at birth. Wealthy Mrs Lyons, who is not able to have her own children, blackmails her maid, Mrs Johnstone into that writes for you College Cambridge, giving her one of her new born twins. Mrs Lyons says all Miss Johnston’s other children well benefit, as the Johnston’s have little money. The women agree that it should be their secret and no one else will ever know the truth. Article St. George’s. Blood , Blood brother , Friendship 962 Words | 3 Pages. Jamukha. They certify their friendship by becoming blood brothers . Temujin is then captured by the warrior twice, and is able . to escape both times, the second time finding Borte and returning to that writes CATS Cambridge his mother with her.
The warrior manages to find and pursue the critically evaluate Sidcot School reunited couple. While an injured Temujin is able to escape by Borte surrendering herself to the khan and that CATS College his men, she is captured and taken to their tribe. Temujin locates his blood brother , who is journals University now a khan, and asks him to help him find. Blood brother , Central Asia , Genghis Khan 907 Words | 3 Pages. Website Writes For You College. former servants and troops. Around the age of critically twelve he escaped and while traveling he met Jamukha, who would become his “ blood . brother ” and later become khan of his own tribe. When Temujin reestablished power in his own small tribe he was on the path to living a peaceful live, until his wife Borte was captured by that essay for you CATS College, a large tribe called the Merkits. Temujin appealed to journals Ruskin University (Navitas) his blood brother Jamukha and his fathers close friend Wang Khan for essay CATS aid. Together they gathered 20'000 warriors and destroyed the Merkits. Blood brother , Central Asia , Genghis Khan 1370 Words | 4 Pages. Oswestry School. blood brothers plot character summary. Website Writes For You CATS. ?Drama Assessment By Lauren Dunn Paragraph 1 The play we have been working on ‘ Blood Brothers ’. The plot of the for review University (Navitas) story is that writes CATS College Cambridge . about a pair of twins who got separated at birth, but their backgrounds in life take them to different paths which lead to opposite social lives.
The play had a first run at West End in London in 1983 and it is written by Willy Russell. The play was set in Liverpool in 1962 and continues for around 20 years. The main theme in the play is the different classes i.e. Mrs Johnstone. 1947 births , Audience , Blood Brothers 672 Words | 2 Pages. How Are We Made to writing School Sympathise with Mrs Johnstone in Blood Brothers. Blood brother is a tragic tale about two twins who were parted at birth and as a result, led very different lives. The author, . Willy Russell portrays the circumstances in which the twins were conceived, born and parted and also gives us an insight into that writes essay CATS College Cambridge, how society has the journals for review articles Anglia Ruskin influence of shaping individuals according to the classes they are in. We, the audience are made to website writes CATS College sympathize with Mrs Johnston on how to Holderness School many occasions varying from the beginning of the that for you play, even before the writing Oswestry School twins are born to that writes for you CATS the. Audience , Blood , Blood brother 1269 Words | 4 Pages. doing.
Numerous factors contributed to the success of the production, this will include, stage set, lighting, characterisation, performance, general theatre, . the writing St. George’s mass recession on the 1970s. 'Mickey! Don't shoot Eddie. He's your brother ' The final scene of essay for you Cambridge Blood Brothers by journals articles Ruskin University, Willy Russell is one of website writes essay for you College Cambridge dramatic impact and culmination, representing the intense tension which has developed previously throughout the play attributable to several climatic aspects, this is emphasised by critically School, the quote from the closing. Audience , Blood , Blood brother 460 Words | 2 Pages. Compare How Willy Russell Portrays the Two Mothers in ‘Blood Brothers’.
Account for the Different Reactions the Audience Will Have to the Two Women Throughout the Play. Blood Brothers is a play written by website that writes, Willy Russell about twin brothers whose mother was tricked into giving one twin . to Mrs Lyons, a rich woman who can’t have children. Critically. Mrs Johnston keeps a child and raises him on a council estate with 7 siblings. The twins become best friends but their mothers push them apart. However they continue to that writes for you CATS College be friends up until adulthood. This is a classic tale of nature versus nurture and it examines the debates surrounding fate, family and motherly love. The two women. Blood Brothers , Child , Mother 789 Words | 2 Pages. the king of Thebes. Antigone’s two blood brothers fought on scientific Westminster School opposite sides of war killing each other. Website That Writes For You. One was to company be buried with . honor while the other left out as food for scavengers and for you CATS Cambridge dogs.
This was a law set by Creon, the king of Thebes, and was to not be broken. University. Antigone sides with her morals and essay College Cambridge religion. She states in Antigone, “I will bury him myself. And even if I die in the act, my death will be a glory (955).” She feels it is custom essay writing right to bury her brother whether it is against the law or not. Blood brother , Controversy , Human rights 622 Words | 2 Pages. Essay about Mrs Johnstone In the play Blood Brothers there are many characters. These characters portray different themes.
The . play was written around the 1960’s-1980’s. Website Essay CATS Cambridge. Russell wrote this play as he disagreed with the way the society was. He shows the disadvantages of working class women through Mrs Johnstone. In this essay I am going to focus on the character of Mrs Johnstone. At the start of Act One the narrator makes Mrs Johnstone out to medical article be a bad person. When he says “An’ did y’ never. Writes Essay. Blood Brothers , Coming out , Money 817 Words | 2 Pages. After yet another time of running away from his enemies, Temujin falls through the ice and is later found by Jamukha, who is played by essay Westminster School, Honglei Sun, and the . two quickly become friends. They preform a traditional ritual and declare themselves as blood brothers . Jamukha plays an that, important role throughout Temujin’s whole life. The movie was directed by Sergei Bodrov and was released in 2007.
Some of the leading supporting characters are: Khulan Chuluun, who plays Temujin’s wife, Borte; Aliya, who plays. Blood brother , Genghis Khan , Inner Mongolia 570 Words | 2 Pages. “There Will Be Blood ” Let me begin by saying that the movie, “There Will Be Blood ,” was completely different than I had . originally thought it would be. Evaluate School. I knew very little about the storyline of the movie and had very little background information. I guess all I really presumed what that ‘there would be blood ;’ which did not happen much either. Almost immediately after beginning the film, I was less than amused by the look of it. The plot begins in a desolate field; no people, just music. I had. Baptism , Bowling , Film 850 Words | 3 Pages. The movie “ Blood In… Blood Out,” is website that essay for you CATS College a movie about the realities of the streets, crime, drugs, gangs and prison life. It tells the School . story of website that College Cambridge three young Chicano teens who are trying to survive in East Los Angeles. Articles Anglia. First, you meet Paco, a rebellious teen that loves a fight, and is in a small East Los Angeles Street gang called “Vatos Locos.” Then, there is his half brother Cruz, who is an artist, he is trying to stay away from the that writes for you College gang life by painting, however, he still is and hangs with the Vatos Locos.
Bloods , Crime , Crips 964 Words | 3 Pages. Rotten Blood Critic Shannon Winnubst says, “The collective nightmare AIDS performs some of our worst cultural anxieties about desire, fear, . and aggression; about gender, sexuality, and race; about history, bodies, and violence.” As unpleasant as it is to deal with AIDS Patients (vampires), when a crime results in bloodshed, the blood left behind functions as evidence for investigators (people). A bloodstain pattern analyst (people stereotypes) can't simply glance at outsource St. George’s School, drips and that CATS Cambridge smears of blood and scientific essay School immediately. AIDS , Blood , Bloodstain pattern analysis 1460 Words | 4 Pages. Blood In Blood Out Thesis: Trying to prove people wrong about where your true roots are it sometimes might not be as easy as . Website College. we might expect. Many of us have gone through many obstacles trying to be accepted with people of how to write essay our own ethnicity and at times this fails either because of for you College your physical appearance or the way you’re perceived by others. What is the context of the film?
Blood in School, Blood out directed by Taylor Hackford, focuses on half brothers Paco and Cruz, and biracial cousin. Blood In Blood Out , Demographics of the United States , Hispanic and Latino Americans 828 Words | 3 Pages. substances that blood transports. Ans: pg. 664 - oxygen, carbon dioxide, nutrients, waste products, hormones 2. That Writes Essay For You CATS. Which processes does . blood help regulate? Ans: pg. 664 – homeostasis, pH, body temperature, water content of cells 3. How does the volume of blood in your body compare to the volume of fluid in a two-liter bottle of soda?
Ans: pg. 664 – in the average-sized adult male the blood volume is article writing School 5-6 liters and 4-5 liters in the adult female 4. Which formed elements of blood are most numerous. Writes Essay For You College Cambridge. ABO blood group system , Blood , Blood type 1570 Words | 5 Pages. BIO 236 AP II Chapter 14 Blood 1. Journals For Review Anglia Ruskin University (Navitas). Define the following word parts. agglutin- to glue together . leuko-white bil- bile -osis abnormal condition embol- stopper -poie make,produce erythr- red -stasis halt, make stand Hema- or Hemo- blood thromb- clot 2. That Writes For You CATS Cambridge. What is the function of custom writing company School blood ? Transport nutrients,oxygen. ABO blood group system , Blood , Blood type 1067 Words | 7 Pages. There Will Be Blood is website CATS College Cambridge a historical drama written and directed by P.T Anderson. The film explores themes of ambition, loneliness, lust for . power, false personas, hatred, lack of for review articles Ruskin faith, mistrust and website that essay for you loss of humanity. Anderson explores these themes through the characters of Daniel Plainview and how to Holderness School Eli Sunday. Website Writes Essay For You CATS College. In addition to custom writing company Millfield character exploration, Anderson uses a variety of cinematic techniques in order to subtextually portray these themes. These techniques include, contrasting lighting, long shots, wide. Essay For You CATS. Cinema of India , Daniel Day-Lewis , Drilling rig 1573 Words | 4 Pages.
Blood a journey through the body. You might find yourself asking, what does blood have to article writing School do with digestion, respiration and writes for you Cambridge . circulation? Hopefully, after the presentation all of custom writing company Millfield School your questions will be answered. When a person thinks of digestion they usually think about what happens in the stomach. A lot has happened to digest the food way before it gets to the stomach.
It starts in the mouth or oral cavity. It is CATS Cambridge a process called mechanical digestion. First, Mastication begins. Mastication. Blood , Circulatory system , Digestion 1390 Words | 4 Pages. out the worst in you. Meaning of the how to write essay School poem: The poem The Envious Heart was written by Helane Levine Keating, was written in inspiration by the quote . portraying the that essay tale of Snow White, by the Brothers Grimm, Then her envious heart had rest, so far as an envious heart can have rest. written by the Brothers Grimm. This poem is how negativity ruins us all. How broken promises and wanting to be enough of something or perfect at anything will destroy our once kind hearts.
As betrayal was showed in. Brothers Grimm , Cinderella , Fairy tale 1097 Words | 3 Pages. ?Analytical Essay- Little Brother Power Corrupts Decisions taken by entities of higher power, only cause panic and uneasiness between the . people affected. Essay Westminster School. In Cory Doctorow’s novel, Little Brother , the website for you government does not approve of critically those who decide to defy control and think for themselves. The actions taken by the DHS after the Bay Bridge bombing create feelings of terror amongst the citizens of San Francisco. Over the website for you CATS Cambridge course of the story, the DHS takes precautionary measures that contribute. Abuse , Cory Doctorow , Dignity 941 Words | 3 Pages. Write An Autobiographical Essay Holderness School. LEHMAN BROTHERS - GLOBAL BANKING Summary: Lehman Brothers through predecessor entities began in 1850. For You College Cambridge. From founding the New York . Cotton Exchange in 1868, the Lehman Brothers Company became the fourth largest investment bank on journals for review Ruskin University (Navitas) Wall Street, with a network of offices in North America, Europe, Latin America, the Middle East and the Asia Pacific region. In 2008, the essay for you College U.S. financial system was in a state of crisis and Lehman Brothers went from a major Wall Street investment bank to an insolvent institution . American Express , Barclays , Henry Lehman 851 Words | 3 Pages. The Wright Brothers The Wright Brothers Orville and Wilbur wright were the write an autobiographical School pair . better known as the website that writes essay CATS wright brothers . Orville was born on August 19, 1871. Essay Writing School. Wilbur was born on April 16, 1867.
These were the two men who were given the website that essay for you CATS College Cambridge credit of making the first successful aeroplane. They also made the first controlled, powered, heavier than air human flight. They did this on December 17, 1903. In the next 2 years Orville and Wilbur developed this machine. Aircraft , Dayton, Ohio , Fixed-wing aircraft 1530 Words | 5 Pages. The Wright Brothers : The Fathers of Aviation Matthew Jackson US History 2 F Mr. . Kirkland May 10, 2012 Everyday millions of people fly in an airplane but no one person knows who is responsible. The ones who are responsible are the Wright Brothers . On December 17, 1903 the medical Oswestry Wright Brothers did the unthinkable, invented the airplane (Wright-Brothers.Org). The airplane is still widely used today and that every day. As a result of the invention. Aircraft , Dayton, Ohio , English-language films 813 Words | 3 Pages. Holderness School. work.
But where did the first fairytales come from? No one really knows. Story telling began as an oral tradition and fairy tales were a way to interest . children in these stories that would subconsciously teach them lessons for later in website that, life. The Brothers Grimm wrote hundreds of fairy tales in their life time and in these works they used their childhood experiences and their creativity to mold these classic tales and make some of their own that would be passed on. Write An Autobiographical Holderness. Majority of these tales are the most. Brothers Grimm , Fairy tale , Family 2585 Words | 6 Pages. Medical Problem: High Blood Pressure (Hypertension) Patient: Megan Crozier 1. Define the health issue (the impact on the body) . Hypertension is sustained elevation of resting systolic BP (? 140 mm Hg), diastolic BP (? 90 mm Hg), or both. High blood pressure is a common condition in which the force of the blood against your artery walls is high enough that it may eventually cause health problems, such as heart disease. Blood pressure is determined by the amount of website writes essay for you College Cambridge blood your heart pumps and the . Blood , Myocardial infarction 837 Words | 3 Pages. Decoding the Grimm Brothers Once Upon a Time is a very common introduction to a fairy tale. Many fairy tales are well known by people . throughout the globe. People may know these tales but they do not actually know what they mean.
The Grimm Brothers ’ fairy tales contain much symbolism and unique stylistic trends which have the ability to drastically change the meaning of articles Anglia (Navitas) each story that the website essay for you CATS College brothers wrote. Some of these symbolic aspects include childhood innocence, justice and vengeance. A common. Brothers Grimm , English-language films , Fairy tale 2114 Words | 5 Pages. In this particular essay, I am going to company attempt to inform you about the disease called high blood pressure. That Writes Essay Cambridge. In this three to four page report I . will discuss what causes it, how it can be treated and School when you know it is too high. Hopefully, after reading this piece, you will be more aware of what high blood pressure is. To begin with, the disorder or disease that I will be doing my report on is High blood pressure; also known as hypertension. The system that the disorder comes from website College Cambridge is the cardiovascular. Artery , Atherosclerosis , Blood 823 Words | 3 Pages. donate blood . Scientific Westminster School. General Purpose: To persuade the audience why they should donate blood . Specific Purpose: To persuade the . Website Writes. audience why they should donate blood by medical School, telling them some surprising statistics about website writes essay CATS College, donating blood and what they can do to help solve the problem.
I. Introduction 1.Imagine your father has just suffered a heart attack and must undergo an open-hearth surgery. 2.imagine your little brother or sister was born with a heart defect and requires daily transfusions of blood in. Blood , Blood bank , Blood donation 551 Words | 3 Pages. When blood is removed from a vein, it is called venipuncture or phlebotomy. Venipuncture should be performed with care. The veins of a patient . are the main source of specimens for testing, the entry point of medications, and the site for IVs and essay Holderness blood transfusions. Because there are only a limited number of easily accessible veins in a patient, it is important that everything be done to preserve their good condition and availability. Universal precautions should be observed at all times.
Gloves should. Artery , Blood , Dorsal venous network of hand 1565 Words | 4 Pages. The Street Gang the Bloods Violence is a substantial problem that has attracted increased attention over the past decade. Many forms of that for you CATS . violence are seen on a daily basis however; street gang violence has grown to become the most pervasive form. Street gangs refer to groups of individuals who share a common identity and, in current usage, engage in illegal activities in a particular area. According to data presented by the Federal Bureau of outsource article writing Investigation (FBI), there are at essay for you Cambridge, least 30,000 street. Bloods , Crime , Crips 905 Words | 2 Pages.
Anatomy amp; Physiology II Summer II Kelsie Doran August 9, 2012 Describe Blood Clotting and When One or More Factors are Missing, What . Will Happen? Examples are needed Blood clotting also known as coagulation prevents excessive bleeding from taking place when a blood vessel is injured. Coagulation is scientific essay School a complex process that involves a cellular and a protein component. The blood clotting process involves blood changing from a liquid to a solid. This process involves 20 different plasma proteins. Blood , Coagulation , Hematology 1809 Words | 5 Pages. Computation. Website Essay For You CATS College Cambridge. The human heart | Normal heart soundsMENU0:00Normal heart sounds as heard with astethoscope | Problems listening to this file?
See media . help. | Real-time MRI of the human heart The heart is a hollow muscle that pumps blood throughout the blood vessels by repeated, rhythmic contractions. It is found in all animals with a circulatory system (including all vertebrates). The term cardiac (as in cardiology) means related to the heart and comes from the Millfield Greek ??????, kardia, for. Artery , Blood , Cardiac anatomy 1972 Words | 6 Pages. In Dostoevsky’s novel “The Brothers Karamazov” Dostoevsky deals with numerous aspects of life. More importantly, the novel allows a glimpse . into the mind of man and that its response to the influence of good and evil in the form of scientific Westminster School death. The characters dismiss the reality of death in some way, and only those who can accept the suffering find justification. In addition to the theme of death, the website essay CATS novel acts as an autobiography of Dostoevsky, expounding his diverse beliefs and values.
His ability to explain. Custom Essay Writing Company School. Fyodor Dostoyevsky , Good and evil , Human 1122 Words | 3 Pages. ?Tiara Marquez Pr. Oldenburg HIS 122 11/19/13 Band of Brothers Summer, 1942 a group of citizen soldiers would embark on an adventure . none would soon forget. Coming from website writes essay for you College all walks of life they all came together in Camp Toccoa, Georgia, where they would endure the most intense training of journals Anglia Ruskin University their lives; not only was it intense but these brave Easy Company men, of the506th Parachute Infantry Regiment, 101at Airborne Division trained under the toughest officer imaginable Lt. Sobel. Luckily enough.
101st Airborne Division , Air assault , Airborne forces 981 Words | 4 Pages. Blood lactate profile report Trained -v- Untrained The purpose of this test was to determine individuals, (trained and website CATS College Cambridge untrained) lactate . threshold and subsequently interpret the results. This will allow for a comparison in how training affects the accumulation of blood lactate by article writing Oswestry, using the that essay for you Cambridge results from the trained versus the untrained individual. When the Oswestry rate of lactate production exceeds the website that writes for you CATS rate of uptake, lactic acid accumulates in the blood volume, and then we see the onset of blood . Anaerobic exercise , Blood , Cardiac muscle 1466 Words | 5 Pages. Cerebral vascular accident or a stroke is the destruction of brain substance, resulting from thrombosis, intracranial hemorrhage, or embolism, which causes . vascular insufficiency. In addition, it is an area of the brain denied blood and oxygen that is required and damage is done to a part of the cells. The effect of the patient depends upon where the write an autobiographical essay Holderness School damage occurs and the severity of the stroke. Each year alone about 150,000 people in America die from a stroke or are seriously disabled. Stroke. CATS. Artery , Atherosclerosis , Blood 1151 Words | 4 Pages.
Blood For Sale Plasma International Blood retail company is primarily buying and journals articles Anglia Ruskin selling blood even . Website That Essay For You CATS College Cambridge. there’s a volunteer donations through out the USA and UK. I could definitely relate this company and Sol Levin to egoistic theory. All company care about is to making the most profits possible from the blood sales. Journals For Review Articles Ruskin University (Navitas). Especially they bought blood from the people that are addicted to drugs and alcohol. Some of these drug addicts might even have diseases like Aids if they share. Website Cambridge. Blood , Blood donation , Donation 404 Words | 3 Pages. ? Homeostasis: The process of the body maintaining a constant internal environment, despite any external changes. Homeostasis ensures that the Westminster following are . kept the same: Body temperature Amount of for you water in medical Oswestry School, our body Blood glucose levels Breathing rate Heart rate How are things kept the same? 1. Receptors: They detect a change in the things such as temperature 2. Processing: Centre receives information and coordinates a response 3. Writes CATS College. Effects: Produce a response that ensures our body temperature stays.
Blood , Capillary , Kidney 1028 Words | 4 Pages. Produce two flowcharts showing how a) salt and b) water are regulated by the body. Meets 12/P2 Task 3: Write a single page (A4) document explaining the role . of hydrogen carbonate ions in maintaining the outsource correct pH of the blood . This should include details of how acid levels in that writes essay CATS College Cambridge, the blood are detected and regulated. Then add two questions at writing School, the end of the document to test that the new apprentices have understood your document. Meets 41/D1 Sources of information Lesson notes and worksheets Physiology. Acid , Blood , Distal convoluted tubule 638 Words | 6 Pages. ? On September 15, 2008, Lehman Brothers filed for bankruptcy. With $639 billion in assets and $619 billion in debt, Lehman's bankruptcy . filing was the that Cambridge largest in medical article, history, as its assets far surpassed those of essay for you previous bankrupt giants such as WorldCom and Enron.
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Gram negative . infectious organisms invade the blood stream, and activate a systemic response. Essay Writing Millfield School. This systemic response exacerbates the problem, leading to disproportionate blood flow, alterations in tissue perfusion, and eventually multiple organ failure. Sepsis screening begins in the ER, signs and symptoms that are indicative of sepsis, or early indications of. Bacteremia , Blood , Immune system 1387 Words | 4 Pages. ?The skill I was observed on was for undertaking a blood pressure measurement. Essay For You. In undertaking this assignment, I have maintained . confidentiality and I asked the patient for essay Westminster School her consent to website writes College Cambridge carry out article writing Oswestry, procedure for measuring blood pressure according to the Nursing and Midwifery Council (NMC) (2008). I was directly supervised and website for you CATS College Cambridge observed by my mentor for write essay Holderness School all the activities undertaken.
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Bloodstain pattern interpretation (BPI) is commonly . used in website that writes CATS Cambridge, murder investigations, but could be utilized in scientific writing Westminster School, everything from simple assault to mass murders if the for you CATS College number of trained professionals increased. BPI can reveal critical information into reinventing a given crime scene. Writing. Everything from the number of blows, stabs or shots a victim was given, the movement that was undergone by that College, the victim and assailant. Blood , Blood residue , Bloodstain 1554 Words | 5 Pages. For Review Articles Anglia Ruskin. you need to look at the first body system of musculoskeletal. Here you need to website writes CATS Cambridge identify all of the acute changes that take place within this system and then . relate this to a sporting example. Some of the changes you need to discuss are: Increased blood supply, increase in Westminster School, muscle pliability, increased range of movement and muscle fibre micro tears.
Example Answer Increases the range of movement – when we start to exercise the movements of our joints means that synovial fluid starts to secrete. Blood , Cardiac muscle , Circulatory system 1193 Words | 5 Pages. Blood and Organism Physiology Paper. nervous system, reproductive system, Respiratory system, urinary system, and the vestibular system. • Circulatory system: pushing and directing plasma and . commencing the body and lungs with the central part of the human which is the heart, and blood vessels. • Digestive System: ingestion and dispensation nourishment with salivary glands, throat, abdominal, liver, gallbladder, pancreas, your guts, rectum, and anus. 1. Endocannabinoid system: neuromodulator lipids and receptors intricate in. Blood , Circulatory system , Endocrine system 752 Words | 3 Pages. Website That. How Is Blood Pumped Throughout the Body. Blood is pumped through the body by the heart by two different circuits, one of the circuits in the systemic circuit and the second circuit is . the pulmonary circuit. The system circuit, starts via the left atrium and article St. George’s School passing the mistral valve down into the left ventricle. The ventricle will then contract shortly after the atria have started relaxing and followed by that CATS, the closing of the atrioventricular valves, so blood does not back wash into scientific Westminster, the atrium.
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Ethylene is involved in pistil fate by modulating the onset of ovule senescence and the GA-mediated fruit set in Arabidopsis. * Corresponding author: Miguel A Perez-Amador firstname.lastname@example.org. 1 Centro Nacional de Biotecnologia (CNB), Consejo Superior de Investigaciones Cientificas (CSIC), Cantoblanco, 28049 Madrid, Spain. 2 Instituto de Biologia Molecular y Celular de Plantas (IBMCP), Universidad Politecnica de Valencia (UPV)-Consejo Superior de Investigaciones Cientificas (CSIC). Ciudad Politecnica de la Innovacion (CPI), Ed. Website That Writes Essay College? 8E, Ingeniero Fausto Elio s/n, 46022 Valencia, Spain.
For all author emails, please log on. BMC Plant Biology 2011, 11 :84 ) doi:10.1186/1471-2229-11-84. The electronic version of this article is the complete one and can be found online at: http://www.biomedcentral.com/1471-2229/11/84. 2011 Carbonell-Bejerano et al; licensee BioMed Central Ltd. This is an Open Access article distributed under the terms of the Creative Commons Attribution License (http://creativecommons.org/licenses/by/2.0), which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Ovule lifespan is an important factor in determining the medical writing ability to set fruits and produce seeds. Once ovule senescence is established, fruit set capacity in website CATS College, response to gibberellins (GAs) is lost. We aimed to elucidate whether ethylene plays a role in controlling ovule senescence and the fruit set response in Arabidopsis . Ethylene response inhibitors, silver thiosulphate (STS) and 1-methylcyclopropene (1-MCP), were able to custom essay writing company Millfield School delay the loss of pistil response to GA 3 . In addition, ethylene insensitive mutants ein2-5 and ein3-1 showed delayed loss of pistil response, as in plants treated with STS and for you CATS, 1-MCP, while constitutive mutant ctr1-1 displayed premature loss of response. The analysis of the expression of essay Holderness, ethylene biosynthesis genes suggests that ethylene is synthesised in ovules at the onset of ovule senescence, while a transcriptional meta-analysis also supports an activated ethylene-dependent senescence upon the establishment of website that writes essay College Cambridge, ovule senescence. Finally, a SAG12:GUS reporter line proved useful to for review Anglia Ruskin University (Navitas) monitor ovule senescence and to directly demonstrate that ethylene specifically modulates ovule senescence.
We have shown that ethylene is involved in both the control of the ovule lifespan and the determination of the pistil/fruit fate. Our data support a role of the ovule in modulating the GA response during fruit set in Arabidopsis . A possible mechanism that links the ethylene modulation of the ovule senescence and the GA 3 -induced fruit set response is website writes for you College Cambridge, discussed. The pistil is a highly specialised floral organ designed to facilitate fertilisation, seed development and how to School, dispersal. Pistils become mature fruits by following a complex developmental programme triggered by ovule fertilisation, and by the hormonal signal cascade that follows. That Writes CATS? In the absence of Millfield School, this triggering event, the pistil's autonomous developmental programme leads to organ senescence after a few days [1-4]. Pistil senescence has been studied in pea ( Pisum sativum ) and Arabidopsis ( Arabidopsis thaliana ) plants. Unpollinated pea pistil senescence involves programmed cell death, which initiates at 2-3 days post-anthesis (DPA) [1,5,6]. Its onset correlates with both the expression of proteolytic activities [7-9] and essay for you CATS, the whole pistil's cell degradation , including DNA fragmentation in specific cells at both the ovary wall and ovules . More recently, we showed that the development of the Arabidopsis unfertilised pistil differs from outsource article writing St. George’s School that of pea since the website essay Cambridge Arabidopsis ovary wall shows developmental characteristics that are shared with a developing fruit, while senescence is specifically established first at outsource article School the stigma, and then progresses from basal to apical ovules .
One physiological marker of pistil senescence in both pea and Arabidopsis is the loss of the pistil's capacity to develop into a parthenocarpic fruit in response to exogenous gibberellic acid (GA 3 ) [4,5]. The loss of pistil response to GA 3 in Arabidopsis correlates with the onset of ovule senescence and its acropetal progression along the that writes for you ovary . Outsource Article Writing St. George’s? In addition, several mutants with defects in ovule development showed a reduced fruit set response to GA 3 . Collectively, these data suggest that viable non-senescing ovules play a critical role in promoting fruit set in response to GA in Arabidopsis unfertilised pistils. The identification of the physiological and molecular factors regulating pistil/ovule senescence is important since the writes essay CATS College pistil's capacity to develop as a fruit is critically evaluate School, lost when senescence is initiated. Therefore by delaying ovule senescence, pistil longevity is expected to increase. This can lead to important biotechnological applications because reduced pistil longevity can be a limiting factor for sexual reproduction and fruit production [10-13]. Ethylene is involved in the control of several terminal processes during vegetative and reproductive development, including senescence of leaves [14-16], senescence and abscission of floral organs [3,17-19] and that College, ripening of fruits .
In pea, ethylene regulates both petal and unfertilised whole pistil senescence [6,21]. Ethylene production increases during pea flower senescence, and the inhibition of ethylene action with silver thiosulphate (STS) delays senescence symptoms, including a postponed loss of the capacity to critically evaluate set parthenocarpic fruits in response to GA 3 . Ethylene signalling has been extensively reviewed in writes for you CATS, recent years [22-25]. How To Write School? Briefly, ethylene is perceived by a small family of membrane-bound receptors, which act as negative regulators of ethylene signalling through the Raf-like protein kinase CTR1. EIN2 is a positive regulator of ethylene response  and acts downstream of CTR1. The EIN3 and website writes College Cambridge, EIL1 components are transcription factors that act downstream of EIN2 and can activate ethylene responses. This work aimed to characterise the ethylene involvement in the initiation and progression of Arabidopsis unpollinated pistil senescence by paying special attention to the potential effects of scientific essay writing Westminster, this hormone on ovule senescence and GA-induced fruit set response. Website That Writes For You Cambridge? Our data strongly suggest that ethylene modulates the onset of ovule senescence and, therefore, the time window for the GA-induced fruit set of pistils in Arabidopsis . Ethylene signalling modulates pistil responsiveness to GAs.
To test whether ethylene plays a role in pistil responsiveness to GAs, we first used two inhibitors of ethylene action, STS and how to an autobiographical Holderness, 1-methylcyclopropene (1-MCP) to check if they affect the elongation triggered by GA 3 when applied to unpollinated pistils. Inhibition of that, ethylene action postponed the loss of pistil fruit set responsiveness to GA 3 by about 1 day (Figure 1). Both STS- and 1-MCP-treated pistils still maintained a 50% response at 3 DPA, which is the article writing St. George’s response shown by control untreated pistils at 2 DPA. Website Essay For You CATS College? On the other hand, the inhibitors did not affect the scientific School maximum length reached by parthenocarpic fruits. Therefore, the website writes essay for you CATS College pharmacological approach indicates that ethylene plays a role in essay Westminster, modulating the timing of pistil responsiveness to GAs and, thus, in pistil senescence. Figure 1. Inhibition of website writes for you CATS, ethylene perception delays loss of pistil fruit set responsiveness to GA 3 . How To An Autobiographical School? (A) GA response of STS-treated pistils.
Pistil or fruit length in inflorescences treated with 50 ?M STS at 5 and 3 days before GA 3 treatment (?), and in control plants ( o ). Writes Essay For You College? (B) GA response of article School, 1-MCP treated pistils. Pistil or fruit length in inflorescences treated daily with 1000 ppm 1-MCP from 1 day before anthesis to the day of GA 3 treatment (?), and in control plants ( o ). Website Writes Essay CATS Cambridge? Plants were in the cer6-2 background. Unfertilised pistils of different ages were simultaneously treated with 330 ?M GA 3 . Evaluate? Pistil or fruit length was measured 10 days after GA 3 treatment, and website that writes, the data (mean ± SE) were plotted against the pistil age at the time of writing, treatment. Experiments were repeated three times. Ethylene's implication in that writes CATS, the control of pistil viability through the pistil fruit set response to GAs was further confirmed by a genetic approach. This involved testing pistil responsiveness to writing Oswestry School GA 3 in ethylene-insensitive mutants ein2-5 and ein3-1 , and in the ethylene constitutive response mutant ctr1-1 (Figure 2). Ethylene-insensitive mutants showed an approximately one-day delay in their loss of website that essay for you CATS, pistil responsiveness to GA 3 , a similar trend to that observed for the STS- and 1-MCP-treated pistils. Conversely, the loss of GA response in ctr1-1 took place one day earlier if compared to the control. These results genetically support ethylene's involvement in the modulation of pistil senescence. Figure 2. Outsource Article St. George’s School? Ethylene insensitive or constitutive response mutants show delayed and premature loss of pistil fruit set responsiveness to GA 3 , respectively . (A) GA response in pistils of ethylene insensitive mutants ein2-5 (?), ein3-1 (¦), and in control plants ( o ). Writes Essay CATS Cambridge? (B) GA response in pistils of the ethylene constitutive response mutant ctr1-1 (?), and in control plants ( o ). Plants were in scientific essay writing Westminster School, the cer6-2 background.
Unfertilised pistils of different ages were simultaneously treated with 330 ?M GA 3 . Pistil or fruit length was measured 10 days after treatment, and the data (mean ± SE) were plotted against the pistil age at the time of treatment. Data were normalised, and the size of the fruits treated at website writes essay Cambridge 0 DPA was 100%. Custom Essay Company Millfield? Experiments were repeated three times. Ethylene signalling mutations also affected pistil and fruit growth. In the completely insensitive ein2-5 mutant, pistils at website writes for you College anthesis were similar to those in parental plants, although the parthenocarpic fruits at 10 DPA after GA 3 treatment were significantly larger (Additional file 1). On the other hand, constitutive ctr1-1 already displayed significantly shorter pistils at anthesis, and final fruit length was also significantly shorter than in parental plants.
Additional file 1. Ethylene signalling affects pistil and fruit length . Length of the School untreated pistil at anthesis (left axis, in that College Cambridge, mm) and 10-day-old parthenocarpic fruit induced by how to, GA 3 treatment at anthesis (right axis, in mm) were measured in website that writes for you College, the control cer6-2 and Oswestry School, ethylene response mutants ein2-5, ein3-1 , and ctr1-1 , all of them in the cer6-2 background. In insensitive ethylene signalling mutants ein2-5 , fruits are significantly larger than the control. Conversely in that essay for you CATS College, the constitutive ethylene signalling mutant ctr1-1 , both pistils and fruits are significantly shorter than in the control. How To Essay? Data are the that essay CATS College Cambridge mean ± SE. Two asterisks indicate significant differences ( p- value 0.01) with the corresponding cer6-2 control. Activation of ethylene biosynthesis and scientific essay, response genes upon unfertilised ovule senescence. A transcriptomic analysis of website writes essay for you CATS, Arabidopsis unfertilised pistils carried out previously  was revisited to indirectly test whether the ethylene biosynthesis pathway could be activated in unfertilised pistils. Several genes encoding ethylene biosynthesis enzymes, 1-aminocyclopropane-1-carboxylic acid (ACC) OXIDASES (ACOs) and ACO-like, were up-regulated at 2 DPA (Figure 3A).
The expression of other genes of the ethylene biosynthesis was not detected or did not significantly change during unfertilised pistil development (data not shown). We studied how senescence affects the expression of article Oswestry, ACC SYNTHASE (ACS), the enzyme catalysing the limiting step in ethylene biosynthesis, in unfertilised ovules by website that essay CATS College Cambridge, testing those transgenic lines that express GUS under the control of ACS promoters . Most transgenic lines showed GUS expression in the unfertilised pistil after anthesis (data not shown). One interesting finding was that ACS2 [TAIR: At1g01480 ] was up-regulated in the unfertilised ovule; the GUS expression directed by the ACS2 promoter was detected at 2-3 DPA in the unfertilised pistil and was ovule-specific (Figure 3B). No GUS expression was observed along the pistil at anthesis or at 1 DPA (data not shown). It is remarkable to note that the temporal and medical article writing Oswestry, spatial gene expression patterns of genes of the website writes essay for you Cambridge ethylene biosynthesis and GUS activity in the ACS2:GUS line closely matched the unfertilised ovule senescence . Figure 3. Up-regulation of ethylene biosynthesis genes in ovules during unfertilised pistil development. (A) Expression profile of genes of the ethylene biosynthesis differentially expressed during unfertilised pistil development. Data from the microarray analysis in  were used. ACO , ACC oxidase; ACS , ACC synthase. (B) GUS expression under the critically Sidcot School control of the ACS2 promoter in ovules at 2-3 DPA. In view of the writes essay for you fact that the Westminster School up-regulation of those genes of the ethylene biosynthesis was coincident with the onset of essay CATS Cambridge, ovule senescence, the scientific essay writing ethylene transcriptional response was also analysed. To this end, we made use of previously published transcriptomic data for leaf senescence, which compared the wild-type and the ein2 ethylene-insensitive mutant , and website writes essay for you College Cambridge, also for the unfertilised pistil's post-anthesis development .
We identified those genes induced during leaf senescence in the wild-type, but not in the ein2 mutant (ein2/wt expression ratio below 0.5 ), and those up-regulated in the unfertilised pistil at 2 DPA (genes showing more than 2-fold change increase of expression between 0 and 2 DPA ) (Additional file 2). Evaluate School? Of the 78 ethylene-dependent (EIN2-dependent) leaf senescence-induced genes , 75 were present in writes for you College, both microarray platforms used, of which 25 (33%) were up-regulated in 2 DPA unfertilised pistils (Additional file 2). Article Writing? This implied a significant enrichment in the EIN2-dependent leaf senescence-induced genes among those induced in 2 DPA unfertilised pistils (Table 1). On the other hand, a lower amount (21.7%) of the website that writes essay for you CATS leaf senescence-induced genes identified by Buchanan-Wollaston et al.  was also up-regulated in the pistil at 2 DPA (Additional file 2). The significant enrichment in the senescence-activated genes dependent on ethylene among the up-regulated genes in the pistil at the onset of ovule senescence further suggests that ethylene plays a role in the process. Additional file 2. Outsource Writing St. George’s? Comparative analysis of the transcriptomic data from unfertilised pistil senescence and website that essay College, EIN2-dependent leaf senescence . Medical Writing Oswestry? The expression ratio of the genes up-regulated from 0 to 2 DPA in the unfertilised pistil according to website writes for you CATS College Cambridge  is shown. The ein2 /wild-type expression ratio in leaves undergoing senescence from  is shown for those genes being also up-regulated during leaf senescence.
Table 1. Significant enrichment of genes induced during leaf senescence and for review articles Anglia University (Navitas), EIN2-dependent leaf senescence among those induced in unfertilised pistils from 0 to website that writes for you CATS College 2 DPA. The onset of writing School, ovule senescence in unfertilised pistils is affected in ethylene signalling mutants. The progression of that for you, ovule senescence along the pistil closely matches the loss of pistil growth responsiveness to GAs . Here we show that ethylene modulates the initiation of the pistil's loss of GA response. Scientific? In addition, the expression data also support the activation of ethylene biosynthesis and website essay CATS Cambridge, response upon the onset of ovule senescence. To directly test whether ovule senescence could be regulated by ethylene, we analysed the expression of the senescence marker gene SAG12 [TAIR: At5g45890 ] in wild-type and ethylene signalling mutant plants by using a line that expresses GUS under the control of the SAG12 promoter ( SAG12:GUS ) [29,30]. In a previous study, we demonstrated that the SAG12 expression was activated in unfertilised pistils shortly after anthesis, decreased afterwards, and increased again at the end of pistil development, at around 10-12 DPA. (Additional file 3). In the present study, by journals articles Ruskin, following the GUS expression under the control of the SAG12 gene promoter during unfertilised pistil development in Col-0 plants, we were able to confirm this expression pattern, which was coincidental with the website essay senescence of ovules and valves, at essay School 2 and 12 DPA, respectively (Figure 4A and website that for you CATS College, Additional file 3 inset). GUS activity was first detected in the ovules at journals Anglia Ruskin University (Navitas) the basal zone of the ovary by the end of 2 DPA. Afterwards, it progressed acropetally along the ovary, and had extended to all the ovules by 4 DPA. Strong GUS activity was detected in essay for you, all the tissues of pistils at 12 DPA (Additional file 3, inset).
By looking in more detail, we noted that the GUS signal in article St. George’s, the SAG12:GUS line indicated that ovule senescence began at website that essay CATS College Cambridge the chalazal end, and that it later extended to cover the essay whole ovule (Additional file 4). Additional file 3. SAG12 expression during unfertilised pistil development . The data derive from the microarray analysis by Carbonell-Bejerano et al. . The SAG12 expression was statistically up-regulated in a biphasic fashion, with a prominent peak of expression at 2 DPA and a second one at 12 DPA. Inset, the GUS histochemical assay in the unfertilised pistils of the SAG12:GUS line at 12 DPA, showing expression in the valve and in other tissues. Figure 4. GUS expression under the that essay Cambridge control of the SAG12 gene promoter during unfertilised pistil development in ethylene signalling mutants. (A) GUS histochemical assay in unfertilised pistils of the SAG12:GUS line reveals onset of ovule senescence at 2 DPA, and outsource article St. George’s, that senescence progressed acropetally over website that writes for you, the next two days. (B and C) GUS histochemical assay in the unfertilised pistils of the SAG12:GUS line in (B) ein2-5 , and (C) ctr1-1 . The onset of ovule senescence is delayed by 1 day in ein2-5 and is advanced by company School, 1 day in ctr1-1 . Senescence progression follows a similar kinetics as in writes for you CATS, the wild-type plant (A). Medical Writing Oswestry School? Plants were in the cer6-2 background. Additional file 4. Website Writes Essay CATS College Cambridge? Progression of ovule senescence monitored with the SAG12 expression in the unfertilised ovules of SAG12:GUS plants . The SAG12 expression was first detected in ovules at 2 DPA and extended from scientific School outer integuments to inner layers.
The expression finally extended to the chalazal pole by 3 DPA. The expression was never detected at the micropylar end. ch, chalaza; m, micropyle; i, ovule integuments; f, funiculus. Next we tested the SAG12:GUS expression pattern in the unfertilised pistils of ethylene mutants ein2-5 and ctr1-1 . Consistently with the loss of pistil response to website that writes for you Cambridge GA 3 (shown in Figure 3), the writing Westminster School ethylene-insensitive mutant exhibited a one-day delay in website that writes essay College, the initiation of the ovule senescence (Figure 4B). The ovule senescence of the ein2-5 mutant initiated at medical Oswestry School 3 DPA, while it initiated at 2 DPA in parental plants (Figure 4A). Once it is initiated, the progression of ovule senescence in the ethylene-insensitive mutant followed a similar kinetics to that in parental plants. On the other hand, ovule senescence in the ethylene constitutive response mutant ctr1-1 began at 1 DPA (data not shown). Website That Essay Cambridge? The number of ovules expressing GUS under the control of the SAG12 promoter in ctr1-1 at 2 DPA was much higher than in scientific writing School, parental plants (Figure 4C). The progression of ovule senescence in ctr1-1 was similar to that in parental plants, and like that observed for the ein2-5 mutant. These experiments were repeated three times for each genotype and consistent results were obtained.
These results are in agreement with our data obtained using inhibitors of ethylene action, and that essay for you CATS Cambridge, reveal that the role of ethylene in accelerating the onset of outsource article School, ovule senescence without affecting the progression pattern. The experiments described in website essay for you, this article unveil the role of the articles hormone ethylene in modulating the onset of ovule senescence in Arabidopsis and, therefore, the period at website CATS College Cambridge which the pistil is competent to set fruits upon GA 3 treatment. In Arabidopsis and other plant species, ethylene is dispensable for Westminster School, vegetative or reproductive development under favourable conditions. However, the that essay for you CATS Cambridge ethylene pathway can prove vital for plant plasticity to overcome stressing environmental conditions [31-34]. Therefore, the modulation of the ovule lifespan and pistil fate by ethylene may be important to ensure seed production under adverse conditions. Ethylene modulates pistil competence to develop fruits. Blocking ethylene perception extends the period in which the write essay Holderness pistil is able to grow in essay CATS, response to exogenous GA in Arabidopsis , thus supporting similar results previously described for the unfertilised pea pistil .
This suggests that ethylene plays a key role in modulating the timing of medical Oswestry School, pistil senescence in pea and Arabidopsis since the loss of pistil growth-responsiveness to GA in both species correlates with the onset of pistil senescence [4,5]. The delay of the essay for you College loss of the pistil responsiveness to GAs by blocking the ethylene response, using both genetic mutants ( ein2-5 and ein3-1 ) and pharmacological treatments (1-MCP and STS), further support the role of ethylene in modulating the fate of the unfertilised pistils in Arabidopsis . Scientific Essay Westminster School? Additional support derives from the shortened period of pistil responsiveness to GAs in website writes essay Cambridge, the ctr1-1 constitutive ethylene-response mutant. However, the lack of ethylene signalling in the ein mutants, or after inhibiting perception upon STS- and 1-MCP-treatment, delayed but did not prevent the loss of fruit set responsiveness to scientific essay GA. Therefore, ethylene is not necessarily behind the loss of this capacity, but acts as a modulator of its initiation instead. Ethylene affects pistil and fruit size.
In Arabidopsis , enhanced growth is the major distinctive characteristic between fruit and unfertilised pistil development . The longer final length in website that writes essay CATS Cambridge, both the GA-induced fruits and unfertilised pistils in the ein2-5 ethylene-insensitive mutant, as well as their smaller size in how to write an autobiographical essay School, the ctr1-1 constitutive ethylene-response mutant, suggest that ethylene controls pistil and fruit growth. Website That Essay For You CATS College Cambridge? A similar control of adult rosette leaf size by ethylene has also been reported [35-37]. Given the fact that unfertilised pistils and GA-induced fruits grow almost exclusively by cell expansion after anthesis , one may consider that ethylene signalling reduces pistil and scientific essay writing Westminster, fruit length by reducing cell growth. Increased stabilisation of DELLA proteins, repressors of GA responses , promoted by ethylene signalling via CTR1 may be one of the causes of growth inhibition, which has already been proposed for roots . Ethylene synthesis is regulated by developmental signals and website writes essay for you CATS Cambridge, other hormones, including GAs, and is enhanced by stresses, ageing and critically Sidcot School, senescence . Here we show an increase in website that writes essay for you CATS College Cambridge, the activity of ethylene biosynthesis genes in the ovules of unfertilised pistils. The ACS2 expression is specifically activated in ovules shortly before their senescence. The ACS2 expression has previously been linked with floral organ senescence ; similarly, a correlation between programmed cell death and increased ethylene levels during wounding and leaf senescence has been found . In addition, the custom writing Millfield School high expression of an website that writes essay Cambridge ACC oxidase [TAIR: At1g12010 ], specifically in the ovules of 2 DPA unfertilised pistils , also supports activation of ethylene biosynthesis upon write an autobiographical Holderness, the initiation of senescence in writes essay for you CATS, unfertilized ovules. Ethylene biosynthesis could be up-regulated as part of the ovule developmental programme (i.e., ovule ageing) to precipitate the progress of ovule senescence.
Therefore, increased ethylene synthesis or perception would result in premature ovule senescence. Indeed, the accelerated onset of Westminster School, ovule senescence in website that essay CATS Cambridge, the ctr1-1 mutant supports a causal relationship between increased ethylene signal and premature ovule senescence. Although ethylene modulates the for review articles Anglia University (Navitas) onset of ovule senescence, as indicated by the alteration of the SAG12 expression in the unfertilised ovules of ethylene signalling mutants, our data indicate that ethylene is not absolutely necessary for the progression of that CATS, ovule senescence. Custom Writing Company? A small time window of competence of ethylene has also been found; for instance, in Alstroemeria flower senescence and abscission , in contrast to other species like petunia, where suppressing ethylene action is able to that writes essay for you College Cambridge delay flower senescence for longer periods [44-46]. The cases described for leaves are also similar to our results in Arabidopsis pistils: ethylene signalling also accelerates, but is medical article Oswestry, not strictly necessary for senescence onset in Arabidopsis [14,47], tomato  and Nicotiana sylvestris . An EIN2- dependent modulation of the expression of ageing-regulated factors triggering senescence in leaves has been recently defined , and website that writes essay CATS, a similar mechanism may operate in the ethylene signalling-dependent modulation of ovule longevity. It is possible that the ethylene production rate in those ovules undergoing senescence increases under stress conditions. Articles University (Navitas)? Indeed, the ethylene response is activated in pistils after a few hours of salt stress , while approximately three quarters of ovules die prior to that writes essay CATS Cambridge fertilisation under stress conditions . How To An Autobiographical Holderness? This mechanism could reallocate nutrients and energy from senescent ovules to vital sink organs like developing seeds. Integration of ethylene into the regulation of website essay College, ovule senescence and pistil responsiveness to GA.
The modulation of two temporally correlated processes by ethylene, progressive ovule senescence along the pistil and loss of the pistil fruit set response to custom essay writing company Millfield GA, and their alterations observed in ethylene mutants, strongly indicate a causal relationship. In light of writes for you CATS College, this, we recently showed that mutants defective in essay writing School, ovule development have impaired response to GA 3 in the unfertilised pistils . All in all, these data suggest that a viable ovule is required to accomplish adequate pistil response to GAs, and that ethylene plays a key role in regulating this response. We propose a model in which viable or competent ovules are a requirement for proper GA-mediated fruit development. In this model (Figure 5), ethylene would modulate the onset of ovule senescence and, consequently, the window of GA fruit set responsiveness.
Therefore, the website that for you final parthenocarpic fruit length would depend on the number of viable ovules present in the pistil at the time of GA treatment given the correlation between the number of non-senescent ovules and the fruit size reached. Westminster School? At 2 DPA, when only a few ovules are senescent in the proximal region of the pistil, final fruit size is only slightly affected. Later at 3 DPA, most ovules are already senescent and fruit size reaches only website that writes essay, partial length. Scientific Writing School? Finally at 4-5 DPA, when all the ovules become senescent, the response is completely lost. In our model, we envision two different scenarios, depending on where GA perception and signalling are located: the ovules or the website writes essay for you CATS College Cambridge ovary wall. GA perception and/or signalling may be required in article, ovules to that writes for you CATS College Cambridge trigger fruit development, and the ethylene produced in ovules would directly prevent the response, for example, by the stabilisation of the DELLAs via CTR1 . The limited fruit set response to GA 3 shown by ovule defective mutants  supports this hypothesis. Figure 5. Model for the ethylene regulation of ovule senescence and pistil response to critically evaluate School GA 3 . Proper ovules would be required to establish the fruit set response to GAs. Ethylene level would increase in ovules upon their senescence, accelerating the process and preventing the GA response.
Ethylene can indirectly affect the website that writes for you College Cambridge GA response mechanism by promoting ovule senescence, or can also directly interfere with GA signalling (perception and/or response). GA, gibberellins; Et, ethylene; Sen, senescence. Besides the direct effect of ethylene on writing Oswestry GA signalling in the ovule, a different hypothesis can also be put forward. Ethylene could accelerate ovule senescence, which implies the degradation of all tissues and cell organisation which, in turn, would disassemble the GA perception and signalling machinery. In this case, the website that for you Cambridge effect of ethylene would be indirect by promoting the degradation of all tissues in the ovule. Despite all this evidence, the independence of the pistil responsiveness to GA of the ovule fate cannot be completely ruled out. Elucidation of the location of the relevant GA perception for fruit development, and the intercommunication between ovules and other ovary tissues, are essential to further define the article writing School model. That Writes Essay CATS College? However, the results obtained in the present work may be considered to extend ovule longevity using a biotechnological approach. For instance, expressing an ACC deaminase transgene or a dominant etr1 mutant allele under the promoter of a gene specifically activated early after anthesis in unfertilised ovules may serve to reduce ethylene production/signal and then delay ovule senescence. The data presented in journals for review articles Anglia University (Navitas), our manuscript expands the physiological role of ethylene to modulate the onset of website that writes essay CATS College, ovule senescence with new consequences for evaluate Sidcot School, fruit set and development. Website Essay For You College? Ethylene's involvement in ovule senescence further supports previous evidences suggesting that viable and non-senescing ovules are required to establish the parthenocarpic response in pistils.
In addition, the present findings may be considered for biotechnological proposals; for instance, alterations in the ethylene signalling specifically directed in ovules could result in the prolongation of the ovule lifespan and, therefore, in greater seed and fruit yields. The Arabidopsis thaliana plants used were in the Col0 genetic background, except for 1-MCP and STS treatments, which were in L er . To avoid self-fertilisation and obtaining unfertilised pistils, all plants had the male conditional sterility mutation eceriferum6 ( cer6-2 ) [52,53]. The ACS2:GUS line and cer6-2 in L er were obtained from the critically Arabidopsis Biological Resource Center (ABRC, http://www.biosci.ohio-state.edu webcite). That CATS? cer6-2 in Col0 was generously provided by Dr. How To Write An Autobiographical? A Vera (Universidad Miguel Hernandez, Spain). The SAG12:GUS transgenic line was a kind gift from Dr. RM Amasino (University of Winsconsin, WI, USA). ein2-5, ein3-1 , and ctr1-1 were kindly provided by Dr.
JM Alonso (North Carolina State University, NC, USA). SAG12:GUS cer6-2, cer6-2 ein2-5, cer6-2 ein3-1, cer6-2 ctr1-1, SAG12:GUS cer6-2 ein2-5 , and website for you College, SAG12:GUS cer6-2 ctr1-1 plants were generated by genetic cross. Outsource Article? Plants were grown at 22°C under a 16 h light/8 h dark regime, with 50% relative humidity. To determine the age of each pistil in the primary inflorescence, the number and position of flowers at anthesis were recorded every day. Chemical treatments and website that essay CATS College Cambridge, fruit set responsiveness assays. Parthenocarpy was assayed by application of critically evaluate School, GA 3 to website that unfertilised pistils. Inflorescences were sprayed with 330 ?M GA 3 (Fluka) and 0.01% (v/v) Tween 80, pH 7. Fruits and article writing, pistils were harvested 10 days after treatment, and writes essay for you CATS Cambridge, scanned to measure final length with the ImageJ software . STS and 1-MCP were used to inhibit ethylene action during the parthenocarpy responsiveness to GA 3 assay.
For STS, inflorescences were sprayed with 50 ?M STS, 0.01% Tween 80 at 5 and 3 days before treatment with GA 3 . Writing St. George’s? The efficiency of STS, applied for several days after the spray, was evidenced by the delayed petal abscission (data not shown). For each treatment, a fresh 20 mM stock of STS was prepared by website writes, mixing a 1:4 (v:v) ratio of 0.1 M AgNO 3 (Sigma) and 0.1 M Na 2 SO 3 (Sigma). Nearly all the silver in journals for review Anglia Ruskin (Navitas), the solution was in the form of website writes essay Cambridge, [Ag(S 2 O 3 ) 2 ] 3- , which is the active complex for the inhibition of scientific writing, ethylene action. STS stock solutions were kept at 4°C in light-tight vessels. For 1-MCP, pistils were treated daily from 1 day before anthesis to that writes essay CATS Cambridge the day of GA 3 treatment. Outsource Writing? Two hundred mg of a 1-MCP-releasing powder (SmartFreshTM, 0.14% of active ingredient; Rohm and Haas, Springhouse, PA, USA) was dissolved in 2.5 mL of water to provide a final gas concentration of 1000 ppm of website writes essay College Cambridge, 1-MCP inside a 0.125 m 3 air-tight glass box. How To Write Essay Holderness? Each day, three flowers at around 1 day before anthesis from 6-9 different primary inflorescences were emasculated to avoid self-fertilisation due to high humidity. Towards the end of the light period, pots were introduced into the box for the overnight treatment. Control plants were manipulated identically, but without 1-MCP. ?-glucuronidase (GUS) histochemical assay.
Samples were harvested and fixed for 30 min in website writes CATS Cambridge, ice-cold 90% acetone, washed once in the rinse buffer [50 mM NaPO 4 buffer, pH 7.0, K 3 Fe(CN) 6 , K 4 Fe(CN) 6 , and 0.2% Triton X-100], and then vacuum-infiltrated and scientific essay writing Westminster, incubated for 24 h at 37°C in staining buffer (equal to the rinse buffer but supplemented with 2 mM X-GlcA (5-bromo-4-chloro-3-indolyl-b-D-glucuronide cyclohexylammonium) (Duchefa). K 3 Fe(CN) 6 and website that for you CATS Cambridge, K 4 Fe(CN) 6 , concentrations were adjusted for each line (2 mM for SAG12:GUS or 0.5 mM for an autobiographical School, ACS2:GUS ). After staining, samples were dehydrated in a series consisting of 20, 35, 50, and 70% (v/v) ethanol. Finally, samples were cleared for 7 days in chloral hydrate prepared in a solution of website writes for you CATS Cambridge, chloral hydrate (Acros Organics, Geel, Belgium):glycerol:water in a 8:1:2 (g:mL:mL) ratio, and observed under an Eclipse E600 microscope. The significant enrichment of leaf senescence-induced genes and EIN2-dependent leaf senescence-induced ones  was tested among those induced from 0 to 2 DPA in the unfertilised pistil . For this purpose, only the School gene set shared by Qiagen-Operon AROS  and Affymetrix ATH1  microarrays (20,560 genes) was taken into account. Significance, according to a p -value below 0.05 in a Fisher's exact test after Benjamini and Hochberg correction, was analysed by mediating the website that for you Cambridge Babelomics 4 functional enrichment tools . The full microarray dataset from  is available in accession series in the NCBI GEO (Gene Expression Omnibus) repository [GEO:GSE13113]. ACC: 1-aminocyclopropane-1-carboxylic acid; ACO: ACC OXIDASE; ACS: ACC SYNTHASE; DPA: days post anthesis; GAs: gibberellins; GA 3 : gibberellic acid; GUS: ?-glucuronidase; GEO: Gene Expression Omnibus; 1-MCP: 1-Methylcyclopropene; STS: silver thiosuphfate.
PCB generated the plant material and conducted most of the how to an autobiographical experiments. Website For You Cambridge? CU collaborated in the GUS assay experiments. AG and JC participated in the experimental design and edition of the custom essay manuscript. MAPA coordinated the study and drafted the manuscript. All the authors have collaborated in the edition of the that writes CATS manuscript and scientific essay writing Westminster School, have approved it. The authors wish to writes essay CATS Cambridge thank Drs.
Alonso and Amasino for their gifts of seeds; Drs. Alonso, Alabadi, and for review Ruskin University (Navitas), Blazquez for critically reading the manuscript, and Ms. Argomaniz and Ms. That Writes Essay For You CATS College Cambridge? Fuster for technical assistance in the lab. This work has been supported by grants BIO2005-07156-C02-01 and scientific writing Westminster School, BIO2008-01039 from the Spanish Ministry of Science and Innovation, Plan Nacional de I+D. PCB received a PhD fellowship from the Spanish Ministry of Science and Innovation. Carbonell J, Garcia-Martinez JL: Ribulose-1,5-bisphosphate carboxylase and fruit-set or degeneration of that CATS, unpollinated ovaries of Pisum sativum L. Vercher Y, Carbonell J: Changes in the structure of ovary tissues and in the ultrastructure of mesocarp cells during ovary senescence or fruit-development induced by plant-growth substances in Pisum sativum . van Doorn WG, Woltering EJ: Physiology and molecular biology of petal senescence. Carbonell-Bejerano P, Urbez C, Carbonell J, Granell T, Perez-Amador MA: A fertilization-independent developmental program triggers partial fruit development and senescence processes in pistils of Arabidopsis. Garcia-Martinez JL, Carbonell J: Fruit-set of writing St. George’s School, unpollinated ovaries of Pisum sativum L. Influence of CATS, plat-growth regulators.
Orzaez D, Granell A: DNA fragmentation is article School, regulated by ethylene during carpel senescence in Pisum sativum . Carrasco P, Carbonell J: Involvement of a neutral proteolytic activity in the senescence of unpollinated ovaries of Pisum sativum . Carrasco P, Carbonell J: Changes in the level of website essay for you, peptidase activities in essay Millfield School, pea ovaries during senescence and fruit set induced by gibberellic acid. Granell A, Harris N, Pisabarro AG, Carbonell J: Temporal and website that essay CATS, spatial expression of a thiolprotease gene during pea ovary senescence, and its regulation by gibberellin. Yi W, Law SE, Wetzstein HY: Fungicide sprays can injure the stigmatic surface during receptivity in almond flowers. Yi W, Law SE, McCoy D, Wetzstein HY: Stigma development and receptivity in almond ( Prunus dulcis ). Huang Z, Zhu J, Mu X, Lin J: Pollen dispersion, pollen viability and pistil receptivity in Leymus chinensis . Hedhly A, Hormaza JI, Herrero M: The effect of temperature on stigmatic receptivity in sweet cherry ( Prunus avium L.). Grbic V, Bleecker AB: Ethylene regulates the timing of essay writing company Millfield, leaf senescence in Arabidopsis. Lim PO, Kim HJ, Nam HG: Leaf senescence. Lim PO, Nam HG: The molecular and genetic control of leaf senescence and longevity in Arabidopsis.
Lanahan MB, Yen HC, Giovannoni JJ, Klee HJ: The never ripe mutation blocks ethylene perception in tomato. Patterson SE, Bleecker AB: Ethylene-dependent and -independent processes associated with floral organ abscission in that writes for you CATS, Arabidopsis. Butenko MA, Stenvik GE, Alm V, Saether B, Patterson SE, Aalen RB: Ethylene-dependent and -independent pathways controlling floral abscission are revealed to converge using promoter::reporter gene constructs in the ida abscission mutant. Adams-Phillips L, Barry C, Giovannoni J: Signal transduction systems regulating fruit ripening. Orzaez D, Blay R, Granell A: Programme of senescence in petals and carpels of Pisum sativum L. School? flowers and its control by ethylene. Kendrick MD, Chang C: Ethylene signaling: new levels of complexity and regulation. Stepanova AN, Alonso JM: Ethylene signaling and website that writes CATS, response: where different regulatory modules meet. Lin Z, Zhong S, Grierson D: Recent advances in ethylene research. Yoo SD, Cho Y, Sheen J: Emerging connections in the ethylene signaling network.
Alonso JM, Hirayama T, Roman G, Nourizadeh S, Ecker JR: EIN2, a bifunctional transducer of Sidcot, ethylene and stress responses in Arabidopsis. Tsuchisaka A, Theologis A: Unique and website that writes essay for you College Cambridge, overlapping expression patterns among the Arabidopsis 1-amino-cyclopropane-1-carboxylate synthase gene family members. Buchanan-Wollaston V, Page T, Harrison E, Breeze E, Lim PO, Nam HG, Lin JF, Wu SH, Swidzinski J, Ishizaki K, Leaver CJ: Comparative transcriptome analysis reveals significant differences in gene expression and signalling pathways between developmental and critically evaluate Sidcot School, dark/starvation-induced senescence in Arabidopsis. Lohman KN, Gan S, John MC, Amasino RM: Molecular analysis of writes for you College, natural leaf senescence in Arabidopsis thaliana . Noh YS, Amasino RM: Identification of a promoter region responsible for the senescence-specific expression of SAG12 . Knee EM, Hangarter RP, Knee M: Interactions of article, light and ethylene in hypocotyl hook maintenance in Arabidopsis thaliana seedlings. Achard P, Cheng H, De Grauwe L, Decat J, Schoutteten H, Moritz T, Van Der Straeten D, Peng J, Harberd NP: Integration of website that writes CATS, plant responses to environmentally activated phytohormonal signals. Achard P, Baghour M, Chapple A, Hedden P, Van Der Straeten D, Genschik P, Moritz T, Harberd NP: The plant stress hormone ethylene controls floral transition via DELLA-dependent regulation of floral meristem-identity genes.
Pierik R, Sasidharan R, Voesenek LACJ: Growth control by ethylene: Adjusting phenotypes to the environment. Bleecker AB, Estelle MA, Somerville C, Kende H: Insensitivity to Ethylene Conferred by a Dominant Mutation in critically evaluate Sidcot School, Arabidopsis thaliana . Guzman P, Ecker JR: Exploiting the triple response of Arabidopsis to identify ethylene-related mutants. Kieber JJ, Rothenberg M, Roman G, Feldmann KA, Ecker JR: CTR1, a negative regulator of the ethylene response pathway in Arabidopsis, encodes a member of the raf family of website that essay CATS College, protein kinases. Vivian-Smith A, Koltunow AM: Genetic analysis of article writing Oswestry, growth-regulator-induced parthenocarpy in Arabidopsis. Fleet CM, Sun TP: A DELLAcate balance: the role of for you CATS College, gibberellin in custom essay writing company School, plant morphogenesis. Achard P, Vriezen WH, Van Der Straeten D, Harberd NP: Ethylene regulates Arabidopsis development via the modulation of DELLA protein growth repressor function. Okushima Y, Mitina I, Quach HL, Theologis A: AUXIN RESPONSE FACTOR 2 (ARF2): a pleiotropic developmental regulator. Kim CY, Bove J, Assmann SM: Overexpression of wound-responsive RNA-binding proteins induces leaf senescence and hypersensitive-like cell death. Wagstaff C, Chanasut U, Harren FJ, Laarhoven LJ, Thomas B, Rogers HJ, Stead AD: Ethylene and flower longevity in Alstroemeria : relationship between tepal senescence, abscission and ethylene biosynthesis.
Wilkinson JQ, Lanahan MB, Clark DG, Bleecker AB, Chang C, Meyerowitz EM, Klee HJ: A dominant mutant receptor from website that writes essay CATS Cambridge Arabidopsis confers ethylene insensitivity in heterologous plants. Langston BJ, Bai S, Jones ML: Increases in article School, DNA fragmentation and induction of a senescence-specific nuclease are delayed during corolla senescence in website that College, ethylene-insensitive ( etr1-1 ) transgenic petunias. Jones ML, Chaffin GS, Eason JR, Clark DG: Ethylene-sensitivity regulates proteolytic activity and cysteine protease gene expression in petunia corollas. Oh SA, Park JH, Lee GI, Paek KH, Park SK, Nam HG: Identification of three genetic loci controlling leaf senescence in Arabidopsis thaliana . Yang TF, Gonzalez-Carranza ZH, Maunders MJ, Roberts JA: Ethylene and the regulation of how to School, senescence processes in transgenic Nicotiana sylvestris plants. Kim JH, Woo HR, Kim J, Lim PO, Lee IC, Choi SH, Hwang D, Nam HG: Trifurcate feed-forward regulation of age-dependent cell death involving miR164 in Arabidopsis. Sun K, Cui Y, Hauser BA: Environmental stress alters genes expression and induces ovule abortion: reactive oxygen species appear as ovules commit to abort. Sun K, Hunt K, Hauser BA: Ovule abortion in Arabidopsis triggered by writes essay for you Cambridge, stress. Preuss D, Lemieux B, Yen G, Davis RW: A conditional sterile mutation eliminates surface components from Arabidopsis pollen and disrupts cell signaling during fertilization. Fiebig A, Mayfield JA, Miley NL, Chau S, Fischer RL, Preuss D: Alterations in CER6 , a gene identical to CUT1 , differentially affect long-chain lipid content on the surface of critically evaluate, pollen and stems. Abramoff MD, Magelhaes PJ, Ram SJ: Image Processing with ImageJ.
Medina I, Carbonell J, Pulido L, Madeira SC, Goetz S, Conesa A, Tarraga J, Pascual-Montano A, Nogales-Cadenas R, Santoyo J, Garcia F, Marba M, Montaner D, Dopazo J: Babelomics: an integrative platform for the analysis of writes essay for you CATS College, transcriptomics, proteomics and genomic data with advanced functional profiling. 2017 BioMed Central Ltd unless otherwise stated. Part of scientific writing, Springer Science+Business Media.
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