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"A tool good enough to be so used and not too good"C.S. Lewis That Hideous Strength
As regards the OP, in another thread (one of his first on TM) he told us that he was a 17-year-old student from New Zealand just started in trumpet (that was perhaps six months ago). So he has tons of youthful zeal and (until we taught him, sometimes the hard way) no knowledge to speak of. I think we can be quite proud of ourselves that we crammed that much knowledge into him in a few short weeks. But it's far too short a time for him to absorb and adjust all that knowledge. But that is the advantage of youth... let him have the benefit of his exuberant enthusiasm. Better that than a disillusioned hard-bitten klapper of a guy...
That is a lot of trumpets for a newcomer!
My experience is that a horn can be designed heavy and be just fine in all registers, or get pimped and play miserably. George makes an interesting point with the vented tuning slide. The weight is in the wrong place for a standard trumpet.
Still, if we look at the repertory, who needs the help that those keys "could" provide? Certainly most amateurs are not playing the symphonies that they were designed for. Those that are playing this stuff professionally have the opportunities to find "their" instrument and for sure do not need internet advice on why and how to pick it.
We cannot guess the motives of a school kid or even university player for collecting anything. We don't know their playing use case, so we voice either our opinion, or experience for what it is worth.
With a username like brass band major, we can guess on the true focus here, and a rotary trumpet in any form is not going to meet that goal. I prefer someone to come here and ask a thousand questions rather than come on like they know it all but not be able to deliver content. Granted, the questions could be consolidated to a couple of threads instead of spread all over. That is a learning process.
Whenever I feel blue, I start breathing again.
I believe that you're just being hard on him. So he asks a lot of questions, so what?! beeeeeebop
That's good for the forum right? Having traffic and all that
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subset: commoncrawl


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5221 Photographers
Photographers operate still cameras to photograph people, events, scenes, materials, products and other subjects. Photographers are employed by photographic studios, newspapers, magazines, museums and government, or they may be self-employed.
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aerial photographer
commercial photographer
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Main duties
Photographers perform some or all of the following duties:
Study requirements of a particular assignment and decide on type of camera, film, lighting and background accessories to be used
Determine picture composition, make technical adjustments to equipment and photograph subject
May operate scanners to transfer photographic images to computers
May operate computers to manipulate photographic images
May adapt existing photographic images and create new digitized images to be included in multimedia/newmedia products
May use delicate instruments, such as optical microscopes attached to cameras
May process exposed film
May use airbrush, computer or other techniques to retouch negatives
Medical photographers may work closely with medical illustrators.
Photographers may specialize in areas such as portrait photography, commercial photography, scientific photography, forensic photography, medical photography, digitized photography, multimedia photography or photojournalism.
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text:
The tank finally went empty.
NUs season finally comes to an end in loss to Lobos
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text:
GKN's technology is shaping the future of aviation. Take a look how we are showcasing our technology at FIA 2016.
GKN Aerospace is a leader in the manufacture of highly complex composite and metallic aerostructures and engine products.
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6th ceremony at Farnborough airshow commends 6 suppliers for their performance excellence - ...READ MORE


text:
Loyalsock, Canton headed to PIAA baseball finals
Loyalsock and Canton earned spots in the PIAA baseball championships with semifinal victories Tuesday at Bloomsburg University.
The PIAA baseball finals are Friday at Penn State's Medlar Field at Lubrano Park
Loyalsock beat Salisbury Twp., 8-1, in the Class AA semifinal. Up next is District 7 champion Beaver at 3:30 on Friday in the final.
Canton beat Old Forge, 2-1. It will play the Johnsonburg-California winner at 10:30 a.m. on Friday in the final. Johnsonburg and California were scheduled for 6 p.m. Tuesday evening in the other semifinal.
Loyalsock will be chasing its second PIAA baseball title, having won in 2008.
Canton has never won a PIAA baseball title, but the school did win a PIAA softball title in 2000.


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Bachelor of Social Work (Honours)
Bachelor Degree (Honours)
|OPRank, ATAR Cutoff
|Mid Year Intake?
|Study Mode
|Entry Requirements
|CQUniversity Bundaberg
|Full-time internal, Part-time internal
Year 12 or equivalent; QTAC: English (4/SA); UAC: English (Standard) (Band 2); VTAC: Units 3 and 4: a study score of at least 25 in any English
|CQUniversity Rockhampton North
|Full-time internal, Part-time internal
|Distance Education
|Full-time internal, Part-time internal
Course Description
This course includes a range of units focusing on social work knowledge, values, and skills with cognate disciplines also enhancing each student's social work education such as psychology, sociology and Indigenous studies. It is accredited and recognised in every state in Australia and will qualify graduates to work in several other countries without seeking additional accreditation.
Subjects you can Study
Social work
^ Shows the minimum tertiary entrance ranking needed by Australian school leavers to get into each CSP-based course. Cut-offs are not determined in advance. Course data and cut-off scores published on Good Universities Guide are indicative of the 2016 academic year.
About CQUniversityCQUniversity Australia has one of the largest and fastest-growing footprints of any university in Australia. Already renowned as one of the nation's most inclusive universities, CQUniversity is now also a more comprehensive university following the merger with CQ TAFE in 2014. They have opened the doors wider, making university more accessible and applicable to more people.
Students can choose from hundreds of education and training options across a variety of study areas, including Apprenticeships, Trades and Training, Business, Accounting and Law, Creative, Performing and Visual Arts, Education and Humanities, Engineering and Built Environment, Health, Information Technology and Digital Media, Psychology, Social Work and Community Services, Science and Environment and Work and Study Preparation.
Students can study on-campus at one of the many locations around Australia, or by distance education and benefit from ongoing support and access to dedicated Distance Education Study Centres and hubs located across Australia.
The programs and courses are designed alongside industry, to prepare students with up-to-date, work-ready knowledge and skills.
Provider CRICOS: 00219C
Bachelor Degree (Honours)
CQUniversity Bundaberg
- Full-time internal = 4 years
- Part-time internal = 8 years
CQUniversity Rockhampton North
- Full-time internal = 4 years
- Part-time internal = 8 years
Distance Education
- Full-time internal = 4 years
- Part-time internal = 8 years
28 units (192 credit points). Students are required to attend a minimum of 20 days residential school attendance throughout the course. The course incorporates two compulsory 13 week and 16 week unpaid field education experiences in Years 3 and 4.
Entry Requirements
Year 12 or equivalent; QTAC: English (4/SA); UAC: English (Standard) (Band 2); VTAC: Units 3 and 4: a study score of at least 25 in any English
How does this course perform?
How do study fields for Bachelor of Social Work (Honours) at CQUniversity perform?
Life after Study
Related Careers
Social Worker
Social workers help people to deal with personal and social problems, either directly or by planning or carrying out programmes that benefit groups or communities. They are interested in challenging the structural barriers that contribute to inequality, discrimination, exploitation and oppression within communities.


text:
realmagick.com The shrine of knowledge.
An obscure Deity, perhaps a son of Zeus, Associated with Demeter in some ways (one tradition holds Him to be Her son), His chief
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bearing a torch and leading worshippers into the Sanctum.
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The most popular use for coffee syrup is coffee milk, the official state drink of Rhode Island. But its rich taste and texture adds more possibilities to the mix. Pour it over ice cream, add it to a cocktail, or use on French toast. This syrup is an original recipe that adds cocoa powder for an extra layer of chocolately richness. Coffee syrup made with simple ingredients and quality coffee beans was an idea that came out of Dave Lanning's love of java. A Rhode Island native, craft coffee roaster and owner of Dave's Coffee, Lanning was looking for a way to branch out with coffee products after years of roasting his own beans. He and his wife realized there was no coffee syrup available on the market that didn't include artificial ingredients, so using their coffee expertise, they set out to be the first to create it. A real series of trial and error tests ensued. And then their coffee syrup was born. Lanning is as devoted to the natural ingredients in the syrup as the quality of the coffee. Which is why he employs only expert roasters to log every minute of the roasting process for the beans; masters who can use their keen senses to know when the flavors have been brought out and a delicious coffee bean is ready to be ground. To achieve a unique flavor, Dave teamed up with Carissa Albergaria, a student at Johnson & Wales who is now the manager at Dave's, to find the perfect ingredients and ratios to make the syrup. They found that cold-brewed Brazilian coffee - selected exclusively for low acidity and a nutty, sweet, buttery profile - mixed with cane sugar, was the best combination. Founded in the state where coffee syrup was first invented, Dave's coffee syrup has added quality to tradition. "We're definitely coffee roasters first," he said. But ones who happen to make a killer syrup.
This syrup is nice and thick so you can pour it over ice cream. It's also great in a coffee-inspired cocktail, or combined with milk for the classic Rhode Island "coffee milk." Directions: Fill an eight ounce glass with ice cold whole milk. Add three generous tablespoons of coffee syrup, stir and enjoy.
Dave's coffee syrup begins with nutty, sweet, smooth Brazilian coffee beans, roasted in small batches by hand in a small gas fired roaster - a process that is monitored closely by Dave's roast masters, who follow the progress of every batch, logging the time and temperature details every minute. The finished roast rests for two days before being ground and cold-brewed for 18 hours in a modified stainless steel kettle. After the brewing, the coffee is drained and combined with pure cane sugar, then boiled for over an hour. It's during the boiling that the syrup reduces and the sugar begins to caramelize. At the perfect consistency, the syrup is hot bottled into amber glass bottles with metal screw caps. The result not only tastes delicious in milk, on ice cream, or added to a cocktail, but the cocoa adds an extra layer of richness that tastes great in baking. The syrup is packaged in amber glass bottles, chosen for their ability to protect the product inside from light and UV rays, which can change the components of products, especially food. When you protect the ingredients from harmful rays you extend the shelf life and quality of the ingredients. Dave's Coffee is a member of the Farm Fresh RI, an organization that unites local Rhode Island food producers with consumers, restaurants and markets. 700 pounds of coffee are roasted each day in 30 pound batches in their facility, using a small gas fired roaster that was custom made for the company by US Roaster Corp of Oklahoma. To keep the product 100 percent Rhode Island-made, Dave's uses a bottler in Warwick to produce the syrup's vintage packaging.
Coffee syrup made from hand-roasted, cold-brewed Brazilian beans simmered with cane sugar and combined with cocoa. Made in Rhode Island. Comes in 16 ounce glass bottle. (more)
16 ounce amber glass bottle
- Cold Brewed Coffee
- Cane Sugar
- Cocoa
Made in
Rhode Island
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Cochise County, AZ Sheriff killed in car crash
Discussion in 'Cop Talk' started by blueiron, Sep 19, 2012.
Page 1 of 2
Page 1 of 2


text:
Dom Camerada, BobbyJoe Dellavalle and Brandon Sach teamed for 24 points as Our Lady Queen of Peace Gold captured the Catholic Youth Organization High School varsity Red division title with a 40-38 win over St. Joseph/St. Thomas.
Our Lady Star of the Sea topped Our Lady of Pity 55-37 to win the HS varsity White division. Andrew Russo, Tom Maher and James Fusillo combined for 34 points.
Anthony Paladino scored 13 and Nick Gonchar added 11 as St. Joseph/St. Thomas won the High School JV title by defeating Our Lady Star of the Sea 40-37.


text:
If like the rest of the world you didn't notice and/or care, Boy Better Know's Jammer has been absent from Twitter recently and has had his "management" *side eye* tweeting on his behalf. But on Tuesday he announced that he is back to take over our timelines.
Did the east London MC really think he had fooled us into thinking for a second that his "management" were really responsible for the tweets coming from his account? Sorry Jammer, but we weren't convinced. We can all refer to ourselves in the third person.
To add to the hilarity he declared that he was only going to be back on Twitter for a day to wreak havoc and then log off for a year, but unsurprisingly Jammer is still tweeting.
Jammer initially left the social networking site following his tantrum over Ghetts not showing enough love to the grime scene and rumours about Ghetts punching him. Since he's been back he's been heavily promoting his work on Lord Of The Mics 3, as the official single has just dropped. Hmm coincidence? We think not.
We're glad to see Jammer promoting his new material, but we think all this attention seeking needs to stop.


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Dare Dukes: Ballad of Darius McCollum
From the Album "+–Prettiest Transmitter of All: Nov. 18, 2008–+"
|The songs that make up Dare Dukes' new album, Prettiest Transmitter of All, are a striking combination of incisive intelligence and sweet, doleful hooks. Looking past the shiny surfaces of American life, the Savannah, GA-based singer-songwriter chronicles the everyday world, mining the margins for the eccentric characters and bizarre events that are the heart of his music. [MySpace]


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Social and personal
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Do be aware though, that we have certain important style rules that need to be adhered to. Some, such as the way we spell certain words, are there to ensure clarity and consistency. But others, such as the way we address the deceased and use non-Jewish terms may delay the publication of your announcement if we have to call to discuss amendments.


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Greenland is Losing More Ice Than Scientists Thought
Bad news keeps flowing for the icy landscapes of the world. Rising temperatures are melting ice and sending it to the ocean, a process that is pushing sea levels higher and altering the landscape at both poles. The latest news comes from Greenland, where researchers have used high-tech satellite and GPS measurements to see how much mass the ice … Read More
These Images Show Near-Record Low 2016 Arctic Sea Ice
Arctic sea ice is one of the grandaddy's of climate indicators. And this grandaddy isn't doing so good these days. Ice is disappearing at a quickening clip as rapidly warming air and water eats away at the briny, frozen mantle on the top of the planet. In the nearly four decades of satellite monitoring, sea ice has disappeared at a clip of 13.4 … Read More
Despite Major Melt, Arctic Sea Ice Will Miss Record Low
As the sun begins its seasonal descent in the Arctic sky and temperatures drop, the summer melt of sea ice is slowing down. In the next few weeks, the span of the Arctic Ocean covered by ice will reach its annual low. But despite beginning the summer at unprecedentedly low levels, this year's minimum won't break the stunning record of 2012, expert… Read More
This Is What the Ice-Free Northwest Passage Looks Like
An ice-free Northwest Passage was once the stuff of legend. But it's now becoming the norm thanks to global warming, and commercial freighters to luxury cruise ships are racing to turn a profit off the newest frontier on earth. An image published by NASA Earth Observatory shows a nearly ice-free path from the North Atlantic to the Pacific… Read More
Greenland Melt Could Expose Hazardous Cold War Waste
When the U.S. military abandoned Camp Century, a complex of tunnels dug into the ice of northwest Greenland, in the mid-1960s, they left behind thousands of tons of waste, including hazardous radioactive and chemical materials. They expected the detritus would be safely entombed in the ice sheet for tens of thousands of years, buried ever deeper… Read More
Arctic Sea Ice Crashes to Record Low for June
The summer sea ice cover over the Arctic raced towards oblivion in June, crashing through previous records to reach a new all-time low. The Arctic sea ice extent was a staggering 100,000 square miles below the previous record for June, set in 2010. And it was 525,000 square miles below the 1981-2010 long-term average, according to the National … Read More
Scientists Urge Obama to Halt Oil Leasing in the Arctic
In a letter being delivered to the White House on Thursday, nearly 400 scientists from more than a dozen countries are urging President Obama to stop future oil and gas drilling in the Arctic Ocean because of the significant environmental and climate risks associated with further fossil fuels exploration there. "No new oil and gas leasing or … Read More
Melting Arctic Could Supercharge Climate Feedback Loop
As global warming heats the Arctic, carbon dioxide emissions from melting permafrost could play a bigger role in worsening climate change than previously thought, according to a new study. Scientists have long considered methane emissions to be the biggest climate threat posed by thawing permafrost. Methane is more potent than carbon dioxide in … Read More


text: Bomb Suspects Appear Driven by Religion: Officials But Dzhokhar, Tamerlan Tsarnaev had no apparent ties to Islamic terrorist orgs. By Newser Editors and Wire Services Posted Apr 22, 2013 6:26 PM CDT 85 comments Comments Tamerlan Tsarnaev, 26, left, and Dzhokhar Tsarnaev, 19. (AP Photo/The Lowell Sun & Robin Young, File) (Newser) – The two brothers suspected of bombing the Boston Marathon appear to have been motivated by their religious faith but do not seem connected to any Muslim terrorist groups, US officials said today after interrogating the severely wounded Dzhokhar Tsarnaev. Two US officials said preliminary evidence from an interrogation suggests Dzhokhar and brother Tamerlan were motivated by religion but were apparently not tied to any Islamic terrorist organizations. The officials spoke on condition of anonymity because they were not authorized to publicly discuss the investigation.


text:
Michigan State's football team held the No. 1 total defense rank from the beginning of the season up until the bowl game.
The No. 3-ranked Spartans (13-1) ran into Pac-12 champion Stanford (11-3) in the Rose Bowl and prevailed, 24-20.
But Michigan State was not statistically dominant enough against the Cardinal to maintain the No. 1 total defense rank and fell to No. 2.
Louisville (12-1), which defeated Miami, Fla. (9-4) in the Dec. 28 Florida Citrus Bowl, 36-9, slipped by the Spartans in the total defense category by allowing just 174 yards.
The Spartans allowed 305 yards to Stanford.
Louisville finished the season allowing 251.5 yards per game, while Michigan State allowed 252.2 yards per game.
The argument could still be made that the Spartans had the best defense in the nation, as they had the only unit to finish ranked in the top four in every major statistical category, and they allowed the fewest yards per play (4.04) and were No. 1 in pass efficiency defense.
The Spartans' offense improved throughout the season, however, and finished 37th in the nation in third-down efficiency which opened the door to being seventh in the country in time of possession (33 minutes, 19 seconds).
Michigan State finished with the No. 51 schedule strength in the nation, according to the Sagarin computer rankings published in the USA Today and used in the BCS formula throughout its lifespan (1998-2013).
Here's a look at how Michigan State compared to the top-ranked units in the nation, and a look at the top-ranked opponent the Spartans played in each category:
Total Offense
1. Baylor, 618.8 yards per game
7. Ohio State, 511.9
80. Michigan State, 385.5
Rushing Offense
1. Auburn, 328.3
5. Ohio State, 308.6
58. Michigan State, 173.8
Passing Offense
1. Fresno State, 394.8
17. Indiana, 306.7
82. Michigan State, 211.7
Scoring Offense
1. Baylor, 52.4 points per game
3. Ohio State, 45.5
64. Michigan State, 29.4
Total Defense
1. Louisville, 251.5 yards per game
2. Michigan State, 252.2
6. Iowa, 303.1
Rushing Defense
1. Louisville, 80.7
2. Michigan State, 86.6
3. Stanford, 89.4
Passing Defense
1. Florida State, 156.6
3. Michigan State, 165.6
4. Western Michigan, 169.2
Scoring Defense
1. Florida State, 12.1 points per game
3. Michigan State, 13.2
9. Iowa, 18.9
Special Teams
Kick Return Defense: 59. Michigan State, 21.18 yards per return
Kick Returns: 99. Michigan State, 19.17
Net Punting: 26. Michigan State, 38.59
Punt Return Defense: 45. Michigan State, 6.67
Punt Returns: 43. Michigan State, 9.81
Time of Possession: 7. Michigan State, 33:19
Turnover Margin: T-10. Michigan State, Plus 0.9 per game
Pass Efficiency: 63. Michigan State, 129.97
Pass Efficiency Defense: 1. Michigan State, 92.28
Third-Down Conversion Defense: 2. Michigan State, .279
Third-Down Conversions: 37. Michigan State, .441
Completion Percentage: 73. Michigan State, .577
Sacks Allowed: 15. Michigan State, 1.21 per game
Team Sacks: T-46. Michigan State, 2.29 per game
Passing Yards Per Completion: 67. Michigan State, 11.95
Fewest Penalty Yards Per Game: 78. Michigan State, 50.43
• Download the MSU basketball on MLive app for iPhone and Android
• Download the MSU football on MLive app for iPhone and Android
• Follow Diamond Leung and Mike Griffith on Twitter
• Like MLive's Michigan State Spartans Facebook page


text:
Snail bob 2 is a game about hunting the hamster and it has come back with the bang. It is now offering more fun and more puzzles and it has new graphics that are incredible. You can now play the new game with more fun and new saga. Snail Bob starts with the present of the birthday of Grandpa. However, the road to take snail bob to his grandfather is full of dangers. It is important to solve the puzzles in order to figure out what it is going to take place so that he can reach to the birthday party.
The instructions of the game are about pressing the keys using the keyboard and to pull or to rotate levers. When you press at 1, you are going to make Bob to move using the normal speed while when you press on 2, you are going to make his speed double. When you click at the snail, you are going to make him to move even further. You may use the time given to figure out what it is going on yandere simulator. You may use many tools that can help snail Bob to go to his way.
You should clear the 25 levels so that you may help Snail Bob to be able to reach to the destiny he had. However, to be able to get the present of grandpa, you should clear all the 8 levels in order to get the present to give to small Snail Bob.
Within the game, the amusing sound track can keep the person entertained. You may turn it off if you are not feeling to listen to it. The developers had done the best job when it comes to using their graphics. You are going to laugh out when you see how the characters express themselves. Creative minds for the developers can give the miraculous touch to all the characters in the entire game. You may play snail bob in order to reach to a new gaming experience.
Game Play
Snail bob 2 is unique puzzle game based on the flash. The game is about a snail and sometime you may think that it is boring to deal with the snail but difficult obstacles, interactive puzzles, interesting enemies make the game interesting. As you continue to play, the game is going to get even harder and range of the challenges may include launching of the snails within the air and pushing the button or pulling of the levers in order to get more pathways while avoiding the dangers. The game can be played by new people or the people who are experienced. Sometime the children can also play it. In order to support this, the controls are too simple. The controls do revolve around the use of the keys in order to slow down or speed up the snail movement. You can also use the left mouse button so that you can interact with the new objects within the environment. When you click at Bob, you will be able to make him to stay while you will be thinking through the puzzle and what dangers he is about to face.


text:
Jason Bentley takes his audience to Critical Mass Studios in Santa Monica for an inside look at the process of remixing a piece of music.
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Appearance of a ventricular 5-HT4 receptor-mediated inotropic response to serotonin in heart failure
Background: Current pharmacological treatment of congestive heart failure (CHF) addresses changes in neurohumoral stimulation or cardiac responsiveness to such stimulation. Yet, undiscovered neurohumoral changes, adaptive or maladaptive, may occur in CHF and suggest novel pharmacological treatment. Serotonin [5-hydroxytryptamine (5-HT)] enhances contractility and causes arrhythmias through 5-HT4 receptors in human atrium and ventricle but not through rat ventricular 5-HT4 receptors.
Objective: We investigated whether CHF could induce ventricular responsiveness to serotonin.
Methods: Postinfarction CHF was induced in male Wistar rats by coronary artery ligation. Contractility was measured in left ventricular papillary muscles 6 weeks after infarction. Messenger RNA was quantified by RT-PCR and cAMP by RIA.
Results: Serotonin caused positive inotropic (−logEC50=7.5) and lusitropic effects in CHF but not Sham papillary muscles. The inotropic effect of 10 μM serotonin in CHF (31.3 ± 2.2%) was of similar size as the effect of 10 μM isoproterenol (34.0 ± 1.7%). The effects of serotonin were antagonised by GR113808 (0.5–5 nM), consistent with mediation through 5-HT4 receptors. This was further supported by positive inotropic effects of the 5-HT4-selective partial agonist RS67506. Carbachol blunted the serotonin responses and serotonin increased ventricular and cardiomyocyte cAMP, consistent with coupling to Gs and adenylyl cyclase. Quantitative RT-PCR revealed fourfold increased 5-HT4(b) mRNA expression in CHF vs. Sham ventricles.
Conclusion: Functional ventricular 5-HT4 receptors are induced by myocardial infarction and CHF of the rat heart. We propose that they are a model for ventricular 5-HT4 receptors of human failing heart and may play a pathophysiological role in heart failure.
- Serotonin (5-HT)
- Heart failure
- 5-HT4 receptor
- Ventricle
- Contractile function
1. Introduction
Our understanding of congestive heart failure (CHF) pathophysiology has changed dramatically from a hemodynamic model with treatment aimed at correcting hemodynamic defects, to a neurohumoral model with treatment aimed at preventing maladaptive, biological changes [1,2]. In addition to the changes in, e.g., the adrenergic system, there may be changes, adaptive or maladaptive, in other neurohumoral systems during CHF. Exploration of such possible changes may increase our understanding of CHF.
The neurohumoral system related to the neurotransmitter and vasoactive indoleamine serotonin is one candidate, and increased plasma levels and activity of serotonin have been reported in human CHF [3,4]. Circulating serotonin, mainly derived from enterochromaffin cells of the gastrointestinal tract, is taken up, stored, transported by and released from platelets [5]. Serotonin can also be captured and released by sympathetic nerve endings [6] and activate 5-hydroxytryptamine (5-HT) receptors in the human atrium [7,8]. Serotonin exerts diverse effects through at least 14 different receptors, named 5-HT1 to 5-HT7 with subgroups, of which 5-HT4, 5-HT6 and 5-HT7 receptors increase cAMP-formation through the G-protein Gs, similar to β-adrenoceptors [9].
Serotonin causes cardiostimulation either directly or indirectly through modulation of norephinephrine release from sympathetic nerve terminals [6]. Direct inotropic effects of serotonin mediated through 5-HT4 receptors have been observed in atrium [6–8] and ventricle [10] of man and pig, and through atrial 5-HT2A receptors in rat [11], and 5-HT4 receptors have been implicated in arrhythmia [8,12,13]. In addition, 5-HT4 receptor mRNA was detected in rat atrium, but not ventricle [14]. Early reports indicated lack of inotropic effects of serotonin in human [15,16] and porcine [17,18] ventricle. However, 5-HT4(a) and 5-HT4(b) receptor mRNA have been detected in human left and right ventricle [19,20]. Moreover, we recently demonstrated that phosphodiesterase inhibition uncovers functional ventricular 5-HT4 receptors in porcine and failing human ventricle and 5-HT4(b) mRNA was increased fourfold in failing human hearts compared to donor hearts [10]. The aim of the present study was to explore the possibility that serotonergic signalling through 5-HT4 receptors may become apparent in the remaining ventricular myocardium of the well-characterised model of postinfarction CHF in rats, following coronary artery ligation [21–23]. Our results revealed the appearance of 5-HT4 receptor-mediated inotropic and lusitropic effects of serotonin in CHF rat papillary muscle, accompanied by a fourfold increase in 5-HT4 receptor mRNA.
2. Materials and methods
2.1. CHF model
The investigation conforms with the Guide for the Care and Use of Laboratory Animals published by the U.S. National Institutes of Health (NIH Publication No. 85-23, revised 1996). Two animals per cage in a temperature-regulated room on a 12:12-h day/night cycle were given access to food and water ad libitum. As described [24], an extensive myocardial infarction (MI) was induced in 320 g male Wistar rats under anaesthesia (68% N2O/29% O2/2–3% Isofluran) by proximal ligation of the left coronary artery. Six weeks later, CHF rats were included in the study if left ventricular end-diastolic pressure (LVEDP, measured by catheterisation as described [24]) was ≥ 15 mm Hg and the rats had clinical signs of CHF. Rats with LVEDP<15 mm Hg without symptoms of CHF were included in the myocardial infarction nonfailing (MInf) group [23]. Sham-operated animals (Sham) underwent identical surgical procedure without coronary artery ligation. The endocardial surface of CHF and MInf hearts was digitally photographed and infarct size (% of inner surface) was traced on screen (KS100, Kontron electronics, Germany).
2.2. Isolated papillary muscles
Posterior left ventricular papillary muscles (diameter<1.0 mm) were prepared, mounted in 31 °C organ baths at 1.8 mM Ca2+, equilibrated and field-stimulated at 1 Hz [24] and the contraction–relaxation cycles (CRCs) were recorded and analysed as described [24,25] with respect to maximal developed force (Fmax, mN), maximal development of force [(dF/dt)max], time to peak force (TPF), time to 80% relaxation (TR80) and relaxation time (t80=TR80−TPF). Inotropic responses were expressed as increase of Fmax (% and mN) and of (dF/dt)max (%). Lusitropic effects were expressed as decrease of t80. Blockers of α1-adrenoceptors (prazosin 1 μM), β-adrenoceptors (timolol 1 μM), muscarinic cholinergic receptors (atropine 1 μM) and, where indicated, 5-HT2A receptors (ketanserin 0.1 μM) were added 90 min before agonist. Serotonin was added to the organ bath cumulatively (concentration–response curves) or as a bolus (10 μM). The apparent inhibition constant Kb for the selective 5-HT4 blocker GR113808 was calculated from ratios of serotonin EC50 values with and without GR113808. GR113808 did not influence basal CRC characteristics or electrical stimulation threshold (not shown). Concentration–response curves were constructed by estimating centiles (EC10 to EC100) and calculating the corresponding means, and the horizontal positioning expressed as −logEC50 values [24].
2.3. cAMP content in perfused hearts
Hearts were excised and retrogradely perfused [24] at 31 °C with the additional presence of the 5-HT7 receptor antagonist SB269970 (0.1 μM) to eliminate stimulation of cAMP production by vascular 5-HT7 receptors [26]. Spontaneously beating hearts were equilibrated for 30 min before 1-min perfusion with 5-HT (10 μM). Infarcted area was quickly removed and remaining heart was freeze-clamped. Frozen heart powder (300 mg) was homogenised at 4 °C in 5 ml 5% trichloroacetic acid and cAMP content was determined by radioimmunoassay [27].
2.4. cAMP accumulation in cardiomyocytes
Hearts were perfused (Langendorff setup) with Buffer-1 (mM): NaCl–130, Hepes–25, d-glucose–22, KCl–5.4, MgCl2–0.5, NaH2PO4–0.4, insulin–0.01 μg/ml (pH 7.4), then with Buffer-1 containing 200 U/ml collagenase Type-II and 0.1 mM Ca2+ until the aortic valves were digested. Left ventricle free wall and septum were minced and gently shaken at 37 °C for 3–4 min in the same solution plus 1% BSA. Following filtration (nylon mesh–200 μm) and sedimentation (37 °C), the cell pellet was washed three times in Buffer-1/1% BSA/0.1 mM Ca2+, resedimented in Buffer-1/1% BSA with first 0.2 mM, then 0.5 mM Ca2+and incubated 2 h (37 °C) in laminin-coated six-well plates. Plated cells were washed in Buffer-1/1 mM Ca2+ and incubated 2 h. Receptor blockers (as in contractility experiments) were added 10 min prior to 500 μM 3-isobutyl-1-methylxanthine, 10 min prior to 15-min incubation with 5-HT (10 μM). Incubations were stopped by trichloroacetic acid (5% final), and intracellular plus extracellular cAMP (expressed as % increase above basal) was determined by radioimmunoassay [27].
2.5. Qualitative and quantitative RT-PCR
Fresh noninfarcted left ventricle tissue or papillary muscle, collected after completion of functional analyses, was stored in RNAlater (Ambion) until use, then prepared and qualitative and quantitative RT-PCR performed as described [20]. Sets of primers (targeted to intron/exon boundaries to avoid genomic DNA signals) and probes (Double-Dye Oligonucleotide, 5′-labeled with the fluorescent reporter dyes FAM (5-HT4(b) and 5-HT2A), JOE [glyceraldehyde-3-phosphate dehydrogenase (GAPDH)] or Yakima Yellow [YY; atrial natriuretic peptide (ANP)] and quenched with TAMRA (5-HT4(b), 5-HT2A and GAPDH) or Dark Quencher (DQ; ANP) for quantitative PCR were designed as described [28]. The names and sequences of upper (U) and lower (L) primers and probes (P) used were (5′-3′): 5-HT4(b): ON283(U), CATGTGCTAAGGTATACAGTGGAATGT; ON284(L), GCAGCCACCAAAGGAGAAGTT; TM14(P), FAM-CTGTGAGGTGACACCGACTCTCCCATT-TAMRA; 5-HT2A: ON273(U), TTCACCACAGCCGCTTCAA; ON274(L), ATCCTGTAGTCCAAAGACTGGGATT; TM9(P), FAM-ATGGATATACCTACAGATATGGTCGTCCACACGGCAAT-TAMRA; ANP: ON285(U), ATCTGATGGATTTCAAGAACC; ON286(L), CTCTGAGACGGGTTGACTTC; TM16(P) YY-CGCTTCATCGGTCTGCTCGCTCA-DQ; GAPDH: ON279(U), CCTGCACCACCAACTGCTTA; ON290(L), GGCATGGACTGTGGTCATGA; TM12(P), JOE-TGGCCAAGGTCATCCATGACAACTTTG-TAMRA.
2.6. Statistics
All results are expressed as mean ± S.E.M. unless otherwise indicated and statistical significance assessed with unpaired and paired Student's t-tests or nonparametric Mann–Whitney test.
3. Results
3.1. Characteristics of CHF animals
All rats in the CHF group had large anterolateral infarcts and signs of CHF including tachypnea, pleural effusion and pulmonary congestion, while the rats in the MInf group had infarcts of variable size and no signs of CHF. For animal characteristics and hemodynamic data, see Table 1.
|Sham (n=10)
|MInf (n=58)
|CHF (n=36)
|Body weight, g
|426 ± 9.2
|416 ± 3.8
|394 ± 4.3*,†
|Heart weight, g
|1.50 ± 0.03
|1.85 ± 0.03‡
|2.45 ± 0.04*,†
|Heart wt/body wt, g/kg
|3.5 ± 0.1
|4.5 ± 0.1‡
|6.3 ± 0.1*,†
|LVEDP, mm Hg
|5.5 ± 0.6
|8.7 ± 0.5‡
|26 ± 1*,†
|LVSP, mm Hg
|125 ± 4.2
|112 ± 2.8‡
|95 ± 2.8*,†
|Lung weight, g
|1.6 ± 0.02
|1.6 ± 0.02
|4.2 ± 0.14*,†
|Basal Fmax, mN
|5.7 ± 0.6
|5.5 ± 0.2a
|5.6 ± 0.2
3.2. Inotropic effects of serotonin
Serotonin (10 μM) elicited a monophasic positive inotropic response in CHF papillary muscles in contrast to the complete lack of mechanical response in Sham (n=6; Figs. 1 and 2A). With a diffusion delay of about 5 s, the time from serotonin addition to 50% and maximal inotropic response was 22 ± 2 s and 1–2 min, respectively (Fig. 1–inset). Serotonin (10 μM) increased Fmax in CHF by 31.3 ± 2.2% (n=10, p<0.0001; basal: 5.7 ± 0.2 mN; 5-HT: 7.3 ± 0.3 mN) and (dF/dt)max by 44.5 ± 2.9%, similar to the effect of 10 μM isoproterenol in CHF [Fmax increase: 34.0 ± 1.7%, n=10, p<0.0001; basal: 5.4 ± 0.4 mN; Iso: 7.3 ± 0.5 mN; Fig. 2A; (dF/dt)max increase: 58.5 ± 7.0%]. Isoproterenol increased Fmax in Sham by 113.4 ± 4.8% [n=6, p<0.0001; basal: 5.7 ± 0.6 mN; Iso: 12.2 ± 1.3 mN; Fig. 2A; (dF/dt)max increase: 111.6 ± 8.6%]. After stabilisation of the serotonin response in CHF, isoproterenol did not cause a significant further increase in Fmax (Iso: 7.7 ± 0.4 mN; n=10, nonsignificant vs. 5-HT) or (dF/dt)max (60.4 ± 5.8%). The concentration–response curve for serotonin exhibited a −logEC50M value of 7.49 ± 0.08 (n=5) with 12.1 ± 1.2% (p=0.0018) increase in Fmax [basal: 5.6 ± 0.5 mN; 5-HTmax: 6.3 ± 0.5 mN obtained at about 1 μM; (dF/dt)max increase:18.1 ± 3.5%]. The lower maximal inotropic effect in these experiments compared to single-dose (10 μM serotonin) experiments, e.g., Fig. 2A, may reflect desensitisation as well as more complete activation of relaxing components during cumulative agonist addition in the concentration–response experiments.
3.3. Lusitropic response to serotonin
Serotonin reduced TR80 more than TPF, resulting in reduced t80 in papillary muscles from CHF but not Sham rats (Table 2, Fig. 2B and C). Similar to β-adrenoceptor stimulation [25], these contraction–relaxation cycle changes demonstrate lusitropic effects in addition to inotropic effects of serotonin, consistent with common signalling pathways.
|Control (ms)
|5-HT (ms)
|5-HT (% of control)
|Iso (ms)
|Iso (% of control)
|125 ± 5.5
|124 ± 5.5
|99.0 ± 0.8
|109 ± 2.7
|87.4 ± 2.3*
|234 ± 11.2
|231 ± 11.0
|99.0 ± 0.7
|174 ± 4.9
|74.5 ± 1.7*
|108 ± 6.2
|107 ± 5.9
|99.0 ± 0.5
|75 ± 2.5
|69.4 ± 3.2*
|148 ± 3.9
|134 ± 3.5
|90.5 ± 1.4*,†
|123 ± 1.6
|83.2 ± 1.7*
|254 ± 6.3
|225 ± 5.4
|88.3 ± 1.4*,†
|204 ± 2.9
|80.3 ± 1.9*
|106 ± 2.5
|91 ± 2.2
|85.2 ± 1.5*,†
|81 ± 1.8
|76.2 ± 1.8*
3.4. The inotropic response to serotonin is inhibited by cholinergic stimulation and associated with increased cAMP levels
In the absence of atropine, stimulation of muscarinic acetylcholine receptors by carbachol (30 μM) reversed the positive inotropic response to 10 μM serotonin (n=5, p=0.002; Fig. 3A) and partially reversed TPF, TR80 and t80 after 1–2 min (not shown). In contrast, carbachol alone did not cause a reduction in basal; instead, on occasion, a small increase in basal force was observed (not shown). Atropine (1 μM) restored the serotonin response in the presence of carbachol and reestablished the contraction–relaxation cycle characteristics. This functional antagonism by cholinergic stimulation, as expected from stimulation of Gi-coupled M2 muscarinic acetylcholine receptors, is consistent with involvement of the Gs-cAMP pathway. Accordingly, cAMP levels in perfused hearts increased by 14.8 ± 4.6% (n=5, p=0.02; Fig. 3B) after 1 min serotonin (10 μM) stimulation (basal: 0.415 ± 0.017; serotonin: 0.476 ± 0.019 pmol/mg wet weight), compared to about 100% increase by 0.1 μM isoproterenol [29]. To eliminate a possible influence of noncardiomyocytes, cardiomyocytes isolated from failing hearts were stimulated for 15 min with 10 μM serotonin, resulting in increased cAMP levels by 43.1 ± 23.6% (n=8, p=0.05; Fig. 3B), which was 5% of that obtained with 10 μM isoproterenol (not shown).
3.5. Evidence for involvement of 5-HT4 receptors
Given subsequently to serotonin, the 5-HT4-selective antagonist GR113808 (1 μM) completely reversed the inotropic response and partially reversed the lusitropic effect, when measured after 1–5 min (Fig. 3C). The apparent incomplete reversal of the lusitropic effect most likely reflects slow reversal kinetics explained by slow dephosphorylation of some proteins involved in the lusitropic effect, e.g., troponin-I [30–32]. Serotonin concentration–response curves without and with 0.5 or 5 nM GR113808 were nearly parallel with −logEC50M values of 7.62 ± 0.06 (n=6), 7.32 ± 0.06 (n=6) and 6.37 ± 0.10 (n=6), respectively (Fig. 3D), and no difference in basal and maximal Fmax. The pooled average values for basal and maximal Fmax were 5.4 ± 0.4 and 6.1 ± 0.4 mN (n=18), respectively. The average apparent GR113808 inhibition constant (Kb) was 0.4 nM (−logKb=9.4), consistent with the reported affinity of GR113808 at 5-HT4 receptors [33]. Since GR113808 displays at least 1000-fold selectivity for the 5-HT4 receptor over all other receptors examined [33], we conclude that the effect is 5-HT4 receptor mediated. We also tested the ability of the 5-HT4 selective partial agonist RS67506 [34] to elicit a positive inotropic effect. RS67506 (1 μM) increased Fmax by 3.9 ± 0.4% (n=3, p=0.013, Fig. 3F) in CHF in contrast to no effect in Sham (data not shown). An increase to 10 μM RS67506 did not cause any additional augmentation of Fmax (Fig. 3F). Subsequent addition of 10 and 100 μM of 5-HT increased Fmax further to 9.9 ± 0.3% (n=3) and 16.8 ± 0.9% (n=3), respectively, above basal. The low effects of serotonin at these concentrations reflect the antagonist activity of the partial agonist RS67506 towards the full agonist serotonin in the failing hearts (Fig. 3F). When added subsequently to 5-HT (10 μM), RS67506 (10 μM) effectively reversed the developed positive inotropic effect (5-HT: 36.3 ± 6.5%, n=3 vs. RS67506: −3.8 ± 2.6%, n=3; Fig. 3E). Accordingly, RS67506 acted as a partial 5-HT4 agonist with an intrinsic activity around 10% compared to the full agonist serotonin in ventricle from failing hearts.
A 5-HT2A receptor-mediated inotropic effect of serotonin was demonstrated in rat atrium [11]. However, in CHF rat papillary muscles, ketanserin (0.1 μM) did not shift the concentration–response curve for serotonin to higher agonist concentrations [−logEC50M value with ketanserin 7.62 ± 0.06 (n=6) vs. 7.49 ± 0.08 (n=5) without, nonsignificant], demonstrating that the 5-HT2A receptor is not involved (Fig. 3E). The maximal inotropic responses with and without ketanserin were similar [Fmax increase without: 12.1 ± 1.2%; basal: 5.6 ± 0.5 mN, 5-HTmax: 6.3 ± 0.5 mN (n=5); with: 12.2 ± 1.2%; basal: 5.0 ± 0.4 mN, 5-HTmax: 5.7 ± 0.5 mN (n=6), nonsignificant; (dF/dt)max increase: 18.1 ± 3.5% vs. 17.0 ± 2.3%; nonsignificant].
3.6. Induction of 5-HT4(b) receptor mRNA in CHF
Quantitative RT-PCR was used to determine the mRNA level for 5-HT4(b) receptor, 5-HT2A receptor, and the heart failure-marker ANP normalised to GAPDH mRNA. In left ventricle and papillary muscle, respectively, 5-HT4(b) mRNA levels were 4- and 18-fold higher in CHF vs. Sham, whereas 5-HT2A mRNA levels were unchanged and ANP mRNA levels increased (Fig. 4). The ratio between normalised 5-HT4(b) and 5-HT2A mRNA levels was increased in CHF in both left ventricle and papillary muscle, confirming increased 5-HT4(b) mRNA level relative to 5-HT2A, independent of GAPDH (Table 3).
|Left ventricle
|0.87 ± 0.33 (n=7)
|3.20 ± 1.09 (n=11)*
|Papillary muscle
|0.85 ± 0.64 (n=4)
|7.48 ± 2.44 (n=5)†
3.7. Maximum inotropic response to serotonin and 5-HT4(b) receptor mRNA related to infarction size
In papillary muscles from infarcted nonfailing (MInf) rats, serotonin (10 μM) elicited an inotropic response qualitatively similar to the response observed in CHF animals. The magnitude of the response correlated positively (r2=0.62, p<0.0001) with infarction size up to 30–40% of the inner myocardial surface in MInf animals (Fig. 5A). In MInf hearts with infarct size 30–40%, the inotropic effect was of the same magnitude as in CHF hearts which all had infarct size >40%. An increase with infarction size was also observed for 5-HT4(b) and ANP mRNA expression, whereas 5-HT2A mRNA levels were unchanged (Fig. 5B). 5-HT4(b) and ANP mRNA levels correlated positively (r2=0.37, p<0.01). These findings are consistent with a gradual transition of phenotype related to the extent of myocardial changes secondary to the infarction.
4. Discussion
We demonstrate for the first time the induction of a ventricular inotropic response to serotonin in infarcted hearts and CHF. The mechanical response, absent in Sham, is of similar size as the maximal inotropic response to β-adrenoceptor stimulation in CHF. The response to serotonin is mediated by Gs-coupled 5-HT4 receptors and accompanied by small increases of cAMP levels. The 5-HT4 receptor mRNA expression is also increased in CHF.
We recently reported functional 5-HT4 receptors in failing human ventricle and an increase in 5-HT4 mRNA in failing human hearts [10]. The appearance of functional 5-HT4 receptors in failing rat ventricle, together with the evidence in human heart, adds new dimensions to neurohumoral changes in CHF and serotonergic signalling in the heart. Since no inotropic effect of serotonin was previously observed in human [15,16] or porcine [17,18] ventricles, administration of 5-HT4 receptor antagonists to prevent arrhythmia [12] was thought of as an atrial-specific intervention [13]. We now propose that the appearance of ventricular 5-HT4 receptor function after myocardial infarction and increased ventricular 5-HT4 mRNA in rats with developing and manifest heart failure may provide an experimental model for 5-HT4 receptor function in human ventricle. 5-HT4 receptor antagonists may therefore target not only atrial 5-HT4 receptors [8,12], but also ventricular 5-HT4 receptors [10].
5-HT4 receptor mRNA was detectable in the Sham rat ventricles (this study) and in rat atria [11,14], but functional responses could not be elicited in these tissues, possibly due to low or absent receptor protein expression. Robust inotropic responses have previously been detected in human and porcine atria despite a very low 5-HT4 receptor density [35,36]. This is apparently also the case in the failing rat ventricle, as we found the 5-HT4 receptor density to be below the limit of detection with both [3H]GR113808 and [125I]SB207710 as radioligands.
Although more than one mechanism of action may be involved, we hypothesise, based on several lines of evidence, that the inotropic effect of serotonin in CHF is at least partly cAMP mediated: (1) Serotonin increased cAMP levels, although to a small extent, in perfused hearts and isolated cardiomyocytes. (2) Serotonin elicited fast-developing inotropic and lusitropic effects–characteristics similar to responses to isoproterenol. (3) The pharmacological criteria for the involvement of Gs-coupled 5-HT4 receptors were fulfilled, and 5-HT4 receptor mRNA was induced in CHF. (4) The inotropic effect of serotonin was attenuated by muscarinic receptor activation. (5) In isoproterenol-stimulated muscle, serotonin did not cause additional response, as would be expected if different signalling pathways were used. Thus, in addition to the measured increase in cAMP levels, all the functional and pharmacological characteristics observed in relation to the serotonin response are to be expected for cAMP-mediated mechanical responses in mammalian heart muscle [25]. Furthermore, activation of 5-HT4 receptors in human atria increases cAMP levels, accompanied by increased PKA activity and phosphorylation of L-type calcium channels, phospholamban and troponin-I, similar to β-adrenoceptor activation [7,8].
Although the lack of an additional effect of serotonin in the presence of isoproterenol and the similarity of the time courses strongly indicate that both agonists fully activate a common signalling pathway in a functionally important compartment, the increase in total cAMP content induced by serotonin was small compared to that induced by isoproterenol. Compartmentation of cAMP may result in differing responses (mechanical and biochemical) to activation of cAMP production by different agonists and different receptor types [37–39]. Correlating increases of cAMP in different subcellular compartments with the functional response indicated that cAMP bound in a particulate fraction was of special relevance [39]. This fraction amounted to about 20% of the total cAMP content in rat heart. Thus, a small increase in total tissue level of cAMP may reflect a sufficient increase in a functionally important compartment. The present results do not, however, exclude that other mechanisms of action may be involved in the response to serotonin.
The mechanism of induction of the 5-HT4-mediated inotropic response is unknown. The results from MInf rats indicate that manifest CHF is not a prerequisite. The lack of a further increase in inotropic response from MInf rats with infarction size 30–40% to CHF rats may reflect that the inducing mechanism(s) are fully activated in this situation or that there may be a desensitisation of the Gs-mediated responses in CHF (as for β-adrenoceptor signalling) superimposed upon the induction. The trigger mechanism for induction of the serotonin response should be sought among changes related to the extent of myocardial damage. Similar studies with different CHF models will be helpful to further address this question. Previous studies did not detect 5-HT4 mRNA in rat ventricle [11,14,40]. Although 5-HT4 mRNA is present in atria from normal rats [11,14,40], no functional response was detected [11]. Although speculative, it is tempting to suggest that 5-HT4 receptor-expression in CHF ventricle, like ANP, reflects some kind of "atrialisation" of compromised ventricular myocytes. The positive correlation between 5-HT4 mRNA and ANP mRNA in MInf hearts is in line with this.
A crucial question is whether the endogenous serotonin level is sufficient to activate myocardial 5-HT4 receptors in vivo. Plasma serotonin concentration in rats was estimated to approximately 0.3 μM [11], about 10-fold higher than the EC50 (0.02–0.03 μM) for serotonin in the present study, at recombinant 5-HT4 receptors [19] and in human atrium [7,8]. In addition, sympathetic nerve endings have been shown to capture and release 5-HT analogous to norepinephrine, thus stimulating myocardial 5-HT4 receptors [6], consistent with an activation of myocardial 5-HT4 receptors in vivo.
The induction of the ventricular serotonin effect in infarcted hearts and CHF may be considered compensatory, e.g., to rescue contractile function. In CHF, sympathetic nerve activity and plasma levels of norepinephrine are increased to levels shown to cause desensitisation in the β-adrenoceptor signalling cascade and to induce cardiomyocyte injury [41]. Overexpression of cardiac β1-adrenoceptors or Gαs in transgenic mice also produces cardiomyopathy [42,43]. In this light, chronic induction of a cAMP-mediated serotonin response may appear maladaptive, considering that although β-adrenoceptor signalling is desensitised in CHF, treatment with β-blockers is beneficial [2]. The plasma norepinephrine level in CHF is positively correlated to mortality. Corresponding data for serotonin are lacking. There are reports of increased plasma levels and activity of serotonin in human CHF [3,4], and increased left ventricular serotonin concentrations were reported in hypertensive heart disease [44]. We have recently found functional 5-HT4 receptors in the ventricular myocardium of patients with terminal heart failure, including ischaemic heart disease and reported that serotonin can elicit experimental ventricular arrhythmias [10]. It is therefore conceivable that 5-HT4-selective antagonists may prevent life-threatening arrhythmias in terminal heart failure. Although speculative, if serotonin stimulates the ventricular 5-HT4 receptors in vivo in CHF, blockade of these receptors may have a beneficial effect, in analogy with β-adrenoceptor-blockade.
5. Conclusion
A 5-HT4 receptor-mediated inotropic response to serotonin, associated with increased 5-HT4 receptor mRNA levels, is induced in ventricular myocardium of infarcted and subsequently failing rat heart. The 5-HT4 receptor and β-adrenoceptors apparently share a common signalling pathway in CHF rats. Future challenges comprise exploring the trigger mechanisms responsible for induction of the serotonin response and exploring the pathophysiological and potential therapeutic importance of this novel neurohumoral change in CHF. We propose that the infarcted failing rat heart is an experimental model for investigation of ventricular myocardial 5-HT4 receptors, present also in the failing human heart.
Supported by The Norwegian Council on Cardiovascular Diseases, The Research Council of Norway, Anders Jahre's Foundation for the Promotion of Science, The Novo Nordisk Foundation and The Family Blix foundation.
↵1 EQ and TB contributed equally to the present work.
Time for primary review 30 days
- Copyright © 2004, European Society of Cardiology


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