document_title: To the Manor Born

question_text: where did they film to the manor born

question_tokens: where did they film to the manor born

text: To the Manor Born

text: Production[edit]

text: Development[edit]

text: Peter Spence first thought of the idea behind To the Manor Born in the early 1970s when he was working for BBC Radio as a gag writer.[2] One of the programmes that Spence wrote for featured a Cockney comedian, who had recently bought a manor house in an English country village.[citation needed] When holding a housewarming party, the comedian invited the previous occupant, a widow who could not afford to keep the house up and had moved to a smaller house in the village.[citation needed][2] The comedian's account of the lady, and the conversation he had with her, Spence later described as a "perfect description" of Audrey.[citation needed]

text: A few years later, following the success of The Good Life, Spence was asked by BBC Radio to come up with an idea for a programme to feature Keith.[citation needed][2] Thinking of Keith's character in The Good Life, Spence had the idea of an upper-class version of Margo Leadbetter and, from the account from the comedian, came up with Audrey fforbes-Hamilton.[citation needed] Instead of a Cockney comedian as the new owner of the manor, Spence decided on an American who sees the manor while in England looking for his roots. The American later discovers he is descended from the fforbes-Hamiltons. This was made into a radio pilot in 1976 with Bernard Braden as the American, but was never broadcast because of the interest to make it into a TV series.[citation needed][3]

text: When writing the TV series, Christopher Bond was brought in as the script associate and helped to adapt the series from radio to television.[3] The American character was changed, and the idea of a character who appeared to be an English gentleman but turns out not to be, was thought of.[citation needed] It was then decided that this character needed someone who could reveal his real background, and his mother was created, although it was not until Spence had nearly finished the first series that she was written in from the beginning.[citation needed] The characters of Brabinger and Marjory followed soon after.[citation needed] The first episode aired on 30 September 1979, a year after Keith had played Margo Leadbetter for the last time in The Good Life.[4]

text: Filming[edit]

text: To the Manor Born was filmed on location in Cricket St Thomas, Somerset, and in studio one, BBC Television Centre, London.[5] The Manor, Cricket House, was at the time of the original series owned by the father-in-law of Peter Spence, the show's creator and writer.[2] Some interior scenes were also filmed inside Cricket House. The Old Lodge, which on screen was at the end of the Manor's drive, is in fact about one mile away and called West Lodge.[6][7] A false gatepost was installed to help the illusion that the two are close together.[6] The 2007 special was, like the original series, filmed on location in Cricket St Thomas in Somerset.[8] This filming took place for about a week starting on 28 October 2007.[9] The studio footage was filmed in front of a live audience at Pinewood Studios in Buckinghamshire on 25 November 2007.

text: All the episodes were directed by Gareth Gwenlan, who also produced the original series.[10][11] The 2007 special was co-produced by Gwenlan and Justin Davies.[10]

text: During the filming, many cars were used; but the BBC continued to use the same vintage Rolls Royce motorcar through the entire series, including the opening titles for the show. The Rolls Royce (BMG443) has also been used in other BBC programmes such as Dad's Army.

text: Cast[edit]

text: The cast is led by Penelope Keith as Audrey Fforbes-Hamilton and Peter Bowles as Richard. The other main cast members in the original series are Angela Thorne (playing Audrey's old friend Marjory), Daphne Heard (Richard's mother, Mrs. Polouvicka), John Rudling (Brabinger the butler), Michael Bilton (Ned, the odd-job man) and Gerald Sim (the Rector). Rudling was absent in the 1979 Christmas special and for much of the second series due to his ill health; his character was temporarily replaced as butler by Ned.[5][12] Rudling died in 1983.[12] Angela Thorne had worked with Keith before when she had played Lady "George" Truscott in a 1977 episode of The Good Life.[5] Michael Bilton played Basil Makepeace, a main character in the first four series of the sitcom Waiting for God in the 1990s before his death in 1993.[12]

text: Peter Bowles had been asked some years earlier to play Jerry Leadbetter, the husband of Keith's Margo, in The Good Life.[5][13][14] Owing to prior commitments, Bowles turned down the role, but had he played Jerry, it is unlikely he would have been offered the part of Richard; as Bowles later said, "there's no way I could have played Penelope Keith's screen husband in two separate sitcoms".[14] However, in 1987 Bowles replaced Geoffrey Palmer as Keith's onscreen husband in the ITV sitcom Executive Stress.[3]

text: The series also features many recurring characters, who are either estate workers or members of Audrey's social circle. Celia Imrie, who makes two appearances as Polly, later became known for her work with Victoria Wood, including a lead role in Dinnerladies, as well as lead roles in Kingdom and After You've Gone.[12][15]

text: The 2007 Christmas special features four of the original cast members: Penelope Keith, Peter Bowles, Angela Thorne, and Gerald Sim.[10] New characters include Alexander Armstrong as Adam fforbes-Hamilton, Audrey's nephew-by-marriage and godson; Alan David as Emmeridge, Audrey and Richard's outspoken butler; and Michael Cochrane as Archie Pennington-Booth, a neighbouring landowner.[10][12] While the Rector makes only a brief appearance, the other three original characters are main characters throughout and are credited in the opening credits.[16]

text:
Regular characters in the original series

Actor
Character

Penelope Keith
Audrey fforbes-Hamilton

Peter Bowles
Richard DeVere (Bedrich Polouvicek)

Angela Thorne
Marjory Frobisher

Daphne Heard
Mrs. Maria Polouvicka (Mrs. Poo)

John Rudling
Brabinger (the Butler)

Michael Bilton
Old Ned

Gerald Sim
The Rector

Daphne Oxenford
Mrs. Patterson

text:
Recurring characters in the original series

Actor
Character

Jonathan Elsom
J.J. Anderson

Dennis Ramsden
Arnold Plunkett

Betty Tucker
Mrs. Beecham

Jayne Lester
Linda Cartwright

Nicholas McArdle
Miller

Joyce Windsor
Mrs. Dorothy Plunkett

Anthony Sharp
Brigadier Lemington OBE

Celia Imrie
Polly

John Gleeson
Mr. Purvis

Ben Aris
Mr. Spalding

text:
Characters in the 2007 Christmas special

Actor
Character

Penelope Keith
Audrey DeVere

Peter Bowles
Richard DeVere

Angela Thorne
Marjory Frobisher

Gerald Sim
The Rector

Alexander Armstrong
Adam fforbes-Hamilton

Alan David
Emmeridge

Michael Cochrane
Archie Pennington-Booth

text: Plot[edit]

text: Original series[edit]

text: To the Manor Born is set in the fictional village of Grantleigh, in Somerset near the fictional town of Marlbury. The series begins with the funeral of Marton (sic) fforbes-Hamilton, the Lord of the Manor of Grantleigh.[17] Audrey, his widow (and apparently also his cousin), is far from distraught, as she now has control of the Grantleigh estate, which her family the fforbes-Hamiltons have controlled for 400 years.[6][18] Her joy is short-lived though, as her solicitor Arnold Plunkett informs her that Marton was bankrupt and that the manor will have to be sold to pay off the debts.[5][6] Audrey tries to buy back the manor at auction, but is outbid.[5] The new Lord of the Manor is Richard DeVere, a recently widowed self-made millionaire who started his career on an East End fruit barrow and founded the "Cavendish Foods" supermarket chain, who to Audrey is simply a "grocer".[6][19] DeVere brings his domineering mother, who is soon nicknamed by Audrey 'Mrs. Poo', her full name being Maria Jaroslava Vladimira Martinka Polouvicka.[16] She reveals to Audrey, who does not like foreigners, that she and her son came to Britain in 1939 from Czechoslovakia, and that Richard, whose real name is Bedrich Polouvicka, is half-Polish (on his late father Lazlo's side) and half-Czechoslovakian.[6][12][16][20] Mrs. Polouvicka tries hard to fit into British country life, but her accent often leads to many mispronunciations and she frequently comes out with sayings from her "old Czechoslovakia"[21] (even though Czechoslovakia is not so old for her; she was 15 years old when the country was created in 1919). From early on in the series, she encourages her son to propose to Audrey.[5][22]

text: Audrey moves into the "Old Lodge", at the end of the drive, where she can see most of what goes on at the manor. Living with her is her elderly and loyal butler, Brabinger, who has worked for the fforbes-Hamiltons his whole life, and her beagle Bertie. Audrey's supportive best friend, and a frequent visitor, is the well-meaning Marjory Frobisher, who quickly develops a crush on DeVere.[21] Marjory, who was at school with Audrey, is still a schoolgirl at heart and a committed countryside campaigner.[5][21] Marjory has no organisational ability, which leads to her being nicknamed "Muddlesome Marj" by some.[23] Despite Marjory's encouragement, Audrey refuses to get a job, saying her only job could be running the Grantleigh estate and doing other public service, which includes her being a magistrate.[22][23] This means that money is a constant problem.[22] During the second series, Brabinger is away ill; and Ned, whose tied cottage is being renovated by Richard, takes over as butler to Audrey. Ned, a gardener and "outside man", has worked on the estate all his life and finds working indoors difficult to get used to.[21] Other estate staff include the estate foreman Mr. Miller, the estate manager Mr. Spalding and the cook Mrs. Beecham.[5] Another character, Polly, originally works at the local "Cavendish Foods" store and then at the doctors' surgery, while Linda Cartwright works in the stables and as a domestic.[19][22][24][25] The postman, Mr. Purvis, appears in two episodes.[22][25]

text: Other characters include the local rector, who is not unhappy when Audrey loses the manor, hoping that it will prevent the dominating Audrey from having a say in the running of the Church and local causes.[21] The rector favours Richard, who is frequently generous when it comes to donations to church funds.[21] The village shop is run by Mrs. Patterson, the local gossip who also favours Richard and does not get on with Audrey.[21] The typically English Brigadier Lemington, of the Somerset Rifles, is another local landowner and friend of Audrey's and the Master of Foxhounds and, like Audrey and Marjory, has an interest in nature conservation.[21][23] Arnold Plunkett and his wife Dorothy are friends of Audrey's, while Arnold is also the family solicitor.[26] Grantleigh's local estate agent is J.J. Anderson, of "Anderson and Fish", who appears at manor parties.[26]

text: During the course of the three series, Audrey and Richard grow closer. Hostile to each other at the start, they grew to understand and accept each other, as both try to adapt to each other's ways.[6] The last two 1981 episodes show Richard having problems at Cavendish. With his board of directors refusing to let Richard buy a refrigerated plant in Argentina, he seriously considers selling the manor to raise the money to buy the plant himself. Trying to help Richard, Audrey asks her uncle, a well-respected and connected member of the financial community, to lend Richard a hand. Thanks to him, things begin to go Richard's way. Unfortunately, Audrey's uncle dies before the deals are signed. Richard decides to sell the manor to pay for the refrigerated plant. By a twist of fate, Audrey inherits her uncle's fortune and buys the manor back. Now back home and on her own turf, Audrey asks Richard to marry her. Taken aback, Richard says yes and they are married in the final episode, broadcast on 15 October.

text: Based on facts presented in the series, the characters of Audrey fforbes-Hamilton and her first husband Marton fforbes-Hamilton appear to have been distant relations who married each other. Grantleigh Manor has belonged to the fforbes-Hamiltons for "400 years" (series one, episode one, and repeated references in numerous other episodes), and Audrey makes repeated references to having grown up on there (series two, episode five, among others).

text: 2007 Christmas special[edit]

text: As Richard and Audrey plan to celebrate their silver wedding anniversary, each planning a surprise party for the other, Richard confesses to Audrey that he owns "Farmer Tom", a company that has been putting the neighbouring estates out of business. Audrey leaves him. Richard visits his mother's grave to develop a plan to win Audrey back. He decides to allow a rock concert to be held on the estate. When the licence application is up before the Magistrates' Court, Richard changes his mind. The following day, their anniversary, Marjory leads Audrey to a surprise party organised by Richard.

text: New characters in the 2007 Christmas special include Emmeridge, the outspoken butler replacing Brabinger;[11][16] Adam fforbes-Hamilton, Marton's nephew and Audrey's godson, who has recently moved to Grantleigh to learn how to run an estate; and Archie Pennington-Booth, a neighbouring landowner driven to bankruptcy by "Farmer Tom".[16] Marjory lives in the Old Lodge, and she develops a crush on Adam.[16] A small acknowledgement of Bertie and Brabinger appears in the special, consisting of a photo of Brabinger and a small beagle statuette on a side table.[16] Richard's mother, aka "Mrs. Poo," is acknowledged by a marble headstone in the church graveyard and a framed photograph on the mantel of the manor's sitting room.[27]

text: Episodes[edit]

text: To the Manor Born ran for three series and one Christmas special from 30 September 1979 to 29 November 1981. Each episode is thirty minutes long. The first series aired for seven episodes on Sundays at 8.45 pm, the second series for six episodes on Sundays at 8.35 pm and the seven-episode Series Three on Sundays at 7.15 pm.[3] The Christmas special aired at 8.00 pm. All episodes aired on BBC1.[3] A second Christmas special, announced on 24 October 2007, aired at 9.30 pm on BBC One on 25 December 2007.[10][28][29]

text: Several episodes received extremely high audience figures, especially the first series. This was partly due to the fact that the main rival channel to BBC One channel in the UK, ITV, was in the middle of a long protracted strike which started in early August 1979, which saw nearly all of the local ITV companies go off air and no programmes being aired from the network (except Channel Television in the Channel Island), leaving only BBC One and BBC Two as the channels left for the British public to watch until Wednesday 24th October 1979 when ITV returned to air, with the strike ending in a pay agreement. However many viewers remained with the BBC and shows such as To The Manor Born, as ITV took some months to get going again with new programmes. For the first month after the strike had ended, ITV found it hard to win back viewers who preferred to remain with the BBC, as ITV was airing repeats and filler programmes until new content was made from November 1979. This gave To The Manor Born a huge advantage with the viewing public.[30][31][32] In 1979, the last episode of the first series received 23.95 million viewers, the fourth-highest figures for any programme in the UK in the 1970s, and the highest for a non-live event.[33] The following year, 21.55 million people watched the last episode of series two, the fifth-highest viewing figure for the 1980s.[34] The final episode in 1981, when Audrey and Richard marry, received 17.80 million viewers.[13][35] The 2007 Christmas special garnered 10.25 million viewers and ranked as the 6th most watched programme in the UK for that week.[36]

text: The theme tune for To the Manor Born was composed by Ronnie Hazlehurst, the BBC's Light Entertainment Music Director who composed the theme tunes for many sitcoms around this time.[37]

text: Reception[edit]

text: 2007 Christmas special[edit]

text: Writing in the Sunday Express on 30 December 2007, Marshall Julius described the 2007 Christmas special as "so cosy and old-fashioned that I could easily have dismissed it with a cynical wave". However, he says he found himself enjoying it "about halfway through" and said "it was a real pleasure to see Peter Bowles and Penelope Keith, for both of whom I feel great affection, once again sparring on the small screen". Julius finished his review by commenting "Not that I'm saying I'd like a whole series of it but as a one-off event it was surprisingly welcome".[38]

text: The British Comedy Guide was more critical saying that "whilst it gained great viewing figures it really wasn't a patch on the original episodes."[39] This opinion was reflected in the voting for The Comedy.co.uk Awards 2007, in which the show picked up the title "The Worst British TV Sitcom Special 2007".[40]

text: In other media[edit]

text: Novelisations[edit]

text: The writer Peter Spence wrote two books that accompanied To the Manor Born. While they were based on the TV episodes, both books did differ with added conversations, changed storylines and the characters being given different personality traits. The first book, published in 1979 by Arrow Books, is entitled To the Manor Born and is based on the first series episodes 1, 2, 4, 6 and 7. In this book, Brabinger's first name, which is never mentioned in the TV series, is revealed as Bertram and Ned's surname revealed as Peaslake. The second book, titled To the Manor Born Book 2, was published by the same publisher in 1980. This is a novelisation of the 1979 Christmas special and episodes 5, 4, 3, 2 and 6 from the second series, plus an original story.

text: Film version[edit]

text: A film version was announced by the Rank Organisation at the Cannes Film Festival in 1980 but then was abandoned when Rank pulled out of filmmaking.[41]

text: Radio version[edit]

text: In 1997, 16 years after the original series of To the Manor Born ended on television, ten 30-minute episodes were recorded for radio, the programme's original intended medium.[5] The episodes, originally aired on Saturdays on BBC Radio 2 from 25 January to 29 March 1997, were written by Peter Spence, six adapted from TV episodes and four new episodes.[5]. Keith and Thorne returned as Audrey and Marjory, while Keith Barron replaced Bowles as Richard. Other actors, including Frank Middlemass and Nicholas McArdle, who played Miller in the TV version, replaced many of the by then deceased original television cast members. The series has subsequently been repeated on BBC 7.[42]

text: Radio cast[edit]

text:
Actor
Character/s

Penelope Keith
Audrey fforbes-Hamilton

Keith Barron
Richard DeVere (Bedrich Polouvicek)

Angela Thorne
Marjory Frobisher

Margery Withers
Mrs. Maria Polouvicka (Mrs Poo)

Nicholas McArdle
Brabinger

Frank Middlemass
Ned/Arnold Plunkett

Zulema Dene
Mrs. Plunkett

Jonathan Adams
Brigadier Lemington, OBE

text: Radio episodes[edit]

text:
#
Title
Original airdate

1
"Rhythms of the Earth"
25 January 1997 (1997-01-25)

Audrey is unhappy that Richard has not attended the Sunday church service, and goes to the Manor to tell him of his duties as Lord of the Manor, which include reading the lesson. He promises to attend church the following Sunday and read one lesson, while she will read the other. However, she fails to turn up because the clocks have gone forward.

2
"The Grapevine"
1 February 1997 (1997-02-01)

At a party at the Manor, Audrey tells Richard that she is going on holiday to Spain. However, she actually spends her holiday hiding at the Lodge. During her "holiday" she finds out that Richard and Marjory have been spending time in the woods behind the Lodge. Unknown to her, they are badger watching but when Audrey confronts him at a post-holiday party, he tells her he knows where she spent her holiday.[citation needed]

3
"What's in a Name"
8 February 1997 (1997-02-08)

Audrey is having problems with her heating, and Old Ned is trying to fix it but keeps needing new parts. Meanwhile, Mrs. Poo is bored and enters a "Cavendish Foods" competition, and upon Richard's insistence does so under an assumed name. She wins the competition, to Audrey's good fortune.[citation needed]

4
"Vive Le Sport"
15 February 1997 (1997-02-15)

Audrey believes she is the only healthy person on the estate, but then hurts her back and gets little help from Dr. Horton (Geoffrey Whitehead). Audrey enjoys Richard coming round to see her, and pretends her back is worse than it is to ensure the visits continue. However, when he asks her to go skiing with him, she makes a quick recovery only for Richard to break his leg while practising skiing.[citation needed][citation needed]

5
"Sons of the Fathers"
22 February 1997 (1997-02-22)

Audrey, a local magistrate, encourages Richard to join the Marlbury bench as they are short of magistrates. Richard meanwhile tries to persuade her to go to the race course with him. However, she refuses as she wants to spend time looking for any close relatives as she is worried about being the last of the fforbes-Hamilton line.[citation needed]

6
"A Wife's Prerogative"
1 March 1997 (1997-03-01)

When Richard gets in a difficult situation with a female business associate Mlle Dutoit (Rula Lenska), he tells her he is married. However, Mlle Dutoit then arranges to visit the Manor, and Richard persuades Audrey to pretend to be his wife for the weekend. Audrey uses the situation to move a piece of modern art she dislikes. The deception goes well until Mrs. Poo returns home early from London.[citation needed][citation needed]

7
"The Spare Room"
8 March 1997 (1997-03-08)

An old friend of Audrey and Marjory, Podge Hodge (Zulema Dene), comes to visit, Audrey has to ask Richard to provide a bedroom as the Lodge has a leaking roof. Podge and Richard soon start getting on well, to Audrey and Marjory's horror, and when they spend the whole day together after going hunting, Audrey and Marjory ensure Podge does not spend a second night at the Manor.[citation needed][citation needed]

8
"An Englishman's Home"
15 March 1997 (1997-03-15)

Richard has a security system installed at the Manor and Audrey is concerned that the CCTV will allow Richard to see into her bedroom. Meanwhile, the estate workers go on strike, fearing they are being watched, and Mrs. Poo has problems with the system, causing the Police to be called to the Manor frequently.[citation needed]

9
"The Honours List"
22 March 1997 (1997-03-22)

Audrey lectures Richard for not attending Lord Allenshaw's (Frank Middlemass) lecture on nature conservation. She then attacks him for his proposed destruction of the estate's hedgerows. However, when Audrey hears that a local person is to receive an honour, she believes it will be Richard, and she changes her mind and agrees to host his New Year party. However, it is the Brigadier who gets an honour, being made an OBE.[citation needed]

10
"Plenty More Fish"
29 March 1997 (1997-03-29)

Audrey and Marjory fall out when Audrey develops a close friendship with Commander Jeremy French-Liversey (Doyne Byrd) and does not help organise the county show. Richard gets jealous, and he finds out the Commander has a "roving eye" for women. Meanwhile, Mrs. Poo and Ned get stuck on a runaway traction engine.[citation needed][citation needed]

text: DVD releases[edit]

text: All three series, including the 1979 and 2007 Christmas specials, of To the Manor Born have been released in the UK (Region 2). The first series and the 1979 Christmas special were released on 5 May 2003, followed by Series Two on 29 December 2003 and Series Three on 14 June 2004.[43] The DVDs feature an interview with Peter Spence.[43] The 2007 Christmas special was released on Region 2 DVD on 11 February 2008.[44]

text: The complete original series was released in a boxset in the United States and Canada (Region 1) on 15 June 2004.[45]


document_title: Hypothalamus

question_text: what part of the brain controls the endocrine system

root

text: Hypothalamus

text: Structure[edit]

text: The hypothalamus is a brain structure made up of distinct nuclei as well as less anatomically distinct areas. It is found in all vertebrate nervous systems. In mammals, magnocellular neurosecretory cells in the paraventricular nucleus and the supraoptic nucleus of the hypothalamus produce neurohypophysial hormones, oxytocin and vasopressin. These hormones are released into the blood in the posterior pituitary.[3] Much smaller parvocellular neurosecretory cells, neurons of the paraventricular nucleus, release corticotropin-releasing hormone and other hormones into the hypophyseal portal system, where these hormones diffuse to the anterior pituitary.

tag: p

text: Nuclei[edit]

text: The hypothalamic nuclei include the following:[4][5][6]

text:
Region
Area
Nucleus
Function[7]

Anterior
Preoptic
Preoptic nucleus

Medial
Medial preoptic nucleus

Regulates the release of gonadotropic hormones from the adenohypophysis
Contains the sexually dimorphic nucleus, which releases GnRH, differential development between sexes is based upon in utero testosterone levels
Thermoregulation[8]

Supraoptic nucleus

Vasopressin release
Oxytocin release

Paraventricular nucleus

thyrotropin-releasing hormone release
corticotropin-releasing hormone release
oxytocin release
vasopressin release
somatostatin release

Anterior hypothalamic nucleus

thermoregulation
panting
sweating
thyrotropin inhibition

Suprachiasmatic nucleus

Circadian rhythms

Lateral

Lateral nucleus
See Lateral hypothalamus § Function – primary source of orexin neurons that project throughout the brain and spinal cord

Tuberal
Medial
Dorsomedial hypothalamic nucleus

Blood Pressure
Heart Rate
GI stimulation

Ventromedial nucleus

satiety
neuroendocrine control

Arcuate nucleus

Growth hormone-releasing hormone (GHRH)
feeding
Dopamine-mediated prolactin inhibition

Lateral
Lateral nucleus
See Lateral hypothalamus § Function – primary source of orexin neurons that project throughout the brain and spinal cord

Lateral tuberal nuclei

Posterior
Medial
Mammillary nuclei (part of mammillary bodies)

memory

Posterior nucleus

Increase blood pressure
pupillary dilation
shivering
vasopressin release

Lateral
Lateral nucleus
See Lateral hypothalamus § Function – primary source of orexin neurons that project throughout the brain and spinal cord

Tuberomammillary nucleus[9]

arousal (wakefulness and attention)
feeding and energy balance
learning
memory
sleep

tag: table

text: See also: ventrolateral preoptic nucleus, periventricular nucleus.

text:
A cross section of the monkey hypothalamus displays 2 of the major hypothalamic nuclei on either side of the fluid-filled 3rd ventricle.

Hypothalamic nuclei

Hypothalamic nuclei on one side of the hypothalamus, shown in a 3-D computer reconstruction[10]

tag: ul

text: Neural connections[edit]

text: The hypothalamus is highly interconnected with other parts of the central nervous system, in particular the brainstem and its reticular formation. As part of the limbic system, it has connections to other limbic structures including the amygdala and septum, and is also connected with areas of the autonomous nervous system.

text: The hypothalamus receives many inputs from the brainstem, the most notable from the nucleus of the solitary tract, the locus coeruleus, and the ventrolateral medulla.

text: Most nerve fibres within the hypothalamus run in two ways (bidirectional).

text:
Projections to areas caudal to the hypothalamus go through the medial forebrain bundle, the mammillotegmental tract and the dorsal longitudinal fasciculus.
Projections to areas rostral to the hypothalamus are carried by the mammillothalamic tract, the fornix and terminal stria.
Projections to areas of the sympathetic motor system (lateral horn spinal segments T1-L2/L3) are carried by the hypothalamospinal tract and they activate the sympathetic motor pathway.

text: Sexual dimorphism[edit]

text: Several hypothalamic nuclei are sexually dimorphic; i.e., there are clear differences in both structure and function between males and females.[citation needed]

text: Some differences are apparent even in gross neuroanatomy: most notable is the sexually dimorphic nucleus within the preoptic area. However most of the differences are subtle changes in the connectivity and chemical sensitivity of particular sets of neurons.[citation needed]

text: The importance of these changes can be recognized by functional differences between males and females. For instance, males of most species prefer the odor and appearance of females over males, which is instrumental in stimulating male sexual behavior. If the sexually dimorphic nucleus is lesioned, this preference for females by males diminishes. Also, the pattern of secretion of growth hormone is sexually dimorphic, and this is one reason why in many species, adult males are much larger than females.[citation needed]

text: Responsiveness to ovarian steroids[edit]

text: Other striking functional dimorphisms are in the behavioral responses to ovarian steroids of the adult. Males and females respond to ovarian steroids in different ways, partly because the expression of estrogen-sensitive neurons in the hypothalamus is sexually dimorphic; i.e., estrogen receptors are expressed in different sets of neurons.[citation needed]

text: Estrogen and progesterone can influence gene expression in particular neurons or induce changes in cell membrane potential and kinase activation, leading to diverse non-genomic cellular functions. Estrogen and progesterone bind to their cognate nuclear hormone receptors, which translocate to the cell nucleus and interact with regions of DNA known as hormone response elements (HREs) or get tethered to another transcription factor's binding site. Estrogen receptor (ER) has been shown to transactivate other transcription factors in this manner, despite the absence of an estrogen response element (ERE) in the proximal promoter region of the gene. In general, ERs and progesterone receptors (PRs) are gene activators, with increased mRNA and subsequent protein synthesis following hormone exposure.[citation needed]

text: Male and female brains differ in the distribution of estrogen receptors, and this difference is an irreversible consequence of neonatal steroid exposure. Estrogen receptors (and progesterone receptors) are found mainly in neurons in the anterior and mediobasal hypothalamus, notably:[citation needed]

text:
the preoptic area (where LHRH neurons are located)
the periventricular nucleus (where somatostatin neurons are located)
the ventromedial hypothalamus (which is important for sexual behavior).

text: Development[edit]

text: In neonatal life, gonadal steroids influence the development of the neuroendocrine hypothalamus. For instance, they determine the ability of females to exhibit a normal reproductive cycle, and of males and females to display appropriate reproductive behaviors in adult life.[citation needed]

text:
If a female rat is injected once with testosterone in the first few days of postnatal life (during the "critical period" of sex-steroid influence), the hypothalamus is irreversibly masculinized; the adult rat will be incapable of generating an LH surge in response to estrogen (a characteristic of females), but will be capable of exhibiting male sexual behaviors (mounting a sexually receptive female).[citation needed]
By contrast, a male rat castrated just after birth will be feminized, and the adult will show female sexual behavior in response to estrogen (sexual receptivity, lordosis behavior).[citation needed]

text: In primates, the developmental influence of androgens is less clear, and the consequences are less understood. Within the brain, testosterone is aromatized to (estradiol), which is the principal active hormone for developmental influences. The human testis secretes high levels of testosterone from about week 8 of fetal life until 5–6 months after birth (a similar perinatal surge in testosterone is observed in many species), a process that appears to underlie the male phenotype. Estrogen from the maternal circulation is relatively ineffective, partly because of the high circulating levels of steroid-binding proteins in pregnancy.[citation needed]

text: Sex steroids are not the only important influences upon hypothalamic development; in particular, pre-pubertal stress in early life (of rats) determines the capacity of the adult hypothalamus to respond to an acute stressor.[11] Unlike gonadal steroid receptors, glucocorticoid receptors are very widespread throughout the brain; in the paraventricular nucleus, they mediate negative feedback control of CRF synthesis and secretion, but elsewhere their role is not well understood.

text: Function[edit]

text: Hormone release[edit]

text: The hypothalamus has a central neuroendocrine function, most notably by its control of the anterior pituitary, which in turn regulates various endocrine glands and organs. Releasing hormones (also called releasing factors) are produced in hypothalamic nuclei then transported along axons to either the median eminence or the posterior pituitary, where they are stored and released as needed.[12]

text: In the hypothalamic–adenohypophyseal axis, releasing hormones, also known as hypophysiotropic or hypothalamic hormones, are released from the median eminence, a prolongation of the hypothalamus, into the hypophyseal portal system, which carries them to the anterior pituitary where they exert their regulatory functions on the secretion of adenohypophyseal hormones.[13] These hypophysiotropic hormones are stimulated by parvocellular neurosecretory cells located in the periventricular area of the hypothalamus. After their release into the capillaries of the third ventricle, the hypophysiotropic hormones travel through what is known as the hypothalamo-pituitary portal circulation. Once they reach their destination in the anterior pituitary, these hormones bind to specific receptors located on the surface of pituitary cells. Depending on which cells are activated through this binding, the pituitary will either begin secreting or stop secreting hormones into the rest of the bloodstream. (Bear, Mark F. "Hypothalamic Control of the Anterior Pituitary." Neuroscience: Exploring the Brain. 4th ed. Philadelphia: Wolters Kluwer, 2016. 528. Print.)

text:
Secreted hormone
Abbreviation
Produced by
Effect

Thyrotropin-releasing hormone
(Prolactin-releasing hormone)
TRH, TRF, or PRH
Parvocellular neurosecretory cells of the paraventricular nucleus
Stimulate thyroid-stimulating hormone (TSH) release from anterior pituitary (primarily)
Stimulate prolactin release from anterior pituitary

Corticotropin-releasing hormone
CRH or CRF
Parvocellular neurosecretory cells of the paraventricular nucleus
Stimulate adrenocorticotropic hormone (ACTH) release from anterior pituitary

Dopamine
(Prolactin-inhibiting hormone)
DA or PIH
Dopamine neurons of the arcuate nucleus
Inhibit prolactin release from anterior pituitary

Growth-hormone-releasing hormone
GHRH
Neuroendocrine neurons of the Arcuate nucleus
Stimulate growth-hormone (GH) release from anterior pituitary

Gonadotropin-releasing hormone
GnRH or LHRH
Neuroendocrine cells of the Preoptic area
Stimulate follicle-stimulating hormone (FSH) release from anterior pituitary
Stimulate luteinizing hormone (LH) release from anterior pituitary

Somatostatin[14]
(growth-hormone-inhibiting hormone)
SS, GHIH, or SRIF
Neuroendocrine cells of the Periventricular nucleus
Inhibit growth-hormone (GH) release from anterior pituitary
Inhibit (moderately) thyroid-stimulating hormone (TSH) release from anterior pituitary

text: Other hormones secreted from the median eminence include vasopressin, oxytocin, and neurotensin.[15][16][17][18]

text: In the hypothalamic-neurohypophyseal axis, neurohypophysial hormones are released from the posterior pituitary, which is actually a prolongation of the hypothalamus, into the circulation.

text:
Secreted hormone
Abbreviation
Produced by
Effect

Oxytocin
OXY or OXT
Magnocellular neurosecretory cells of the paraventricular nucleus and supraoptic nucleus
Uterine contraction
Lactation (letdown reflex)

Vasopressin
(antidiuretic hormone)
ADH or AVP
Magnocellular and parvocellular neurosecretory cells of the paraventricular nucleus, magnocellular cells in supraoptic nucleus
Increase in the permeability to water of the cells of distal tubule and collecting duct in the kidney and thus allows water reabsorption and excretion of concentrated urine

text: It is also known that hypothalamic-pituitary-adrenal axis (HPA) hormones are related to certain skin diseases and skin homeostasis. There is evidence linking hyperactivity of HPA hormones to stress-related skin diseases and skin tumors.[19]

text: Stimulation[edit]

text: The hypothalamus coordinates many hormonal and behavioural circadian rhythms, complex patterns of neuroendocrine outputs, complex homeostatic mechanisms, and important behaviours. The hypothalamus must, therefore, respond to many different signals, some of which generated externally and some internally. Delta wave signalling arising either in the thalamus or in the cortex influences the secretion of releasing hormones; GHRH and prolactin are stimulated whilst TRH is inhibited.

text: The hypothalamus is responsive to:

text:
Light: daylength and photoperiod for regulating circadian and seasonal rhythms
Olfactory stimuli, including pheromones
Steroids, including gonadal steroids and corticosteroids
Neurally transmitted information arising in particular from the heart, the stomach, and the reproductive tract[citation needed]
Autonomic inputs
Blood-borne stimuli, including leptin, ghrelin, angiotensin, insulin, pituitary hormones, cytokines, plasma concentrations of glucose and osmolarity etc.
Stress
Invading microorganisms by increasing body temperature, resetting the body's thermostat upward.

text: Olfactory stimuli[edit]

text: Olfactory stimuli are important for sexual reproduction and neuroendocrine function in many species. For instance if a pregnant mouse is exposed to the urine of a 'strange' male during a critical period after coitus then the pregnancy fails (the Bruce effect). Thus, during coitus, a female mouse forms a precise 'olfactory memory' of her partner that persists for several days. Pheromonal cues aid synchronization of oestrus in many species; in women, synchronized menstruation may also arise from pheromonal cues, although the role of pheromones in humans is disputed.

text: Blood-borne stimuli[edit]

text: Peptide hormones have important influences upon the hypothalamus, and to do so they must pass through the blood–brain barrier. The hypothalamus is bounded in part by specialized brain regions that lack an effective blood–brain barrier; the capillary endothelium at these sites is fenestrated to allow free passage of even large proteins and other molecules. Some of these sites are the sites of neurosecretion - the neurohypophysis and the median eminence. However, others are sites at which the brain samples the composition of the blood. Two of these sites, the SFO (subfornical organ) and the OVLT (organum vasculosum of the lamina terminalis) are so-called circumventricular organs, where neurons are in intimate contact with both blood and CSF. These structures are densely vascularized, and contain osmoreceptive and sodium-receptive neurons that control drinking, vasopressin release, sodium excretion, and sodium appetite. They also contain neurons with receptors for angiotensin, atrial natriuretic factor, endothelin and relaxin, each of which important in the regulation of fluid and electrolyte balance. Neurons in the OVLT and SFO project to the supraoptic nucleus and paraventricular nucleus, and also to preoptic hypothalamic areas. The circumventricular organs may also be the site of action of interleukins to elicit both fever and ACTH secretion, via effects on paraventricular neurons.[citation needed]

text: It is not clear how all peptides that influence hypothalamic activity gain the necessary access. In the case of prolactin and leptin, there is evidence of active uptake at the choroid plexus from the blood into the cerebrospinal fluid (CSF). Some pituitary hormones have a negative feedback influence upon hypothalamic secretion; for example, growth hormone feeds back on the hypothalamus, but how it enters the brain is not clear. There is also evidence for central actions of prolactin.[citation needed]

text: Findings have suggested that thyroid hormone (T4) is taken up by the hypothalamic glial cells in the infundibular nucleus/ median eminence, and that it is here converted into T3 by the type 2 deiodinase (D2). Subsequent to this, T3 is transported into the thyrotropin-releasing hormone (TRH)-producing neurons in the paraventricular nucleus. Thyroid hormone receptors have been found in these neurons, indicating that they are indeed sensitive to T3 stimuli. In addition, these neurons expressed MCT8, a thyroid hormone transporter, supporting the theory that T3 is transported into them. T3 could then bind to the thyroid hormone receptor in these neurons and affect the production of thyrotropin-releasing hormone, thereby regulating thyroid hormone production.[20]

text: The hypothalamus functions as a type of thermostat for the body.[21] It sets a desired body temperature, and stimulates either heat production and retention to raise the blood temperature to a higher setting or sweating and vasodilation to cool the blood to a lower temperature. All fevers result from a raised setting in the hypothalamus; elevated body temperatures due to any other cause are classified as hyperthermia.[21] Rarely, direct damage to the hypothalamus, such as from a stroke, will cause a fever; this is sometimes called a hypothalamic fever. However, it is more common for such damage to cause abnormally low body temperatures.[21]

text: Steroids[edit]

text: The hypothalamus contains neurons that react strongly to steroids and glucocorticoids – (the steroid hormones of the adrenal gland, released in response to ACTH). It also contains specialized glucose-sensitive neurons (in the arcuate nucleus and ventromedial hypothalamus), which are important for appetite. The preoptic area contains thermosensitive neurons; these are important for TRH secretion.

text: Neural[edit]

text: Oxytocin secretion in response to suckling or vagino-cervical stimulation is mediated by some of these pathways; vasopressin secretion in response to cardiovascular stimuli arising from chemoreceptors in the carotid body and aortic arch, and from low-pressure atrial volume receptors, is mediated by others. In the rat, stimulation of the vagina also causes prolactin secretion, and this results in pseudo-pregnancy following an infertile mating. In the rabbit, coitus elicits reflex ovulation. In the sheep, cervical stimulation in the presence of high levels of estrogen can induce maternal behavior in a virgin ewe. These effects are all mediated by the hypothalamus, and the information is carried mainly by spinal pathways that relay in the brainstem. Stimulation of the nipples stimulates release of oxytocin and prolactin and suppresses the release of LH and FSH.

text: Cardiovascular stimuli are carried by the vagus nerve. The vagus also conveys a variety of visceral information, including for instance signals arising from gastric distension or emptying, to suppress or promote feeding, by signalling the release of leptin or gastrin, respectively. Again this information reaches the hypothalamus via relays in the brainstem.

text: In addition hypothalamic function is responsive to—and regulated by—levels of all three classical monoamine neurotransmitters, noradrenaline, dopamine, and serotonin (5-hydroxytryptamine), in those tracts from which it receives innervation. For example, noradrenergic inputs arising from the locus coeruleus have important regulatory effects upon corticotropin-releasing hormone (CRH) levels.

text: Control of food intake[edit]

text:
Peptide hormones and neuropeptides that regulate feeding[22]

Peptides that increase
feeding behavior
Peptides that decrease
feeding behavior

Ghrelin
Leptin

Neuropeptide Y
(α,β,γ)-Melanocyte-stimulating hormones

Agouti-related peptide
Cocaine- and amphetamine-regulated transcript peptides

Orexins (A,B)
Corticotropin-releasing hormone

Melanin-concentrating hormone
Cholecystokinin

Galanin
Insulin

Glucagon-like peptide 1

text: The extreme lateral part of the ventromedial nucleus of the hypothalamus is responsible for the control of food intake. Stimulation of this area causes increased food intake. Bilateral lesion of this area causes complete cessation of food intake. Medial parts of the nucleus have a controlling effect on the lateral part. Bilateral lesion of the medial part of the ventromedial nucleus causes hyperphagia and obesity of the animal. Further lesion of the lateral part of the ventromedial nucleus in the same animal produces complete cessation of food intake.

text: There are different hypotheses related to this regulation:[23]

text:
Lipostatic hypothesis: This hypothesis holds that adipose tissue produces a humoral signal that is proportionate to the amount of fat and acts on the hypothalamus to decrease food intake and increase energy output. It has been evident that a hormone leptin acts on the hypothalamus to decrease food intake and increase energy output.
Gutpeptide hypothesis: gastrointestinal hormones like Grp, glucagons, CCK and others claimed to inhibit food intake. The food entering the gastrointestinal tract triggers the release of these hormones, which act on the brain to produce satiety. The brain contains both CCK-A and CCK-B receptors.
Glucostatic hypothesis: The activity of the satiety center in the ventromedial nuclei is probably governed by the glucose utilization in the neurons. It has been postulated that when their glucose utilization is low and consequently when the arteriovenous blood glucose difference across them is low, the activity across the neurons decrease. Under these conditions, the activity of the feeding center is unchecked and the individual feels hungry. Food intake is rapidly increased by intraventricular administration of 2-deoxyglucose therefore decreasing glucose utilization in cells.
Thermostatic hypothesis: According to this hypothesis, a decrease in body temperature below a given set-point stimulates appetite, whereas an increase above the set-point inhibits appetite.

tag: ol

text: Fear processing[edit]

text: The medial zone of hypothalamus is part of a circuitry that controls motivated behaviors, like defensive behaviors.[24] Analyses of Fos-labeling showed that a series of nuclei in the "behavioral control column" is important in regulating the expression of innate and conditioned defensive behaviors.[25]

text: Exposure to a predator (such as a cat) elicits defensive behaviors in laboratory rodents, even when the animal has never been exposed to a cat.[26] In the hypothalamus, this exposure causes an increase in Fos-labeled cells in the anterior hypothalamic nucleus, the dorsomedial part of the ventromedial nucleus, and in the ventrolateral part of the premammillary nucleus (PMDvl).[27] The premammillary nucleus has an important role in expression of defensive behaviors towards a predator, since lesions in this nucleus abolish defensive behaviors, like freezing and flight.[27][28] The PMD does not modulate defensive behavior in other situations, as lesions of this nucleus had minimal effects on post-shock freezing scores.[28] The PMD has important connections to the dorsal periaqueductal gray, an important structure in fear expression.[29][30] In addition, animals display risk assessment behaviors to the environment previously associated with the cat. Fos-labeled cell analysis showed that the PMDvl is the most activated structure in the hypothalamus, and inactivation with muscimol prior to exposure to the context abolishes the defensive behavior.[27] Therefore, the hypothalamus, mainly the PMDvl, has an important role in expression of innate and conditioned defensive behaviors to a predator.

text: Likewise, the hypothalamus has a role in social defeat: Nuclei in medial zone are also mobilized during an encounter with an aggressive conspecific. The defeated animal has an increase in Fos levels in sexually dimorphic structures, such as the medial pre-optic nucleus, the ventrolateral part of ventromedial nucleus, and the ventral premammilary nucleus.[31] Such structures are important in other social behaviors, such as sexual and aggressive behaviors. Moreover, the premammillary nucleus also is mobilized, the dorsomedial part but not the ventrolateral part.[31] Lesions in this nucleus abolish passive defensive behavior, like freezing and the "on-the-back" posture.[31]

text: Sexual orientation[edit]

text: According to D. F. Swaab, writing in a July 2008 paper, "Neurobiological research related to sexual orientation in humans is only just gathering momentum, but the evidence already shows that humans have a vast array of brain differences, not only in relation to gender, but also in relation to sexual orientation."[32]

text: Swaab first reported on the relationship between sexual orientation in males and the hypothalamus's "clock", the suprachiasmatic nucleus (SCN). In 1990, Swaab and Hofman[33] reported that the suprachiasmatic nucleus in homosexual men was significantly larger than in heterosexual men. Then in 1995, Swaab et al.[34] linked brain development to sexual orientation by treating male rats both pre- and postnatally with ATD, an aromatase blocker in the brain. This produced an enlarged SCN and bisexual behavior in the adult male rats. In 1991, LeVay showed that part of the sexually dimorphic nucleus (SDN) known as the 3rd interstitial nucleus of the anterior hypothalamus (INAH 3), is nearly twice as large (in terms of volume) in heterosexual men than in homosexual men and heterosexual women. However, a study in 1992 has shown that the sexually dimorph nucleus of the preoptic area, which include the INAH3, are of similar size in homosexual males who died of AIDS to heterosexual males, and therefore larger than female. This clearly contradicts the hypothesis that homosexual males have a female hypothalamus. Furthermore, the SCN of homosexual males is extremely large (both the volume and the number of neurons are twice as many as in heterosexual males). These areas of the hypothalamus have not yet been explored in homosexual females nor bisexual males nor females. Although the functional implications of such findings still haven't been examined in detail, they cast serious doubt over the widely accepted Dörner hypothesis that homosexual males have a "female hypothalamus" and that the key mechanism of differentiating the "male brain from originally female brain" is the epigenetic influence of testosterone during prenatal development.[35][36]

text: In 2004 and 2006, two studies by Berglund, Lindström, and Savic[37][38] used positron emission tomography (PET) to observe how the hypothalamus responds to smelling common odors, the scent of testosterone found in male sweat, and the scent of estrogen found in female urine. These studies showed that the hypothalamus of heterosexual men and homosexual women both respond to estrogen. Also, the hypothalamus of homosexual men and heterosexual women both respond to testosterone. The hypothalamus of all four groups did not respond to the common odors, which produced a normal olfactory response in the brain.