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Hi All,

Discovery of a hormone related to hunger are in the pdf-available below from

Science, not yet in Medline.

There appears to be amply explanation of the articles in the below popular press

report, reviews and article (1-5).

Two-day fasting is a long time for rats, is it not?

1. Appetite-suppressing hormone identified

Last Updated Thu, 10 Nov 2005 19:16:44 EST

CBC News

Scientists used clues from the Human Genome Project to find a hormone that seems

to

suppress appetite in rats.

Three hormones, leptin, melanocortin and ghrelin, are known to act on appetite

and

weight.

The discovery of each raised interest in weight-control strategies, but

scientists

say dozens of hormones are likely involved. Little is understood about the

stomach

and brain's complex system of regulating weight.

Researchers at Stanford University School of Medicine identified the latest

hormone,

obestatin, by studying human genome data for hormone receptors.

The hunger hormone ghrelin was found to have obestatin attached to it.

The team then found obestatin was present in rat stomach tissues and brains.

When obestatin was injected into rats, it suppressed food intake in the animals

by

about half compared to those not given the hormone, the team reports in the Nov.

11

issue of the journal Science.

Obestatin seems to act by putting a brake on ghrelin, say the researchers, who

found

it also slowed the movement of digested food from the stomach into the

intestines.

The normal rats lost 20 per cent of their weight in eight days. Researchers plan

to

test its effect on obese rats next.

They also need to test if obestatin made the rats eat less by suppressing

appetite

directly or by making the rodents feel ill.

& Pharmaceutical Research & Development, LLC sponsored the

research

into the newly identified hormone. The company has licence rights on the

discovery.

http://www.cbc.ca/story/science/national/2005/11/10/appetite-hormone051110.html

2. Dueling Hunger Hormones?

Science 11 November 2005:?

Perspectives

BIOMEDICINE:

Separation of Conjoined Hormones Yields Appetite Rivals

Nogueiras and Matthias Tschöp

In their Perspective, Nogueiras and Tschöp discuss the study of Zhang et al.

that

has identified obestatin, a new hormone regulator of energy balance. Obestatin

suppresses food intake, body weight gain, and gastrointestinal motility in

rodents.

The paradox is that obestatin is derived from a precursor peptide that also

yields

ghrelin, a hormone that increases appetite and body weight.

3. This Week in Science

Science 11 November 2005:939

Dueling Hunger Hormones?

Ghrelin, a circulating peptide hormone produced in the stomach, has attracted

much

attention because of its stimulatory effect on food intake, but the effect of

ghrelin may represent only half of the story. Using a bioinformatics approach,

Zhang

et al. (p. 996; see the Perspective by Nogueiras and Tschöp) show that ghrelin

encodes a second peptide hormone that is processed from the same protein

precursor

as ghrelin. In rodents, a synthetic version of this hormone, obestatin, has the

opposite physiological effect as ghrelin--it suppresses food intake. Obestatin

mediates its actions through an orphan G protein-coupled receptor, GPR39, which

shares sequences with, but is distinct from, the receptor targeted by ghrelin.

4. Science 11 November 2005: 985-986

Perspectives

BIOMEDICINE:

Separation of Conjoined Hormones Yields Appetite Rivals

Nogueiras and Matthias Tschöp

When we refer to our " gut feelings, " not many of us actually visualize how the

gastrointestinal tract spills myriads of small peptide hormones into our

bloodstream

to activate defined circuits of the central nervous system. Nevertheless, that

picture does reflect a current scientific concept called the " gutbrain axis. "

This

model consists of a complex network of hormonal and neuronal signaling pathways

that

is believed to balance numerous homeostatic and behavioral processes (1, 2). In

this

context, our stomach does not just collect, process, and transport ingested

food,

but it also represents a multileveled conversational partner of the central

nervous

system. A key element of this communication process is the hunger-inducing

hormone

ghrelin, which is believed to convey information about nutrient availability

from

the stomach to the brain (3, 4).

Zhang and colleagues (5) now report on page 996 of this issue that ghrelin not

only

has a sibling derived from the same peptide precursor (preproghrelin), but also

that

this new ghrelin-associated peptide behaves as a physiological opponent of

ghrelin.

Guided by bioinformatics-based predictions for typical enzymatic cleavage sites,

they identified a 23-amino acid region of preproghrelin that is highly conserved

across species, suggesting a relevant biological function. The authors purified

a

secreted peptide of the predicted size and sequence from rat stomach tissue and

also

detected it in rat blood. Similar to ghrelin, which requires posttranslational

modification close to its amino terminus by acylation (6), the biological

activity

of the ghrelin- associated peptide also depends on modification, but by much

more

common amidation at its carboxyl terminus.

The ghrelin-motilin receptor family and their ligands. Each of these

gastrointestinal hormones acts on a specific G protein-coupled receptor from the

same family to affect food intake and gastrointestinal motility (9-11). Similar

dual

effects on satiety and gastrointestinal motility are known for glucagon-like

peptide

1, cholecystokinine, or peptide YY. Collectively, these peptides may serve to

couple

meal termination with inhibition of upper gastrointestinal function to prevent

malabsorption and postprandial metabolic disturbances (1, 2, 8).

The surprising finding is the pharmacological effects of the newly identified

peptide in comparison with the known actions of ghrelin. Whereas ghrelin

increases

food intake and body weight (7), the ghrelin-associated peptide decreases food

intake and body weight gain in rodents. Moreover, Zhang et al. observed that the

new

peptide decelerates gastric emptying and decreases intestinal contractility in

mice,

both of which counteract the well-defined effects of ghrelin (8). Through a

targeted

screen of mammalian orphan receptors and subsequent analyses in cultured

mammalian

cells, Zhang et al. show that the ghrelin-associated peptide binds to and

activates

the orphan receptor GPR39 (9). This G protein-coupled receptor has been mapped

to

human chromosome 2 and is expressed in multiple tissues, including the stomach,

intestine, and hypothalamus. This localization is consistent with a role in

energy

balance regulation (10). GPR39 is a member of a family that includes the

receptors

for ghrelin and motilin, another gastrointestinal hormone that stimulates food

intake, gastric emptying, and gut motility (9, 11). These facts support a

somewhat

counterintuitive, but nevertheless intriguing, relationship between ghrelin and

the

ghrelin-associated peptide.

To denote its anorexigenic actions, Zhang and colleagues named this new gastric

hormone obestatin (from the Latin term obedere, meaning to " devour " ).

Inevitably,

the terms " obesity " and " statins, " a class of lipid-lowering drugs, come to

mind.

However, obestatin has not been tested in animal models of obesity and there is

no

evidence for a lipid-lowering effect. Furthermore, even its effect on body

weight

appears to be very subtle. The failure of obestatin treatment to decrease leptin

levels in mice may indicate lack of lipolytic potency. Effects of obestatin on

food

intake regulation following administration to peripheral circulation or directly

into the brain of mice suggest the typical action profile of a gastrointestinal

satiety hormone. However, it is possible that obestatin may simply suppress

appetite

by triggering nausea or visceral illness. Recent examples have emphasized the

importance of excluding nonspecific appetite suppression when examining

anti-obesity

drug candidates (12). Furthermore, despite sequence homologies between rodent

and

human obestatin (87%) and GPR39 (93%) sequences (5, 9), data from rodents cannot

always be translated to humans, where the effects of obestatin have yet to be

determined.

Another concern regarding a role for obestatin in energy balance regulation

arises

from its quantification in blood. Although Zhang et al. confirmed earlier

findings

that the level of plasma ghrelin increases upon fasting and decreases following

nutrient ingestion (5, 11), they did not observe any changes in circulating

obestatin upon fasting or feeding in rodents. Detection methods for

differentiating

between circulating amidated and nonamidated obestatin are not yet available,

but

could still reveal an association with nutrient availability. Nevertheless,

total

plasma obestatin generally appears to be a fraction of the level of plasma

ghrelin.

Should hormones derived from the same prepropeptide not circulate in an

equimolar

ratio?

The Yin and Yang personalities of ghrelin and obestatin. Both hormones derive

from

the same precursor protein and are predominantly secreted by the stomach and

released into the blood. Each acts on a different receptor (GPR39 and GHS-R, as

shown) and has an opposite effect on food intake, body weight, and

gastrointestinal

motility.

CREDIT: K. SUTLIFF/SCIENCE

Another peptide precursor that gives birth to antipodal regulators of food

intake

may provide some answers. The neuropeptide proopiomelanocortin is cleaved into

several active fragments that include the appetite-suppressing - and

melanocyte-stimulating hormones (-, -MSH) and the appetite-stimulating hormone

-endorphin (13). Tissue-specific enzymes determine which of these are generated.

A

similar scenario could determine how and where preproghrelin is fragmented into

bioactive peptides. An earlier study postulated one other circulating

preproghrelin

fragment, a 13-amino acid peptide called C-ghrelin (14). In addition, turnover

rates

of ghrelin and obestatin may differ appreciably, according to their acylation or

amidation rates, which again would be a parallel to the acetylation of the

proopiomelanocortin derivative -MSH (15). Dissecting the posttranslational

cleavage,

activation, or degradation processes of peptide hormones may reveal elegant

enzymatic drug targets: Simultaneous activation of an agonist and deactivation

of

its endogenous functional antagonist could provide a powerful strategy for

homeostatic control.

If obestatin lives up to its name as a circulating hormone with a

physiologically

relevant anorectic as well as an obesity-preventing function, the puzzling

discrepancy between the very mild phenotype of mice lacking ghrelin (16, 17) and

the

unsurpassed pharmacological effects of ghrelin on energy balance would receive

an

unexpected--but logical--explanation. The absence of an orexigenic hormone may

be

counterbalanced by the simultaneous deletion of an equally potent satiety

factor.

Targeted mouse mutagenesis is widely used as a strategy to unmask or validate

the

biological function of a gene product. An obvious abnormality of such a knockout

mouse is usually interpreted as a reliable indicator of the target's

physiological

role. However, subtle or absent differences between gene-disrupted mice and

their

wild-type littermates are often regarded as evidence of negligible biological

relevance. Such conclusions should be regarded with caution because

developmental

compensation may mask loss of function. However, rarely has such compensation

been

defined on a molecular level. The Zhang et al. findings caution against the

interpretation of results based exclusively on gene disruption or messenger RNA

quantification due to an additional level of complexity represented by

posttranslational processing of proteins.

The discovery of obestatin leaves several questions unanswered. Why does a mouse

that is deficient for the ghrelin receptor not exhibit an impressive phenotype?

Should the absence of ghrelin action in the presence of an intact obestatin

signaling pathway not generate a robust negative energy balance? Why does

obestatin,

unlike ghrelin, not affect growth hormone secretion from the pituitary gland,

despite the presence of the obestatin receptor in this organ? Although the

adversarial relationship between ghrelin and obestatin certainly is an important

contribution to our understanding of body weight regulation, the search for a

magic

bullet against obesity is likely to continue--admittedly, a gut feeling.

5. Obestatin, a Peptide Encoded by the Ghrelin Gene, Opposes Ghrelin's Effects

on

Food Intake

Jian V. Zhang, Pei-Gen Ren, Orna Avsian-Kretchmer, Ching-Wei Luo, Rami Rauch,

Klein, and J. W. Hsueh

Science 11 November 2005: 996-999.

Ghrelin, a circulating appetite-inducing hormone, is derived from a prohormone

by

posttranslational processing. On the basis of the bioinformatic prediction that

another peptide also derived from proghrelin exists, we isolated a hormone from

rat

stomach and named it obestatin—a contraction of obese, from the Latin " obedere, "

meaning to devour, and " statin, " denoting suppression. Contrary to the

appetite-stimulating effects of ghrelin, treatment of rats with obestatin

suppressed

food intake, inhibited jejunal contraction, and decreased body-weight gain.

Obestatin bound to the orphan G protein–coupled receptor GPR39. Thus, two

peptide

hormones with opposing action in weight regulation are derived from the same

ghrelin

gene. After differential modification, these hormones activate distinct

receptors.The increasing prevalence of obesity is a global problem. Body weight

is

regulated in part by peptide hormones produced in the brain or gut or both (1).

Earlier studies on synthetic and peptidyl growth hormone (GH) secretagogues

(2–4)

led to the identification of a specific G protein–coupled receptor (GPCR), the

GH

secretagogue receptor (GHSR) (5, 6), and subsequently to the discovery of its

endogenous ligand, ghrelin (7), a gut-derived circulating hormone that

stimulates

food intake (4, 8).

Human ghrelin, a 28–amino acid peptide, is derived by posttranslational cleavage

from a prepropeptide of 117 residues. On the basis of bioinformatic searches of

putative hormones derived from the prepropeptides of known peptide hormones, we

identified a ghrelin-associated peptide. We searched GenBank for orthologs of

the

human ghrelin gene and compared preproghrelin sequences from 11 mammalian

species.

In addition to the known ghrelin mature peptide, which immediately follows the

signal peptide, we identified another conserved region that was flanked by

potential

convertase cleavage sites (fig. S1, underlined). This region encodes a putative

23–amino acid peptide, with a flanking conserved glycine residue at the C

terminus,

suggesting that it might be amidated (9). We named this ghrelin-associated

peptide

obestatin.

Characterization of endogenous obestatin.

To detect endogenous obestatin, we prepared a synthetic obestatin peptide and

performed radioimmunoassays on rat-tissue extracts with obestatin-specific

antibodies. As shown in Fig. 1A, the stomach extract displaced I125-obestatin

binding to the obestatin antibodies. Obestatin-like activities from stomach

extracts

were purified. Immunoreactive (ir) obestatin was eluted in a Sephadex G-50 gel

permeation column (Amersham Biosciences, Piscataway, NJ) with estimated sizes of

2.6

and 1.5 kilo-daltons (kD), distinct from the elution position of mature ghrelin

(Fig. 1B). We subjected peak 1 (2.6 kD) fractions to ion-exchange fast protein

liquid chromatography (FPLC). A single peak of ir obestatin was eluted (Fig. 1C)

and

shown by mass spectrometry and Edman sequencing to contain a peptide with a

molecular mass of 2516.3 (Fig. 1D) and with a sequence of

FNAPFDVGIKLSGAQYQQHG-XX

(10). Combined with molecular-weight determination, the full sequence of the

purified peptide was predicted to be FNAPFDVGIKLSGAQYQQHGRALNH2, consistent with

the

obestatin sequence deduced from rat ghrelin cDNA. In addition, mass

spectrometric

analyses suggested that peak 2 (1.5 kD) represented the last 13 residues of

amidated

obestatin, indicating further processing.

To investigate differential secretion of ghrelin and obestatin in vivo, we

fasted

adult male rats for 48 hours before refeeding. Consistent with earlier findings

(11), fasting led to a major increase in serum ghrelin levels, whereas

subsequent

refeeding for 2 hours by allowing animals free access to food or drinking water

containing dextrose decreased circulating ghrelin (Fig. 1E). In contrast, serum

levels of obestatin determined by a radioimmunoassay were constant in the

different

treatment groups.

Obestatin suppression of food intake and gastrointestinal functions. We next

synthesized amidated human obestatin and tested its effect on food intake in

adult

male mice. Intraperitoneal injection of obestatin suppressed food intake in a

time-

and dose-dependent manner (Fig. 2A). Intracerebroventricular treatment with

obestatin also decreased food intake (Fig. 2B), similar to the anorexigenic

effect

of the synthetic melanocortin agonists MTII (12). In contrast, treatment with

the

nonamidated obestatin (NA-obestatin) was less effective. We also investigated

the

effect of obestatin, ghrelin, or vehicle alone on body weight in adult male

rats.

Treatment with ghrelin (1 µmol per kg body weight, three times daily) increased

body

weight, whereas the same dose of obestatin suppressed body-weight gain (Fig.

2C).

Serum leptin levels were not affected after treatment with either obestatin or

ghrelin (fig. S2), suggesting minimal modulation of body-fat content.

Furthermore,

treatment with obestatin led to a sustained suppression of gastric emptying

activity

(Fig. 2D). In vitro, isometric force measurement demonstrated that obestatin

treatment decreased the contractile activity of jejunum muscle strips and

antagonized the stimulatory effect of ghrelin (Fig. 2E) (13). The observed

inhibition of jejunal contraction may trigger an afferent vagus signal to induce

a

central satiety response. Unlike ghrelin, obestatin did not increase GH

secretion by

cultured rat pituitary cells (fig. S3).

Obestatin is the cognate ligand for GPR39. Experiments with crude

plasma-membrane

preparation of rat jejunum revealed that I125-obestatin bound to jejunal

preparations with a high affinity (dissociation constant Kd = 4 nM), and this

binding was not competed by ghrelin, motilin, neurotensin, or neuromedin U (fig.

S4). Furthermore, NA-obestatin and truncated (des1-10)obestatin showed a lower

binding affinity than did obestatin. I125-obestatin also bound to the pituitary,

stomach, ileum, and hypothalamus, but less so to other tissues (fig. S4).

We hypothesized that obestatin interacts with an orphan GPCR, and we tested

obestatin binding to Chinese hamster ovary (CHO) cells transfected with 30

individual orphan receptor cDNAs. I125-obestatin interacted with high affinity

(Kd =

1 nM) to the orphan receptor GPR39, which belongs to the ghrelin receptor

subfamily

(Fig. 3A) (14, 15). I125-obestatin binding to GPR39 was competed by obestatin

but

not by ghrelin or several other brain/gut hormones including motilin,

neurotensin,

or neuromedin U (Fig. 3B). In addition, NA-obestatin and truncated

(des1-10)obestatin had a lower affinity for GPR39 than did obestatin. In CHO

cells

overexpressing GPR39, treatment with obestatin stimulated cyclic adenosine

monophosphate (cAMP) production, whereas treatment with ghrelin or motilin was

ineffective (Fig. 3C). Consistent with the reported activation of the serum

response

element (SRE) by constitutive active GPR39 (14), hormonal treatment of CHO cells

co-transfected with GPR39 and a SRE promoter-luciferase construct led to

obestatin

but not ghrelin or motilin signaling (Fig. 3D). Similar stimulation of cAMP

production and the SRE promoter by obestatin was found when GPR39 was

overexpressed

in HEK293T cells (fig. S5). Although CHO cells expressing GHSR did not respond

to

treatment with obestatin or ghrelin, cotransfection with a chimeric Gsq protein,

which is capable of switching Gq-mediated signaling to Gs proteins (16), led to

cAMP

increases induced by ghrelin but not obestatin (Fig. 3E). Likewise, cells

expressing

the Gsq protein and the motilin receptor responded to treatment with motilin but

not

obestatin (Fig. 3F). Cross-linking studies further demonstrated that

I125-obestatin

bound to recombinant GPR39, forming a high–molecular-weight complex (fig. S6).

Real-time reverse-transcription polymerase chain reaction (RT-PCR) analyses

indicated that GPR39 is expressed in the jejunum, duodenum, stomach, pituitary,

ileum, liver, hypothalamus, and other tissues (Fig. 3G), consistent with

obestatin

binding studies.

Discussion.

Ghrelin is implicated in meal initiation and body-weight regulation. Chronic

ghrelin

administration increases food intake and decreases energy expenditure, thus

causing

weight gain. In contrast to ghrelin, which causes hyperphagia and obesity in

rats

(17), obestatin appears to act as an anorexic hormone by decreasing food intake,

gastric emptying activities, jejunal motility, and body-weight gain. Mutant mice

with a deletion of the ghrelin gene did not show impaired growth or appetite (6,

18), most likely because these animals lacked both orexigenic ghrelin and

anorexic

obestatin. Indeed, transgenic mice bearing the preproghrelin gene under the

control

of the chicken ß-actin promoter produced high levels of inactive des-acyl

ghrelin

but exhibited lower body weights (19), most likely due to excessive obestatin

biosynthesis.

The discovery of amidated obestatin and its cognate receptor underscores the

power

of comparative genomic analyses in the postgenomic era. A peptide derived from

the

66 C-terminal amino acids of proghrelin, named C-ghrelin, was detected in human

circulation, and its serum levels were elevated in patients with heart failure

(20).

Although the antibodies used to detect C-ghrelin overlap with obestatin by 13

residues, the exact chemical nature and function of the circulating C-ghrelin

remain

unclear.

Our finding that two peptide hormones derived from the same proprotein act

through

distinct receptors and exert opposing physiological actions highlights the

importance of posttranslational regulatory mechanisms. Thus, monitoring of

ghrelin

transcript levels does not accurately reflect the secretion of these two

polypeptides. After removal of the signal peptides from prepropeptides,

convertases

cleave prohormones at mono- or dibasic residues (21). In processed peptides with

a

C-terminal glycine, the residue is further amidated (9). Similar to the

importance

of posttranslational amidation for obestatin bioactivity, ghrelin also requires

acylation on its serine-3 residue for bioactivity (7).

Ghrelin binds to GHSR, which belongs to the subgroup of type A GPCRs consisting

of

GPR39 and receptors for ghrelin and motilin (22). Our discovery that obestatin

is

the cognate ligand for GPR39 suggests that GHSR and GPR39 could have evolved

from a

common ancestor but diverged in their functions, thus maintaining a delicate

balance

of body-weight regulation. This scenario is similar to the divergent and

sometimes

opposing actions of two paralogous corticotropin-releasing hormone receptors and

their ligands in the regulation of adaptive stress responses (23–25).

In addition to roles in meal initiation, weight regulation, and gastrointestinal

activity, ghrelin also regulates the pituitary hormone axis, carbohydrate

metabolism, and various functions of the heart, kidney, pancreas, adipose

tissues,

and gonads (26). Because ghrelin mRNA was found in almost all human tissues

analyzed

(27), the identification of obestatin derived from the same gene product as

ghrelin

provides a basis for future elucidation of the differential posttranslational

processing and modification of these two peptides. A better understanding of the

roles of ghrelin and obestatin in the intricate balance of energy homeostasis

and

body-weight control may be essential for the successful treatment of obesity.

Al Pater, PhD; email: old542000@...

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