Brage NP

Trollposten 12/2005

Pedersen, Christina A.

Winther, Jan-Gunnar

English

Intercomparison and validation of snow albedo parameterization schemes in climate models

Isaksson, Elisabeth

Kohler, Jack

Pohjola, Veijo

Moore, John

Igarashi, Makoto

Karlöf, Lars

Martma, Tonu

Meijer, Harro

Motoyama, Hideaki

Vaikmäe, Rein

van de Wal, Roderik S.W.

Two ice-core d18O records from Svalbard illustrating climate and sea-ice variability over the last 400 years

Våge, Dag Inge

Fuglei, Eva

Snipstad, Janne

Landsem, Veslemøy Malm

Klungland, Helge

Peptides 26 (2005) 1814–1817
Two cysteine substitutions in theMC1Rgenerate the blue variant
of the arctic fox (Alopex lagopus) and prevent expression
of the white winter coat
Dag Inge V̊agea,∗, Eva Fugleib, Kristin Snipstada, Janne Beheima,
Veslemøy Malm Landsemc, Helge Klunglandc
a Norwegian University of Life Sciences (UMB), Department of Animal and Aquacultural Sciences,
P.O. Box 5003, N-1432̊As, Norway
b Norwegian Polar Institute, The Polar Environmental Centre, N-9296 Tromsø, Norway
c Department of Laboratory Medicine, Children and Women’s Health, Faculty of Medicine,
NTNU, Morfologibygget, St. Olav Hospital, N-7006 Trondheim, Norway
Received 6 August 2004; accepted 6 November 2004
Available online 27 June 2005
A
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d
bstract
We have characterized two mutations in theMC1Rgene of the blue variant of the arctic fox (Alopex lagopus) that both incorporate a nov
ysteine residue into the receptor. A family study in farmed arctic foxes verified that the dominant expression of the blue color p
osegregates completely with the allele harboring these two mutations. Additionally to the altered pigment synthesis, the blue
uppresses the seasonal change in coat color found in the native arctic fox. Consequently, these findings suggest that the M
egulatory system is involved in the seasonal changes of coat color found in arctic fox.
2005 Elsevier Inc. All rights reserved.
eywords:Pigmentation; MC1R; Blue fox; Winter coat; Arctic fox; Polar fox;Alopex lagopus
. Introduction
At high latitudes, mammals have to face seasonal changes
n temperature, light, food availability, and colors in their
urroundings. According to the species, there are various
hysiological adaptations to these environmental changes
ike hibernation, storage of energy via fat deposition and
odification in pelage. The suprachiasmatic nucleus (SCN)
f the hypothalamus is the principal component of the mam-
alian biological clock. The pacemaker of the SCN oscillates
ith near 24 h period, and coordinates a broad spectrum of
hysiological, endocrine and behavioral circadian rhythms.
onsistent with its role in circadian timing, several experi-
ents have provided evidence that SCN is the locus of the
∗ Corresponding author. Tel.: +47 64 96 52 81; fax: + 47 64 96 51 01.
E-mail address:daginge.vage@umb.no (D.I. Våge).
brains endogenous calendar, enabling organisms to anti
seasonal changes (for review see[8]). Mechanisms of how
the circadian rhythm of SCN is translated into seasonal
ation and how seasonal variation in coat color is function
regulated are not well understood.
The arctic fox (Alopex lagopus) is a circumpolar inhab
tant of the Arctic, and occurs naturally in two color morp
white and blue. In the white morph the production of pigm
is dramatically changed with season. The summer pe
is characterized by a dark dorsal and a light ventral pat
while the white winter coat is produced as an adaptatio
snow and icy environments. A seasonal pelage shift oc
with molting of the highly insulative white winter fur in Ma
to a shorter brown/grey summer fur. The initial growth
the winter fur occurs in September, and continues until e
December. The winter fur is 200% thicker than the sum
fur [21]. The exact mechanism of the down regulation
196-9781/$ – see front matter © 2005 Elsevier Inc. All rights reserved.
oi:10.1016/j.peptides.2004.11.040
D.I. Våge et al. / Peptides 26 (2005) 1814–1817 1815
Fig. 1. Illustrations show the blue variant of the arctic fox in (A) and the winter white variant in (B). Both pictures show the summer coat of a puppy. In (C),
an adult blue fox is seen to the left, whereas the winter white variant is seen to the right. Illustrations are of individuals that were included in the stu y (for
interpretation of the references to color in this figure legend, the reader is referred to the web version of this article).
pigment synthesis connected with the change to winter fur
is not known.
Blue fox is a color variant of arctic fox, showing a uniform
dark grey/blue coat color during the summer. The dominant
blue coat color is inherited like a simple Mendelian trait
caused by a single gene. In the winter season, the blue fox
develops a coat color that is lighter than the summer coat but
far more colored compared to the winter white native arctic
fox (Fig. 1). At Svalbard, as in several other natural habi-
tats of the arctic fox, 3–5% of the population is of the blue
variant.
Dominant acting mutations in theMC1Rcausing a gen-
eral eumelanizing effect are well known from several mam-
malian species including mouse, cattle, red fox and sheep.
These have been functionally tested in cell assays, and have
been found to produce a constitutively active receptor caus-
ing eumelaninization mediated through the cAMP pathway
[11,14,19,25,26]. Except for the silver fox, these mutations
are almost completely epistatic to theagouti gene, even in
the heterozygous state.
In the present study we have identified two substitution
mutations in the melanocortin 1 receptor (MC1R) in the blue
fox, that co segregate completely with the blue fox phenotype
in crosses between blue fox and winter white arctic fox. In
addition to causing a uniform dark summer pelage, the two
mutations also prevent the development of a completely white
w C1R
r coat
c
2. Materials and methods
2.1. Animals
The family study with farmed blue foxes was done based
on a cross between a heterozygous female blue fox and a
white male in the research farm at the Norwegian University
of Life Sciences (Fig. 1). Among ten offspring, two were blue
foxes and the remaining eight were of the winter white phe-
notype. In addition to the family study, blood samples were
drawn from seven wild arctic foxes (of the white morph) that
were caught near Ny-Ålesund (78◦55′N, 11◦56′E), Svalbard,
Norway. These animals were a part of a larger project at the
Norwegian Polar Institute, Tromsø, Norway[6,7]. Verifica-
tion of the mutations found in the blue fox was also done in
a blue fox originating from Finland.
2.2. PCR amplification and sequencing
DNA was isolated from blood samples following standard
protocols. The complete coding region of theMC1Rgene was
amplified with primers AL1 and AL2 (Table 1). All primers
are designed based on the red foxMC1Rsequence (X90844).
The PCR reaction was carried out in a 20l reaction contain-
ing 20 ng genomic DNA, 10 pmol of primers AL1 and AL2,
200M dNTP, standard buffer conditions and 1.5 U Taq poly-
m n
f
3 withinter pelage. These findings indicate that the agouti/M
egulatory system is involved in seasonal changes in
olor found in arctic fox.erase. DNA was denatured for 3 min at 94◦C, and PCR ru
or 35 cycles at 95◦C for 15 s, 60◦C for 15 s, and 72◦C for
0 s. Sequencing of the complete region was carried out
1816 D.I. Våge et al. / Peptides 26 (2005) 1814–1817
Table 1
Primers used in the study to amplify, sequence and genotype the arctic fox
MC1Rgene
Primer Position Direction Sequence of primer in 5′–3′ direction
AL1 ÷54 to÷35 Forward GAA CTG AGC GAG ACA CCT GA
AL2 980–997 Reverse ATC ACC ACC TCC CTT TGC CCA
AL3 192–211 Forward CAA GAA CCG CAA CCT GCA CT
AL4 42–61 Reverse TGG TTG GGG AGG TGG CAA TG
Positions are given relatively to the translational start ATG of the red fox
MC1Rgene (X90844).
primers AL1, AL2 and AL3 using the protocol of Applied
Biosystems.
2.3. RFLP analysis
Following the detection of two mutations in theMC1R
gene of arctic fox, a RFLP reaction was developed to dis-
tinguish between the two variants. Shortly, primer AL1 and
AL4 were used to amplify a 116 bp region containing the first
of two linked mutations of interest localized to amino acid
positions 5 and 280. PCR was carried out in an 20l reaction
containing 10 pmol of each primer and 20 ng DNA, and was
run for 35 cycles at 95◦C for 15 s, 60◦ for 15 s, and 72◦ for
15 s. The PCR fragment was cut withApaIat 25◦C over night.
If the wild type allele is present, the reaction will produce two
fragments of 71 and 45 bp. If the blue fox allele is present,
ApaI will not cut the fragment. The mutation localized to
amino acid position 280 was genotyped by DNA-sequencing
(Applied Biosystems) in the present study.
3. Results and Discussion
By sequencing the complete coding region of theMC1R
gene of the winter white arctic fox (AJ786717), as well as
t sing
a , the
g fox,
w ition
2 tra-
c ry of
t ased
o nd a
w g-
r rctic
f e
w s a
b l
t
nt
e 2nd
a acid
l bil-
i s, a
c pa-
Fig. 2. Sequence comparison forMC1R regions that span amino acid 5
(amino acid 1–9 in the left box) and 280 (region 276–284, or 274–282 in
mouse which has a two aa deletion). For the regions illustrated, additional
species have been shown to be identical to species included in this figure
[5,10–12,16,18,25,26].
rable to the TM2/TM3 – mutations might be foreseen. Such
interactions could be formed either between the two new cys-
teine residues, or to cysteine residues localized to any of the
extracellular regions of the receptor. Both alterations in the
MC1Rgene of the blue fox are found within relatively well
conserved amino acid positions of the receptor (Fig. 2). The
5th position is conserved in all mammalians characterized so
far, except for mouse which has a glutamine substituting the
glycine. In position 280, all species are identical except for
pig which has a valine substituting the phenylalanine. Based
on the present data, it is not possible to conclude whether
one or both of these mutations are involved in the observed
phenotypic change.
The darker dorsal and lighter ventral pattern found in
the summer pelage of the arctic fox is comparable to dif-
ferentagouti alleles found in mouse, affecting the dorsum
and ventrum independently. This phenomenon is explained
by alternative promoter use in theagoutigene[4,15,20,23].
In this respect, the blue fox mutation act as a classical con-
stitutively activatedMC1Rmutation in being epistatic to the
agoutigene. Inter-species crossing between the winter white
arctic fox and the standard silver variant (aaE+E+) of the
red fox (Vulpes vulpes) provide firm evidence for a func-
tional agouti protein in the arctic fox. The standard silver
fox do not carry a functional agouti protein, and the red
color found in the offspring (named Golden island) must
t tein
[
s
p e
f oat
o ults,
t t. A
f ium
b g
t
d tein
d ould
e hite
o ivityhe blue color variant (AJ786718), two mutations cau
mino acid substitutions were discovered. In position 5
lycin residue is substituted by a cysteine in the blue
hile a phenylalanine is replaced by a cysteine at pos
80. These two mutations are localized to the initial ex
ellular region of the receptor and the external bounda
ransmembrane domain 7, respectively. A family study b
n a cross between a heterozygous female blue fox a
hite male fox (Fig. 1) verified that the two mutations cose
egated with the dominant blue phenotype. The wild a
oxes from Svalbard have aMC1Rgenotype identical to th
inter-white domestic arctic foxes from Norway, wherea
lue fox of Finnish origin have aMC1Rgenotype identica
o the Norwegian blue fox.
Previously reportedMC1Rmutations causing domina
umelanizing effects are localized within or close to the
nd 3rd transmembrane (TM) domains of the 317 amino
ong receptor[3,13]. However, since cysteine has the capa
ty of forming disulfide bridges with other cysteine residue
onformational change of the receptor with an effect comherefore be ascribed the functional “arctic” agouti pro
17,25].
Based on classical genetic studies[2] a hypothesis wa
ut forward that a dominantagoutiallele (Aw) is responsibl
or gradually eliminating all eumelanin from the birth c
f the pups and from the colored summer coat of ad
hus replacing the summer color by the white winter coa
unctional explanation of this could be that the equilibr
etween the agouti protein and-MSH is shifted accordin
o the photoperiods, with a higher concentration of-MSH
uring summer, and a relatively higher level of agouti pro
uring the winter season. According to this model one sh
xpect a yellow/red winter coat instead of the observed w
ne. However, a general reduction in melanocyte act
D.I. Våge et al. / Peptides 26 (2005) 1814–1817 1817
combined with a rapid growth of the winter coat may dilute
the pheomelanin in a level giving this white appearance.
-MSH and UV radiation are well known as major phys-
iological signals that modulate skin pigmentation[1,22].
Melanocytes react to these signals by proliferation, increased
dendriticity and elevated pigment synthesis. Prolonged day
length during summer increases the UV radiation and the
local -MSH production in the skin is increased. Although
the fox skin is not directly subjected to sunlight, it has been
proposed that the hair shaft may function as a fiber-optic sys-
tem for guiding UV light to the hair follicle[9]. This has
of course to be verified experimentally, but should not yet
be exclude as a possible contribution to the eumelanization
seen in the summer pelage. In addition to a local reaction in
the skin, due to increased day length, SCN guided signals do
most likely play an important role in forming a coordinated
reaction to seasonal changes. Changes in circulating levels
of -MSH synthesized in the pituitary gland is one possible
SCN-guided signal.
Based on our results we do assume that the two muta-
tions found in theMC1Rgene of the blue fox are acting as
a classical dominant mutation, causing a constitutive active
receptor. Such mutations are capable of activating the recep-
tor in the absence of the-MSH ligand[14]. These processes
may explain both the general darker pelage of the blue fox,
the lack of a light ventrum and a lesser responsiveness to
s a-
s
A
Fur
B
R
s on
tured
ight.
rela-
fects
tion
ture
gene
lor.
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[ olor
[ s in
P1)
quus
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[ enet
[ rctic
[ s
and
ytes.
[ s
on of
–71.
[ RD.
lpes
[ a-
eep.hifts in the agouti/-MSH equilibrium influencing the se
onal changes in coat color.
cknowledgement
The Research Council of Norway and the Norwegian
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Two cysteine substitutions in the MC1R generate the blue variant of the arctic fox (Alopex lagopus) and prevent expression of the white winter coat

Gundestrup, Niels

Mulvaney, Robert

Pourchet, Michael

Hofstede, Coen

Lappegard, Gaute

Pettersson, Rickard

van den Broeke, Michiel

Accumulation variability over a small area in east Dronning Maud Land, Antarctica, as determined from shallow firn cores and snow pits: some implications for ice-core records

Strøm, Hallvard

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De marine fuglene
Sjøfuglene i Antarktis er nært knyttet til det marine miljø og henter all 
sin næring her. De tilbringer hele livet på havet, og er kun på land i 
hekketiden. I Antarktis er det særlig tre artsgrupper som dominerer blant 
sjøfuglene. Dette er pingvinene (sju arter), albatrossene (seks arter) og 
petrellene (23 arter). Pingvinene utgjør 85 % av sjøfuglenes totale 
kroppsmasse (biomasse), og er derfor viktige konsumenter i 
økosystemet.
Tilpasninger
De fleste sjøfuglene hekker langs kysten av kontinentet eller på øyer i 
havområdene rundt. Bare noen få arter har tilpasset seg det ekstreme 
miljøet inne på selve kontinentet, og de sjøfuglene som har gjort dette 
er de eneste virveldyrene du finner her inne. Blant disse finner vi flere 
arter av pingviner og petreller. Tykke lag med underhudsfett, mye varm-
ende dun og en tett fjærdrakt er tilpasninger som gjør livet mulig. 
Underhudsfettet gir isolasjon, men utgjør også en viktig matreserve i 
perioder hvor det er lite næring å finne.
Hekkesesongen
Hekkesesongen i Antarktis strekker seg fra slutten av november til 
slutten av februar. Hekkeforløpet er lagt opp slik at næringstilgangen skal 
være best mulig når ungene blir uavhengige av foreldrene. For mange 
arter er den lille rekelignende krillen det viktigste byttedyret, sammen 
med ulike arter av fisk.
Artene ved Troll
Rundt den norske forskningsstasjonen Troll finner vi tre fuglearter. 
Antarktispetrellen er trolig den mest tallrike. Den hekker i store og tette 
kolonier på barmark. Det finnes flere kolonier i området sørøst for Troll 
(Jutulsessen). Antarktispetrellen oppholder seg hele året i drivisen, men 
bruker fjellene langs kysten og nunatakene i innlandet som hekkeplass 
om sommeren. Koloniene i Dronning Maud Land ligger ca. 200 km inn 
fra kysten (iskanten). Fuglene må derfor gjøre lange turer for å skaffe 
mat til seg selv og ungene. Spesielt gjelder dette tidlig i hekkesesongen, 
når drivisen gjør avstanden ut til åpent hav enda større. Fugler fra Svart-
hamaren utstyrt med satellittsendere har vist at Antarktispetrellen kan 
gjøre enkeltturer på opptil 11 000 km! De søker da over isfylte havom-
råder som er flere ganger større enn Barentshavet!
Snøpetrellen er den andre av petrellene som har tilhold rundt Troll. Den 
er mindre enn Antarktispetrellen og umiskjennelig med sin hvite fjær-
drakt, svarte øyne, nebb og føtter. Snøpetrellen kan også opptre i store 
kolonier, men foretrekker grov blokkmark hvor reiret kan plasseres trygt 
i hulrom under steinene. Tettheten av reir blir derfor sjelden høy. Kolonier 
av arten forekommer spredt i fjellområdene over hele Dronning Maud 
Land.
Sørjoen er rovfuglenes representant i Dronning Maud Land. Sørjoen 
livnærer seg i hekketiden utelukkende på egg, unger og voksne fugler 
av de to petrellartene. Den hekker også nesten bare i tilknytning til 
petrellkoloniene, og da i størst antall ved de store koloniene. Reiret 
legges på barmark, og det forsvares aktivt med angrep fra lufta dersom 
Det finnes omtrent 45 arter av sjøfugler i Antarktis, men bare noen få 
har tilpasset seg det ekstreme miljøet inne på kontinentet. Her lever de 
på grensen av hva som er fysiologisk mulig og kan fly turer på opptil 
11 000 km for å finne mat. 
Sjøfuglene i Antarktis
Snøpetrellen er umiskjennelig med sin hvite fjærdrakt. Den hekker i spredte kolonier rundt Troll.
Gulltoppingviner på Bouvetøya.
noen skulle bevege seg innenfor ”territorialgrensene”. Sørjo er en kraftig 
og robust art som på imponerende vis – men også brutalt – kan angripe 
og drepe voksne antarktis- og snøpetreller i flukt.
Forskning og overvåking
Det meste av norsk forskning på sjøfugl i Antarktis har foregått i 
Svarthamaren i Dronning Maud Land og på Bouvetøya. Svarthamaren 
er en nunatak som ligger ca. 100 kilometer øst for Troll. Her finnes den 
største kjente antarktispetrellkolonien i verden. Kolonien teller nærmere 
250 000 hekkende par. I tillegg hekker også mange par av snøpetrell 
og sørjo i samme område. Stasjonen Tor ble i 1992 etablert ved foten 
av fjellet som base for arbeidet. Her har forskerne først og fremst 
konsentrert seg om fuglenes tilpasninger til det ekstreme miljøet de 
lever i, og hvilke hekkestrategier de har utviklet for å kunne overleve i 
dette barske miljøet. Det gjennomføres også langsiktig overvåking av 
sjøfuglbestandene i Svarthamaren for å øke kunnskapen om hvordan de 
påvirkes av endringer i miljøet. Slike lange tidsserier er også viktige for å 
kunne forstå og forutsi effekter av klimaendringer.
Bouvetøya
Den lille sub-antarktiske Bouvetøya er interessant i forsknings- og 
overvåkingssammenheng på grunn av sin isolerte beliggenhet. Øya har 
store bestander av blant annet ringpingvin, gulltoppingvin og pelssel. 
Siden 1976 har Norsk Polarinstitutt drevet forskning og overvåking på 
disse artene, og siden sesongen 1996/97 har dette arbeidet vært et 
ledd i et internasjonalt program for overvåking av økosystemet under 
konvensjonen for bevaring av marine levende ressurser i Antarktis 
(CCAMLR). 
Nye muligheter
Ved utvidelsen av Troll til helårsstasjon og åpningen av flystripe på blåisen 
i februar 2005, er det åpnet for nye muligheter innen overvåking og 
forskning på sjøfugler i Dronning Maud Land. En interessant mulighet 
er å etablere et forsknings- og overvåkingsprogram knyttet til de store 
petrellkoloniene i Jutulsessen-området. Dette vil kunne bli svært 
verdifulle dataserier som støtte til aktiviteten som allerede pågår i 
Svarthamaren. Ved at det nå skal være bemanning på Troll hele året, kan 
tid frigjøres til mer forskning ved at f.eks. overvintrerne kan ha praktisk 
ansvar for overvåkingsaktiviteter.
Norsk Polarinstitutt 2005. Tekst: Hallvard Strøm, strom@npolar.no. Foto: Bjørn Krafft og Hallvard Strøm. Design: Jan Roald
Antarktispetrellen hekker på barmark i tette kolonier.
Sørjoen lever av egg, unger og voksne fugler av snø- og antarktispetrell. 
Forskningsstasjonen Tor ble etablert ved Svarthamaren i 1992. Den store antarktispetrellkolonien ligger i fjellskråningen bak.


Norwegian

Sjøfuglene i Antarktis

Sander, Gunnar

Effekter av miljøgifter i Arktis : resultater av Transport- og effektprogrammet

Holmén, Janet

Polar Research in Tromsø 2005

Hanssen-Bauer, Inger

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Prestrud, Pål

Norwegian Bokmål

Miljøovervåking av Svalbard og Jan Mayen – MOSJ : en dokumentasjon av systemet og den første vurderingen av miljøstatus

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Growth and population parameters of the world’s northernmost harbour seals Phoca vitulina residing in Svalbard, Norway

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