全 文 ：广 西 植 物 Guihaia 30(3)：377— 385 2010年 5月
彩色马铃薯色素相关基因座的
种类、功能与染色体定位
赵昶灵，李孙文 ，张维明
(云南农业大学 农学与生物技术学院，昆明 650201)
摘 要 ：综述了与彩色马铃薯色素产生与分布相关基因座 的观念起源、种类、功能和染色体定位。与彩色马
铃薯色素相关基因座的观念起源于试 图解释四倍体和二倍体马铃薯块茎和其他部位颜色呈现遗传行为的两
个遗传模式 。与彩色马铃薯色素相关的 13个基因座可划分为 4类，第 1、第 2和第 3类分别与马铃薯花色苷
的合成、酰化和分布有关 ，第 4类与马铃薯类胡萝 卜素的产生相关。基因座 J，P，R和 y分别编码一个 MYB
结构域转录因子 、类黄酮 3 ，5 一羟化酶、二氢黄酮醇4一还原酶和p一胡萝 卜素羟化酶。基因座之间复杂多样的互
作综合决定了彩色马铃薯色素特别是花色苷的产生与分布。基因座 D 和R定位在马铃薯的 2号染色体上，
E，F，I和PSC在 10号染色体上，P在 l1号染色体上，y在3号染色体上。可为彩色马铃薯颜色呈现的遗传
机理探索提供参考 。
关键词 ：彩色马铃薯；色素；基因座；种类 ；功能；染色体定位
GLC Number：Q343．1 Document Code：A Article ID：1000—3142(2010)03—0377—09
Category，functions and chromosomal locations of
the gene loci related to colored potato pigments
ZHAO Chang-Ling，LI Sun-Wen ，ZHANG Wei—Ming
(Colege o厂Agricultural Sciences and Biotechnology，Yunnan Agricultural University，Kunming 650201，China)
Abstract：This paper summarized the idea origin，category，functions and chromosomal locations of the gene loci in—
volved in the production and distribution of the pigments in colored potatoes．The idea of the gene loci related to the
pigments in colored potatoes originated from the two genetic models which attempted to explain the inheritant behav—
iors of the colorations of the stem tubers and other parts of tetraploid and diploid potatoes．The overal thirteen gene
loci related tO the pigments in colored potatoes could be classified into four kinds．The first，second and third kinds
were related to the synthesis，acylation and distribution of the potato anthocyanins respectively，and the fourth kind
was related tO the production of the potato carotenoids．Locus，，P，R and Y encoded a MYB-domain transcription
factor，flavonoid 3 ，5-hydroxylase，dihydroflavonol 4-reductase and la-carotene hydroxylase respectively．Various and
complicated interactions among the gene loci synthetically determined the production and distribution of the pigments，
especially the anthocyanins，in colored potatoes．Locus D and R were located on Chromosome 2 of potato，E，F，I and
～ PSC on、Chromosome 10，P on Chromosome 1 1，and Y on Chromosome 3．This paper could provide a reference for
the exploration on the genetic mechanism of the colorations Of colored potatoes．
Key words：colored potato；genetic locus；category ifunction；chromosomal location
Received date：：2008-08—04 Accepted date：：2009-04-i9
Foundation item：Supported by grants from the Provincial Department of Science and Technology of Yunnan(2006CO030Q)；the Startup Fund for Doctor of
Yunnan Agricultural University(A2002096)
Biography：ZHA0 Chang—I ing(1969一)．Male．Born in Dujiangyan City of Sichuan Province，Doctor of Science，Full Professor．working in Plant
physiology，Phytochemist ry，Bioinformatics and Plant biochemistry and molecular biology，(E-mail)zhaoplumblossom7@ 163．com．
Author for corrcsp0ndence．E—mail：swl68@ yahoo．en
378 广 西 植 物 3O卷
Generally，the skins and／or fleshes of the stem tu—
bers of Dotato(Solanum tuberosurn)are white，yellow or
safifon vellow． Worldwide，the potato cuhivars in
which the skins and／or fleshes of the stem tubers are
red r purDle，blue or orange are intuitively denominated
co1ored potatoes(Brownet a1．，2003；Brown，2004；
Jansen& Flamme，2006)．
The coloration patterns of the skins and fleshes of
the stem tubers of colored potatoes are changeable and
fascinating．Not all of the colors of the skins are consist—
ent with those of the fleshes(Oroza et a1．，2004)．Col—
ored skins do not mean the fleshes are definitely colored．
However，if the fleshes are colored，the skins are usually
colored，e．舀 the red or purple fleshes are often accom—
Danied by the red or purple skins respectively(Brown et
． ，
2003)，and red or blue colored fleshes have identical～
lv colored skins(Brown et a1．，2003；de Jong et a1．，
2003a)．Furthermore，the tuber skins of colored potatoes
are uniformly colored，but the colorations of the fleshes
are diverse，i．e．the fleshes may range from partial pig—
mentation to complete pigmentation(Brown et a1．，
2003)。resuhing in the colorful arcs，rings or radiating
stars in the fleshes．Cases in point are the potato culti—
vars native to the Andes of South America and the local
Dotato cuhivar‘Zhuanxinwu’planted in Baoshan Town
of Xuanwel‘City in Yunnan Province of China(Brown et
“Z．，2003；Zhao et a1．，2007)．
Various pigmentations of colored potatoes result
basica11v from the accumulation of anthocyanins in the
sDecific parts of the stem t ubers．It has been proved
that the skin and flesh colors of the stem tubers of po—
tatoes are mainly determined by two different classes of
Digment，i．e．carotenoids and anthocyanins(Lewis et
“Z．，1998)． Carolenoids lead to the white，yellow or
saffron velow of the skins and／or fleshes(Lewis et
“￡．，1998；Brown et“ ．，1993，2003；Morris et a1．，
2004)．Anthocyanins lead to the red，purple or blue of
the skins and／or fleshes，fundamentally producing col一
。red Dotatoes(Hung et a1．，1997；Fossen et a1．2003)．
In the stem tubers of colored potatoes,anthocyanins al—
wavs accumulate principally in the periderms and pe—
ripheral cortexes(Howard et a1．，1970；Burton et a1．，
1989)．In fact，anthocyanins may produce anywhere In
the plants of colored potatoes，e．g．，in stem tubers，
flowers，sprouts or stems(Jung et a1．，2005)，and they
also result in the red，purple or blue of the upground
Darts of the potatoes(Harborne，1960；Hung et a1．，
1997：Lewis eta1．，1998；Brown eta1．，2003)．
This summary deh with the idea origin，category，
functions and chromosomal locations of the gene loci
involved in the production and distribution of the pig—
ments，i．e． anthocyanins and carotenoids，in colored
potatoes，attempting to provide a reference for the fur—
ther molecular biological studies on the pigments，espe—
cially the anthocyanins，of colored potatoes．
1 Establishment of the idea of the
gene loci involved in the production
and distribution of colored potato pig—
m ents
The idea of the gene loci involved in the produc—
tion and distribution of the pigments in colored pota—
toes was established primarily on the two classic genet—
ic mode1s which tried to explain the inheritant behav—
iors of the colorations of the stem tubers and other
parts of tetraploid and diploid potato plants．
The first genetic model was originaly raised by
Salarnan(1910)to elucidate the color inheritance of the
stem tuber of tetraploid European potato varieties．In
this model，the purple，red and white of the tuber were
postulated to be controlled by three independent gene lo—
ci，i．e．D，R and P．The red of the tuber skin results
from the complementary action of the dominant D and
R,and，together wi th P，D and R make the skin purple
(Salaman，1910)．Afterwards，Locus E was proposed to
be related to the accumulation of the anthocyanins in the
tuber epidermi s and bud eye，R was believed to express
in the tuber cortex(Salama n，1926；Lunden，1937)．
The second genetic model was raised by Dodds&
L0ng(1955，1956)to elucidate the color inheritance of
the stem tuber of diploid cultivated potato cuhivars．In
this model，Locus P，R／R ，E，J and F were intro—
duced to explain the various accumulations of the an—
thocyanins in different pari s of the potato plant s．
3期 赵昶灵等：彩色马铃薯色素相关基因座的种类、功能与染色体定位 379
However，in the 1990s，the above two models
were united to a great extent since they were compared
each other by Howard(1970)and de Jong(1991)．A1一
though，for the diploid and tetraploid potato cultivars，
the number and functions of the genes involved in the
production and distribution of the anthocyanins were
tremendously different，the genes coding the similar
traits in these cultivars were thought to belong to the
same(de Jong，1991；van Eck et a1．，1994)．van Eck et
a1．(1994)further evaluated the two genetic models in
detail and proposed to unify some of the gene locus
names properly．R／R and PSC were revoked．R／R舡’
was replaced by D．R，E，J and PSC were thought to
be actually a same locus on Chromesome 10 of potato，
and h1vo1ved in the tuber skin coloration(van Eck et
ⅡZ．，1994)．
2 Category and functional character—
istics of the gene loci involved in the
oduc‘ and tistribution of coloredpr tion d l l l
potato pigments
2．1 Coloration-related functions of the gene loci
There are 1 3 gene loci related to colored potato
pigments，i．e．anthocyanins and carotenoids，in all，and
they can be devided into 4 kinds(Table 1-3)．
Table 1 Functions of the gene loci involved in the synthesis and acylation of the anthocyanidins
and the production of the carotenoids in colored potatoes
Function Required for the
synthesis of red an—
thocyanins in each
part of the plant
【 iterature Swaminathan &
Howard (1953)；
Harborne (1960)
I unden (197,1)；
van Eck el a1．
(1993，1994)
Encoding the basic factors re—
quired for the production of
blue／purple petunins in any
tissue of the plant and eontro-
ling the formation of methylie
delphinin
Dodds＆ Long(1955，1956)；
Harborne (1960)：I unden
(1974)； van Eck et a1．
(1993，1994)；de Jong el a1．
(2004a)；Jung el a1．(2005)
Encoding a basic factor required Controlling the Required for the
for the production of red pelar— acylation of the accumulation of
gonins in any tissue of the plant anthocyanins yellow carote—
and controling the production of holds in tuber
acylated pelargonin in tuber and flesh
the cyanin in flower
Dodds＆ Long(1955，1956)； Dodds & Long Bonierbale et a1．
Harborne (1960)； I unden (1955)； Har一 (1988)；de Jong
(1960)：van Eck et aZ．(1993， borne(1960) et a￡．(2004a)
1994)：de Jong el a1． (2003b，
2004a)
2．1．1 The gene loci related to the synthesis of colored
potato anthocyanidins These loci，including D，P and
R，decide the kind of anthocyani dins synthesized lead—
ing to the specific coloration of the stem tuber or other
parts of the potato plants(Table 1)．D is the dominant
gene deciding the synthesis of red anthocyanins in each
part of the potato plants． Genetype dd leads to the
pink tubers and seedlings．and white flowers(Dodds
Long，1955，1956)． P is the dominant gene deciding
the synthesis of purple anthocyanins in each part of po—
tato plants，especially in the embryonal axes and bud
tips(Lunden，1937)．Moreover，P controls the appear—
ance of the blue pigments in diploid potatoes(Dodds＆
Long，1955，1956)．Dominant R regulates not the color
of the tuber skin，but that of the outer 1ayers of the tu—
ber cortex(Lunden，l937)．
2．1．2 Th e gene locus related to the acylation of colored
potato anthocyanins The locus in point is Ac(Table
1)．It controls the acylation of the anthocyanins，actu—
ally deciding the existent states of anthocyanins in the
specific parts of potatoes． Swaminathan & Howard
(1953)found diploid potatoes display both acylated and
non-acylated anthocyanins while only acylated forms
are present in the tetraploid potatoes．Harborne(1960)
and Rodriguez_Saona et a1．(1 998)discovered the pig—
ments resulting in colored potatoes are varied types of
acylated anthocyanidin glycosides．Brown et a1．(2003)
testified red-fleshed potatoes contain predominantly ac—
ylated glycosides of pelargonidin while the purple-
fleshed potatoes contain predominantly acylated glyco—
38O 广 西 植 物 3O卷
sides of petunidin and peon!din． In addition，Ac also
controls the linkage of glucose residue at the C5 of the
anthocyanidins，and the methylation of the delphinidin
or cyanidin derivates(Harborne，1960)．
2．1．3 The gene loci related to the distribution of colored
potato anthocyanins These loci decide whether the an—
thocyanins appear in the specific parts of tuber and leaf
or in the flowers and stems(Table 2，3)．
Table 2 Functions of the gene loci deciding the anthoeyanins appear
in the specific parts of the tuber in colored potatoes
Table 3 Functions of the gene loci deciding the anthocyanins appear in
the flower and the specific parts of the leaf in colored potatoes
(t)The gene loci deciding the anthocyanins appear
in the specific parts of tuber are B，E，I，Pf and PSC
(Table 2)．B and 1 were first raised by Dodds& Long
(1955)to control the distribution of the anthocyanins in
the tuber．E was initially used by Lunden(1960)to ex—
plain the phenotype of the skin color．It participates in
the pigmentations of epidermis and bud eyes．j is re—
quired for the tissue-specific expression of anthocyanins
in tuber skin(Dodds& Long，1956)，which is proved by
de Jong(1991)and de Jong et“Z．(2004a)．It controls
whether the anthocyanins appear in the skin and flesh
(Harborne，1960)．Genotype ii causes the absence of the
tuber pigmentation(Dodds＆ Long，1956)．Pf was o—
riginally proposed by Harborne(1 960)to control the
presentation of anthocyanins in the interior tissues of the
tuber beyond the periderm．It decides the distribution of
the anthocyanins in the tubers of diploid potatoes，and
controls the flesh color(de Jong et a1．，1987)．PSC was
introduced by Gebhardt et“ ．(I989)to control the put—
pie skin color in diploid potatoes．
(2)The genetic lOCUS deciding the anthocyanins ap—
pear in the flowers is F(Ta ble 3)．Like B and I，F were
first raised by Dodds& Long(1955．1956)to control the
distribution of the anthocyanins． Lunden(1 937，1 974)
thought F is related to the specific expression of the an—
thocyanins in the flower，and it is involved in the contrast
between selbcoloured flowers and flecked ones．
(3)The gene loci deciding the anthocyanins appear
in the specific parts of the 1eaf are Pd and Pv(Table
3)．Pd and Pv were formally put forward by Kessel
& Rowe(1974)and Garg et“Z．(1981)．They are both
the single dominant genes which respectively control
the production of the anthocyanins on the back and the
abdominal part of 1ear． Koopmans et aZ．(1951)ever
thought Pv is recessive．but Kessel& Rowe(1974)and
Garg et a1．(1981)thought Pv is dominant．
3期 赵昶灵等 ：彩色马铃薯色素相关基因座的种类、功能与染色体定位 38l
(4)The genetic locus related to the production of
colored potato carotenoids：The locus in point is Y(Ta—
ble 1)． It is required for the accumulation of yellow
carotenoids in the tuber fleshes(de Jong et a1．，2004a)．
2．2 Enzyme or protein factors encoded by the specified
gene loci
For the gene loci related to colored potato pig—
merits，only l，P，R and Y have been definitely proved
to encode specific enzymes or protein factors，i．e．』en—
codes a transcription factor(TF)，P，R and Y encode an
enzyme respectively(Table 4)．
(1)J encodes a MYI3-domain 了、F，i．e．anthocya—
nins 2(AN2)of Petunia spp．(de Jong et a1．，2003a，b，
2004a，b；Jung et a1．，2005)(Table 4)． AN2 is a
R2R3一type MYB factor，and is involved in the activa—
tion of anthocyanin coloration．MYB proteins contain a
conserved DNA-binding domain(the MYB domain)
with 1-3 imperfect repeats(Rl—R3)which define their
binding specificity to the target gene promoters(Martin
& Paz-Ares，1997)．R2R3～type proteins form the lar—
gest class of MYB factors in plants．and among them
are the factors involved in the activation of anthocyanin
pigmentation in various plants(Elomaa et a1．，2003)．
As the homologous gene of C1 of Maize(Zea mays)and
the third regulator of the anthocyanin pathway in Pe—
tunia spp．，AⅣ2 acts in concert with ANl and AN1l
to activate the promoter of dihydroflavonol 4-reduetase
gene(D )in the petal limbs(Quattrocehio et口Z．，
1998；Quattrocchio et a1．，1999)．It can interact with
either of two distinct basic he1i loop-helix HLH)
factors，JAF1 3 or AN1，and functions upstream of
ANl but not of JAF13(Elomaa et a1．，2003)．
Table 4 Enzymes or protein factors encoded
involved in the pigment synthesis in
bythe
colored
specified gene loci
potatoes
(2)P encodes flavonoid 3 ，5-hydroxylase(F3 ，5
H)(de Jong et a1．，2003a，b，2004a；Jung et a1．，2005)
(Table 4)．F3 ，5 H has been regarded as the“blue
gene”in higher plants(Okinaka et a1．，2003)．In plant
organs or tissues whose colors are determined by an—
thocyanins，the hydroxylation pattern of the B-ring of
dihydrokaempferol(DHK)is the most important step to
decide the organ color(Stafford，1990)．If the B-ring of
DHK is hydroxylated at the 3 position due to the cata—
lyzing of flavonoid 3-hydroxylase(F3 H)，DHK wil
be transferred to dihydroquercetin(DHQ)，leading to
the production of the cyanins which are responsible for
the red color．If t he B-ring of DHK is hydroxylated at
both the 3 and 5 positions due to the catalyzing of
F3 ，5 H，DHK wil be transferred to dihydromyrieetin
(DHM )，leading to the production of the delphinins
which are responsible for the blue／violet color(Lewis，
l996；Brugliera et a1．，1999)．
(3)R encodes dihydroflavonol 4-reductase(DFR)
(de Jong et“f．，2003，b，2004a，b；Jung et nf．，2O05)(Ta—
ble 4)．DFR catalyzes the reduction of DHK into leuco—
pelargonidin，ultimately producing pelargonidins(Lewis，
1996；de Jong et a1．，2004a)．So，R is necessary for the
production of red pelargonins in any tissue of all red po—
tatoes(de Jong，1991；Lewis，1996)．Overexpression of a
DNA encoding DFR in sense orientation has been proved
to result in a 4-fold increase in petunidin and pelargoni—
din derivatives in the tubers(Stobiecki et a1．，2003)
．
(4)Y encodes l~-carotene hydroxylase(BCH)(de
Jong et a1．，2004b)(Table 4)．BCH catalyses the con—
version of~-earotene into~-cryptoxanthin，and further
into zeaxanthin(Sun et a1．，1996；Hirschberg，2001；Tian
& DelaPenna，2001)，contributing to the synthesis of
yellow carotenoids in the tuber flesh．It is found that si—
lencing of BCH can increase the total carotenoid and
carotene 1evels in the tubers(Diretto et a1．，2007)．
2．3 Interactions among the gene loci
2．3．1 Interactions among D or P and E，F，R D or P
is complementary with E，F and Rwhich determine the
colors of the flower and tuber skin(Lunden，1937；van
382 广 西 植 物 3O卷
Ec 以 “Z．，j994)．
(1)P，R and D control the purple，red and white of
the tuber skins of tetraploid potatoes respectively，the
appearance of red pigments in the tuber is due to the
complementary effect between the dominant D and don>
inant R，and，if D and R interact simultaneously with P，
the tuber skin will be purple(Salaman。1910)．The corn—
bination of R wi th D or P can deepen the skin color，
namely make the skin black(Lunden，1 937)，producing
red or blue／purple tuber skin(van Eck et a1．，1994)．
Genotype D-R—makes the skin red(Swaminathan ＆
Howard，1953)，and D-P-R—res；ult in an intensely col—
ored tuber cortex with a black skin(Lunden，1937；van
Eck et a1．，1994)．The almost white tuber skin pheno—
type of ddppR-is difficult to distinguish from white-
skinned D rr or P-rr genotypes(Lunden，1937)．In the
absence of P，D controls the red of the tubers，flowers
and buds(Dodds& Long，1955，1956)．Genotype dd
procuces pink tubers，pink sprouts and white flowers
(Dodds& Long，1955)．Harborne(1960)thought P and
R al1 detemaine the kind of the anthocyanins．However，
the epistatic interactions among D，P and R &re not elu—
cidated completely．P is epistatic to R(Dodds& Long，
1955)．In tubers，P is also epistatic to D，but in flowers，
P is incompletely epistatic to D(Dodds＆ Long，1955，
1956)．Lunden(1974)thought，for the flower COlOrS in
diploid potatoes，P is epistatic to D．However，van Eck
et“Z．(1993)believed D is epistatic to P．
(2)In the absence of dorninant E，F and R，the
presence of a dominant D can be reflected by the weak
brownish—red of the sprout tips，stems，flower stalks and
calyx(Lunden，1937)．
(3)When E is dominant，ii regulates the presence of
red or purple hue in t uber epidermis in combination with
D or P，respectively(van Eck et a1．，1994)．Genotype
ppddE-produces the slender reddish color of the tuber，
with stronger pigm entation in the bud eyes and at the
sprout bases(Lunden，1937；van Eck et a1．，1994)．
(4)D、F and P are unlinked，they control the in—
heritance of the flower colors in diploid potatoes，and F
is regulaled by D and P(Lunden，1974)．Genotype D
ppF—provides redish purple flowers，daPLF_light blue
f]owers。 P—F- blueish purple flowers and ddppF-
or—一 ff white flowers(Lunden，1937)．Genotype DDff
produces anthocyanins throughout the plant，DDF-and
DD厂厂lcads to white flowers(van Eck et a1．，1994)．
(5)F-R—leads to self—coloured flowers， R—
leads to flecked flowers which are white flowers
with some pigmentation due to leaky aleles of ff
(van Eck et a1．，1994)．
2．3．2 Interactions among J，P，R and Y I，R and P are
the three classical jocj which are involved in the the
coloration of the tuber skin in diploid potatoes(Dodds
& Long，1955，1956)．The diploid I，R and P mimic
functionally and are likely alelic to the tetraploid D，R
and P，respectively，which was originaly described by
Salaman(1 9 1 0)． Genotype iippR—offer red sprouts
with white tuber skin(Dodds& Long，1955)．de Jong
(1 9 9 1)further summarized the variations of the red，
purple and white tubers in diploid and tetraploid pota—
toes are all controlled by the unlinked R，P and J．ru—
bets with genotype I-PpRr or J P r7．have purple skin
while those with genotype I-ppR—are red．Potatoes
lacking a functional allele at J produce white tubers，ir—
respective of the aleles present at R and P． On the
other hand，j ust because the stem tubers of colored po—
tatoes contain carotenoids as well as anthocyanins
(Lewis et a1．，1998；Brown et a1．，1993，2003；Brown，
2004)，de Jong et a1．(2oo4b)believed the variations of
the tuber coloration is synthetically controlled by four
independent and unlinked loci，i．e．R，P，I and y．
2．3．3 Interactions among D ，P and J J is epistatic to D
and P(Harborne，1960)． It epistaticaly controls the
presence and absence of the pigm entation of the tuber
skin and flesh even when D and P are present(de Jong et
a1．，2004b；Jung et a1．，2005)．In the absence of a domi —
nant D．dominant J can lead to the pink of tuber skin
(van Eck et a1．，1994)． Genotype DDiiP-offer purple
sprouts wi th white tuber skin(Dodds Long，1955)．
2．3．4 Interactions among D，P and Ac Generally，D，P
and Ac colectively control the production of the antho—
cyanins(Harborne，1960)．
2．3．5 Interactions among B，F and J B，F and J are
tightly linked，they all control the distribution of the
pigments in potato plants(Dodds＆ Long，1955)．
2．3．6 Interaction between Pf and I Pf is linked to I
3期 赵昶灵等：彩色马铃薯色素相关基因座的种类 、功能与染色体定位 383
(Harborne，1960；de Jong，1987)．For the anthoeyanin
distribution in the tuber of diploid potatoes，Pf con—
trols the flesh color(de Jong，1987)．
2．3．7 Interaction between Pd and Pv Pd and Pv are
both dominant genes．They are linked，and the distance
between them is 4O mapping units(Kessel& Rowe，
1974；Garg et a1．，1981)．
3 Chromosomal locations of the gene
loci involved in the production and dis—
tribution of colored potato pigments
Along with the development of Amplified Frag—
ments Length Polymorphism(AFLP)technology and the
establishment of the genetic map of potato(Bonierbale et
a1．，1988；Gebhardt et aZ．，1989)，Locus D，E，F，I，P，
PSC。R and Y have been detected to locate on the specif—
ic chromosome of potato，and E，F，I and PSC al con—
formably 1ocate on Chromosome 10(Table 5)．
％ ble 5 Chromosomal locations of the gene loci
involved in the production and distribution
of colored potato pigments
4 Discussion
In recent years，colored potato anthocyanins have
displayed a broad applying perspective in modern socie—
ty．On one hand，as natural colorants，the anthocyanins
not only endue the tubers with various and peculiar
coloration patterns but also are regarded as the good
alternatives to synthetic dyes(Francis，1989；Opheim ＆
Andersen，1992；Bridle& Timberlake 1997；Brown et
a1．，2003；Brown，2004)．On the other hand，the an—
thocyanins have been proved to be provided with mul—
tiple pharmacological activities，such as antioxidant
(Brown，2004，2005；Lachman& Hamouz，2005；Reyes
et a1．，2OO5)，antivirus(Hayashi et a1．，2003)and anti—
cancer(Hayashi et a1．，2006；Reddivari et a1．，2007)．
Therefore，increasing the anthocyanin content has been
one of the important targets in the breeding practice of
potatoes(Brown et a1．，2003；Brown，2005)．
A full understanding of the gene loci involved in
the production and distribution of the pigments，espe—
cially the anthocyanins，of colored potatoes will not on—
ly provide a profound level to explore the mechanism
by which the anthocyanins synthesize and accumulate
in vivo but also underlie the work to create the potato
cuhivars with new genotypes．The previous work that
the coloration phenotypes of the stem tuber and other
parts of potatoes were due to the extence of the parti—
cular gene loci is necessary and elementary，but it is su—
perficial and insufficient．The elucidation of the chro—
mosomal locations of the gene loci and the enzymes and
ptotein factors encoded by the loci is the further and
critical step to explain the color-form ing of potatoes．
Later researches on the sequence structures and ex—
pression properties of the genes should be the cogent
basis to create the new genotypes of potatoes with high
functional and commercial values for the food，nutra—
ceutica1。cosmetic and medicinal industries．
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