Plantactinospora mayteni gen. nov., sp. nov., a member of the family Micromonosporaceae

Abstract

A novel Gram-positive, aerobic, spore-forming, endophytic actinomycete, designated strain YIM 61359(T), was isolated from the roots of Maytenus austroyunnanensis plants collected from tropical rainforest in Xishuangbanna, Yunnan Province, south-west China. The strain formed single or cluster spores with smooth surfaces from substrate mycelia. The strain contained meso-diaminopimelic acid in the cell wall and arabinose, xylose, galactose and glucose in whole-cell hydrolysates. The acyl type of the cell-wall polysaccharides was glycolyl. MK-10(H(6)), MK-10(H(8)) and MK-10(H(4)) were the predominant menaquinones. The polar lipids were phosphatidylethanolamine, diphosphatidylglycerol, phosphatidylinositol and several unknown phospholipids. The major fatty acids were iso-C(15 : 0), anteiso-C(15 : 0), C(17 : 0), anteiso-C(17 : 0) and iso-C(16 : 0). The DNA G+C content of strain YIM 61359(T) was 69.7 mol%. These chemotaxonomic data indicated that the strain belongs to the family Micromonosporaceae. Phylogenetic analysis based on 16S rRNA gene sequences also suggested that strain YIM 61359(T) fell within the family Micromonosporaceae. On the basis of morphological and chemotaxonomic data, phylogenetic analysis and characteristic patterns of 16S rRNA gene signature nucleotides, strain YIM 61359(T) is considered to represent a novel species of a new genus within the family Micromonosporaceae, for which the name Plantactinospora mayteni gen. nov., sp. nov. is proposed. The type strain of Plantactinospora mayteni is YIM 61359(T) (=CCTCC AA 208022(T)=DSM 45238(T)).

Full text

Plantactinospora mayteni gen. nov., sp. nov., a
member of the family Micromonosporaceae
Sheng Qin,
1
Jie Li,
1
Yu-Qin Zhang,
1,2
Wen-Yong Zhu,
1
Guo-Zhen Zhao,
1
Li-Hua Xu
1
and Wen-Jun Li
1
Correspondence
Wen-Jun Li
wjli@ynu.edu.cn or
liact@hotmail.com
1
The Key Laboratory for Microbial Resources of the Ministry of Education, PR China and Laboratory
for Conservation and Utilization of Bio-resources, Yunnan Institute of Microbiology,
Yunnan University, Kunming, Yunnan 650091, PR China
2
Institute of Medicinal Biotechnology, Chinese Academy of Medical Sciences and Peking Union
Medical College, Beijing 100050, PR China
A novel Gram-positive, aerobic, spore-forming, endophytic actinomycete, designated strain YIM
61359
T
, was isolated from the roots of Maytenus austroyunnanensis plants collected from
tropical rainforest in Xishuangbanna, Yunnan Province, south-west China. The strain formed
single or cluster spores with smooth surfaces from substrate mycelia. The strain contained meso-
diaminopimelic acid in the cell wall and arabinose, xylose, galactose and glucose in whole-cell
hydrolysates. The acyl type of the cell-wall polysaccharides was glycolyl. MK-10(H
6
), MK-10(H
8
)
and MK-10(H
4
) were the predominant menaquinones. The polar lipids were
phosphatidylethanolamine, diphosphatidylglycerol, phosphatidylinositol and several unknown
phospholipids. The major fatty acids were iso-C
15 : 0
, anteiso-C
15 : 0
,C
17 : 0
, anteiso-C
17 : 0
and
iso-C
16 : 0
. The DNA G+C content of strain YIM 61359
T
was 69.7 mol%. These
chemotaxonomic data indicated that the strain belongs to the family Micromonosporaceae.
Phylogenetic analysis based on 16S rRNA gene sequences also suggested that strain YIM
61359
T
fell within the family Micromonosporaceae. On the basis of morphological and
chemotaxonomic data, phylogenetic analysis and characteristic patterns of 16S rRNA gene
signature nucleotides, strain YIM 61359
T
is considered to represent a novel species of a new
genus within the family Micromonosporaceae, for which the name Plantactinospora mayteni gen.
nov., sp. nov. is proposed. The type strain of Plantactinospora mayteni is YIM 61359
T
(5CCTCC
AA 208022
T
5DSM 45238
T
).
The family Micromonosporaceae was first described by
Krasil’nikov (1938), and its description has been subse-
quently emended by Goodfellow et al. (1990), Koch et al.
(1996) and Stackebrandt et al. (1997) on the basis of
chemotaxonomic data and 16S rRNA gene sequence
analysis. At the time of writing, the family Micro-
monosporaceae comprises 19 genera: Micromonospora
(Ørskov, 1923), Actinoplanes (Couch, 1950), Pilimelia
(Kane, 1966), Dactylosporangium (Thiemann et al., 1967),
Catellatospora (Asano & Kawamoto, 1986), Catenuloplanes
(Yokota et al., 1993), Couchioplanes (Tamura et al., 1994),
Spirilliplanes (Tamura et al., 1997), Verrucosispora (Rheims
et al., 1998), Virgisporangium (Tamura et al., 2001), Asanoa
(Lee & Hah, 2002), Longispora (Matsumoto et al., 2003),
Salinispora (Maldonado et al., 2005), Actinocatenispora
(Thawai et al., 2006), Polymorphospora (Tamura et al.,
2006), Luedemannella (Ara & Kudo, 2007a), Krasilnikovia
(Ara & Kudo, 2007b), Planosporangium (Wiese et al., 2008)
and Pseudosporangium (Ara et al., 2008).
Endophytic actinomycetes have attracted increasing atten-
tion in recent years, but they remain relatively unexplored
as potential sources of novel species and novel natural
products for medical and commercial exploitation. Some
species of the plant genus Maytenus are known to produce
the anti-cancer compound maytansine (a 19-membered
macrocyclic lactam) (Reider & Roland, 1984), and
endophytic micro-organisms and related secondary meta-
bolites have been investigated (Zhao et al., 2007). During
our investigations of the diversity and taxonomy of rare
actinomycetes associated with tropical rainforest medicinal
plants from Xishuangbanna, Yunnan Province, south-west
China, strain YIM 61359
T
was isolated from the roots of
healthy Maytenus austroyunnanensis plants. On the basis of
its 16S rRNA gene sequence, this isolate falls phylogeneti-
The GenBank/EMBL/DDBJ accession number for the 16S rRNA gene
sequence of strain YIM 61359
T
is FJ214343.
A table comparing 16S rRNA gene signature nucleotide positions of
strain YIM 61359
T
and related genera is available with the online version
of this paper.
International Journal of Systematic and Evolutionary Microbiology (2009), 59, 2527–2533 DOI 10.1099/ijs.0.010793-0
010793 G2009 IUMS Printed in Great Britain 2527

cally within the family Micromonosporaceae adjacent to the
genera Salinispora and Micromonospora. Data from the
present polyphasic taxonomic study indicate that this
strain represents a novel species of a new genus within the
family Micromonosporaceae.
Healthy root samples of Maytenus austroyunnanensis,a
traditional Chinese medicinal plant, were used as the
source for isolation of endophytic actinomycetes. Samples
were air-dried for 48 h and then washed via an ultrasonic
step to remove surface soil. Subsequently, samples were
subjected to a modified five-step surface sterilization
procedure according to Qin et al. (2008b): an 8-min wash
in 5 % NaOCl, followed by a 10-min wash in 2.5 %
Na
2
S
2
O
3
, a 5-min wash in 75 % ethanol, a wash in distilled
water and a final rinse in 10 % NaHCO
3
for 10 min. After
air-drying at room temperature, surface-sterilized stems
were subjected to air-drying at 80 uC for 30 min, then
aseptically crumbled into small fragments and processed
with a calcium carbonate enrichment method as described
by Otoguro et al. (2001) for 3 weeks. The enriched samples
were serially diluted in sterile distilled water, spread-plated
onto modified cellulose-proline agar [per litre tap water:
2.5 g cellulose, 2.0 g sodium pyruvate, 0.25 g KNO
3
, 1.0 g
proline, 0.2 g MgSO
4
.7H
2
O, 0.2 g K
2
HPO
4
, 0.5 g CaCl
2
,
10 mg FeSO
4
.7H
2
O and 15 g agar (pH 7.2–7.4)]and
incubated at 28 uC for 2 weeks. The isolate was routinely
cultured on International Streptomyces Project medium 2
(ISP 2) (Shirling & Gottlieb, 1966) and maintained as a
glycerol suspension (20 %, w/v) at 280 uC.
Cultural characteristics of strain YIM 61359
T
were tested
on ISP 2, ISP 3, ISP 4, ISP 5, nutrient agar, potato-glucose
agar (PDA) and Czapek’s agar plates, commonly used for
characterization of Streptomyces species (Shirling &
Gottlieb, 1966; Waksman, 1967) at 28 uC for 2–6 weeks.
Colony colour was determined based on ISCC–NBS colour
charts (Kelly, 1964). Observations of mycelia and spores of
strain YIM 61359
T
grown on ISP 2 for 21–45 days were
made by light microscopy (BH 2; Olympus) and scanning
electron microscopy (JSM5600LV; JEOL). Growth was
tested at 0, 4, 10, 15, 20, 28, 37, 40, 45 and 55 uConISP2.
The ability of the strain to grow at different pH and NaCl
concentrations was examined according to Qin et al.
(2008a). Media and procedures used for the determination
of physiological characteristics, carbon source utilization
and acid production from carbohydrates were those
described by Gordon et al. (1974).
Biomass for most of the chemotaxonomic studies was
obtained after cultivation at 28 uC for 7–14 days in shaken
cultures with yeast extract-malt extract broth (ISP 2) or
trypticase soy broth (for fatty acids). Amino acid and sugar
analysis of whole-cell hydrolysates was performed accord-
ing to the procedures described by Hasegawa et al. (1983).
The N-acyl group of muramic acid was determined by the
method of Uchida & Aida (1984). Cellular menaquinones
were extracted and purified as described by Collins et al.
(1977) and were analysed by HPLC (Groth et al., 1997).
Polar lipids were extracted, examined via two-dimensional
TLC and identified by using established procedures
(Minnikin et al., 1979; Collins & Jones, 1980). Analysis of
whole-cell fatty acids followed the instructions of the MIDI
System (Microbial ID) (Sasser, 1990; Ka
¨mpfer &
Kroppenstedt, 1996) by using exponential-phase cultures.
Determination of the DNA G+C content was performed
according to Mesbah et al. (1989).
Genomic DNA extraction, PCR-mediated amplification of
the 16S rRNA gene and sequencing of the PCR products
were carried out as described by Li et al. (2007). 16S rRNA
gene sequence similarities among closely related sequences
obtained from the GenBank/EMBL/DDBJ databases were
calculated manually after pairwise alignments by using the
CLUSTAL_Xprogram, v. 1.8 (Thompson et al., 1997). A
phylogenetic tree was constructed with the neighbour-
joining method (Saitou & Nei, 1987) by using the MEGA 3.1
software package (Kumar et al., 2004). For construction of
the maximum-likelihood tree, the online version of
PhyML (Guindon et al., 2005) was used (http://
www.atgc-montpellier.fr/phyml/). The stability of relation-
ships was assessed by performing bootstrap analyses
(Felsenstein, 1985) of the neighbour-joining data based
on 1000 resamplings.
Strain YIM 61359
T
was an aerobic, Gram-positive, non-
acid-fast actinomycete that formed extensively branched
substrate mycelia and sparse aerial mycelia. Non-motile
spores were borne singly or in clusters from substrate
mycelia. The spore surface was smooth (Fig. 1). Strain YIM
61359
T
grew well on ISP 2, ISP 3 and Czapek’s agar,
moderately well on ISP 4 and PDA, but poorly on ISP 5
and nutrient agar. No soluble pigments were produced on
any of the media tested. Colonies on ISP 2 were orange–
yellow and lacked aerial hyphae initially; after 4 weeks
growth colonies were black, raised and folded. The colour
of colonies on other media was orange–yellow (nutrient
agar/Czapek’s agar), yellowish (ISP 5), pale orange–yellow
to black (ISP 3/PDA) and black (ISP 4). Strain YIM 61359
T
grew well at 15–37 uC and pH 6.0–8.0 but could not grow
in the presence of 4 % NaCl. Other physiological and
biochemical characteristics are presented in the species
description below.
The novel isolate contained meso-diaminopimelic acid as
the diagnostic diamino acid in the cell wall, and the whole-
cell sugars detected were arabinose, xylose, galactose and
glucose. The acyl type of the cell-wall polysaccharides was
glycolyl. The predominant menaquinones were MK-10(H
6
)
(53 %), MK-10(H
8
) (23 %) and MK-10(H
4
) (12 %), with
MK-10(H
2
) (6 %) and MK-9(H
6
) (6 %) as minor compo-
nents. Mycolic acids were not detected. The polar lipids
detected were phosphatidylethanolamine, diphosphatidyl-
glycerol and phosphatidylinositol, corresponding to phos-
pholipid type PII of Lechevalier et al. (1977). The fatty acid
profile of strain YIM 61359
T
comprised iso-C
15 : 0
(20.3 %
of the total), anteiso-C
15 : 0
(13.8 %), C
17 : 0
(12.5 %),
anteiso-C
17 : 0
(12.1 %), iso-C
16 : 0
(10.2 %), C
16 : 0
(6.3 %),
S. Qin and others
2528 International Journal of Systematic and Evolutionary Microbiology 59

C
18 : 1
9c(6.1 %), iso-C
17 : 0
(3.6 %), C
18 : 0
(3.2 %), iso-C
16 : 1
G (2.5 %), iso-C
15 : 1
G (1.8 %), C
17 : 1
9c(1.6 %), anteiso-
C
17 : 1
A (1.5 %), iso-C
14 : 0
(1.0 %), C
19 : 0
(1.0 %), C
15 : 0
(0.9 %), C
16 : 1
9c(0.7 %), iso-C
18 : 0
(0.6 %), C
14 : 0
(0.3 %)
and C
16 : 1
B (0.3 %), which corresponds to fatty acid type
2d of Kroppenstedt (1985). The G+C content of the DNA
of strain YIM 61359
T
was 69.7 mol%.
The almost-complete 16S rRNA gene sequence (1427 bp)
of strain YIM 61359
T
was used for phylogenetic analysis
together with those of members of the family
Micromonosporaceae. 16S rRNA gene sequence compar-
isons revealed that strain YIM 61359
T
was most closely
related to the type strains of members of the genera
Micromonospora (96.6–98.1 % similarity) and Salinispora
(97.7 %) and Polymorphospora rubra (97.6 %).
Phylogenetic analysis based on 16S rRNA gene sequences
showed that strain YIM 61359
T
belonged to a separate
cluster within the family Micromonosporaceae and formed a
distinct monophyletic clade that was different from each of
the 19 recognized genera in this family. It did not cluster
within the genus Micromonospora, but was related to the
genus Salinispora (Fig. 2). The topology of the tree
generated with the maximum-likelihood algorithm was
similar (data not shown). The 16S rRNA gene signature
nucleotide positions of strain YIM 61359
T
were compared
with those of its closest phylogenetic relatives (see
Supplementary Table S1 in IJSEM Online).
Strain YIM 61359
T
formed single or cluster spores from
substrate mycelia, similar to members of the genus
Salinispora, but was clearly different from members of
the genera Micromonospora and Polymorphospora, which
formed only single spores or short spore chains from
substrate mycelia (Ørskov, 1923; Tamura et al., 2006).
Members of the genus Salinispora have MK-9(H
4
) as the
major menaquinone and fatty acid profile type 3a.
Members of the genus Micromonospora have MK-
10(H
4,6
) and MK-9(H
4,6
) as major menaquinones and
fatty acid profile type 3b. The genus Polymorphospora has
MK-10(H
6,4
) and MK-9(H
6,4
) as major menaquinones and
fatty acid profile type 2a. Strain YIM 61359
T
can be
distinguished from the genera Salinispora,Micromonospora
and Polymorphospora based on the major menaquinones
and the fatty acid profile (Table 1). In addition, the 16S
rRNA gene signature nucleotides clearly distinguished the
new isolate from members of these three genera.
Strain YIM 61359
T
was not affiliated with any other
recognized genus of the family Micromonosporaceae based
on the distinctness of its 16S rRNA gene sequence and
phylogenetic position. On the basis of morphological,
chemotaxonomic and physiological data and the signature
nucleotide pattern of the 16S rRNA gene, strain YIM
61359
T
is readily distinguishable from the genera
Salinispora,Micromonospora and Polymorphospora.
Therefore, we suggest that strain YIM 61359
T
represents
a novel species of a new genus within the family
Micromonosporaceae, for which the name Plantactinospora
mayteni gen. nov., sp. nov. is proposed.
Description of Plantactinospora gen. nov.
Plantactinospora (plan.tac.ti.no.spo9ra. L. n. planta a plant;
Gr. n. actis actinos a ray; Gr. n. spora a seed, and in biology
a spore; N.L. fem. n. Plantactinospora pertaining to a spore-
forming actinomycete isolated from plant tissues).
Aerobic, Gram-positive and non-acid-fast actinomycetes.
Cells form extensively branched substrate mycelia (0.16–
0.23 mm in diameter), which carry smooth-surfaced spores
(0.63–1.10 mm in diameter) borne singly or in clusters.
Spores are non-motile. White aerial mycelia are sparse. The
cell wall contains meso-diaminopimelic acid as the
diagnostic diamino acid. Arabinose, xylose, galactose and
glucose are detected as whole-cell sugars. The acyl type of
the cell-wall polysaccharides is glycolyl. Polar lipids
detected are phosphatidylethanolamine, diphosphatidyl-
glycerol and phosphatidylinositol and several unknown
(a)
(b)
(c)
Fig. 1. Scanning electron micrographs of cells of strain YIM
61359
T
growing on ISP 2 after incubation for 5 weeks at 28 6C.
Bars, 10 mm (a), 2 mm (b) and 5 mm (c).
Plantactinospora mayteni gen. nov., sp. nov.
http://ijs.sgmjournals.org 2529

phospholipids, corresponding to phospholipid type PII.
Mycolic acids are absent. The predominant menaquinones
are MK-10(H
6
), MK-10(H
8
) and MK-10(H
4
); MK-10(H
2
)
and MK-9(H
6
) are present as minor components. The
major cellular fatty acids are iso-C
15 : 0
, anteiso-C
15 : 0
,
C
17 : 0
, anteiso-C
17 : 0
and iso-C
16 : 0
(fatty acid type 2d). The
Fig. 2. Neighbour-joining phylogenetic tree
derived from 16S rRNA gene sequences
showing the relationship between strain YIM
61359
T
and the type strains of species of related
genera in the family Micromonosporaceae.
Numbers at branch nodes are bootstrap per-
centages (based on 1000 replications; only
values .50%aregiven).Bar,0.01substitutions
per nucleotide position.
S. Qin and others
2530 International Journal of Systematic and Evolutionary Microbiology 59

Table 1. Differential characteristics between strain YIM 61359
T
and recognized genera in the family Micromonosporaceae
Taxa: 1, strain YIM 61359
T
;2,Salinispora;3,Micromonospora;4,Polymorphospora;5,Pseudosporangium;6,Couchioplanes;7,Krasilnikovia;8,Actinoplanes;9,Luedemannella; 10, Longispora;
11, Actinocatenispora; 12, Asanoa; 13, Catellatospora; 14, Catenuloplanes; 15, Dactylosporangium; 16, Pilimelia; 17, Spirilliplanes; 18, Verrucosispora; 19, Virgisporangium; 20, Planosporangium. Data
for reference genera were taken from Ørskov (1923), Couch (1950), Kane (1966), Thiemann et al. (1967), Asano & Kawamoto (1986), Yokota et al. (1993), Rheims et al. (1998), Kudo et al. (1999),
Tamura et al. (1994, 1997, 2001, 2006), Lee & Hah (2002), Matsumoto et al. (2003), Maldonado et al. (2005), Thawai et al. (2006), Ara & Kudo (2006, 2007a, b), Wiese et al. (2008) and Ara et al.
(2008). +, Present; 2, absent; +/2, variable; ND, no data available.
Characteristic 1 2 3 4 5 6 7 8 9 10 11 12 13 14 15 16 17 18 19 20
Sporangium 222 2 222++22 2 2 2++ 22 ++
Spore motility 222 2 2+2+22222++++22+/2
Diamino acid(s)* m-DAP m-DAP m-DAP m-DAP m- and 3-
OH-DAP
L-Lysine m-DAP m-DAP m-DAP m-DAP m-DAP m-DAP m-DAP L-Lysine m-DAP m-DAP m-DAP m-DAP m-DAP m-DAP
Diagnostic
sugarsD
Ara, Gal,
Xyl
Ara, Gal,
Xyl
Ara, Xyl Xyl Gal, Glu,
Rib, Man,
Xyl, Ara
Ara, Gal,
Xyl
Gal, Man,
Xyl, Ara,
Rib
Ara, Xyl Xyl, Gal,
Man Rha,
Rib, Ara
Ara, Gal,
Xyl
Ara, Gal,
Xyl
Ara, Gal,
Xyl
Xyl, Man,
Gal, Ara,
Rha, Rib
Xyl Ara, Xyl Ara, Xyl Ara, Xyl Man,
Xyl
Ara, Gal,
Xyl
Ara, Xyl
Fatty acid type 2d 3a 3b 2a 2d 2c 2d 2d 2d 2d 3b 2d 3b 2c 3b 2d 2d 2b 2d 3b
Major mena-
quinone(s)
MK-
10(H
6,8,4
)
MK-
9(H
4
)
MK-
10(H
4,6
),
MK-
9(H
4,6
)
MK-
10(H
6,4
),
MK-
9(H
6,4
)
MK-9(H
6
) MK-
9(H
4
)
MK-
9(H
6,4,8
)
MK-
9(H
4
),
MK-
10(H
4
)
MK-
9(H
6,4,2,8
)
MK-
10(H
4,6
)
MK-
9(H
4,6
)
MK-
10(H
6,8
)
MK-
9(H
4,6
)or
MK-
10(H
4,6
)
MK-
9(H
8
),
MK-
10(H
8
)
MK-
9(H
4,6,8
)
MK-
9(H
4,2
)
MK-
10(H
4
)
MK-
9(H
4
)
MK-
10(H
4,6,8
)
MK-
9(H
4
),
MK-
10(H
4
)
Phospholipid
type
PII PII PII PII PII PII PII PII PII PII PII PII PII PIII PII PII PII PII PII PII
DNA G+C
content (mol%)
69.7 70–73 71–72 71 73 70–72 71 72–73 71 70 72 71–72 70–71 71–73 71–73 ND 69 70 71 71.4
*m-DAP, meso-diaminopimelic acid.
DAra, Arabinose; Gal, galactose; Glu, glucose; Man, mannose; Rha, rhamnose; Rib, ribose; Xyl, xylose.
Plantactinospora mayteni gen. nov., sp. nov.
http://ijs.sgmjournals.org 2531

DNA G+C content is approximately 69–70 mol%. The
type species is Plantactinospora mayteni.
Description of Plantactinospora mayteni sp. nov.
Plantactinospora mayteni (may.te9ni. N.L. n. Maytenus a
botanical genus name; N.L. gen. n. mayteni of the plant
genus Maytenus).
Has the following characteristics in addition to those
described for the genus. Colonies develop well on ISP 2,
ISP 3 and Czapek’s agar, moderately well on ISP 4 and PDA,
but poorly on ISP 5 and nutrient agar. Soluble pigments are
not formed. Colony colour varies from yellowish, orange–
yellow, pale orange–yellow to black. Colonies can become
darkened, raised and folded during sporulation. Grows at
15–37 uC and pH 6–8.0; unable to grow in the presence of
4 % NaCl. Optimal temperature and pH for growth are
28 uC and pH 7.0. Degrades adenine, hypoxanthine, starch
and xanthine. Milk is not coagulated or peptonized. Nitrate
and H
2
S are reduced. Utilizes D-arginine, L-asparagine, L-
phenylalanine and L-proline as nitrogen resources. Utilizes
L-arabinose, cellobiose, dulcitol, erythritol, D-fructose, D-
galactose, glucose, inositol, lactose, maltose, mannitol,
mannose, raffinose, L-rhamnose, D-ribose, sorbitol, sucrose,
trehalose and D-xylose as carbon resources. Acid is not
produced from L-rhamnose, inositol or D-fructose.
The type strain, YIM 61359
T
(5CCTCC AA 208022
T
5
DSM 45238
T
), was isolated from surface-sterilized roots of
Maytenus austroyunnanensis collected from tropical rain-
forest in Xishuangbanna, Yunnan Province, south-west
China. The DNA G+C content of the type strain is
69.7 mol%.
Acknowledgements
This research was supported by National Basic Research Program of
China (Project no. 2004CB719601) and Yunnan Provincial Education
Commission Scientific Research Foundation (No.08J0008). W.-J. L.
was supported by Program for New Century Excellent Talents in
University.
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