全 文 ：珍贵树种红椿与毛红椿遗传生
理生态研究
赵伯金 1，甄爱国 2，贺润轩 1，程德华 3，汪洋 4*
（1.建始县林业局，湖北建始 445300；2.英山县
林业局，湖北英山 438700；3.湘潭第一中学，
湖南湘潭 411100；4. 湖北生态工程职业技术
学院，湖北武汉 430200）
摘 要 红椿和毛红椿都是国家二级保护用
材树种。 其在我国零星分布，种群规模小，但发
展潜力巨大。 对红椿和毛红椿的遗传表现性状
选优和毛红椿的分子遗传标记以及红椿和毛
红椿的生理与生态研究方面进展进行了概述，
对相关研究利用提出了建议和前景展望。
关键词 红椿；毛红椿；遗传；生理；生态
基 金 项 目 湖 北 省 公 益 性 研 究 项 目
（402012DBA40001）。
作者简介 赵伯金（1966-），男，湖北建始人，
工程师， 主要从事林业技术工作及研究，E-mail：
289355637@qq.com。 *通讯作者，汪洋（1968-），
男，湖北武汉人，副教授，硕士，主要从事生态
恢复与植物种质资源教学与研究 ，E-mail：
13986200308@163.com。
收稿日期 2015-03-05
修回日期 2015-04-15
Research on Genetic Physiological Ecology of
Precious Species: Toona ciliata and Toona ciliata
var. pubescens
Bojin ZHAO1, Aiguo ZHEN2, Runxuan HE1, Dehua CHENG3, Yang WANG4*
1. Jianshi Forestry Administration, Jianshi 445300, China;
2. Yingshan Forestry Administration, Yingshan 438700, China;
3. Xiangtan First Middle School, Xiangtan 411100, China;
4. Hubei Vocational College of Ecological Engineering, Wuhan 430200, China
Supported by Public Welfare Research Projects of Science and Technology Bureau of
Hubei Province (40 2012DBA40001).
*Corresponding author. E-mail: 13986200308@163.com
Received: February 5, 2015 Accepted: April 15, 2015A
Agricultural Science & Technology, 2015, 16(5): 997-1001
Copyright訫 2015, Information Institute of HAAS. All rights reserved Forestry
T oona ciliata belongs to genusToona (Meliaceae family).Toona ciliata is a deciduous or
semi-evergreen megaphanerophyte
and is a heliophyte with straight trunk,
tender and straight tree performance.
Toona ciliata var. pubescens belongs
to genus Toona (Meliaceae family) as
well. Toona ciliata var. pubescens is a
deciduous megaphanerophyte with
straight trunk and obvious natural
training. Toona ciliata and Toona cilia-
ta var. pubescens are precious and
fast-growing timber species in tropic
and subtropics of China. Toona ciliata
and Toona ciliata var. pubescens are
key protected wild plants of national
levelⅡ, which are called “Chinese ma-
hogany” [1 -2], and the timbers of which
are superior for furniture. Toona ciliata
distributes naturally in India, Burma,
Laos, Pakistan, Thailand, Malaysia,
Indonesia, Australia, etc. Toona ciliata
distribute in low mountains and hills in
South China and the geographic coor-
dinate range is 100°16′-119°40′ east
longitude and 24° 21′ -30° 31′ north
latitude[3]. Toona ciliata is found to be
distributed in Nanjiang, Bazhong,
Sichuan, which locate at 106°93′ lon-
gitude and 32° 42′ latitude with the
elevation of 620 m[4]. This area is the
natural distribution area of Toona cili-
ata with the highest latitude. In 2013, A
well-preserved natural population of
Toona ciliata with more than one thou-
sand trees was found in Gucheng
County, Hubei Province located at
111° 24′ east longitude and 32° 10′
north latitude. Toona ciliata var.
pubescens is an endemic tree of
China and distributes in Sichuan,
Guizhou, Fujian and Anhui with a ver-
tical distribution of 500-2 500 m[5].
The distribution of Toona ciliata
and Toona ciliata var. pubescens are
not wide in China and they distribute
sporadically with small population size
but with great development potential [6].
The number of Toona ciliata and
Toona ciliata var. pubescens decreas-
es gradually because of overexploita-
tion and slow natural regeneration[3].
Germplasm resource, genetic
Abstract Toona ciliata and Toona ciliata var. pubescens are timber species of na-
tional second-class protective plants. In China, Toona ciliata and Toona ciliata var.
pubescens distribute sporadically with small population size but with great develop-
ment potential. Plus selection on phenotype of Toona ciliata and Toona ciliata var.
pubescens, molecular genetic marker of Toona ciliata var. pubescens, research
progress in physiology and ecology of Toona ciliata and Toona ciliata var.
pubescens were summarized in this paper. Suggestions and prospect forecast were
proposed for relative research and utilization.
Key words Toona ciliata; Toona ciliata var. pubescens; Heredity; Physiology; Ecology
DOI:10.16175/j.cnki.1009-4229.2015.05.030
Agricultural Science & Technology 2015
structure, plant physiology, ecology,
breeding, cultivation, afforestation,
pharmaceutical, chemical constituent
and resource protection of Toona cili-
ata and Toona ciliata var. pubescens
were widely studied by more and more
domestic scholars in recent years be-
cause they were endangered species
with excellent varietal characteristics.
The research progresses of Toona cil-
iata var. pubescens and Toona ciliata
were combined in the paper for the
supplementation research of Toona
ciliata because the biological charac-
teristics of Toona ciliata were the same
with as Toona ciliata var. pubescens [5]
and the research on Toona ciliata was
relative insufficient.
Research on the Genetics
Direction of Toona ciliata
The methods for the detection of
plant genetic diversity are improved
and perfected ceaselessly with the de-
velopment of biology, especially ge-
netics and molecular biology. The
methods have developed from the ini-
tial phenotypic variation, subsequent
chromosomal polymorphism and pro-
tein polymorphism to present DNA
polymorphism.
Genetic expressed characters
Using morphology or phenotypic
character to detect genetic variation
are the most direct, simple and accu-
rate methods. Liu et al. studied the
variation patterns of geographic
source of seedling period and
tanezane growing characters accord-
ing to the analysis of 11 phenotypic
characters such as fruit quality, fruit
length and two growth characters in-
cluding seedling height and ground di-
ameter in 13 provenances in the natu-
ral distribution areas of Toona ciliata
var. pubescens. Growth characters at
seedling period of Toona ciliata var.
pubescens were mainly controlled by
the level of above or medium on the
provenance level; broad-sense heri-
tability and variable coefficient of the
height of one-year-old seedlings were
the biggest; broad-sense heritability
and variable coefficient of ground di-
ameter were the second while the
broad-sense heritability and variable
coefficient of tanezane were not obvi-
ous. Proposing that Toona ciliata var.
pubescens could be preliminarily
selected in different provenances with
the seedling height and ground di-
ameter, and ideal results could be
obtained[7].
Zhang et al. used six half-sib fam-
ilies of Toona ciliata in Enshi, Hubei as
study materials; significant differences
existed in seedling height, ground di-
ameter, leaf length, leaf width, ratio of
length to breadth of one-year-old
seedlings and elite families were se-
lected. Introduction adaptability in the
area with similar latitude and different
longitude was tested based on differ-
ent characters and overall perfor-
mance[8]. Song et al. used 46 half-sib
families of Toona ciliata in Jiangxi,
Yunnan, Chongqing, Sichuan etc. as
study objects for seedling stage ex-
periment. Six characters including
seedling height, ground diameter,
crown breadth, main root length, main
root width, number of tertiary root of
one-year-old seedlings were mea-
sured to provide experimental founda-
tion for genetic selection. The research
indicated that highly significant differ-
ences existed in seedling height,
ground diameter and crown breadth
among Toona ciliata families and the
three characters were controlled by in-
tension inheritance. Biggish genetic
improvement effect could be obtained
by combined selection of seedling
height and ground diameter[9]. To some
extent, root parameters are important
indicators for the evaluation of
seedling stage growth and genetic
breeding of half-sib families of Toona
ciliata.
Selection of superior tree is one of
the most effective and reliable meth-
ods for elite breeding of forest. Plus
selection on progenies of elite trees is
the guarantee for sustainable heredity
of excellent provenances. The space
and potential for genetic improvement
and elite breeding are big because of
the abundant variations among half-sib
families in germplasm families. Wen et
al. studied the growth expression of
seedling height and ground diameter
at seedling stage of progeny seedlings
of 20 one-year-old elite Toona ciliata.
The results showed that highly signifi-
cant differences existed in the growth
of seedling height and ground diame-
ter among the 20 families of Toona cil-
iata. Fast-growing families of Toona
ciliata with good quality can be
screened by selective breeding etc.
Twenty percent fast-growing elite fam-
ilies of Toona ciliata during seedling
stage were selected based on the
analysis results of the growth of
seedling height and ground diameter
at seedling stage of one-year-old
seedlings of Toona ciliata [10]. Liu used
progenies of elite Toona ciliata var.
pubescens as objects for introduction
research and seedling height, ground
diameter, dry weight of root, stem
mass, ratio of root and stem, total
length of root, root surface area, root
volume were analyzed at seedling
stage. Highly significant differences ex-
isted in seedling height, ground diam-
eter, dry weight of root, stem mass and
ratio of root and stem, which indicated
that there were big differences among
various trait heritability of Toona ciliata
var. pubescens. The heritability of
seedling height was the highest, and
the heritabilities of ground diameter,
dry weight of root, stem mass and ratio
of root and stem were medium while
the heritabilities of total length of root,
root surface area, root volume were
relative low[11].
Modern digital technology has the
advantages of high efficiency and con-
venience for the plus selection of
Toona ciliata at seedling stage. Fer-
reira et al. used best linear unbiased
prediction(BLUP) at the individual level
with multiplicating and combined tradi-
tional prediction technology with digital
image software (Image-j) to estimate
genetic parameter and genetic value
of Toona ciliata by the processing of
complex and unstable data, which had
practicability, speediness and high effi-
ciency for genetic breeding[12].
Molecular marker technique
Morphological marker, a simple
method, can be easily affected by en-
vironmental conditions and dominance
or recessive of genes, and the genetic
expression of which is not stable
sometimes. DNA molecular marker
technique can fast and accurately an-
alyze plant genetic diversity for more
genetic information, which overcomes
the long period and unstabilizing fac-
tors of traditional analysis method.
In the area of molecular marker
technique, traditional magnetic beads
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Agricultural Science & Technology2015
enrichment (a method for the isolation
of microsatellite DNA in genome) had
been improved and microsatellite DNA
had been isolated from the genome of
Toona ciliata var. pubescens, and the
microsatellite DNA had been se-
quenced; SSR primers were designed
and synthesized, and SSR reaction
system was optimized to provide tech-
nical support for further research of
genetic structure and genetic variation
of Toona ciliata var. pubescens popu-
lations [13]. SSR molecular marker had
been applied to the analysis of genetic
diversity of Toona ciliata var.
pubescens populations. The genetic
differentiation coefficient of Toona
ciliata var. pubescens populations
was 0.185 4 and the gene flow (Nm)
among populations was 1.098 3, which
illustrated that the genetic variations
among Toona ciliata var. pubescens
populations were low; the genetic dis-
tances among populations were sig-
nificantly correlated with geographical
distances [14]. The research of genetic
structures of three natural populations
of Toona ciliata var. pubescens with
microsatellite marker indicated that the
genetic diversities among the natural
populations of Toona ciliata var.
pubescens were low; the gene flow
was higher than the average level of
contrastive perennial plant while the
genetic differentiation coefficient was
lower than the average level; one of
the three natural populations had spa-
tial genetic structure while the genetic
variation of alleles of the other two
natural populations lacked spatial
genetic structures[15]. Genetic diversity
research of central populations and
peripheral populations indicated that
the genetic diversities of peripheral
populations of Toona ciliata var.
pubescens were higher than the cen-
tral populations; the genetic differenti-
ation degrees of central populations
were obviously smaller than the pe-
ripheral populations; the genetic dis-
tances of populations were not signifi-
cantly related with geographical dis-
tances and the genetic differentiations
among populations might resulted
from terrain[16].
The conservation of biological di-
versity namely to protect the genetic
diversity because the genetic diversity
of a species is closely related with its
viability and competitiveness[6, 17]. The
plant population sizes of natural re-
served Toona are very small with
scattered distribution and the charac-
teristics of the populations include very
little seedlings, low genetic diversities,
small variations among natural plants
and bad renewal ability, which implies
serious recession of inbreeding[18]. The
low level of genetic diversities of natu-
ral populations of Toona ciliata var.
pubescens might be related with fre-
quent human activities on the popula-
tions and the severe human interven-
tion resulted in decreased genetic di-
versity [14]. Therefore, the conservation
of natural populations of Toona ciliata
var. pubescens should be enhanced.
Research on Physiology
and Ecology
Plant physiology is a science for
the research of laws governing life ac-
tivities of plants including plant growth
and development, morphogenesis,
conversion of matter and energy, infor-
mation transfer, signal transduction
and the influence of environment con-
ditions in vivo and in vitro on plant vital
activity. Physiological studies of Toona
ciliata are mainly focused on the area
of stress, photosynthesis and nutrient.
Physiological study
Great progress has been
achieved for physiological study of
Toona ciliata in recent years. Drought
stress is mainly studied in the area of
stress resistance research. Wu et al.
performed drought stress experiment
with different periods on one-year-old
potted seedlings of five clones of
Toona ciliata in April; the leaf relative
water content of young clone
seedlings of Toona ciliata was the
lowest under the severe stress(water-
break for 17 d after normal water ap-
ply); the differences of relative water-
ing content among different clones
were not significant; leaf chlorophyll
content of young seedlings was the
highest and the differences of leaf
chlorophyll content of seedlings a-
mong different clones were not signifi-
cant; leaf chlorophyll content of full pe-
riod clone seedlings of Toona ciliata
showed an increasing tendency [19].
Research on leaf net photosynthetic
rate, stomatal conductance, intercel-
lular CO2 concentration, chlorophyll
content and leaf area of Toona ciliata
var. pubescens seedlings under the
treatments of three soil moistures
(high, medium, low) with full exposure,
60% shading and 80% shading re-
vealed that daily photosynthetic
progress of Toona ciliata var.
pubescens under full exposure and
moderate shading in summer at
seedling stage were “two peaks”; high
luminous intensity and high tempera-
ture at noon contributed to obvious
phenomenon of photosynthesis “noon
break”; soil moisture content was pro-
portional to leaf area under the same
light condition; seedlings of Toona cili-
ata var. pubescens increased leaf area
and improved chlorophyll content to
effectively use light radiation and to
form ecology survival countermeasure
for the adaption to shading condition[20].
Water stress experiment with the
clones of Toona ciliata seedlings indi-
cated that clones with strong resistant
ability to drought could be screened by
the variation tendency of chlorophyll
content, MDA content, proline content,
SOD activity and POD activity of clone
seedlings [21]. Plus selection of promo-
tion lines with the combination of sur-
vival rate of cuttage, rooting percent-
age during summer and autumn and
amount of growth of one-year-old
seedlings was a feasible method for
Toona ciliata breeding [21]. Foliar SOD
and POD were significantly different
among different clones with different
drought stress treatment. Foliar SOD
contents of clone seedlings of Toona
ciliata were the highest (606.83 U/g)
under medium stress (water-break for
12 d after normal water apply) and fo-
liar SOD contents of different clone
seedlings were not significantly differ-
ent under various periods; foliar POD
contents of seedlings were the highest
under severe stress (water-break for
17 d after normal watering apply) and
foliar POD contents of clone seedlings
were not significantly different, and fo-
liar POD content decreased following
the test period[22].
Shortage of microelement was
against the growth of height and di-
ameter, dry matter accumulation of
Toona ciliata M. Roem var. australis
and the most direct effect was resulted
from the boron deficiency. Boron defi-
ciency resulted in withered tender
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Agricultural Science & Technology 2015
leaves and morphological change of
shoot and root. Manganese deficiency
resulted in foliar roll-up and slight yel-
lows; copper deficiency resulted in
blue point on leaves and wilted leaves;
zinc deficiency resulted in shortened
internode and decrescent lanceolate
leaves; iron deficiency resulted in
slow growth of plants and yellows of
tender leaves[23]. Leaf length, leaf area,
biomass, content and accumulation of
Pb in each organ were different when
potted seedlings of one-year-old
Toona ciliata grew on acid purple soil,
calcareous purple soil and atteration in
the Southwest exposed to Pb stress
with different concentrations (0, 200,
450 mg/kg and 2 000 mg/kg), and the
resistances and transfer efficiencies of
Pb contamination by Toona ciliata
were different. As a fast-growing tim-
ber species with resistance to Pb,
Toona ciliata could be used as a pio-
neer species for the remediation of Pb
contaminated soil in the Southwest
because of the ability of Pb absorption
and Pb accumulation[24].
Ecological study
Ecology is a science to analyze
and solve environmental problems
faced by species and adaptability of
plants under different environments.
Toona ciliata is a deciduous
species and new shoot has not
sprouted on leafless culm top in early
March; it lasts 260 d from germination
to leaf fall. Toona ciliata seedling is
shade tolerant and updated well under
crown canopy, and root sprouting
ability of Toona ciliata is strong. Toona
ciliata sprouts in late March, blooms in
June to July, matures in mid to late
October, falls leaves in late November
and it takes 260 d in all. Toona ciliata
often mixes plant with Machilus lepto-
phylla, Quercus dentata, Lisea core-
ana, Liquidambar formosana, Maackia
amurensis , Pteroceltis tatarinowii [ 25 ]
and also mixes with Toona ciliata var.
pubescens. Toona ciliata grow well on
fertile, moist, well drained open forest
land, forest edge and ravined valley
with deep soil layer[26].
The size of plant population af-
fects its life potential and small popu-
lation size is one of the characteristics
of endangered populations [27]. Toona
plants distribute dispersedly with bad
update ability and the natural reserved
population sizes are very small. In or-
der to reveal interrelationships among
populations in Toona ciliata var.
pubescens community, to reveal the
structures and functions of popula-
tions, to understand the development
laws of populations, typical sample
plots of different geographic positions,
of different soil conditions and different
community compositions were select-
ed to measure various ecological fac-
tors and to set quadrat for the analysis
of inter specific associations of main
species in Toona ciliata var. pube-
scens community; Fu et al. found that
the overall association of multiple
species in Toona ciliata var. pube-
scens community was significant posi-
tive correlation but the inter specific
association between Toona ciliata var.
pubescens and plants of tree layer
was not significant, and the popula-
tions were facing interspecific com-
petition, selection and succession of
environment[28].
The threatening factors of Toona
ciliata are shown as follows: low germi-
nation rate of seed, slow growth at
seedling stage, small population size,
serious inbreeding depression, seri-
ous human disturbance, disruptive
biotope[18]. Shen investigated Toona cil-
iata resources, especially Toona ciliata
var. pubescens in Huaping, Yunnan.
Quadrat points were randomly select-
ed and quadrats were extracted in
each quadrat points based on different
elevations (locations); quantity, age
group, ecological regime of Toona cili-
ata in quadrats were recorded. The
relative density d(d=n/s) of Toona cili-
ata was calculated within quadrats to
acquire total distribution[29]. Endanger-
ing factors were proposed based on
the species characteristics and distri-
bution characteristics. In situ conser-
vation was proposed as the main
safeguard procedure and ex situ con-
servation was proposed as the assis-
tant safeguard procedure while scien-
tific research monitoring and protection
publicity should be strengthened[18, 29].
Suggestions and Prospects
Toona ciliata is a national second-
class protective wild plant, a precious
fast-growing timber species and is one
of the preferred species for tree
species structure adjustment of mixed
coniferous broad leaved forests. Cur-
rently, molecular markers are only
used for the research of genetic struc-
ture of Toona ciliata var. pubescens
but not used for Toona ciliata var.
pubescens breeding. Research on ge-
netic characters of Toona ciliata has
still been restricted to phenotypic vari-
ation and genetic research on molecu-
lar level has not been reported. More
abundant genetic information can be
obtained by genetic diversity analysis
of Toona ciliata with DNA molecular
marker technique to study geographi-
cal provenance variation. Molecular
genetic structure and molecular ge-
netic breeding of Toona ciliata are ur-
gent to be researched on.
Ecology research on population
dynamics, on population structure and
spatial pattern, on geographical prove-
nance variation and selection of supe-
rior trees of Toona ciliata has rarely
been reported. The development and
utilization of Toona ciliata and Toona
ciliata var. pubescens, especially uti-
lization research on chemical and
pharmacological have an extensive
foreground.
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