全 文 ：Journal of Forestry Research, 17(3): 171–176 (2006)

171


Significance of the leaf epidermis fingerprint for taxonomy of
Genus Rhododendron

WANG Xiu-wei 1, MAO Zi-jun *1, CHOI Kyung 2, PARK Kwang-woo 2
1 Key Laboratory of Forest Plant Ecology Ministry of Education China, Northeast Forestry University, Harbin 150040, P. R. China
2 National Arboretum, Korea Forest Service, Pochen-Gun, 487-820, Korea

Abstract: Leaf epidermal fingerprints of six species of Rhododendron（Rh. Aureaum, Rh. dauricum, Rh. micranthum, Rh. Mucronulatum,
Rh. Redowskianum, Rh. schlippenbachii）were observed by optical microscope with nail polish expression method in Key Laboratory of
Forest Plant Ecology of Ministry Education China in Northeast Forestry University in 2004. The leaf morphological features including of
stomata types, characters of guard cells, subsidiary cells in lower epidermis were observed. And ordinary cells (in shape and anticlinal walls
feature) as well as the trichomes in both sides of the leaves are described in detail. The results showed that there were three types of stoma in
six investigated Rhododendron species, from which Pericytic stomata type exists in three species (Rh. dauricum, Rh. micranthum, and Rh.
mucronulatum), Anomocytic stomatal type in Rh. Redowskianum, Diacytic stomata type in Rh. aureaum and Rh. schlippenbachii. The sub-
sidiary cells of the Pericytic and Diacytic stomata type are different in shape and surface feature between the species, respectively. The or-
dinary epidermal cells show a variety from quadrangular to hexagonal, polygonal or irregular in surface view, the anticlinal walls are
straight or sinuose. Trichomes (gland scales) are present in the both of the leaf sides in three species (Rh. dauricum, Rh. micranthum, and Rh.
mucronulatum). All of these detail leaf features show specific specificity of leave finger print for 6 rhododendrons.
Keywords: Taxonomy Significance; Epidermal Fingerprint; Stomata; Trichome; Rhododendron.
CLC number: Q949.772.3 Document code: A Article ID: 1007-662X(2006)03--0171-06


Introduction
Rhododendron (Ericaceae) is considered as a large genus (Hu
et al. 1994; Fang 1999) because of having more than 850 differ-
ent natural species (Zhang 2003), of which 8 species distribute in
northeast of China (Liu et al. 1955). These species are native to
the temperate regions of Asia, North America, and Europe, as
well as to the tropical regions of Southeast Asia and northern
Australia, largest number of which are native to Asia, where they
occur in a variety of habitats, and exhibit an enormous diversity
in size and shape.
A classification system of rhododendron first imposed by
Linnaeus in 1764 has been developed and is also constantly be-
ing revised as new discoveries appear. New discoveries lead
botanists to refine the classification system of the plants. Many
studies have been conducted on classification of rhododendrons
by morphological characteristics, such as leaf, flower, trichome
(gland scale), seed, and so on (Hu et al. 1994; Fang 1999; Hede-
gaard 1980a, 1980b). Now the genus Rhododendron has system-
atic problems at various classification levels, owing to the enor-
mous number of species and conflicting ideas of classification
proposed by many taxonomists (Kurashige 2001). Developing
new approaches to identify the species characteristics is neces-
sary for rhododendron taxonomy.

Foundation project: This work was supported by the Korea Research Foun-
dation and The Korean Federation of Science and Technology Socities Grant
funded by Korea Government (MOEHRD, Basic Research Promotion Fund),
(Project No. 032-4-1).
Biography: WANG Xiu-wei (1981-), Ph. D. student in Key Laboratory of
Forest Plant Ecology Ministry of Education China, Northeast Forestry Uni-
versity, Harbin 150040, P. R. China. E-mail: wxgreat@126.com
Received date: 2006-02-23;
Accepted date: 2006-05-19;
Responsible editor: Zhu Hong
*Corresponding author: Email: park1035@foa.go.kr
This article for cuticular analysis provides a proof of biologi-
cal classification. The taxonomic significance of epidermal
morphology is well documented in botanical literature (Dehgan
1980). Some particular groups of plants or taxa seem to be char-
acterized by specific type of epidermal features, which are the
epidermis, stomata, gland and trichomes (Park 1994; Hong et al.
1999; Hong et al. 2000). Many distinct patterns of stomata have
been found in epidermis of different plants. Stomata in epidermal
morphology can be considered as a classified feature depending
on patterns of stomata and subsidiary cells in a particular plant
(Metcalfe 1950; Shtromberg 1956; Van Cotthem 1970b;
Fryns-Claessens et al. 1973; Rasmussen 1981; Prabhakar 2004).
The peculiar types of stomatal in dicotyledons have been reported
in various families such as in Acantheceae (Paliwal 1966),
Leguminosaea (Edeoga et al. 1996), Nyctaginaceae (Edeoga et al.
2001) and Commelinaceae (Edeoga et al. 2001). Stomatal char-
acteristic, dermal appendages have often been used as morpho-
logical markers for plant taxonomy (Paliwal 1969a; Van Cotthem
1970a; Guyot 1971; Leelavathi 1980; Jelani et al. 1990; Ferzana
et al. 1991).
Epidermal cells may potentially adopt certain fates through a
cell lineage based mechanism or a cell interaction mechanism
(Glover 2000). The ordinary epidermal cells can vary in shape,
size, arrangement and anticlinal wall (straight or sinuose, thin or
thick) in different species. Thus, it is possible that different spe-
cies of plant show specific fingerprints due to their different cell
lineages.
Leaf epidermal fingerprint consists of different typical cells,
not only specialized cells, such as stomatal complex cells (guard
cells and subsidiary cells, if present), trichome and other differ-
ent kinds of epidermal appendages, but also a large number of
ordinary cells. We have a hypothesis that these various types of
epidermal cells combine in different ways in different species
and show specific specificity which may be considered as indi-
cators for classification of plant species.
WANG Xiu-wei et al.

172
However, so far no detailed epidermal morphological studies
have been conducted on all of rhododendron species though
some researchers had done some anatomical researches for some
other species of this genus (Copeland 1943; Cox 1948). The
main objective of the present work is to investigate and describe
the epidermal micromorphological fingerprint of 6 Rhododen-
dron species in order to provide more detailed descriptions to
discuss their taxonomic significance.
Material and Methods
Material
Mature leaf materials used for this study were collected from
living trees and herbarium specimens (Table 1). Voucher speci-
mens of 6 species are deposited in the Herbarium of Key Labo-
ratory of Forest Plant Ecology of Ministry Education China at
Northeast Forestry University (HNEFU). The six species (Rho-
dodendron aureum Georgi., Rh. dauricum Linn., Rh. micranthum
Turcz., Rh. mucronulatum Turcz., Rh. redowskianum Maxim.
and Rh. schlippenbachii Maxim.) were native in Northeast of
China, Korea, Far East of Russia and two of them in Japan (Rh.
dauricum, Rh. mucronulatum).

Table 1. A list of the studied species and their origin
Collecting place Species
Specimen Fresh leaf
Specimen
Number*
Rh. aureum HNEFU, 8649
Rh. dauricum HNEFU
HCFPG
DM 600 m a.s.l.
DM, 700m a.s.l.
8654
Rh. micranthum HNEFU HCFPG
Living plant introduced
from Dangong in 2003
Rh. mucronulatum HNEFU HCFPG
Living plant introduced
from Dangong in 2003
Rh. redowskianum HNEFU 1616
Rh. schlippenbachii HNEFU 20294
Notes: HNEFU--- Herbarium of Key Laboratory of Forest Plant Ecology of
Ministry Education China at Northeast Forestry University; HCFPG----Harbin
City Forest Plant Garden; DM---Daxxing’an Mountain
*Voucher specimens of the plants are deposited in the HNEFU

Epidermal impressions
The epidermal impressions were made with nail polish method.
Leaf portions between second-order veins of leaf in width of
0.5−1.0 cm from the middle were selected respectively. The two
sides of the leaf samples were painted with clear nail polish re-
spectively. After the nail polish had dried thoroughly, a square of
very clear strip was selected and was gently peeled from the leaf
completely and attached to a clear plastic package tape piece,
and then a leaf impression was made. The leaf impressions were
taped to a clean glass slide respectively for observation under
microscope.

Observation
Leaf impression was observed for the lower and upper leaf
surface with a light microscope (Olympus BH-2, Japan) fitted
with a digital camera (Olympus C-5060) at a magnification of
200 and 400 times. For accuracy, several fields of the different
leaf impressions were examined. Observed characteristic pa-
rameters are shape, arrangement, Anticlinal wall features of or-
dinary epidermal cell, stomatal complex (guard cells and sub-
sidiary cells), trichome (gland scale).
The terms about stomatal features used in this study are ac-
cording to the nomenclature and classification by Metcalfe (1950)
and Prabhaka (2004) mainly as follows.
Stomatal pore: An opening in the epidermis surrounded by a
pair or more guard cells.
Stoma: Stomatal pore and a pair of guard cells. Stomatal com-
plex: stoma surrounded by subsidiaries (stomatal types).
Subsidiary cells: cells surrounding a pair of guard cells in one
or more cycles; subsidiaries of the cycle abutting on stoma may
or not be distinct from the adjacent epidermal cells.

Results

The observation result shows that, the upper epidermises of
the leaves consist of ordinary epidermal cells in the six species.
Stomatal complex distributed in lower epidermis is found in all
of the 6 species and shows paracytic type, anomocytic type and
diacytic type, respectively. The pavement epidermal cells and
stomatal complex combine in different ways in different species
and show specific conserved in the six Rhododendron species
(Table 2). Trichom (gland scale) was found in three of the spe-
cies (Rh. dauricum, Rh. micranthum, Rh. mucronulatum) in the
both side of the leaves.

Table 2. Epidermal Characteristics of six Rhododendron species
Shape of Epidermal cell / anticlinal wall Species
Upper epidermis Lower epidermis
Stomatal
Type
Trichome Type in both
leaf sides
Rh. aureum Lathy or elongated rectangular/ slight sinuos Irregular / unclear Diacytic Absent
Rh. dauricum Irregular / sinuos Pentagona, quadrangular or hexago-nal / slightly sinuos or nearly straight
Paracytic Multicellular
Circular shape in top view
Rh. micranthum Nearly honeycomb / nearly straight Quadrangular to polygonal/ near straight
Paracytic Multicellular
Circular shape in top view
Rh. mucronulatum Quadrilateral, pentagonal or hexagonal / slight sinuos
Irregular / sinuos Paracytic Multicellular
Circular shape in top view
Rh. redowskianum Rectangular or square / near straight Irregular long shaped / sinuos Anomocytic Absent-
Rh. schlippenbachii Quadrilateral, pentagonal or hexagonal / straight Irregular / sinuos Diacytic Absent-

Rh. aureaum
Lower epidermis: The ordinary epidermal cells are irregular
shape and unsharp image (Fig. 1a). It may be owing to a thick
cerolipoid layer covered in the epidermis. Stomata are broad
elliptical in shape, and are considered as diacytic type according
to Matcalfe and Chalk (1950) and Prabhakar (2004). A pair of
guard cells is surrounded by two distinct subsidiary cells，which
are the same in shape and size (Fig. 1a, Fig. 2a). Occasionally
2−3 stomata may become laterally contiguous (Fig. 2b).
Upper epidermis: The ordinary cells of the epidermis are small
and arranged in irregular lathy in shape or appear to be elongated
rectangular in surface view. The anticlinal walls are most slight
sinuosity or near straight (Fig. 3a.).

Journal of Forestry Research, 17(3): 171–176 (2006)

173
Rh. dauricum
Lower epidermis: The most of ordinary epidermal cells appear
to be pentagonal and rarely quadrangular or hexagonal in surface
view. The anticlinal walls are most slight sinuosity or near
straight (Fig.1b). Stomata are oblong elliptse or circule in shape,
are considered as peracytic type according to Prabhakar (2004).
In this type, a pair guard cells are surrounded by a single narrow
subsidiary cell, irregularly shaped, which is indistinct from the
guard cells (Fig. 2 d,e). But occasionally the guard cell sur-
rounded by a few small subsidiary cells was found, which are
indistinct from the guard cells. Occasionally 2-3 stomata may
show laterally contiguous or polar contiguity. Rarely are the
contiguous stomata arranged at right angles to each other (Fig. 2f)
or irregularly. Gland scales are found in lower epidermis of
Rh.dauricum. These gland scales distribute randomly and circu-
larly shaped. The scale consists of two kinds of cells. The border
cells are long shaped surround several central cells, which are in
irregular shape and smaller than border cells (Fig. 2c).
Upper epidermis: the epidermal cells are arranged randomly,
irregular in shape. The anticlinal walls are mostly sinuous. The
similar Gland scales in lower epidermis distributed randomly and
in upper epidermis also are found (Fig. 3b).

Rh. micranthum
Lower epidermis: the ordinary epidermal cells appear to be
quadrangular to polygonal in shape. The anticlinal walls are near
straight (Fig. 1c). There are lipoid drops accumulating in some of
the cells (Fig. 2g). Stomata are elliptical in shape and show as
Pericytic Type according to Prabhakar (2004). A pair of guard
cells is surrounded by a single distinct narrow subsidiary cell,
which are monocyclic multiangular (Fig. 2g) or radial in shape
(Fig. 2h). Occasionally there is two or more subsidiary cells
around the guard sells. Two stomata arrange laterally contiguous
(Fig. 2i) or irregularly in direction, in which cellulose molecular
bundles in radial are found. It may be owing to the cuticle layer
thickening (Fig. 2g-i). Gland scales distribute in lower epidermis
of Rh. micranthum randomly (Fig. 2l). The composition of these
gland scales is similar with those of Rh. duricum.
Upper epidermis: the epidermis cells arrange regularly and the
shape of the cells are similar with honeycomb shape. The anti-
clinal walls are clearly, honeycomb. Gland scales also distribute
in lower epidermis (Fig. 3c).

Rh. mucronulatum
Lower epidermis: the ordinary epidermal cells appear to be ir-
regular in shape. The anticlinal wall is sinuous (Fig. 1a). Stomata
are elliptical in shape, Pericytic Type according to Prabhakar
(2004). A pair of guard cells is surrounded by a single distinct
narrow subsidiary cell, which are monocyclic irregular in shape.
The subsidiary wall abutting against the stoma is indistinct
(Fig.2j). Occasionally 2−3 stomata may become contiguous ar-
ranged irregularly (Fig. 2k). Gland scales are found in lower
epidermis of Rh.mucronulatum, the composition of which is
similar with those of Rh. duricum.
Upper epidermis: The epidermal cell arranged orderly rela-
tively and appear to be quadrilateral, pentagonal or hexagonal in
surface view. The anticlinal walls are straight and clear. Broad
dark strips impresses are found in the cells (Fig. 3d). The similar
gland scales are found in upper epidermis also (Fig. 3d).



Fig. 1 Lower epidermis impressions of six Rhododendron species.
a. Ordinary epidermal cells, stomatal complex of Rh. aureum b. Ordinary epidermal cells, stomatal complex and cells group of a trichome of Rh.
dauricum. c. Ordinary epidermal cells, stomatal complex and cells group of a trichome of Rh. micranthum. d. Ordinary epidermal cells, stomatal complex
and cells group of a trichome of Rh. mucronulatum. e. Ordinary epidermal cells, stomatal complex of Rh. redowskianum. f. Ordinary epidermal cells,
stomatal complex of Rh. schlippenbachii.

WANG Xiu-wei et al.

174
Rh. redowskianum
Lower epidermis: The ordinary epidermal cells are irregular
pattern in shape, but generally showed long shape. The anticlinal
walls are sinuous (Fig. 1e). Stomata are oblong in shape, are
considered as nomocytic type according to Matcalfe and Chalk
(1950), in which the guard cells are surrounded by a certain
number of cells that do not differ in size and shape from other
epidermal cells (Fig. 2m). The stomata are oriented in the direc-
tion to extend parallel to lateral vine of the leaf (Fig. 1f).
Upper epidermis: The epidermal cells arrange orderly and ap-
pear to be rectangular or square in surface view. Generally two
cells arranged in lateral direction and two in longitudinal direc-
tion alternately. The anticlinal walls are near straight (Fig. 3e).
Rh. Schlippenbachii
Lower epidermis: the ordinary epidermal cells are irregular in
shape. Dark sinuate leptonema strias impress are allover densely
in the cells (Fig. 1f). Stomata are elliptical in shape. The stomatal
type is Diacytic type (Matcalfe and Chalk 1950, Prabhakar 2004).
The two abutting subsidiaries cells, which are not the same in
shape and size (showed no conjoint walls towards both poles),
indistinct, parallel to the guard cell, and subsidiary cells con-
joined with the guard cells with distinct radial cellulose molecu-
lar bundles (Fig. 2n). 2−3 stomata may become laterally con-
tiguous or may show polar contiguity or irregularly arranged
each other (Fig .2o).
Upper epidermis: the epidermal cells arrange closely and ap-
pear to be quadrilateral, pentagonal or hexagonal in surface view.
Intercellular spaces are clearly. Cell anticlinal walls are straight.
Sinuate leptonema strias impress are allover densely in the cells.



Fig. 2 Stomatal complex and trichome impressions of six species Rhododendron.
a. stomata and b. laterally and irregularly contiguous stomata of Rh. aureaum. c. trichome in lower epidermis in top view. d. e. Stomatal complex. f. two
perpendicularly contiguous stomata of Rh. dauricum. g,h. Stomatal complex. i. two laterally contiguous stomata of Rh. micranthum. j. stomatal complex,
k. two laterally contiguous stomata and l. trichome in lower epidermis of Rh. mucronulatun in top view. m. stomata of Rh. redowskianum. n. stomatal
complex and o. four contiguous stomata of Rh. schlippenbachii.
Journal of Forestry Research, 17(3): 171–176 (2006)

175


Fig.3 Upper epidermis impressions of six Rhododendron species.
a. Ordinary epidermal cells of Rh. aureum b. Ordinary epidermal cells and gland scale cells group of Rh. dauricum. c. Ordinary epidermal cells and gland
scale cells group of Rh. micranthum. d. Ordinary epidermal cells and gland scale cells group of Rh. mucronulatum. e. Ordinary epidermal cells of Rh.
redowskianum. f. Ordinary epidermal cells of Rh. schlippenbachii.

Fig.4 Leaf epidermis impressions of Rhododron dauricum from three populations distributed in different locations.
a, d. Show the lower epidermis and upper epidermis from Daxingan mountain (51°N,124°E, 600m hight), respectively. b, e. Showing the lower epidermis
and upper epidermis from Daxingan mountain (51°N, 124°E, 700 m at height). c,f. Showing the lower epidermis and upper epidermis from Harbin city



Discussion

The epidermal fingerprint analysis in this work provides the
first detailed description of leaf features for six species of
Rhododendron. The results show distinct different epidermal
fingerprints among the six species. The features of ordinary epi-
dermal cells, guard cells, subsidiary cells and trichones show
specific specificities and have significance for classification of
Rhododendron in species level.

WANG Xiu-wei et al.

176
According to the morphological classification by Nakai (1917),
the investigated six species belong to 5 sections respectively,
which are section Osmothumnus Maxim. (Rh. micranthum),
section Rhodorastrum Maxim.(Rh. dauricum, Rh. mucronulatum),
Section Therorhodion (Maxim.) Drude.(Rh. redowskianum),
section Eurhododendron (DC.)A. Gray. (Rh. aureaum) and sec-
tion Azalea (Linn.) Maxim. (Rh. schlippenbachii). The obtained
epidermal impression features of the 6 species are coincident
with morphological classification. Rh. dauricum, and Rh.
mucronulatum have similar epidermal fingerprint (stomata are
Pericytic type, subsidiary cells are similar in shape (Fig. 2d,j))
comparing with other four species, indicating the close relation-
ship in the same section. The epidermal fingerprint of Rh. mi-
cranthum is similar with that of Rh. dauricum and Rh. mucronu-
latum in a certain extent (stomata is Pericytic type, ordinary cells
in lower epidermis are similar with those of Rh. dauricum in
shape). It is coincident with morphological classification that the
three species have gland scales in both sides of the leaves.
Among the examined six species, three of them is peracytic
somata type (Rh. dauricum, Rh. micranthum, and Rh. mucronu-
latum), 2 of them is Diacytic type (Rh. aureaum and Rh. Schlip-
penbachii). But the subsidiary cells of them are very different in
shape, size and the surface feature in same stomatal type. So
these detail characters should be considered as indicators of
natural taxonomic affinity for the taxonomical study of Rhodo-
dendron.
Considering the possibility of ecological effect on the leaf
features of the same species between different populations, a
comparative study has been done for the leaf fingerprint between
three populations of Rh. dauricum, distributing in Daxingan
Mountain (51°N, 124°E, at latitude of 500 m and 700 m) and in
Harbin city (45°N, 127°E, at latitude of 400 m), respectively. The
result shows that the qualitative characters of the epidermis (the
shape, anticlinal walls of ordinary cells in the both side, stomatal
types, trichomes) are the similar among the three populations
(Fig. 4 a-f), though some differences in quantity parameters, such
as stomatal size, stomatal density appear to be different between
populations. This result indicates the specific stability in leaf
fingerprint in the six Rhododendron species.
In summary, these comparative data of leaf fingerprint are sig-
nificant to suggest that leaf epidermal fingerprints of Rhododen-
dron have highly conserved specificity and can be considered as
the key character for the classification of Rhododendron. For the
morphological similar species, it will also be significant to com-
bine with evidences from scanning electronic microscope (SEM)
and other studies such as cytology, palynology, anatomy and
molecular biology, etc. in order to arrive at a concrete delimita-
tion of taxa.

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Chinese Abstracts 1
《林业研究》（英文）2006年第 17卷第 3期
中文摘要
(Chinese abstracts attached to Journal of Forestry
Research, Vol. 17, No.3 (2006))

06–03–001
叶片表皮指纹在杜鹃花属分类中的意义/王秀伟，毛子军(东北
林业大学森林植物生态学教育部重点实验室，哈尔滨
150040)，CHOI Kyung，PARK Kwang-woo (National Arbo-
retum, Korea Forest Service, Pochen-Gun, 487-820)// Journal of
Forestry Reseach.–2006, 17(3): 171–176.
本实验于 2004年在东北林业大学进行。用指甲油印记法
在光学显微镜下观察了杜鹃花属 6 个种（牛皮杜鹃 Rh.
aureaum，兴安杜鹃 Rh. dauricum, 照白杜鹃 Rh. micranthum,
迎红杜鹃 Rh. mucronulatum，苞叶杜鹃 Rh. Redowskianum和
大字杜鹃 Rh. schlippenbachii）的表皮指纹特征。对叶片的形
态特征包括叶片下表皮气孔类型、保卫细胞和副卫细胞的特
征、叶片两面的普通细胞（形状和垂周壁特征）和表皮毛进
行了较详细的描述。所研究的 6 种杜鹃花中，有 3 种气孔类
型，其中兴安杜鹃、照白杜鹃、和迎红杜鹃为单细胞型气孔
（Pericytic型），苞叶杜鹃为不规则型气孔（Anomocytic型），
牛皮杜鹃和大字杜鹃是平轴式气孔（Diacytic式）。不同种间，
单细胞型和平轴式气孔的副卫细胞类型在形状和表面特征上
有所不同。普通表皮细胞的表面观为从四边形、六边形, 多
边形或不规则型, 垂周壁呈平直或弯曲状。兴安杜鹃、照白
杜鹃和迎红杜鹃 3 种的叶片两面均有腺鳞。所有这些叶片特
点表明上述 6 种杜鹃花的表皮指纹具有种的特性。图 4 表 2
参 29。
关键词：分类意义；表皮指纹；气孔；毛状体；杜鹃花属
CLC number: Q949.772.3 Document code: A
Article ID: 1007−662X(2006)03−0171–06

06–03–002
沙地樟子松天然林和人工林中针叶养分含量比较/朱教君（中
国科学院沈阳应用生态研究所，沈阳 110016，中国），谭辉，
康宏樟（中国科学院沈阳应用生态研究所，沈阳 110016，中
国；中国科学院研究生院，北京 100039，中国），许美玲（中
国科学院沈阳应用生态研究所，沈阳 110016，中国）//Journal
of Forestry Research.–2006, 17(3): 177–184.
为了诊断沙地樟子松人工林的衰退原因，我们比较分析
了沙地樟子松天然林和人工林中的叶片养分含量。结果表明，
天然林中的樟子松叶片内氮、磷含量均比人工林中低，而钾
含量比人工林高。樟子松人工林叶片中 N: P, P: K和 N: K在
45年前随年龄增加而增加，从植物养分含量的角度来看，这
一结果表明氮或磷可能不是沙地樟子松生长的绝对限制因
子。然而，天然林和人工林叶片中的钾含量比以前报道过的
松属植物中的含量都要低。天然林叶片中的 N: P比值在正常
的范围内，但人工林的则在该范围以外。这些结果表明天然
林具有比人工林更好的养分平衡状况。如果只从樟子松的生
长地来考虑，我们可以认为樟子松人工林的衰退现象可能是
因其它的矿物元素或者氮和磷的有效性，而不是氮和磷的绝
对含量不足造成的。养分的不平衡状况和人工林内针叶快速
分解可能也是衰退的重要因素。图 3表 3参 42。
关键词：衰退现象；沙地森林生态系统；氮磷比；樟子松；
樟子松天然林；樟子松人工林
CLC number: S791.253 Document code: A
Article ID: 1007-662X(2006)03-0177-08

06–03–003
NaCl胁迫对青檀种子发芽率、叶气体交换和苗木生长的影响
/方升佐，宋立奕，洑香香(南京林业大学森林资源与环境学院，
南京 210037, 中国)//Journal of Forestry Reseach.–2006, 17(3):
185–188.
在水培条件下，本文探讨了 NaCl 胁迫对青檀种子发芽
率、叶气体交换和苗木生长的影响。研究结果表明，NaCl胁
迫延长了青檀种子的发芽时间，特别是当 NaCl 浓度超过 17
mM (1.0g/L)时，发芽率明显降低；NaCl胁迫降低了青檀叶的
呼吸速率、气孔导度和净光合速率，并随着 NaCl胁迫浓度的
提高，下降的幅度更大，但 NaCl 胁迫苗木胞间隙 CO2浓度
明显高于对照；与对照相比，在 NaCl胁迫 50天后，青檀苗
木的成活率、苗高和地径生长以及根系、茎和叶生物量显著
降低，但茎和叶生物量下降更为明显。本文的研究结果认为，
青檀是一种对盐胁迫较敏感的植物，一年生幼苗的耐盐阈值
在 34 mM (2.0 g/L)左右。图 2表 3参 27。
关键词：盐胁迫；Hoagland 培养液；发芽率；净光合速率；
成活率；生物量增量。
CLC number: Q945.78 Document code: A
Article ID: 1007−662X(2006)03−0185–04

06–03–004
森林土壤有机氮的 X 光吸收近边结构光谱技术测定/庄舜尧
(中国科学院南京土壤研究所，土壤与农业可持续发展国家重
点实验室，南京 210008), 徐梦洁(南京农业大学公共管理学
院，南京 210095), 胡正义(中国科学院南京土壤研究所，土
壤与农业可持续发展国家重点实验室，南京 210008) //Journal
of Forestry Reseach.–2006, 17(3): 189–192.
本研究通过采用氮的 K边 X光吸收近边结构（XANES）
技术来了解森林土壤有机氮，以此深入研究土壤中有机氮种
类与其转化的定量关系。土壤样品采自台湾中部的云杉、铁
杉林与草地。结果表明，氮的 XANES 可以揭示样品中不同
的氮的种类。在土壤腐植质、可溶性氮及本体土壤中胺态及
吡咯氮占了主要的比率。然而不同处理及植被下的土壤样品
氮的种类分布是不同的。云杉与铁杉土壤可溶性有机氮在
402.3 eV能量处有显著的差异。在 A层土壤中，吡啶类氮含
量要显著高于 O层土壤，说明氮在不同土层中的转化率存在
极大的差异，这种变化对于氮在森林土壤中的循环将起重要
作用。图 3表 1参 8。
关键词: 森林土壤；氮的 X光近边吸收光谱；氮的种类；
有机氮；氮的转化
CLC number: S714.5 Document code: A
Article ID: 1007−662X(2006)03− 0189–04