全 文 ：Received 4 Feb. 2004 Accepted 20 Aug. 2004
Supported by the Natural Science Foundation of Fujian Province (B0110007).
* Author for correspondence. E-mail: .
http://www.chineseplantscience.com
Acta Botanica Sinica
植 物 学 报 2004, 46 (11): 1316-1323
Dynamics of Leaf Mass, Leaf Area and Element Retranslocation Efficiency
During Leaf Senescence in Phyllostachys pubescens
LIN Yi-Ming*, PENG Zai-Qing, LIN Peng
(School of Life Sciences, Xiamen University, Xiamen 361005, China)
Abstract: Dynamics of leaf mass (LM), leaf area (LA) and element retranslocation efficiency during leaf
senescence was investigated in Phyllostachys pubescens Mazel ex H. de Lehaie in Yongchun, Fujian,
China. Comparison of differences in element retranslocation efficiencies (RE) based on per gram leaf dry
weight, per leaf and per LA during leaf senescence was carried out. With leaf senescence, the mean
decreases of LM, LA and specific leaf mass (SLM) were 19.55%, 15.16% and 5.07%, respectively. The
seasonal changes in decrease percentage of LM and LA were similar, indicating that certain mass to area
ratios occurred in P. pubescens leaves. On different bases, RE of N and K was positive, while RE of Ca was
negative, suggesting that with leaf senescence, N and K were translocated out of senescing leaves to
other parts of plant, while Ca accumulated in senescing leaves. For the mean RE of N, P, K, Ca and Mg on
different bases, the rank order was RE2 (mg element/leaf)＞RE3 (mg element/cm2 leaf)＞RE1 (mg
element/g), therefore, RE on the basis of leaf weight or LA would be underestimated.
Key words: Phyllostachys pubescens ; leaf mass (LM); leaf area (LA); element retranslocation efficiency;
leaf senescence
Retranslocation from senescing leaves is the process
by which plants withdraw elements from these leaves, mak-
ing them available for later investment in new structure
(Millard and Neilsen, 1989; Aerts, 1996). This increases the
use of absorbed elements and reduces plant dependence
on soil supply (Pugnaire and Chapin, 1993). The process of
retranslocation is closely associated with leaf senescence
and conservation of elements, and is an important mecha-
nism enabling plants to maintain growth in element-poor
sites (Fife and Nambiar, 1997; Lin and Wang, 2001; Lodhiyal
and Lodhiyal, 2003). Elements may be used more efficiently
in element-poor sites, and this efficient element use could
be important to the survival of individuals in such sites
(Birk and Vitousek, 1986; Aerts, 1995). High element
retranslocation efficiency (RE) and low growth rate are the
characteristics of plants under element-poor conditions
(Boerner, 1984; Lajtha, 1987; Ralhan and Singh, 1987).
However, some researches reported that high RE is not an
important adaptation to low element status, but a charac-
teristic of most plant species with contrasting life histories
(Chapin and Kedrowski, 1983; Miao, 2004). Plants growing
on infertile soils do not retranslocate a greater fraction of
elements from senescing leaves, i.e. RE is independent of
status of individuals (Birk and Vitousek, 1986; Chapin and
Moilanen, 1991; Walbridge, 1991; Helmisaari, 1992). RE was,
however, found to be high under higher element status
(Nambiar and Fife, 1987).
The absence of consistent results could reflect the dif-
ferent ways conducted. Low-element-adapted species on
infertile soils were frequently compared to high-element-
adapted species on fertile soils, which made it difficult to
differentiate phenotypic and genotypic responses to soil
fertility (Pugnaire and Chapin, 1993). Some of the results
reported RE on a concentration basis (mg/g) (Chapin and
Kedrowski, 1983; Cote et al., 1989; Schlesinger et al., 1989;
Lodhiyal and Lodhiyal, 1997; Lodhiyal and Lodhiyal, 2003),
others on a milligrams element per leaf or fascile basis
(mg/leaf) (Nambiar and Fife, 1987; Ralhan and Singh, 1987;
Dalla-Tea and Jokela, 1994) or milligrams per unit leaf area
(mg/cm2) basis (Killingbeck, 1985; Pugnaire and Chapin,
1993), and even on a whole-canopy basis (kg/hm2) (del-
Arco et al., 1991). Few studies reported the differences of
RE using different bases (Birk and Vitousek, 1986; del-Arco
et al., 1991; Delucia and Schlesinger, 1995; Lin and Wang,
2001; Covelo and Gallardo, 2002). And only few report (Lin
and Wang, 2001) recorded changes in leaf area during leaf
senescence. To answer the question whether elements are
retranslocated out of senescing leaves, the changes in the
absolute contents (mg/leaf) of elements during leaf senes-
cence should be measured to assess retranslocation. Our
aim of the present study is to compare the RE on three
bases: mg/g, mg/leaf and mg/cm2.
LIN Yi-Ming et al.: Dynamics of Leaf Mass, Leaf Area and Element Retranslocation Efficiency During Leaf Senescence in
Phyllostachys pubescens 1317
Phyllostachys pubescens, a species native to China and
the giant woody bamboo, has been widely used for food,
handicraft, furniture, and construction materials in many
parts of the world. In recent years, research on P. pubescens
has been focused on the diversity of species, succession,
matter recycling and energy flow in their ecosystems (Li
and Lin, 1995; Isagi et al., 1997; Torii and Isagi, 1997; Li et
al., 2000; Peng et al., 2002). Until now, element
retranslocation efficiency during leaf senescence in P.
pubescens has not been reported. We studied this species
in Yongchun County, Fujian Province, China, concentrat-
ing on the changes in leaf mass, leaf area, and element
retranslocation efficiency during leaf senescence, so as to
have a better understanding of the characters and ecologi-
cal adaptations of P. pubescens and it will be beneficial to
relative nutrient diagnosis. Pugnaire and Chapin (1993)
adopted an instantaneous sampling method to determine
element RE in several Mediterranean evergreen trees. This
method was used by Schlesinger et al. (1989) and Delucia
and Schlesinger (1995). Dalla-Tea and Jokela (1994) and
del-Arco et al. (1991) adopted a periodical sampling method.
The other aim of the current paper is to compare differ-
ences between the two methods.
1 Materials and Methods
1.1 Site
Research was conducted at Yidu Town, Yongchun
County (25°21¢33²-25°31¢33² N, 117°40¢40² E), Fujian,
China. The climate is subtropical monsoon. The average
annual temperature is 18.3℃, the average annual precipita-
tion is 1 724 mm, the frost-free season lasts for 310 d in this
region. The mountain red soil is about 90 cm in depth, with
3-5 cm humus layer, pH 5.0-5.1, N 0.102%-0.293%, P
0.042%-0.075%, and an organic C content 3.45%-3.65%
in the upper of 30 cm. The characteristic of P. pubescens
forest was described in previous report (Peng et al., 2002).
The coverage of dense forest was 0.8, tree density was 26.
8 trees /100 m2, and mean height and base diameter were 12
m and 8 cm, respectively. The understory consisted of
Engelhardtia fenzelii and Ilex pubescens with Pteridium
aquilinum var. latiusculum and Paris polyphylla in herb layer.
1.2 Materials
Fifty individuals of 2-year-old were chosen and labeled.
The height and living conditions of the chosen trees were
similar. From April 2002 to March 2003, ten pieces of mature
leaves and ten pieces of senescent leaf from the same shoot
on the upper canopy of each labeled tree were collected
monthly. “Mature leaf” was a leaf which was fully expanded,
just prior to the onset of the leaf senescence (determined
by appearance). “Senescent leaf” was a leaf which was
ready to abscise. Leaves collected from the same species
were pooled, resulting in one sample for mature leaves and
one sample for senescent leaves each month. Leaves dam-
aged by insects or by mechanical factors were avoided.
1.3 Chemical analysis
The collected leaves were taken to the laboratory and
the leaf area was measured by LI-3000A portal leaf area
instrument (LI-COR). The leaves were washed with dis-
tilled water, dried at 80 ℃, ground in a mill to pass a 1-mm
sieve and stored for chemical analyses.
Subsamples of leaves were digested in sulfuric acid-
hydrogen peroxide, and N was determined by the
microKjeldahl method (Yoshida et al., 1972), P was deter-
mined by acorbic acid-antimony reducing phosphate colo-
rimetric method (Murphy and Riley, 1962). Samples were
digested in nitric acid-perchloric acid, and K, Ca and Mg
determinations were made with Model WFX-ⅠB Atomic
Absorption Spectrophotometers (Beijing Analytical
Instrument). All element determination had two to three
replicates, and the error between two replicates were less
than 5% for K, Ca and Mg, and less than 2% for N and P,
respectively.
1.4 Calculation
Expression of RE on concentration basis (mg element/g,
RE1), content basis (mg element/leaf, RE2), and leaf area
basis (mg element/cm2 leaf, RE3) is as follows (Lin and Wang,
2001):
RE1=（1－ A2/A1×W1/W2）× 100%
RE2=（1－ A2/A1）× 100%
RE3=（1－ A2/A1× S1/S2）× 100%
Where A1 is the mean element content (mg) per mature leaf;
A2 is the mean element content (mg) per senescent leaf; S1
is the mean area (cm2) per mature leaf; S2 is the mean area
(cm2) per senescent leaf; W1 is the mean mass (g) per ma-
ture leaf; and W2 is the mean mass (g) per senescent leaf.
The element contents (mg/leaf) were calculated by mul-
tiplying the element concentrations (mg/g) of the leaves by
the mass (g/leaf) of the leaves. Sometimes RE was negative.
This indicated that the element accumulated in senescing
leaves.
2 Results
2.1 Monthly changes in leaf mass, leaf area and specific
leaf mass
Monthly dynamics in leaf mass (LM), leaf area (LA) and
specific leaf mass (SLM) are shown in Fig.1. There were
seasonal changes in LM and LA in P. pubescens from April
2002 to March 2003. LM of mature leaves was ranged from
Acta Botanica Sinica 植物学报 Vol.46 No.11 20041318
76.89 to 123.06 mg/leaf, averaged (87.28±12.88) mg/leaf;
those of senescent leaves varied from 59.02 to 89.76 mg/
leaf, with the average of (69.96±9.45) mg/leaf. Meanwhile,
LA of mature leaves was from 7.10 to 11.39 cm2/leaf, aver-
aged (8.47±1.12) cm2/leaf. That of senescent leaves was
from 5.74 to 9.06 cm2/leaf, with the average of (7.17±0.98)
cm2/leaf. LM and LA of mature leaves were higher than
those of senescent leaves (P < 0.01), with few exception.
Monthly changes in SLM were not as significant as those
in leaf mass and leaf area. SLM was from 9.68 to 11.17 mg/
cm2 (with the mean of (10.30±0.46) mg/cm2) for mature
leaves, and from 9.07 to 10.52 mg/cm2 (with the mean of
(9.77±0.44) mg/cm2) for senescent leaves. The coefficient
of variation of SLM (4.47% for mature leaves and 4.50% for
senescent leaves) was much lower than those of LM
(14.76% and 13.51%) and LA (13.22% and 13.67%).
2.2 Changes in LM and LA during leaf senescence
With leaf senescence, the mean decreases of LM, LA
and SLM were 19.55%, 15.16% and 5.07%, respectively.
The changes in SLM were less than those in LM and LA.
Decreases in LM, LA and SLM were from 10.19% to
27.06%, from 0.23% to 24.32%, and from –1.51% to 10.14%,
respectively. The seasonal changes in decrease
percentage of LM and LA were similar (Fig.2), indicating
that the certain mass to area ratios occurred in P. pubescens
leaves. As shown in Table 1, there were significant correla-
tions between LM and LA of P. pubescens from April 2002
to March 2003.
2.3 Element concentrations (mg/g DW) in mature leaves
Generally speaking, leaf element concentrations could
reflect the element supply status of trees. But leaf element
concentrations are influenced by various factors, such as
season, and physiological status of trees (Walbridge, 1991;
Cuevas and Lugo, 1998; Lin and Wang, 2001). The element
concentrations in mature leaves of P. pubescens displayed
monthly changes. The highest mean element concentra-
tion in mature leaves was N, and the lowest was Mg. The
rank order of the mean element concentrations in mature
leaves was: N> Ca> K> P > Mg (significantly different at P
<0.05 level) (Fig.3). The result was different from the obser-
vations (N> K >Ca > Mg > P) of Lin et al. (2001) on
Castanopsis eyrei and Pinus taiwanensis leaves. The char-
acteristics of element accumulation were different with
species.
2.4 Changes in element levels during leaf senescence
The monthly mean element levels in mature leaves and
senescent leaves are listed in Fig.3. Statistics results
showed that, the concentrations (mg/g) of N, P and K in
Fig.1. Monthly changes in leaf mass, leaf area and specific leaf
mass of mature leaf (ML) and senescent leaf (SL) of Phyllostachys
pubescens in Yongchun County, Fujian, China. LA, leaf area;
LM, leaf mass; SLM, specific leaf mass.
Fig.2. Decrease in LM, LA and SLM during leaf senescence of
Phyllostachys pubescens in Yongchun County, Fujian, China from
April 2002 to March 2003. The abbreviations are the same as in Fig.1.
LIN Yi-Ming et al.: Dynamics of Leaf Mass, Leaf Area and Element Retranslocation Efficiency During Leaf Senescence in
Phyllostachys pubescens 1319
mature leaves were much higher than those in senescent
leaves (P<0.01), the Ca concentration in mature leaves was
lower than that in senescent leaves (P < 0.000 5), and the
Mg concentration in mature leaves was close to that in
senescent leaves (P = 0.653).
While on content basis (mg/leaf), during leaf
senescence, N, P, K and Mg contents showed the decrease
trends (the differences were all significant at P<0.005 level),
while Ca level was almost unchanged (P = 0.178).
2.5 Differences in mean element RE on difference bases
For the mean RE of N, P, K, Ca and Mg on different
bases, the rank order was RE2 > RE3 > RE1 (Table 2). The
result was accorded with the observation of Lin and Wang
(2001) on Kandelia candel (a mangrove species).
Nevertheless, these relationships were no tenable for the
RE of each month.
2.6 Monthly changes in element retranslocation effi-
ciency on content basis (RE2)
RE2 of the five elements appeared to show significant
monthly variation (Fig.4). The coefficient of variation of N,
P, K, Ca and Mg was 48.61%, 67.80%, 38.68%, 82.74% and
2 561.11%, respectively. The coefficients of variation were
high, especially in Mg. Not only did the degree of element
retranslocation change monthly, but also the direction of
retranslocation varied monthly.
3 Discussion
3.1 Relationship between RE and nutrient concentrations
in mature leaves
As far as the relationship between the RE and nutrient
concentrations in mature leaves was concerned, Chapin
Table 1 Relationship between leaf mass (x) and leaf area (y) of Phyllostachys pubescens from April 2002 to March 2003*
Month Leaf Equation R2
4 Mature leaf y =81.362 x +1.476 1 0.916 5
Senescent leaf y =84.833 x +0.840 6 0.854 0
5 Mature leaf y = 79.568 x + 1.268 5 0.935 6
Senescent leaf y = 81.521 x + 1.341 0 0.899 1
6 Mature leaf y = 89.104 x + 0.590 2 0.887 8
Senescent leaf y = 83.462 x + 0.895 1 0.848 0
7 Mature leaf y = 92.579 x + 0.818 0 0.943 8
Senescent leaf y = 85.116 x + 1.283 6 0.891 7
8 Mature leaf y = 69.891 x + 1.556 8 0.926 3
Senescent leaf y = 89.057 x + 0.361 8 0.893 5
9 Mature leaf y = 84.827 x + 1.010 4 0.973 0
Senescent leaf y = 87.520 x + 1.265 7 0.918 4
10 Mature leaf y = 87.140 x + 1.031 4 0.916 5
Senescent leaf y = 100.690 x + 0.434 3 0.935 2
11 Mature leaf y = 82.580 x + 1.371 2 0.978 5
Senescent leaf y = 79.951 x + 1.637 4 0.916 2
12 Mature leaf y = 80.992 x + 1.105 5 0.863 2
Senescent leaf y = 96.305 x + 0.574 4 0.985 1
1 Mature leaf y = 85.802 x + 1.188 7 0.973 8
Senescent leaf y = 83.367 x + 1.393 2 0.954 5
2 Mature leaf y = 89.992 x + 1.114 1 0.983 3
Senescent leaf y = 91.817 x + 1.141 5 0.864 9
3 Mature leaf y = 82.411 x + 1.244 9 0.966 1
Senescent leaf y = 86.367 x + 1.309 1 0.940 5
*, P < 0.01 and df = 48.
Fig.3. Monthly mean element concentrations (mg/g) and con-
tent (mg/leaf) in mature leaf (ML) and senescent leaf (SL) of
Phyllostachys pubescens in Yongchun County, Fujian, China.
Acta Botanica Sinica 植物学报 Vol.46 No.11 20041320
and Kedrowski (1983) found a direct correlation between
proportional nutrient retranslocation from the leaves dur-
ing senescence and nutrient concentration in tree leaves.
Lodhiyal and Lodhiyal (1997) argued that the higher leaf
tissue nutrient level was , the greater the percent transloca-
tion capacity would be. However, Stapel and Hemminga
(1997) thought that the resorption efficiency was not sig-
nificantly different in seagrass with a relatively high and a
relatively low nutrient concentration, although within-spe-
cies comparison showed that in some cases resorption ef-
ficiency was positively related to the nutrient concentra-
tions of the leaves.
The present observation of N and P is not consistent
with that of Chapin and Kedrowski (1983), but the observa-
tion of K, Ca and Mg supported their views. The correla-
tion equations between RE from the leaves during senes-
cence and nutrient concentration in mature leaves were as
follows:
K: y = －6.1926 x2＋59.64 x－112.94, P < 0.01;
Ca: y = 16.314 x－115.15, P < 0.01;
Mg: y = 80.415 x－119.64, P < 0.01.
3.2 Differences of RE on different bases
As mentioned above, Leaf senescence is not a single
process of element retranslocation, but a process accom-
panied by respiration, hydrolysis of carbohydrate, protein
and the translocation of hydrolyzates such as soluble sugar,
amino acid out of the senescing leaves (He and Jin, 1999;
Lindroth et al., 2002). Certain elements such as Ca accumu-
late in the senescing leaves, too (Ralhan and Singh, 1987;
Lin and Wang, 2001). Present studies indicated that Ca ac-
cumulated in senescing leaves of P. pubescens. During
leaf senescence, there are not only the changes in element
concentrations but also the changes in leaf mass. Chapin
and Kedrowski (1983) reported that there was significant
(≈18%) loss in leaf mass associated with senescence and
retranslocation subsequent to leaf abscission. This study
indicated that during the experiment, the mean decrease of
leaf mass was 19.55% for P. pubescens. Changes in ele-
ment levels on concentration basis were offset by the
Table 2 Nutrient retranslocation efficiency (RE) on the basis of concentration (mg/g DW, RE1), element content (mg/leaf, RE2) and leaf
area (mg/cm2, RE3)(%)

Element
RE1 RE2 RE3
Mean SD Mean SD Mean SD
N 21.71 19.54 36.68 17.83 25.70 18.59
P -3 . 2 1 15.49 17.33 11.75 2.04 14.94
K 6.12 11.08 24.64 9.53 10.68 12.84
Ca -53 .4 8 28.51 -22 .5 4 18.65 -45 .8 8 29.22
Mg -24 .2 5 26.31 0.72 18.44 -18 .2 6 27.29
Fig.4. Changes in element retranslocation efficiency on content
basis (RE2) during leaf senescence of Phyllostachys pubescens in
Yongchun County, Fujian, China.
LIN Yi-Ming et al.: Dynamics of Leaf Mass, Leaf Area and Element Retranslocation Efficiency During Leaf Senescence in
Phyllostachys pubescens 1321
changes in leaf mass. The element concentrations in ab-
scised leaves may be a reflection of the initial element con-
centrations and the changes in leaf mass, rather than the
extent of element retranslocation (Chapin and Kedrowski,
1983). Pugnaire and Chapin (1993) suggested that the con-
centration based on data of RE would be misleading when
significant amounts of leaf mass were lost at leaf senescence.
If there were no changes in leaf area during leaf
senescence, A1 > A2, W1 > W2 and S1 = S2. Thus the result
would be: RE1 < RE2 = RE3. In other words, RE1 underesti-
mate the level of element retranslocation. The result (RE2 >
RE3 > RE1) is consistent with this deduction (Table 2). Birk
and Vitousek (1986) got the same result from their study on
loblolly pine (Pinus taeda). Killingbeck (1985) assumed that
interpreting most of these literatures on a concentration
basis (RE1) would have serious limitations.
Having recognized the limitation of data on a concen-
tration basis (RE1), a few researchers turned to the RE on a
leaf area basis (RE3) (Killingbeck, 1985; Pugnaire and Chapin,
1993; Lin and Wang, 2001). Woodwell (1974) suggested
that least distortion of element relationships occurred when
the leaf area was used as the basis for the element
composition. But there were exceptions, Delucia and
Schlesinger (1995) reported RE1> RE3 in Lyonia lucida.
All of these studies stood on one basis: there were small
or no changes in leaf mass at leaf senescence. When there
were small differences in SLM between mature leaves and
senescent leaves for a species, little accumulation or with-
drawal of carbon compounds occurred during leaf senes-
cence (Schlesinger et al., 1989), and the differences in ele-
ment concentrations were used to calculate RE. The re-
search indicated that though there was relatively smaller
change in SLM for P. pubescens (5.07%) during the
experiment, leaf mass and leaf area decreased significantly,
amounting to 19.55% and 15.16%, respectively. Changes in
SLM were offset by change in leaf area, and that changes
in element concentrations on area basis (RE3) were offset,
too. The changes in SLM were not a reliable parameter for
the changes in LA or LM, but a comprehensive parameter
of them. If the decrease in LM and the decrease in LA oc-
curred at the same time, SLM may change little. In this
study, the changes in SLM were less than those in LM and
LA. Schlesinger et al. (1989) stated that in the absence of
strong changes in SLM with leaf age, the comparison of
element concentrations in mature and senescent leaves were
directly indicative of element retranslocation efficiency.
If the decrease in LM was larger than that in LA during
leaf senescence, in other words, SLM decreased with leaf
age, RE1 would be lower than RE3. In this study, the SLM
of P. pubescens decreased with leaf senescence, RE1The reasons for the decrease of leaf area are still
unknown, but the decrease in leaf area with leaf age is com-
mon in trees (Lin and Wang, 2001). According to the
observation, the decrease of leaf area occurs mainly around
the time that leaf drops. Leaves may contract because of
loss of tissue water during leaf senescence.
3.3 Monthly changes in RE
The element retranslocation efficiency on different ba-
sis showed the different results, not only did the degree of
element retranslocation change monthly, but also the di-
rection of retranslocation varied monthly. As shown in Table
2, RE1 of P was -3.21 (negative), Schlesinger et al. (1989)
also found that RE of N in Aretostaphylos patula was
negative. These situations were impossible physiologically
(Lin and Wang, 2001). P. pubescens, a giant evergreen
woody bamboo, has great economic value. The researches
on RE will be beneficial to relational nutrient diagnosis.
Acknowledgements: We thank LIU Jian-Bin, ZOU Xiu-
Hong and GUO Zhi-Jian for their help in plant sampling and
valuable comments.
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