Acta Botanica Sinica
植 物 学 报
2004, 46 (11): 1316-1323
http://www.chineseplantscience.com
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, making 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 mechanism 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 characteristic 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 different ways conducted. Low-element-adapted species on
infertile soils were frequently compared to high-elementadapted 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) (delArco 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 senescence 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.
Received 4 Feb. 2004 Accepted 20 Aug. 2004
Supported by the Natural Science Foundation of Fujian Province (B0110007).
* Author for correspondence. E-mail: <[email protected]>.
LIN Yi-Ming et al.: Dynamics of Leaf Mass, Leaf Area and Element Retranslocation Efficiency During Leaf Senescence in
Phyllostachys pubescens
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, concentrating 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 ecological 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 differences between the two methods.
1 Materials and Methods
1.1 Site
Research was conducted at Yidu Town, Yongchun
County (25°21′33″-25°31′33″ N, 117°40′40″ E), Fujian,
China. The climate is subtropical monsoon. The average
annual temperature is 18.3℃, the average annual precipitation 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
１３１７
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 damaged 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 distilled 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 acidhydrogen peroxide, and N was determined by the
microKjeldahl method (Yoshida et al., 1972), P was determined by acorbic acid-antimony reducing phosphate colorimetric method (Murphy and Riley, 1962). Samples were
digested in nitric acid-perchloric acid, and K, Ca and Mg
determinations were made with Model WFX-Ⅰ B 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):
RE 1=（1 － A 2/A 1 × W1 /W2 ）× 100%
RE 2 =（1 － A 2 /A 1 ）× 100%
RE 3 =（1 － A 2 /A 1 × S 1 /S 2 ）× 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 mature leaf; and W2 is the mean mass (g) per senescent leaf.
The element contents (mg/leaf) were calculated by multiplying 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
１３１８
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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, averaged (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%,
r esp ectively. The seasonal changes in decr ease
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 correlations 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 concentration 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 observations (N> K >Ca > Mg > P) of Lin et al. (2001) on
Castanopsis eyrei and Pinus taiwanensis leaves. The characteristics 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
Table 1
１３１９
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 content (mg/leaf) in mature leaf (ML) and senescent leaf (SL) of
Phyllostachys pubescens in Yongchun County, Fujian, China.
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 efficiency 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
１３２０
Ａｃｔａ　Ｂｏｔａｎｉｃａ　Ｓｉｎｉｃａ　植物学报　Ｖｏｌ．４６　Ｎｏ．１１　２００４
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
N
P
K
Ca
Mg
RE1
Mean
21.71
-3.21
6.12
-53.48
-24.25
RE2
SD
19.54
15.49
11.08
28.51
26.31
Mean
36.68
17.33
24.64
-22.54
0.72
Fig.4. Changes in element retranslocation efficiency on content
basis (RE2) during leaf senescence of Phyllostachys pubescens in
Yongchun County, Fujian, China.
RE3
SD
17.83
11.75
9.53
18.65
18.44
Mean
25.70
2.04
10.68
-45.88
-18.26
SD
18.59
14.94
12.84
29.22
27.29
and Kedrowski (1983) found a direct correlation between
proportional nutrient retranslocation from the leaves during senescence and nutrient concentration in tree leaves.
Lodhiyal and Lodhiyal (1997) argued that the higher leaf
tissue nutrient level was , the greater the percent translocation capacity would be. However, Stapel and Hemminga
(1997) thought that the resorption efficiency was not significantly different in seagrass with a relatively high and a
relatively low nutrient concentration, although within-species comparison showed that in some cases resorption efficiency was positively related to the nutrient concentrations of the leaves.
The present observation of N and P is not consistent
with that of Chapin and Kedrowski (1983), but the observation of K, Ca and Mg supported their views. The correlation equations between RE from the leaves during senescence 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 accompanied 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 accumulate in the senescing leaves, too (Ralhan and Singh, 1987;
Lin and Wang, 2001). Present studies indicated that Ca accumulated 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 element levels on concentration basis were offset by the
LIN Yi-Ming et al.: Dynamics of Leaf Mass, Leaf Area and Element Retranslocation Efficiency During Leaf Senescence in
Phyllostachys pubescens
changes in leaf mass. The element concentrations in abscised leaves may be a reflection of the initial element concentrations and the changes in leaf mass, rather than the
extent of element retranslocation (Chapin and Kedrowski,
1983). Pugnaire and Chapin (1993) suggested that the concentration 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 underestimate 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 concentration 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 withdrawal of carbon compounds occurred during leaf senescence (Schlesinger et al., 1989), and the differences in element concentrations were used to calculate RE. The research 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 occurred 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
１３２１
of P. pubescens decreased with leaf senescence, RE1<RE3
<RE2.
The reasons for the decrease of leaf area are still
unknown, but the decrease in leaf area with leaf age is common 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 basis showed the different results, not only did the degree of
element retranslocation change monthly, but also the direction 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 XiuHong and GUO Zhi-Jian for their help in plant sampling and
valuable comments.
References:
Aerts R. 1995. The advantages of being evergreen. Trends Ecol
Evol, 10: 402-407.
Aerts R. 1996. Nutrient resorption from senescing leaves of
perennials: are there general patterns? J Ecol, 84: 597-608.
Birk E M, Vitousek P M. 1986. Nitrogen availability and nitrogen
use efficiency in loblolly pine stands. Ecology, 67: 69-79.
Boerner R E J. 1984. Foliar nutrient dynamics and nutrient efficiency of four deciduous tree species in relation to site fertility.
J App Ecol, 21: 1029-1040.
Chapin III F S, Kedrowski R A. 1983. Seasonal changes in nitrogen and phosphorus fractions and autumn retranslocation in
evergreen and deciduous taiga trees. Ecology, 64: 376-391.
Chapin III F S, Moilanen L. 1991. Nutrient controls over nitrogen
and phosphorus resorption from Alaskan birch leaves. Ecology,
72: 709-715.
Cote B C S, Vogel S, Dawson J O. 1989. Autumnal changes in
tissue nitrogen of autumn olive, black alder and eastern
cottonwood. Plant Soil, 118: 23-32.
Covelo F, Gallardo A. 2002. Effect of pine harvesting on leaf
nutrient dynamics in young oak trees at NW Spain. For Ecol
Manag, 167: 161-172.
Cuevas E, Lugo A E. 1998. Dynamics of organic matter and nutrient return from litterfall in stands of ten tropical tree plantation species. For Ecol Manag, 112: 263-279.
Dalla-Tea F, Jokela E J. 1994. Neddle fall returns and resorption
rates of nutrients in young intensively managed slash and
１３２２
Ａｃｔａ　Ｂｏｔａｎｉｃａ　Ｓｉｎｉｃａ　植物学报　Ｖｏｌ．４６　Ｎｏ．１１　２００４
loblolly pine stands. For Sci, 40: 650-662.
del-Arco J M, Escudero A, Garrido M V. 1991. Effects of site
layan Tarai poplar plantations. Ann Bot, 79: 517-527.
Lodhiyal N, Lodhiyal L S. 2003. Aspects of nutrient cycling and
characteristics on nitrogen retranslocation from senescing
leaves. Ecology, 72: 701-708.
nutrient use pattern of Bhabar Shisham forests in central
Himalaya, India. Forest Ecol Manag, 176: 237-252.
Delucia E H, Schlesinger W H. 1995. Photosynthetic rates and
nutrient-use efficiency among evergreen and deciduous shrubs
Miao S L. 2004. Rhizome growth and nutrient resorption: mechanisms underlying the replacement of two clonal species in
in Okefenokee swamp. J Plant Sci, 156: 19-28.
Fife D N, Nambiar E K S. 1997. Changes in the canopy and
Florida Everglades. Aquatic Bot, 78: 55-66.
Millard P, Neilsen G H. 1989. The influence of nitrogen supply
growth of Pinus radiata in response to nitrogen supply. For
Ecol Manag, 93: 137-152.
on the uptake and remobilization of stored N for the seasonal
growth of apple trees. Ann Bot, 63: 301-309.
Helmisaari H S. 1992. Nutrient retranslocation within the foliage
of Pinus sylvestris. Tree Physiol, 10: 45-58.
Murphy J, Riley J P. 1962. A modified single solution method for
the determination of phosphate in natural waters. Ann Chem
He P, Jin J-Y. 1999. Relationships among hormone changes, transmembrane Ca2+ flux and lipid peroxidation during leaf senes-
Acta, 27: 31-36.
Nambiar E K S, Fife D N. 1987. Growth and nutrient retranslocation
cence in Spring Maize. Acta Bot Sin, 41: 1221-1225. (in Chinese with English abstract)
in needles of radiata pine in relation to nitrogen supply. Ann
Bot, 60: 147-156.
Isagi Y, Kawahara T, Ito H A. 1997. Computer-aided management system of Phyllostachys stands based on the ecological
Peng Z-Q , Lin Y-M , Liu J-B, Zou X-H, Guo Z-J , Guo Q-R , Lin
P. 2002. Biomass structure and energy distribution of
characteristics of carbon cycling. Chapman G P. The Bamboos.
London: Academic Press. 125-134.
Phyllostachys pubescens population. J Xiamen Univ (Nat Sci),
41: 579-583. (in Chinese with English abstract)
Killingbeck K T. 1985. Autumnal resorption and accretion of
trace metals in gallery forest trees. Ecology, 66: 283-286.
Pugnaire F J, Chapin III F S. 1993. Controls over nutrient restortion
from leaves of evergreen Mediterranean species. Ecology, 74:
Lajtha K. 1987. Nutrient resorption efficiency and the response
to phosphorus fertilization in the desert shrub Larrea tridentata
124-129.
Ralhan P K, Singh S P. 1987. Dynamics of nutrients and leaf mass
(DC) Cov. Biogeochemistry, 4: 265-276.
Lindroth R L, Osier T L, Barnhill H R H, Wood S A. 2002. Effects
in central Himalayan forest trees and shrubs. Ecology, 68:
1974-1983.
of genotype and nutrient availability on phytochemistry of
Schlesinger W H, Delucia E H, Billings W D. 1989. Nutrient-use
trembling aspen (Populus tremuloides Michx.) during leaf
senescence. Biochem Syst Ecol, 30: 297-307.
efficiency of wood plants on contrasting soils in the Western
Great Basin, Nevada. Ecology, 70: 105-113.
Lin P, Wang W Q. 2001. Changes in the leaf composition, leaf
mass and leaf area during leaf senescence in three species of
Stapel J, Hemminga M A. 1997. Nutrient resorption from seagrass
leaves. Mar Biol, 128: 197-206.
mangroves. Ecol Eng, 16: 415-424.
Lin Y-M, Yang Z-W , Li Z-J. 2001. Research on Evergreen For-
Torii A, Isagi Y. 1997. Range expansion of bamboo species in
southern area of Kyoto Prefecture, Japan. JPN J Ecol, 47:
ests in the Wuyi Mountains. Xiamen: Xiamen University Press.
(in Chinese)
31-41.
Walbridge M R. 1991. Phosphorus availability in acid organic
Li R, Werger M J A, de Kroon H, During H J, Zhong Z C. 2000.
Interaction between shoot age structure, nutrient availability
soils of the lower North Carolina coastal plain. Ecology, 72:
2083-2100.
and physiological integration in the giant bamboo Phyllostachys
pubescens. Plant Biol, 2: 437-446.
Woodwell G M. 1974. Variation in the nutrient content of Quercus
alba, Quercus coccinea, and Pinus rigida in the Brookheaven
Li Z-J, Lin P. 1995. Accumulation and distribution of some elements of Phyllostachys pubescens community in Southern
forest from bud-break to abscission. Am J Bot, 61: 749-753.
Yoshida S, Forno D A, Cock J H, Gomez K A. 1972. Laboratory
Fujian. Chin J Appl Ecol , 6(Suppl.): 9-13. (in Chinese with
English abstract)
Manual for Physiological Studies of Rice. 2nd ed. Philiphines:
The International Rice Research Institute. 7-9, 36-38.
Lodhiyal L S, Lodhiyal N. 1997. Nutrient cycling and nutrient
use efficiency in short rotation, high density central Hima-
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