ORIGINAL ARTICLE
J For Res (2004) 9:205–210 © The Japanese Forestry Society and Springer-Verlag Tokyo 2004
DOI 10.1007/s10310-003-0074-7
Md. Nabiul Islam Khan · Rempei Suwa · Akio Hagihara
Kazuharu Ogawa
Interception of photosynthetic photon flux density in a mangrove stand of
Kandelia candel (L.) Druce
Received: November 10, 2003 / Accepted: December 26, 2003
M.N.I. Khan · R. Suwa · A. Hagihara (*)
Laboratory of Ecology and Systematics, Faculty of Science,
University of the Ryukyus, Okinawa 903-0213, Japan
Tel./Fax +81-98-895-8546
e-mail: amyh@sci.u-ryukyu.ac.jp
K. Ogawa
Laboratory of Forest Ecology and Physiology, Graduate School of
Bioagricultural Sciences, Nagoya University, Nagoya, Japan
Abstract The canopy structure and interception of photo-
synthetic photon flux density (PPFD) in a 10-year-old
Kandelia candel (L.) Druce stand were investigated before
and after artificial defoliation. Leaf and wood areas for
different layers were measured through area–weight rela-
tionships of subsamples. PPFD was measured at specified
heights before and after leaf clipping. The leaf area index
(LAI) and wood area index (WAI) were 4.501 m
2
m
-2
and
1.412 m
2
m
-2
, respectively. There was a strong linear rela-
tionship between the cumulative wood area (C) and leaf
area (F) densities from the top down to a given depth of the
canopy, C = aF (r
2
= 0.950), with a proportional constant a
of 0.096 0.008 (mean SE). The PPFD relative to that
above the canopy (relative PPFD; I
R
) at a given depth of the
canopy was assumed to be given by the equation I
R
=
e
-(K
C
C+K
F
F
) = e
-KF
, where the apparent light extinction
coefficient K (= K
F
+ aK
C
, where K
F
and K
C
are respec-
tively the light extinction coefficient of leaves and woody
organs) was calculated to be 0.502 0.041 (mean SE)
m
-2
m
2
before leaf clipping. After leaf clipping, I
R
C
= e
-K
C
C
is satisfied. As a result, the value of K
C
was estimated
to be 0.785 0.046 (mean SE) m
-2
m
2
. The light
extinction coefficient of leaves K
F
was calculated to be
0.427 m
-2
m
2
using the indirect method, K
F
= K - aK
C
, and
0.432 0.026 (mean SE) m
-2
m
2
using the direct method,
I
R
/I
R
C
= e
-K
F
F
. Of the total PPFD intercepted by the canopy,
the fraction K
F
/K due to leaves alone was estimated to
be 85.0%–86.1% and the rest was contributed by woody
organs.
Key words Apparent light extinction coefficient · LAI ·
Light extinction coefficient of leaves · PPFD, WAI
Introduction
Canopy structure, i.e., spatial distributions of leaf and wood
areas, plays a significant role in the interception of photo-
synthetic photon flux density (PPFD) by leaves, which
strongly affects the gas exchange between forests and the
atmosphere (Kurachi et al. 1993; King and Maindonald
1999; Sterck and Bongers 2001; Law et al. 2001). The pio-
neer study by Monsi and Saeki (1953) assumed that light
interception by plant canopies is caused by leaves alone,
and their model for estimating plant photosynthetic produc-
tion was based on the apparent light extinction coefficient,
K. In forest canopies, however, PPFD is intercepted not
only by the leaves, but also by the woody organs. Some
studies had been performed on the light interception by
forest canopies (Yim et al. 1969; Kurachi et al. 1986;
Hagihara and Yamaji 1993; Maass et al. 1995; Chen et al.
1997; Levy and Jarvis 1999; Green et al. 2001). Among these
studies, only a few considered the contribution of woody
organs to the light attenuation by the canopy (Yim et al.
1969; Kurachi et al. 1986; Hagihara and Yamaji 1993; Chen
1996). Some modifications of the Monsi–Saeki model
(Kurachi et al. 1986, 1993; Hagihara 1987), which incorpo-
rated not only the K but also the light extinction coefficient
of leaves K
F
, suggested that canopy photosynthetic produc-
tion of a Larix leptolepis (Sieb. et Zucc.) Gordon stand
based on K showed approximately 30% overestimation in
comparison with their model based on K and K
F
(Kurachi
et al. 1986). Therefore, the estimation of K and K
F
became
an important issue for forest ecophysiologists. The knowl-
edge of PPFD interception not only by the canopy as a
whole, but also by the leaves and woody organs separately,
is very important in understanding the structural and func-
tional characteristics of forest canopies.
So far, information regarding the light extinction
coefficients of leaves K
F
and of woody organs K
C
remains
inadequate. In addition, similar studies in mangrove forests
have not been conducted. Therefore, the aim of this study
was to elucidate the canopy structure of a Kandelia candel
(L.) Druce stand and to evaluate the pattern of PPFD